Volume 3 NOVON Number 1 1993 Monochaetum amistadense (Melastomataceae): A New Species from the Paramo of Costa Rica Frank Almeda Department of Botany, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118-4599, U.S.A. ABSTRACT. Monochaetum amistadense, a new species from the paramo of Costa Rica, is described, illustrated, and compared to its closest presumed relative, M. compactum. The distributional range of M. trichophvllum is extended from Panama to Costa Rica, and a diagnostic key is provided for the 10 species of Monochaetum now known from Costa Rica. The evolution of morphologically distinct taxa, often of limited geographic extent, has been a re¬ current theme in the evolution of Monochaetum, a montane neotropical genus of some 45-50 species. The most recent regional monograph attributed 18 species of Monochaetum to Mexico and Central America, eight of which were reported from Costa Rica (Almeda, 1978). Collections of Monochaetum made in the Cor¬ dillera de Talamanca during the last decade have added two additional species to the Costa Rican flora. Monochaetum trichophyllum Almeda, which was described from and thought to he endemic to Pan¬ ama, is now known from the Altantic slope of the Talamanca range (Limon: S side of unnamed cor¬ dillera between the Rio Terbi and Rio Sini, 2-4 airline km W of the Costa Rican-Panamanian bor¬ der at 2,300-2,500 m, 1 1 Sep. 1984, Davidse et al. 28946, MO). The second addition, a distinctive novelty described below, is of special interest be¬ cause it is the only known Costa Rican species con¬ fined to paramo habitats. Monochaetum amistadense Almeda, sp. nov. TYPE: Costa Rica. Limon: Cordillera de Tala¬ manca, Cerro Kamuk massif, between Cerro Dudu and Cerro Apri, 9°1 4'30"- 1 5'30"N, 83°03'30"-04'30'/W, 23 and 26 Mar. 1984 (fl, fr), Davidse et al. 25932 (holotype, CAS; isotypes, CR, MO). Figure 1 . Frutex 0.5-2 m altus; ramuli teretes sicut petioli in- florescentiaque dense pilis patulis barbellatis 0.5-1 mm longis induti. Petioli 0.5-1 mm longi; lamina 3-7.5 x 1.5- 3 mm elliptica vel elliptico-ovata apice acuta basi obtusa vel rotundata, supra utrinsecus venas primarias densiuscule strigosa, subtus dense vel modice strigosa pilis 0.5-1 mm longis barbellatis, 3-nervata vel 3-plinervata. Inflorescentia terminalis 1-2 flora, flores 4-meri. Hypan- thium (ad torum) 5-6 mm longum; calycis lobis 4. 5-5. 5 x 2-2.5 mm anguste oblongo-triangulares persistentes. Petala 10-12 x 5-7 mm obovata. Stamina dimorpha glabra, thecis subulatis, poro dorsaliter inclinato. Stamina maiora; thecae 7-8.5 mm longae; appendix dorsalis 3 3.5 x 0.5 mm, apice rotundato. Stamina minora: thecae 4.5- 5 mm longae; appendix dorsalis 3 x 0.25 mm, apice hebeti-acuto. Ovarium 4-loculare apice modice setoso. Semina 0.5 mm longa, oblongo-arcuata. Erect, rigidly branched shrub 0.5 2 m tall. Distal internodes terete, copiously covered with spreading barbellate hairs 0.5-1 mm long, the nodes beset with flexuous spreading distally barbellate hairs 1.5 2 mm long. Older branches glabrate with reddish brown exfoliating bark. Mature leaves equal to some¬ what unequal in size in each pair; petioles 0.5- 1 mm long and 0.5 mm wide; blades membranaceous, entire, 3-7.5 mm long, 1.5-3 mm wide, elliptic to elliptic-ovate, 3-nerved or 3-plinerved, the apex acute, the base obtuse to rounded, the margin entire; adaxial surface copiously beset with a strigose in- dument of barbellate multicellular hairs in 4 longi¬ tudinal belts between the impressed primary veins Novon 3: 1-4. 1993. 2 Novon I m m Figure 1. Monochaetum amistadense Almeda. — A. Habit. --B. Enlargement of cauline node and internode. — C. Representative leaves, adaxial surface (left) and abaxial surface (right). — D. Enlargement of adaxial foliar hair. — E. Fruiting hypanthium. — F. Petal. — G. Antepetalous stamen in profile view. — H. Antesepalous stamen, profile view (left), dorsal view (right). —I. Seeds. (A from Davidse et al. 29023, CAS; B-I from the holotype.) that essentially conceal the blade surface; abaxial surface also copiously beset with a strigose indument of barbellate multicellular hairs largely concentrated on but not restricted to the three elevated primary veins. Inflorescence reduced to solitary or paired 4-merous flowers commonly terminating lateral flo- riferous shoots; bracteoles only slightly reduced in size, otherwise identical to principal leaves in all Volume 3, Number 1 1993 Almeda Monochaetum amistadense 3 respects; pedicels (at anthesis) 2-9 mm long length¬ ening up to 1 .3 cm in fruit, moderately beset with appressed to antrorsely spreading barbellate hairs to 1 mm long. Fruiting hypanthia 5-6 mm long to the torus, 4 mm wide, campanulate to subcylindric, deeply pigmented and moderately covered with an¬ trorsely spreading hairs like those of the pedicels. Calyx lobes (in fruit) persistent, 4. 5-5. 5 mm long and 2-2.5 mm wide at the base, narrowly triangular, adaxial surface glabrous, abaxial surface glabrous or beset with a few scattered barbellate hairs, the margins entire but fringed with barbellate hairs. Petals 10-12 x 5-7 mm, magenta, obovate, gla¬ brous or with 1-3 hairs apically. Stamens strongly dimorphic, with dorsally inclined apical pores; larger (antepetalous) stamens geniculate, filaments 7-8 mm long, anther thecae 7-8.5 mm long, subulate, con¬ nective modified at the filament insertion into an incurved, dorso-basal, linear-oblong appendage 3- 3.5 mm long; smaller (antesepalous) stamens ± erect, filaments 7-9 mm long, anther thecae 4.5- 5 mm long, linear-oblong to subulate; connective modified at the filament insertion into an upturned, dorso-basal narrowly elliptic-lanceolate appendage 3 mm long. Ovary (at anthesis) superior, ovoid to obovoid, the body essentially glabrous but copiously setose at the summit surrounding the stylar scar. Style 6-7 mm long, somewhat declined to one side of the flower opposing the larger stamens. Capsules dry, semiwoody, loculicidal, erect at maturity and enclosed by the enveloping hypanthium; seeds 0.5 mm long, oblong-arcuate, somewhat keeled on the concave side, light brown and essentially smooth with a subdued luster. Distribution. Known only from paramo habitats dominated by Blechnum, Diplostephium, and other shrubs in the southern Cordillera de Talamanca from Cerro Durika to an unnamed cordillera just southeast of the Cerro Kamuk massif between the Rio Sini and the Rio Terbi west of the Panamanian frontier at elevations of 2,400-3,280 m. The range of M. amistadense is contained within the Costa Rican sector of Parque Internacional La Amistad. Monochaetum amistadense, one of six endemic species in Costa Rica, is characterized by unusually small leaf blades that are 3-nerved or 3-plinerved, inflorescences reduced to solitary or paired flowers, and oblong-arcuate seeds that are somewhat keeled on the concave side and not strongly coiled distally. It is most similar to and may be derived from M. compactum Almeda of western Panama: the two have similar habits, stem pubescence, leaf shape, and petal color. Monochaetum compactum, how¬ ever, has modally larger (to 2.8 cm long), 5-plinerved leaf blades, a mixture of glandular and barbellate hypanthial and pedicellar hairs, an inflorescence consisting of compound dichasia with flowers sub¬ tended by reduced bracteoles, and cochleate seeds that are coiled distally and lack a keel on the concave side. Paratypes. Costa Rica, limon: Cordillera de Tala¬ manca, Atlantic slope, unnamed cordillera between Rio Sini and Rio Terbi, 9°00'-9°l 2'N, 82°58'-82°59'W, 13 Sep. 1984 (fl), Davidse et al. 29023 (CAS, CR, MO). PUNTARENAS: Canton de Buenos Aires, Ujarras, El Car¬ men, Cerro Dfirika, 9°22'35"N, 83°18'24"W, 12 Oct. 1989 (fl, fr), A. Chacon 521 (CAS, CR, INBIO, MO). Morphologically, M. amistadense does not ap¬ pear to be particularly close to any of the Costa Rican members of the genus and is readily distin¬ guished from them by the following key. Key to the Species of Monochaetum in Costa Rica la. Distal internode pubescence consisting of widely to retrorsely spreading barbellate hairs. 2a. Sprawling or trailing subshrubs, the lateral branchlets divaricate and held at an angle of 65°-90°; adaxial foliar pubescence not forming well-defined longitudinal belts between the impressed primary veins; seeds dark brown, glossy, semicircular and laterally compressed; petals pink; Costa Rica and Panama . M. trichophyllum Almeda 2b. Erect, laxly to compactly branched shrubs, the lateral branchlets held at an angle of 45° or less; adaxial foliar pubescence forming 4 to 6 well-defined longitudinal belts between the impressed primary veins; seeds orange-brown with a subdued luster, cochleate or oblong-arcuate; petals white or magenta. 3a. Principal leaves lanceolate to elliptic-lanceolate, 1. 4-8.0 cm long, 0.6-2. 9 cm wide, petioles 1- 5(— 11) mm long; canline hairs l-3(-5) mm long; inflorescence a multiflowered dichasial cyme with flowers subtended by sessile bracteoles that differ from principal leaves in size and shape; petals white; seeds distinctly cochleate and strongly coiled at the distal end; Mexico to Panama . M. floribundum (Schlechtendal) Naudin 3b. Principal leaves elliptic to elliptic -ovate, 3-7.5 mm long, 1.5-3 mm wide, petioles 0.5-1 mm long; cauline hairs 0.5-1 mm long; inflorescence reduced to solitary or paired flowers subtended by petiolate bracteoles that are similar to principal leaves in size and shape; petals magenta; seeds oblong-arcuate, not strongly coiled at the distal end; endemic to Costa Rica . . M. amistadense Almeda 4 Novon lb. Distal internode pubescence consisting of appressed, barbellate or smooth hairs, or internodes entirely glabrous. 4a. Sprawling or trailing subshrubs, the lateral branchlets divaricate and held at an angle of 65°-90°; seeds dark brown and glossy, somewhat semicircular, laterally compressed and without pronounced gyrations. 5a. Hypanthial hairs appressed, essentially smooth and eglandular; calyx lobes 6-10 mm long, an- trorsely spreading on fruiting hypanthia; principal leaves 3(-5)-plinerved with appressed hairs on and closely adjacent to the elevated primary veins abaxially; petals 15-19 mm long; white when fresh but drying to shades of pink; endemic to Costa Rica . M. talamancense Almeda 5b. Hypanthial hairs antrorsely to widely spreading, smooth and often partly gland-tipped, calyx lobes 4-6 mm long, widely spreading to reflexed on fruiting hypanthia; principal leaves 5-7-plinerved (infrequently 3-plinerved) with appressed to antrorsely spreading hairs on the abaxial elevated primary veins and pilosulous to glabrate between them; petals 10-14 mm long, pink when fresh but often drying lavender or intense purple; Costa Rica and Panama . M. neglectum Almeda 4b. Erect, laxly to compactly branched shrubs, the lateral branchlets held at an angle of 45° or less; seeds orange-brown with a subdued luster, cochleate, laterally gyrate, and strongly coiled at the distal end. 6a. Distal internodes glabrous; principal leaves sessile, cordate and clasping with 7 — 9( —11) elevated primary veins abaxially; hypanthia mostly glabrous except for a few barbellate hairs along the torus at the abaxial base of the calyx lobes; Costa Rica and Panama . M. cordatum Almeda 6b. Distal internodes sp; rsely to copiously covered with appressed, barbellate hairs; principal leaves petiolate, varying in shape but never cordate or sessile; hypanthia covered with appressed, barbellate hairs occasionally intermixed with smooth, spreading glandular hairs. 7a. Principal leaves linear-oblong, 1.5-4 mm wide, l(-3)-nerved with only the median vein elevated abaxially and extending the entire length of the blade; endemic to Costa Rica . . . . M. linearifolium Almeda 7b. Principal leaves narrowly elliptic, elliptic-ovate, elliptic-lanceolate, ovate, or obovate, (2 )4 - 1 6(— 20) mm wide, 3-5(-7)-plinerved with at least three veins elevated abaxially, the innermost pair of primary veins diverging from the median vein at a point well above the petiole laminar junction. 8a. Bracteoles sessile, narrowly lance-triangular to subulate, commonly navicular and en¬ veloping floral buds or their pedicels; petals white or pink; endemic to Costa Rica .... . M. macrantherum Gleason 8b. Bracteoles distinctly petiolate or tapering basally into a compressed, ill-defined petiolar region, elliptic, elliptic-lanceolate, linear-oblong, oblanceolate, or spatulate but never enveloping floral buds or their pedicels; petals magenta. 9a. Distal internodes quadrangular and strongly carinate at the intersection of adjacent stem faces; principal leaves ovate, elliptic-ovate, or varying to obovate, glabrous adaxially or with pubescence restricted to narrow belts along the distal half or third of the blade; calyx lobes lance-triangular and acute apically; endemic to Costa Rica . M. vulcanicum Cogniaux 9b. Distal internodes terete to subterete; principal leaves narrowly elliptic to elliptic- lanceolate with 4 well-defined adaxial belts of pubescence between the impressed primary veins extending the entire length of the blade; calyx lobes linear -oblong to bluntly deltoid, rounded to obtuse apically; endemic to Costa Rica . . M. amabile Almeda Acknowledgments. 1 thank Sheva Myers for the illustration and the curators and staff's of the herbaria at the Missouri Botanical Garden, Museo Nacional de Costa Rica, and the Instituto Nacional de Bio- diversidad for hospitality during study visits and/ or gifts of specimens. Literature Cited Almeda, F. 1978. Systematics of the genus Mono- chaetum (Melastomataceae) in Mexico and Central America. Univ. Calif. Publ. Bot. 75: 1-134. New Species of Miconia (Melastomataceae) from Costa Rica Frank Almeda Department of Botany, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118-4599, U.S.A. Gina Umana Dodero Herbario Nacional de Costa Rica, Museo Nacional, Apartado 749-1000, San Jose, Costa Rica ABSTRACT. Two new Costa Rican species of Mi¬ conia, M. friedmaniorum and M. pendula, are de¬ scribed and compared with their probable relatives. Discussions, distributional notes, and diagnostic il¬ lustrations are provided for each species. Collecting activity in the mid-elevation montane forests of Costa Rica continues to yield new and unusual species of Miconia. Of the approximately 98 taxa of Miconia known lrom Costa Rica, 12 now appear to be endemic to the country. Shared floral and fruit characters suggest that the two species proposed here are closely related. Both taxa occur in wet primary forests and have similar elevational distributions, but they appear to be allopatric. Miconia friedmaniorum is known from the Cordillera de Tilaran and the Cordillera de Gua- nacaste, whereas M. pendula has been collected at one site in the Tapanti National Wildlife Refuge and Braulio Carrillo National Park. Miconia friedmaniorum Almeda & Umana, sp. nov. TYPE: Costa Rica. Alajuela: Upala, Co- lonia Libertad, subiendo hasta el Llano Agua- catales, 1,500 m, 10°48'25"N, 85°17'50"W, 28 Apr. 1988 (fl), Herrera 1900 (holotype, CR; isotypes, CAS, F, MEXU, MO, PMA, USJ). Figure 1 . Arbor 2-5 m. Ramuli quadrangulati demum teretes sicut petioli foliorum subtus venae primariae inflorescentia hypanthiaque pilis asperis 0.25-0.5 mm longis erectis dense induti demum glabrati. Lamina 10.1-14.4 x 5.5- 7.2 cm elliptica vel elliptico-ovata apice acuminata basi obtusa vel asymmetrice rotundato-obtusa, supra glabra, subtus in venis secundariis tertiariisque pilis asperis vel sparse caduceque granuloso-furfuracea, 5-plinervata, membranacea. Inflorescentia 5-6 cm longa multiflora; flores 5-meri, pedicellis (ad anthesim) 1-1.6 mm longis, bracteolis 0.5- 1.5 x 0.5 mm persistentibus. Hypanthium (ad torum) 1.5-2 mm longum; calyx primum in cono clausus demum in lobos regulares persistentes ruptus, dentibus exterioribus 0.5-0. 6 mm eminentibus. Petala 2.5-3 x 0.9-1 mm oblongo-elliptica glabra. Stamina isomorphica glabra; antherarum thecae 1.2- 1.5 x 0.3- 0.5 mm angustae oblongae, poro dorsaliter inclinato; con- nectivum nec prolongatum nec appendiculatum. Stylus 5 mm glaber; ovarium 5-loculare et % inferum apice spar- siuscule glanduloso-puberulo. Tree 2-5 m tall, young branchlets, distal petioles, elevated primaries on the abaxial foliar surfaces, and hypanthia densely covered with inconspicuously stalked asperous-headed multicellular hairs but gla- brate with age, the distal nodes sparingly beset with spreading simple multicellular hairs. Older branches somewhat quadrate to rounded with age, the nodes inconspicuously beset with interpetiolar lines or ridg¬ es. Mature leaves membranaceous, entire but vary¬ ing to inconspicuously denticulate, 10.1-14.4 cm long, 5. 5-7. 2 cm wide, elliptic to elliptic-ovate, acu¬ minate apically, obtuse to conspicuously oblique basally, glabrous adaxially, moderately to sparsely covered with a mixture of inconspicuously asperous- headed and scalelike multicellular hairs on the sec¬ ondary and higher order veins abaxially, 5-plinerved with uppermost primaries mostly diverging from the median nerve in alternate or subalternate fashion, the secondaries conspicuous abaxially and mostly 3-6.5 mm apart; petioles 2. 1-4.8 cm long, 1-1.5 mm wide. Inflorescence an erect modified cyme 5- 6 cm long, commonly branched from the base, bracts of the rachis nodes paired, linear-oblong to narrowly triangular, 1.5-4 mm long, 1 mm wide, with pu¬ bescence like that of the distal branchlets, bracteoles sessile, persistent, triangular, 0.5- 1.5 mm long, 0.5 mm wide, glabrous. Pedicels 1-1.6 mm long. Hy¬ panthia (at anthesis) campanulate, 1.5-2 mm long to the torus (vascular ring). Calyx fused into a dome in young buds but rupturing at anthesis into 4 or 5 persistent broadly rounded hyaline lobes mostly 0.5- 1.5 mm long and 0.5 - 1 . 1 mm wide basally; exterior calyx teeth 5, narrowly triangular, 0.5-0. 6 mm long, typically exceeding the lobes, torus glabrous and undulately lobed abaxially. Petals 5, glabrous, pale pink, linear-oblong, obtuse to rounded but often Novon 3: 5-10. 1993. 6 Novon F Figure 1. Miconia friedmaniorum Almeda & Umana. — A. Habit. — B. Representative leaf, abaxial surface (left) and enlargement of median vein (right). C. Floral bract (abaxial surface). — D. Bracteoles, adaxial surface (left) and abaxial surface (right). — E. Berry. — F. Petal. — G. Stamens, ventral view (left) and profile view (right). — H. Seeds. (A-G from Almeda & Anderson 5416 ; H from Herrera 1465.) Volume 3, Number 1 1993 Almeda & Umana Dodero Miconia 7 somewhat cucullate and superficially appearing sub¬ acute apically when dry, 2.5-3 mm long, 0.9-1 mm wide. Stamens 10, isomorphic, filaments com- planate, glabrous, constricted and incurved distally, 0.8-1 mm long, 0.25-0.3 mm wide; anthers 1.2- 1.5 mm long, 0.3-0. 5 mm wide, yellow, linear- oblong but laterally compressed and ovate-oblong in profile view, somewhat truncate with a dorsally in¬ clined pore; connective thickened dorsally but not prolonged or appendaged at the filament insertion. Ovary (at anthesis) % inferior, 5-celled, globose, apex gently fluted, sparingly beset with glandular hairs but lacking a stylar collar. Style somewhat curved distally, glabrous, 5 mm long, 0.6 mm wide; stigma truncate and somewhat dilated. Berry globose, red turning purple when mature, 3-3.5 mm long and 3-3.5 mm diam. Seeds angular-pyramidate to some¬ what crescent-shaped in profile view, beige, ca. 0.5 mm long, smooth. Distribution. Known from the Monteverde Cloud Forest Reserve (Cordillera de Tilaran), the adjacent San Ramon F'orest Reserve, and Parque Nacional Rincon de la Vieja (Cordillera de Guanacaste) in northwestern Costa Rica, where it commonly occurs in cloud forests with an Atlantic slope exposure at elevations of (900—) 1 ,350- 1 ,700 m. Miconia friedmaniorum is notable for its brown pubescence, which consists of asperous-headed hairs that commonly obscure the surfaces of young branchlets, elevated primary veins on abaxial foliar surfaces and young hypanthia. These asperous- headed hairs most closely approach the dendritic hairs with short thin-walled (flattened) arms illus¬ trated by Wurdack (1986: 64). T his indument to¬ gether with the basally branched cymose inflores¬ cence, pink, linear-oblong petals, and 5-plinerved leaves with uppermost primaries diverging from the median vein in alternate fashion provide a diagnostic suite of characters distinguishing M . friedmaniorum from its congeners. In overall aspect, M. friedmaniorum is most rem¬ iniscent of M. pendula (also described herein). We interpreted the first fragmentary collection of the latter taxon as a conspecific variant of M. fried¬ maniorum. However, subsequent field study and the collection of good flowering and fruiting material indicate that M. pendula is distinctive and worthy of specific status. Miconia friedmaniorum resem¬ bles M. pendula in foliar shape and venation and in the details of petal, staminal, and seed morphol¬ ogy. Both species also share an inconspicuous calyx dome that ruptures into rounded hyaline lobes that are largely concealed by the prominent exterior ca¬ lyx teeth. Miconia pendula differs consistently in several important characters. It has a complex la- nate (not asperous) indument on young branchlets, petioles, inflorescence branches, and hypanthia, a mostly pendent modified cyme branched well above the node initiating the inflorescence, and bracts and bracteoles that are pustulate abaxially and invested basally with hairs that are often bifid or irregularly barbed at the apex. Unfortunately, the pustulate nature of the bracts and bracteoles is often distorted by pressing and drying and not always readily ob¬ served in herbarium material. This species is named in honor of Nathan Jay and Virginia Friedman, long-time members of the California Academy of Sciences, in recognition of their exemplary interest and support of botanical exploration and conservation in Costa Rica. Paratypes. Costa Rica, alajuela: Monteverde Re¬ serve, Cerro Negro, continental divide, 10°20'N, 84°50'W, 12 Sep. 1985 (fr), Bello 3242 (CAS, CR, MO); Reserva Forestal de San Ramon, camino de entrada cerca de la estacion, 10°13'N, 84°37’W, 6 July 1992 (fl), Gomez- Laurito 12073 (CR, USJ). alajuela-puntarenas bor¬ der: Cordillera de Tilaran, Monteverde Reserve, Sendero Brillante along continental divide, 25 Feb. 1992 (fr), Almeda & Daniel 7074 (CAS, CR). alajuela-guana- CASTE-PUNTARENAS BORDER: Cordillera de Tilaran, Mon¬ teverde Reserve, Sendero El Valle, 7 Mar. 1986 (fl), Almeda & Anderson 5416 (CAS, CR, MO, NY, FIS); Reserve Monteverde, vertiente Pacifico cerca de division continental (ventana), 31 July 1976 (fr). Dryer 535 (CR); Reserva Monteverde, vertiente Atlantico, Sendero El Valle, Dec. 1977 (fr), Dryer 1718 (CR). guanacaste: Parque Nacional Rincon de la Vieja, Cabeceras de Quebrada Provision y Quebrada Rancho Grande, Meseta Aguaca- tales, 10°46'N, 85°49'W, 1 Dec. 1987 (fr), Herrera 1465 (CAS, CR, MO). Miconia pendula Umana & Almeda, sp. nov. TYPE: Costa Rica. Cartago: Refugio Nacional de Vida Silvestre Tapanti, orilla de Sendero Los Palmitos, 1,300 1,400 m, 09°44'00"N, 83°47'00"W. 2 Aug. 1990 (fl), Umana et al. 391 (holotype, CR; isotypes, BM, BR, CAS, COL, CR, F, MENU, MO, NY, PM A, US, USJ). Figure 2. Frutex 4 m. Ramuli sicut petioli foliorum subtus venae primariae inflorescentia hypanthiaque dense setosi pilis 1 - 3 mm longis et modice pilis amorpho-pinoideis ca. 0.1- 0.25 mm longis puberuli. Lamina 10.1-20.3 x 6.4- 12.4 cm elliptica vel elliptico-ovata apice acuminata basi obtusa vel asymmetrice rotundato-obtusa, supra sparse strigosa vel glabrata, subtus in venis secundariis tertiariis- que sparse caduceque granuloso-furfuracea, 7-plinervata, membranacea. Inflorescentia 7-11.2 cm longa (pedun- culo 0.5-4. 7 cm arcuato incluso) multiflora; flores 5(-6)- meri, pedicellis (ad anthesim) 0.5-2. 5 mm longis, brac- teolis 0.2-0. 3 x 0.1 -0.2 cm persistentibus. Hypanthium (ad torum) 2-3 mm longum; calyx primum in cono clausus demum in lobos regulares persistentes ruptus, dentibus exterioribus 0.9-1. 3 mm eminentibus. Petala 2.5-4 x 7m m 8 Novon Figure 2. Miconia pendula Umana & Almeda. — A. Habit. — B. Representative leaf, abaxial surface (left) and enlargement of median vein (right). — C. Floral bract (adaxial surface). — D. Bracteoles, abaxial surface (left) and adaxial surface (right). — E. Petal. — F. Stamens, lateral view (left) and ventral view (right). — G. Berry. — H. Seeds. (A-F from Umana et al. 391\ G, H from Croat <& Grayum 68255.) Volume 3, Number 1 1993 Almeda & Umana Dodero Miconia 9 1 mm oblongo-elliptica glabra. Stamina isomorphica gla¬ bra; antherarum thecae 1.1 -1.5 x 0.4-0. 5 mm anguste oblongae, poro paulo dorsaliter inclinato; connectivum nec prolongatum nec appendiculatum. Stylus 2.5-5 mm gla- ber; ovarium 5-loculare et % inferum apice sparsiuscule glanduloso-puberulo. Laxly branched shrub to 4 m tall, young branch- lets, petioles of young leaves, and inflorescence ra- chis moderately to densely covered with a lanate indument of curly or sinuate barbellate or distally bifid multicellular hairs mostly 1-3 mm long that are intermixed with and seemingly grade into a ground layer of shorter amorpho-pinoid multicellular hairs. Older branches somewhat quadrate to rounded with age, bearing conspicuous interpetiolar lines or ridges. Mature leaves membranaceous, inconspic¬ uously denticulate to subentire, 10.1-20.3 cm long, 6.4 12.4 cm wide, elliptic to elliptic-ovate, acu¬ minate apically, obtuse to conspicuously oblique ba- sally, sparingly strigose to glabrous adaxially, lanate on the elevated primaries abaxially and diminutively furfuraceous on prominulous secondary and higher order veins, 7-plinerved with the uppermost pri¬ maries typically diverging from the median nerve in alternate or subalternate fashion, the secondaries conspicuous abaxially and mostly 4-8 mm apart; petioles 1 .5-9 cm long, 1 -2 mm wide. Inflorescence an arcuate or pendent, modified cyme 7-11.2 cm long (including a peduncle 0.5 4.7 cm long) branched well above the base with flowers aggregated into umbelliform clusters; bracts of the rachis nodes paired below but quaternate above, linear-oblong to broadly triangular, 0.2- 1.4 cm long, 0.1 0.2 cm wide, spar¬ ingly furfuraceous to essentially glabrous or with a few sinuate hairs on the adaxial surface, pustulate proximally on the abaxial surface, bracteoles sessile, persistent, triangular, 0.2-0. 3 cm long, 0. 1-0.2 cm wide, glabrous and pustulate basally on the abaxial surface. Pedicels 0.5-2. 5 mm long, lanate. Hypan- thia (at anthesis) campanulate, 2-3 mm long to the torus (vascular ring) with pubescence like that of the petioles. Calyx fused into a dome in bud but rupturing at anthesis into 5 broadly triangular hy¬ aline lobes 0.25-0.50 mm long and 1.25-1.6 mm wide; exterior calyx teeth 5(-6), green (in fruit), narrowly triangular, 0.9-1. 3 mm long, exceeding and obscuring the lobes, with pubescence like that of the hypanthia; torus glabrous and undulately lobed adaxially. Petals 5(— 6), glabrous, pale pink, linear- oblong, obtuse to rounded apically, 2.5-4 mm long, 1 mm wide. Stamens 1 0(— 12), isomorphic; filaments glabrous, somewhat geniculate distally, 1.2-2 mm long, 0.2-0.25 mm wide; anthers 1 . 1-1 .5 mm long, 0.4-0. 5 mm wide, yellow, linear-oblong, apically truncate with a somewhat dorsally inclined pore; connective thickened dorsally but not prolonged or appendaged at the filament insertion. Ovary (at an¬ thesis) % inferior, 5-celled, globose, apex somewhat fluted and sparsely beset with glandular hairs, collar 0.25 mm high. Style somewhat curved distally, gla¬ brous, 2.5-5 mm long, 0.5-0. 6 mm wide; stigma truncate and somewhat dilated. Berry globose, pink to red but turning purple when mature, 4-5 mm long, 5 mm diam. Seeds angular-pyramidate to somewhat crescent-shaped in profile view, beige, ca. 0.5 mm long, smooth. Distribution. Known only from the Tapanti Na¬ tional Wildlife Refuge on the northeastern slopes of the Cordillera de Talamanca and Braulio Carrillo National Park in the Cordillera Central where it occurs in the understory of wet primary forests near streams at elevations of 1,300-1,800 m. Miconia pendula may be expected elsewhere along the At¬ lantic montane corridor of the Cordillera de Tala¬ manca that extends southward to Panama. Clidemia utleyana Almeda, Gonocalyx almedae Luteyn, Monochaetum cordatum Almeda, and Stemodia re- liquianim D’Arcy, all of which are known in Costa Rica only from the Tapanti region, have also been collected in western Panama. Miconia pendula , which was first collected in 1984, is readily overlooked. The flowers are small, the petals fall away quickly, and the leaves are commonly so ravaged by phytophagous insects that collectors may be inclined to pass it up in hopes of finding undamaged material. Miconia pendula differs from M. friedmaniorum in having modally larger 7-plinerved leaves, but its most striking feature is the complex lanate indument of contorted or sinuate multicellular hairs that grade into a ground layer of amorpho-pinoid hairs. By virtue of their shared seed and petal char¬ acters, M. pendula and M. friedmaniorum appear to be related to a group of three largely Central American species that includes M. iteophylla Al¬ meda, M. ligulata Almeda, and M. peltata Almeda (Almeda, 1989). These three species have rounded- triangular hyaline calyx lobes that are not fused into a rupturing dome as floral buds enlarge. Among this trio of species, the two new taxa described here are perhaps closest to M. peltata , which differs most conspicuously in its elliptic -ovate peltate leaves and indument of rusty brown pinoid hairs. The epithet for this species alludes to the arcuate or nodding posture of the inflorescence. Paratypes. Costa Rica, cartago: Tapanti Wildlife Refuge, Sendero Palmito, 5 Mar. 1992 (fr), Almeda et al. 7245 (AAU, CAS, CR, DUKE, F, MO, NY, US); Tapanti Reserve ca. 1 km S of jet. of Quebrada Salta 10 Novon and Rio Grande de Orosi, 9°43'N, 83°47'W, 29 Sep. 1987 (fr), Croat <£ Grayum 6 8255 (CAS, CR, MO); Tapanti Reserve, ca. 1 km upstream from confluence of Quebrada Salta and Rio Grande de Orosi, 12 July 1984 (fl), Grayum & Sleeper 3486 (CAS, CR, MO). HEREDIA: Parque Nacional Braulio Carrillo, San Rafael de Vara Blanca, 10°13'N, 84°06'W, July 1992 (fl), Ballestero 157 (CR). Acknowledgments. We thank Ellen del Valle for preparing the illustrations, Pablo Sanchez, Manuel Ramirez, Juan Diego Alfaro, and the staffs of the Herbario Nacional de Costa Rica and the lnstituto Nacional de Biodiversidad for logistical support. Per¬ mission to collect specimens in protected areas of Costa Rica was kindly granted by the Tropical Sci¬ ence Center, the Servicio de Parques Nacionales, and the Ministerio de Recursos Naturales, Energia y Minas. Literature Cited Almeda, F. 1989. New species and taxonomic notes on Mexican and Central American Melastomataceae. Proc. Calif. Acad. Sci. 46(9): 209-220. Wurdack, J. J. 1986. Atlas of hairs for neotropical Melastomataceae. Smithsonian Contr. Bot. 63: 1- 80. A New Species of Deprea (Solanaceae) from Venezuela Barbro Axelius Botaniska Tnstitutionen, Stockholms Universitet, S-106 91 Stockholm, Sweden William G. D’Arcy Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A. Abstract. The range of the genus Deprea is dis¬ cussed, and a new species, Deprea hirtiflora, from Venezuela, is described and illustrated. An over¬ looked, second species, Deprea granulosa , is also presented and illustrated. Deprea Rafinesque is a small neotropical genus of Solanaceae. Its separation from Physalis was long ignored after first publication (Rafinesque, 1838), but the genus was reinstated by D’Arcy (1973), who also lectotypified it by D. orinocensis (Humboldt & Bonpland) Rafinesque. This view has been accepted by Hunziker (1977) and Barbosa & Hunziker (1989). The genus now includes eight species: the type spe¬ cies, D. xalapensis (Humboldt & Bonpland) Raf¬ inesque, D. sylvarum (Standley & Morton) Hunzik¬ er, D. glabra (Standley) Hunziker, D. cardenasiana Hunziker, D. granulosa (Miers) Barbosa & Hun¬ ziker, Deprea subtriflora (Miers) D’Arcy, and the species described here. D’Arcy (1979, 1986, 1993) has indicated that although the corolla is more deeply divided in Physalis subtriflora (the type species of Larnax Miers) than in other species now placed in Deprea , this species is actually a member of Deprea and hence Deprea and Larnax are congeneric. A difficulty in assessing the geographic range of Deprea is in the provenance of the specimens on which the original two species were described. In his description of Deprea, Rafinesque included Physalis xalapensis Kunth and Physalis orino¬ censis Kunth, both of which are typified by quite similar specimens in Paris (P). Subsequently, no species of Deprea has been found in the region around Xalapa, Mexico (Nee, 1986), the ostensible source of Physalis xalapensis, nor was any spec¬ imen of Deprea encountered from the region of the Orinoco, Venezuela, by D’Arcy, Benitez, and Nee, who have recently completed a treatment of the Solanaceae for the Flora of the Guayana High¬ lands. It is therefore reasonable to assume that the Kunth material actually came from somewhere other than Mexico or Venezuela. Plants similar to the material at Paris used by Kunth are commonly found in the uplands of Costa Rica and western Panama (D’Arcy, 1973), and also in Colombia. After excluding Mexico and the Orinoco lowlands as part of the range of Deprea, the genus appears to range from Costa Rica to Colombia {D. orini- nocensis, D. glabra, D. granulosa), Ecuador ( D . glabra), Peru (D. subtriflora ), and Bolivia (D. car¬ denasiana). The species described here, D. hirti¬ flora, extends the range of the genus eastward into Trujillo, Venezuela, but not as far as the Orinoco basin. Deprea hirtiflora Axelius & D’Arcy, sp. nov. TYPE: Venezuela. Trujillo: Mpio. de Bocano, camino al Cerro Guaramacal via la laguna de “Los Cedros,’’ cloud forest, 21 Mar. 1981, B. Stergios 2590 (holotype, MO). Figure 1A-C. Species Depreae orinicensi (Kunth) Rafinesque affinis, a qua corolla azurea, extus dense villosa differt. Differt a D. granulosa (Miers) Barbosa & Hunziker colore corollae, pubescentia corollae densiore, tubo corollae breviore et antheris longioribus. Branched, unarmed herb or shrub to 1.2 m tall, twigs terete, pubescent overall with conspicuous, weak, whitish 5-7-celled hairs that dry flattened, ribbonlike. Leaves subentire, ovate; blade mostly 3— 5x1-3 cm wide, apically acuminate, basally ob¬ tuse, cuneate or acuminate, veins arcuate-ascend¬ ing, ca. 5 on each side, not forming an evident submarginal vein, both sides evenly puberulent, slightly denser on the main veins beneath; petioles slender, 3-10 mm long. Inflorescences (1 3)-flow- ered fascicles terminal on peduncles in the leaf axils or branch forks, peduncle obsolete, pedicels slender, ca. 6 mm, puberulent. Flowers with fusiform buds; calyx pilose with elongate hairs, cupular, 5-lobate, lobed '/(-way down, the lobes small and narrowly triangular, ca. 0.5 mm long; corolla blue, tubular- infundibular, lobed ca. '/,-way down, the lobes ca. 3 mm long, narrowly triangular, ciliolate, the tube ca. 5 mm long, slightly expanding upward, ca. 3 mm wide at the widest, villous outside, glabrous within; Novon 3: 11-13. 1993. 12 Novon Figure 1. A-C. Deprea granulosa (Miers) Hunziker & Barbosa. — A. Habit. — B. Flower. — C. Pubescence of leaf. After Zarucchi 5157 (MO). D-H. Deprea hirtiflora Axelius & D’Arcy. — D. Habit. — E. Flower. — F. Flower opened to show stamens and style. — G. Pubescence of leaf. — H. Fruiting calyx with hole made to show fruit. After Stergios 2590 (MO). Drawing by John Myers. Volume 3, Number 1 1993 Axelius & D’Arcy Deprea 13 stamens 5, filaments inserted about '/,-way up the corolla tube, slender, glabrous, anthers oblong, 2 X 0.8 mm, not apiculate; ovary rounded-conical, gla¬ brous, style ca. 8 mm, slender, glabrous, stigma capitate. Fruit a globose berry, 5-9 mm; fruiting calyx subglobose, loosely enveloping the berry, open apically, 10-ribbed, overall pubescent; fruiting ped¬ icels ca. 10 mm. Deprea hirtiflora is only known from two col¬ lections made in paramo and cloud forest in upland Trujillo, Venezuela. This species is distinctive in its dense pubescence of weak hairs that collapse on drying to appear flat and ribbonlike, and in its blue, ca. 5-rnm-long co¬ rolla, which is densely hairy on the outside. Paratype. Venezuela, trujillo: Bocono, paramo Guaramacal, 2,640-2,700 m, 9°13'N, 70°13'W, 14 Mar. 1984, Luteyn & Cotton 9707 (MO). In most species of Deprea corolla tubes are gla¬ brous outside, and the corolla lobes sometimes have granular to pubescent margins. In two species, how¬ ever, the newly described D. hirtiflora and D. gran¬ ulosa (Fig. 1 D-G), the corolla is conspicuously pu¬ bescent with coarse, simple hairs. In D. hirtiflora, the hairs are dense and elongate and the corolla tube has a furry appearance, while Deprea gran¬ ulosa has sparse, scattered hairs on the corolla. The general appearance of trichomes is similar in both species. Deprea hirtiflora is very similar to D. granulosa, both being branched, unarmed herbs with weak, whitish pubescence. The leaves of D. granulosa are somewhat bigger (4 9 x 3-5 cm) than those in D. hirtiflora, it has longer petioles (8-20 mm), and its pedicels are longer (ca. 15 mm). The main differ¬ ences are found in the flower: in D. granulosa the corolla is greenish yellow and much longer (8 mm vs. 5 mm), the anthers are shorter (1.5 mm vs. 2 mm), and the filaments are inserted relatively higher in the corolla tube. Deprea granulosa is also apparently a rare spe¬ cies, though it was reported (Escobar, 1989) to be abundant in the vicinity of Medellin, Colombia. For our comparisons, we have seen three collections of it: Colombia. “N. Grenada, locis frigidis inter Ibaque et Cartago,” Goudot s.n. (BM). Colombia. Antioquia: Mpio. de Medellin, Cerro del Padre Amaya, 8.4 km from Medellin -Santa Fe highway on road to the summit, 6°16'N, 75°41'W, 2,770 m, 30 Mar. 1987, Zarucchi 5157 (HUA, MO). Colombia. Antioquia: Mpio. de Urrao, paramo de Frontino, El Rio, 3,1 15 m, 10 Oct. 1984, Ramira-Londono et al. 369 (MO). The first of these is an isotype of the species. Literature Cited Barbosa, G. & A. Hunziker. 1989. Estudios sobre So- lanaceae XXIX. Sinopsis taxonomica de Athenaea. Bol. Soc. Argent. Bot. 26: 91-105. D’Arcy, W. G. 1973. Solanaceae. In: R. E. Woodson et al.. Flora of Panama. Ann. Missouri Bot. Gard. 60: 573-780. - . 1979. The classification of the Solanaceae. Pp. 3-47 in J. G. Hawkes et al., The Biology and Taxonomy of the Solanaceae. Academic Press, Lon¬ don. - . 1986. The genera of Solanaceae and their types. Solanaceae Newsl. 2: 10-33. - . 1993 A new combination in Deprea (Sola¬ naceae). Novon 3: 22. Escobar, L. 1989. Inventario floristico de un bosque muy humedo montano bajo en el muncipio de Caldas, Antioquia. Actualidades Biologicas 18: 2-44. Hunziker, A. T. 1977. Estudios sobre Solanaceae VIII. Novedades varias sobre tribus, generos, secciones y especies de Sud America. Kurtziana 10: 7-50. Nee, M. 1986. Solanaceae I. Flora of Veracruz 49: 1- 191. Rafinesque, C. S. 1838. Sylva Telluriana. Rafinesque, Philadelphia. A New Combination in Exodeconus (Solanaceae) Barbro Axelius Missouri Botanical Garden and Botaniska Institutionen, Stockholms Universitet, S-106 91 Stockholm, Sweden U illiam G. D'Arcy Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A. Abstract. Cacabus flavus is transferred to the genus Exodeconus. the generic names Exodeconus Rafinesque (1838) and Cacabus Bernhardi (1839) are both based on the same type, Physalis prostrata L’Heritier de Brutelle. Cacabus was for many years erroneously used for this small group of plants from western South America, and not all necessary transfers into Exodeconus have been made. During a revision of the genus by Axelius, we examined the type collec¬ tion of Cacabus flavus and considered it a species belonging to Exodeconus. We make the necessary new combination below. In many species described in the same publication with Cacabus flavus, Johnston used the designation “type” and “isotype,” also noting the herbarium of deposit for material upon which he based his species, a practice that did not come into general use until some years later. In the case of Cacabus flavus, his material was designated as “type. Gray Herb.,” and another collection was cited as well. In accordance with present usage, we refer to the first as the holotype, a duplicate of it (which we have seen) as an isotype, and the second collection cited as a paratype. These were clearly Johnston’s concepts, although all of these terms did not come into cur¬ rency until much later. Exodeconus flavus (I. M. Johnston) Axelius & D'Arcy, comb. nov. Basionym: Cacabus flavus I. M. Johnston, Contr. Gray Herb. 85: 177. 1929. TYPE: Peru. Arequipa: open rocky slopes, Tiabaya, 2,050 m, 8 Apr. 1925, Pen¬ nell 13066 (holotype, GH; isotype, K). Paratype. Peru. MOQUEGUA: dry ravines in hills SE of Moquegua, 1,550 rn, 22-24 Mar. 1925, If eberbauer 7459 (F, K). Literature Cited Bernhardi, J. J. 1839. Ueber Saracha und Physalis. Linnaea 13: 357-362. Rafinesque, C. S. 1838. Sylva Telluriana. Rafinesque, Philadelphia. Novon 3: 14. 1993. Five New Tribes in the Scrophulariaceae Kerry Barringer Brooklyn Botanic Garden, 1000 Washington Avenue, Brooklyn, New York 11225, U.S.A. ABSTRACT. Four new tribes of Scrophulariaceae are described: Alonsoeae, Bowkerieae, Caprarieae, and Freylineae. The Globulariaceae are given new status as a tribe in the Scrophulariaceae. The relationships of many genera within the Scrophulariaceae are poorly understood. Thirty tribes are generally recognized, though some of these are clearly polymorphic and genera exist that do not fit within any recognized tribe. Recognition of the fol¬ lowing tribes clarifies some of the relationships within the family. Alonsoeae Barringer, tribus nov. TYPE: Alonsoa Ruiz & Pavon. Herbae suffruticesve. Folia opposita vel verticillata. Inflorescentia centripeta, uniformis. Corolla explanato-ro- tata, resupinata, tubus nullus, labium posticum exterius. Stamina 4, antheris divaricatis vel in unam confluentes. Staminodia carentes. Capsula septicida, valvis integris bi- fidisve. Semina exarata. Herbs or subshrubs. Leaves opposite or whorled. Flowers in terminal racemes or panicles, resupinate; lower bracts leaflike, the upper reduced. Pedicels ebracteolate. Calyx 5-parted, the lobes imbricate. Corolla rotate, resupinate, the lower lobes (from resupination) outermost in bud. Stamens 4, anthers unithecate and reniform or bithecate and parallel to divaricate; staminodes lacking. Ovary biloculate, with many ovules; stigma subcapitate. Capsule septicidal, the valves entire or bifid. Seed with a solid, longi¬ tudinally furrowed coat. A single genus native to Andean South America, Central America, and South Africa: Alonsoa Ruiz & Pavon, Syst. Veg. FI. Peruv. Chil. 150. 1798. TYPE: Alonsoa caulialata Ruiz & Pavon fide Pennell (1920). The true affinities of this tribe are difficult to discern. Bentham ( 1 846) included it in the tribe Hemimerideae, but it is unlike any of the other genera in the tribe. Its often unithecate anthers, rotate corolla, and furrowed seeds are reminiscent of Verbascum L. It appears to be most closely related to Verbascum and Scrophularia L., which also have harpagioside and related 8/3-8a-methyl substituted iridoids (Nicoletti et al., 1988). These compounds appear to mark a distinct clade within the Scroph- ulariales that also includes Capraria L. and the Pedaliaceae. While most species of Alonsoa are neotropical, Roux (1985) and Steiner (1989) have found South African plants that they refer to the genus. Similar plants were described by Kunze (1841) as Schis- tanthe Kunze, because they differed from Alonsoa in having the corolla slightly saccate, the two low¬ ermost corolla lobes split to the base and the capsules ovate, obtuse, and emarginate (Hilliard & Burtt, 1984). They also differ in having eliaophores in the corolla sacs similar to those found in the South African Hemimerideae and the neotropical Ange- loneae and Melospermeae. Whether the South Af¬ rican species are placed in Alonsoa or Schistanthe, they are closely related to the South American spe¬ cies (Steiner, 1989) and would also be placed in this tribe. Bowkerieae Barringer, tribus nov. TYPE: Bowk- eria Harvey. Frutex arboresve. Folia opposita vel verticillata. Inflo¬ rescentia composita, pedunculis cymoso-multifloris rarius unifloris bibracteatis. Corolla tubo brevi. Stamina 4 vel 2, antherarum loculis apice confluentibus. Staminodia parva vel carentes. Capsula septicida, valvis integris bifidisve. Semina reticulata. Shrubs or small trees. Leaves opposite or verti- cillate. Flowers in terminal or axillary cymes, some¬ times forming panicles; not resupinate; bracts grad¬ ing into leaves. Pedicels ebracteolate. Calyx 5-lobed, rarely 3-lobed, lobes imbricate or valvate. Corolla campanulate to funnelform with a conspicuous sac or pouch in the distal half of the tube, the upper lobes exterior in bud. Stamens 4 with 1 staminode occasionally present, or stamens 2 with 2 or 3 stam¬ inodes, anthers confluent, reniform. Ovary biloculate or rarely triloculate, with many ovules; stigma punc- tiform or slightly bifid. Capsule septicidal, the valves sometimes bifid. Seed with loose, thin reticulate coat. Three genera native to southern Africa: Ixianthes Bentham, Companion Bot. Mag. 2: 54. 1835. TYPE: Ixianthes retzioides Bentham. Novon 3: 15-17. 1993. 16 Novon Anastrabe E. Meyer ex Bentham, Companion Bot. Mag. 2: 54. 1835. TYPE: Anastrabe inte- gerrima E. Meyer. Bowkeria Harvey, Thes. Cap. 1: 24, t. 37. 1859. TYPE: Boivkeria triphylla Harvey. Bentham (1835, 1846) placed the genera in this tribe in the Cheloneae because of their cymose in¬ florescences. Within that tribe, he grouped it with African genera having inconspicuous staminodia or lacking them entirely. These genera are now seg¬ regated into the tribes Teedieae G. Don, Freylinieae Barringer, and Bowkerieae Barringer. The Bowk- erieae differ from the Teedieae in having dehiscent capsules. They differ from the Freylineae in having a short corolla tube, valvate calyx, and confluent anthers. The seeds found in this tribe are similar to those of the Hemimerideae, but the Bowkerieae dif¬ fer from that tribe in their shrubby habit, cymose inflorescences, and confluent anthers. Hilliard (1969) pointed out the close relationship between Anastrabe and Bowkeria. Ixianthes is a rare, monotypic genus of shrubs with whorled, linear leaves, 3- or 5-parted calyx, and flowers with two stamens. It differs from the other two genera in the tribe by having more elongate corollas, but it shares with the others the saccate corolla, shrubby habit, and confluent anthers that distinguish the tribe. Caprarieae Barringer, tribus nov. TYPE: Ca¬ pra ria L. Herbae. Folia alterna. Inflorescentia centripeta. Corolla actinomorpha vel subzygomorpha campanulata lobis ae- qualibus. Stamina quinque vel quattuor aequales, anther- arum loculis apice confluentis. Capsula loculicida. Semina reticulata. Herbs. Leaves alternate. Flowers axillary or in terminal or axillary racemes. Bracteoles absent. Se¬ pals 5, free or fused only at the base. Corolla ac- tinomorphic or slightly zygomorphic, 5-lobed, broad¬ ly campanulate, not saccate or spurred, the lobes imbricate in bud, spreading. Stamens 5, inserted at the base of the corolla tube, the anthers exserted, bithecate, the thecae fused at the tip, divaricate. Stigma exserted, subcapitate to bilobed. Capsule lo- culicidal. Seed reticulate. One genus native to Central America, western South America, and the Caribbean Islands, wide¬ spread as a weed in tropical regions: Capraria L., Sp. PI. 628. 1753. TYPE: Capraria bi flora L. Bentham (1876), Sprague (1921), and Thieret (1954) placed Capraria L. in the tribe Digitaleae near Scoparia L. and Sibthorpia L., two other genera with actinomorphic corollas and alternate leaves. Bentham’s Digitaleae, like his Cheloneae, was a grab bag of genera that are best arranged into different tribes (Thieret, 1967), with Capraria in a tribe of its own. The genus is unusual in the family. The leaves are alternate and spiral, the corolla is campanulate and often radially symmetrical. The pollen is tri- colpo-diorate, an unusual type in the family found only in Capraria L. and a few species of Mimulus L. (Niezgoda & Tomb, 1975). It is one of the few Scrophulariaceae containing harpagide compounds, iridoid glycosides found more commonly in Pedali- aceae but also found in Scrophularia L. and its relatives (Heinrich, 1989). Classification of the ge¬ nus is still unsettled. It appears to have some re¬ lationship to the tribe Leucophylleae and the My- oporaceae (Niezgoda & Tomb, 1975). Freylinieae Barringer, tribus nov. TYPE: Frey- linia Pangella. Frutices suffruticesve. Folia opposita vel verticillata rarissime alterna. Inflorescentia composita, pedunculis cy- moso-multifloris rarius unifloris bibracteatis. Corolla tubo infundibuliformi, labium posticum exterius. Stamina 4, antherarum loculis distinctis. Staminodia parva vel car- entes. Capsula septicida, valvis integris bifidisve. Semina reticulata. Shrubs or subshrubs. Leaves opposite or verti- cillate, rarely scattered along the stem. Flowers in terminal or axillary cymes, often secondarily clus¬ tered into panicles, bracts grading into leaves. Ped¬ icels ebracteolate. Calyx 5-parted, lobes imbricate. Corolla tubular to funnelform, the upper lobes ex¬ terior in bud. Stamens 4, anthers with two parallel thecae. Staminodes short or absent. Stigma capitate, subcapitate, or sometimes slightly bifid. Capsule sep- ticidal, secondarily loculicidal in Freylinia. Seed with thin reticulate exotesta. Four genera native to southern Africa: ]>hygelius E. Meyer ex Bentham, Companion Bot. Mag. 2: 53. 1836. TYPE: Phygelius capensis E. Meyer ex Bentham. Freylinia Pangella ex Colla, Freyl. Gen. Add. Icon. (Hort. Ripul. 56). 1823. TYPE: Freylinia ces- troides Colla. Antherothamnus N. E. Brown, Hooker’s Icon. PI. t. 3007. 1915. TYPE: Antherothamnus pear- sonii N. E. Brown. Manuleopsis Thellung, Vierteljahr Nat. Ges. Zurich 60: 405. 1915. TYPE: Manuleopsis dinteri Thellung. Volume 3, Number 1 1993 Barringer New Tribes in Scrophulariaceae 17 This tribe is related to the Bowkerieae, described above, and is distinguished from that tribe by its anthers with two parallel thecae and its elongate corolla tube. Bentham (1876) described Phygelius as having confluent anthers but, as Dyer (1975) pointed out, the anthers have two parallel thecae. Globularieae (A. DC.) Barringer, stat. nov. Glob- ulariaceae A. DC. in Lamarck & A. DC., fl. Fran^. 1805. TYPE: Globularia L. Herbs or shrubs. Leaves alternate or rosulate. Flowers in globose capitula, terminal or rarely ax¬ illary; not resupinate; lower bracts not leaflike, form¬ ing an involucre. Calyx campanulate, 5-lobed, the lobes imbricate. Corolla tubular to funnelform, the upper lobes outermost in bud. Stamens 4 or rarely 2; staminodes lacking; anthers uniloculate, conflu¬ ent. Ovary uniloculate, uniovulate; stigma subcap- itate to minutely bifid. Fruit indehiscent. Seed with a solid, slightly reticulate coat. Two genera native to the Mediterranean region of the Atlantic islands: Globularia L., Sp. PL 95. 1754. TYPE: Globularia vulgaris L. Poskea Vatke, Linnaea 43: 321. 1882. TYPE: Poskea africana Vatke. Many authors recognize this group as a distinct family following Wettstein (1891), while others com¬ bine it with the genera of the tribe Selaginieae and recognize that group as a family following Bentham (1876). Hallier (1903) was the first to combine these groups with the Scrophulariaceae. There is a very strong relationship among the tribes Manuleae, Se- lagineae, and Globularieae. They appear to form a distinct clade within the family that is characterized by a reduction in ovule and seed number and con¬ solidation of the flowers into more densely packed inflorescences. Acknowledgments. I thank the curators of A, F, GH, MO, and NY for allowing me to study their collections. I also thank the librarians at NY and MO for their help. Literature Cited Bentham, G. 1835. [Classification of the Scrophulari¬ aceae.] Bot. Reg. 21 (post t. 1770): 1-7. - . 1846. Scrophulariaceae. In: A. DeCandolle, Prodromus 10: 186-586. - . 1876. Scrophulariaceae. In: G. Bentham & J. D. Hooker, Genera Plantarum 2: 913-980. Dyer, R. A. 1975. The Genera of South African Plants, vol. 1. Dicotyledons. Department of Agricultural Technical Services, Pretoria. Hallier, H. 1903. IJber die Abgrenzung und Verwand- schaft der einzelnen Sippen bei der Scrophulariaceen. Bull. Herb. Boissier ser. 2, 23: 181-207. Heinrich, M. 1989. Ethnohotanik der Tieflandmixe (Oa¬ xaca, Mexico) und phytochemische Untersuchung von Capraria biflora L. Diss. Bot. 144: 1-81. Hilliard, O. M. 1969. Bowkeria and Anastrabe: Two woody South African Scrophulariaceae. Notes Roy. Bot. Card. Edinburgh 29: 1-14. - & B. L. Burtt. 1984. A revision of Diascia section Racemosae. J. South African Bot. 50: 269- 340. Kunze, G. 1841. Index Seminum Hortus Universitatis Lipsensis. Nicoletti, M., M. Serafina, J. A. Garbarino & V. Gambaro. 1988. A chemosystematic study of Scrophularia¬ ceae: Iridoid glycosides. Giorn. Bot. Ital. 122: 13 24. Niezgoda, C. & S. A. Tomb. 1975. Systematic paly- nology of the tribe Leucophylleae (Scrophulariaceae) and selected Myoporaceae. Pollen & Spores 17: 497- 516. Pennell, F. W. 1920. Scrophulariaceae of Colombia, I. Proc. Acad. Nat. Sci. Philadelphia 72: 136-188. Roux, J. P. 1986. Alonsoa peduncularis rediscovered. South African J. Bot. 52: 7-9. Sprague, T. A. 1921. A revision of the genus Capraria. Kew Bull. 1921: 205-212. Steiner, K. E. 1989. A second species of the Amphi- Atlantic genus Alonsoa (Scrophulariaceae). Ann. Missouri Bot. Card. 76: 1152-1159. Thieret, J. W. 1954. The tribes and genera of Central American Scrophulariaceae. Ceiba 4: 164-184. - . 1967. Supraspecific classification in the Scrophulariaceae: A review. Sida 3(2): 87-106. Wettstein, R. von. 1891. Scrophulariaceae. In: A. En- gler & K. Prantl (editors), Die Natiirlichen Pflan- zenfamilien, ed. 1. 4(3b): 39-107. Orchidaceae Dunstervillorum VI: New Pleurothallidinae from the Venezuelan Guayana German Carnevali Jardm Botanico de Caracas, Herbario Nacional de Venezuela (VEN), INPARQUES, Aptdo. 2156, Caracas 1010-A, Venezuela; present address: Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. and Department of Biology, University of Missouri- St. Louis Gustavo A. Romero Oakes Ames Orchid Herbarium, Harvard University Herbaria, 22 Divinity Avenue, Cambridge, Massachusetts 02138, U.S.A. ABSTRACT. The following two taxa are proposed as new: Pleurothallis erebatensis and Trichosalpinx oxychilos. Both species are illustrated with plates by the late G. C. K. Dunsterville. Comments are provided on their relationships. During preparation of the treatment of the Or¬ chidaceae for the Flora of the Venezuelan Gua¬ yana, we have detected the following undescribed taxa in the subtribe Pleurothallidinae. Pleurothallis erebatensis Carnevali & C. Rom¬ ero, sp. nov. TYPE: Venezuela. Bolivar: Mun- icipio Sucre, alrededores de Santa Maria de Erebato, 4°59'N, 64°49'W, 360 m, feb. 1989, Elio Sanoja 2554 (holotype, PORT; isotypes, MO, VEN). Figure 1. Species haec P. aphtosae Lindley affinis, sed planta minore, ovario glabro, sepalis extus conspicue carinatis, minute papillosis non conspicue pubescentibus, intus gla- brescentibus non verruculosis, dorsali base non atenuato, synsepalo versus apicem connatis, labello angustiore dis- crepat. Epiphytic herbs, caespitose to shortly creeping, erect to subpendent; rhizome covered with scarious sheaths. Stems (1.5-)3-8.5 cm long, terete, the basal clothed in a close-fitting tubular brown sheath. Leaves 5. 5-9. 5 cm long, ( 1 — )1 .3— 1 .8( — 2) cm wide, fleshy-coriaceous when fresh, coriaceous when dry, erect on the stem, narrowly elliptic to oblanceolate, apex obtuse to acute, the midnerve sulcate, dorsally angled. Inflorescences 5-10 mm long, 1-5, successive or 2 or 3 simultaneously, erect or deflexed downward; peduncle very short and com¬ pletely covered by a sheath at the base of the leaf; rachis completely covered by the floral bracts, 4- 7-flowered; floral bracts ca. 2 mm long, fleshy, broadly cupuliform, apex acute, dorsally carinate. Flowers resupinate or not, dark brownish red, open¬ ing ± simultaneously; pedicel 0.9- 1.1 mm long, terete, ovary 1.2- 1.5 mm long, glabrous, terete. Sepals thick-fleshy, 3-nerved, dorsally prominently carinate, finely papillose on both surfaces but more conspicuously so without, apices sharply mucronate; dorsal sepal 4.8-5 mm long, 1.8-2 mm wide, con¬ cave, oblong to elliptic-oblanceolate when flattened, acute, forming a hood over the column; lateral sepals connate for % of their length into a concave synsepal 4.9-5. 1 mm long, when spread 4. 3-4. 4 mm wide, basally produced into a prominent mentum. Petals 2. 4-2. 6 mm long, 0.9- 1 . 1 mm wide, thin but fleshy, 1 -nerved, oblanceolate to obtrullate, obtuse, margins in the apical half finely and irregularly erose, gla¬ brous. Labellum 3-3.2 mm long, 1-1.1 mm wide in natural shape, 1.5 mm wide when flattened, 3-nerved, concave, narrowly oblanceolate in natural shape, becoming oblong-elliptic to elliptic when spread, rounded to obtuse, fleshy, thicker at middle, basally with a pair of minute, retrorse lobules, an¬ terior margin erose-laciniate; disk covered with pu¬ bescence of hairs in transverse lamellalike rows, coarser in the apical half; callus composed of 2 submarginal, parallel keels with erose dentate mar¬ gins, located in the middle % of the labellum. Column 1-1.3 mm long, straight or ± arcuate; margins prominently winged, apically 3-lobed, basally pro¬ duced into a conspicuous foot 11.3 mm long. Fruits 0.9- 1.3 mm long, obliquely ellipsoid. (Subg. Acianthera (Scheidweiler) Luer, sect. Hrachysta- chyae Lindley.) Pleurothallis erebatensis resembles several oth¬ ers in section Brachystachyae, and it is difficult to Novon 3: 18-21. 1993. Volume 3, Number 1 1993 Carnevali & Romero New Pleurothallidinae from the Venezuelan Guayana 19 Figure 1. Pleurothallis erebatensis Carnevali & G. Romero. — A. Flowering habit. — B. Inflorescence. — C. Lateral view of labellum and column. — D. Sepals and petals, spread. — E. Labellum, spread. (Based on Dunsterville & Dunsterville 1311.) ascertain which is its closest relative. It is similar to Pleurothallis aphtosa Lindley, from Colombia, Bo¬ livia, Paraguay, and southern Brazil, from which it is easily distinguished by its smaller vegetative and floral stature, its conspicuously dorsally carinate se¬ pals that are papillose without and glabrescent within (pubescent without and verruculose within in P. aphtosa), the synsepal connate almost to the apex, and a narrower labellum. The collectors of the new species have not described the fragrance of the 20 Novon Figure 2. Trichosalpinx oxychilos Carnevali & G. Romero. — A. Habit. — B. Detail of stem, base of leaf and inflorescence. — C. Lateral view of column. — D. Clinandrium. — E. Labellum and petal. — F. Sepals, spread. — G. Anther and pollinia. (Based on Dunsterville & Dunsterville 1347.) flowers, but it is likely to be similar to that of Pleu- rothallis aphtosa. A plant illustrated as P. aphtosa from Bolivia ( Vazquez 177, Herbarium Vasque- zianum, Dodson & Vazquez, 1990, t. 276) is most likely a variant of Pleurothallis erebatensis. Paratypes. VENEZUELA. BOLIVAR: vie. of Sta. Maria de Erebato, 400 m, Mar. 1974, Dunsterville & Dunsterville 13 1 1 (AMES), tachira: 10 km E of La Fundacion, around Represa Dorada, 600-1,000 m, 10-13 Mar. 1981, Lies- ner ukasovii (Vargas) Gereau & Brako, Hip¬ peastrum cuzcoense (Vargas) Gereau & Brako, Hippeastrum ferreyrae (Traub) Gereau & Brako, Hippeastrum hugoi (Vargas) Gereau & Brako, Hip¬ peastrum intiflorum (Vargas) Gereau & Brako, Hippeastrum leonardii (Vargas) Gereau & Brako, Hippeastrum macbridei (Vargas) Gereau & Brako, Hippeastrum variegatum (Vargas) Gereau & Brako, Ismene hawkesii (Vargas) Gereau & Meerow, Is¬ mene morrisonii (Vargas) Gereau & Meerow, Is¬ mene ringens (Ruiz & Pavon) Gereau & Meerow, Ismene sublimis (Herbert) Gereau & Meerow, and Leptochiton helianthus (Ravenna) Gereau & Mee¬ row. During the inventory of Peruvian Amaryllidaceae for the Catalogue of the Flowering Plants and Gymnosperms of Peru (Zarucchi & Brako, in prep.), it became clear that the following new combinations are needed to treat the Peruvian species in accor¬ dance with current nomenclatural rules and the ge¬ neric concepts employed by Meerow (1990) in the Flora of Ecuador. 1. Hippeastrum Herbert Based on the lectotypification of Amaryllis L. proposed first by Hitchcock (1929: 145) and the lectotypification of its type species A. belladonna L. proposed by Sealy (1939) and accepted by Dandy & Fosberg (1954), Goldblatt (1984) proposed the conservation of Amaryllis for the single South Af¬ rican species. The unanimous acceptance of this proposal by the Committee for Spermatophyta (Brummitt, 1987) resulted in the authorization of its use in this sense under Article 15 of the Inter¬ national Code of Botanical Nomenclature (Greuter et al., 1988: 178) and necessitates the transfer of all New World species formerly included in Ama¬ ryllis to the conserved genus Hippeastrum. The following new combinations are needed for species occurring in Peru: Hippeastrum bukasovii (Vargas) Gereau & Brako, comb. nov. Basionym: Amaryllis bu¬ kasovii Vargas, PI. Life 31: 30. 1975. TYPE: Peru. Puno: Prov. Sandia, bridge at San Jose, 1,400-1,800 m, C. Vargas C. 21882 (holo- type, CUZ). Hippeastrum cuzcoense (Vargas) Gereau & Brako, comb. nov. Basionym: Amaryllis cuz- coensis Vargas, PI. Life 31: 32. 1975. TYPE: Peru. Cuzco: Prov. Galea, Hacienda Vilcabam- ba, 2,800 m, C. Vargas C. 22395 (holotype, CUZ; isotype, USM). Hippeastrum ferreyrae (Traub) Gereau & Brako, comb. nov. Basionym: Amaryllis ferreyrae Traub, PL Life 6: 62. 1950. TYPE: Peru. Loreto: Isla Santa Maria, near Yurimaguas, Huallaga Valley, 150 200 m, R. Ferreyra 4497 (holotype, MO). Hippeastrum hugoi (Vargas) Gereau & Brako, comb. nov. Basionym: Amaryllis hugoi Var¬ gas, Herbertia 40: 126. 1984. TYPE: Peru. La Libertad: Prov. Bolivar, Dist. Bambamarca, 2,800 m, C. Vargas C. 22651 (holotype, CUZ). Hippeastrum intiflorum (Vargas) Gereau & Brako, comb. nov. Basionym: Amaryllis inti- fora Vargas, Bob Fac. Ci. Univ. Nac. Cuzco 1: 2. 1960. TYPE: Peru. Cuzco: Prov. Quis- picanchis, Hacienda Cadena, Valle de Marca- pata, C. Vargas C. 12985 (holotype, CUZ). Novon 3: 28-30. 1993. Volume 3, Number 1 1993 Gereau et al. Hippeastrum, Ismene, and Leptochiton 29 Hippeastrum leonardii (Vargas) Gereau & Brako, comb. nov. Basionym: Amaryllis leonardii Vargas, Herbertia 40: 131. 1984. TYPE: Peru. Puno: Prov. Sandia, San Juan del Oro, C. Var¬ gas C. 21654 (holotype, CUZ). Hippeastrum macbridei (Vargas) Gereau & Brako, comb. nov. Basionym: Amaryllis mac¬ bridei Vargas, Biota 8: 240. 1970. TYPE: Peru. Cuzco: Urubamba (cult, from bulbs col¬ lected in Puno, Sandia), 2,840 m, C. I argas C. 16422 (holotype, CUZ; isotype, USM). Hippeastrum variegatum (Vargas) Gereau & Brako, comb. nov. Basionym: Amaryllis var- iegata Vargas, PI. Life 31: 29. 1975. TYPE: Peru. Puno: Prov. Sandia, near Oconeque, 1,800 m, C. Vargas C. 16423 (holotype, CUZ). 2. Ismene Salisbury Herbert (1837) recognized Elisena Herbert, Hy- menocallis Salisbury, and Ismene as related but separate genera. Baker (1888) maintained Elisena at generic rank, but reduced Ismene to Hymeno- callis subg. Ismene (Salisbury) Baker. Macbride (1936) simply combined Elisena and Ismene with Hymenocallis with no recognition of subgeneric dis¬ tinctions. Meerow (1988, 1990) included Elisena and Pseudostenomesson Velarde in Ismene, and despite considerable polymorphism in flower and pollen morphology, he considered the expanded Is¬ mene a monophyletic group deserving generic status on the basis of vegetative, ovary, and seed mor¬ phology and chromosome number. The following new combinations are needed for species occurring in Peru: Ismene hawkesii (Vargas) Gereau & Meerow, comb. nov. Basionym: Hymenocallis hawkesii Vargas, PI. Life 31: 21. 1975. TYPE: Peru. Cuzco: Prov. Quispicanchis, Lucre, 3,050 m, C. I argas C. 16995 (holotype, CUZ). Ismene morrisonii (Vargas) Gereau & Meerow, comb. nov. Basionym: Stenomesson morrisonii Vargas, Natl. Hort. Mag. 22: 132. 1943. Pseu¬ dostenomesson morrisonii (Vargas) Velarde, Revista Ci. (Lima) 51 : 47. 1949. Hymenocallis morrisonii (Vargas) Traub, PI. Life 21: 96. 1965. TYPE: Peru. Apurimac: Prov. Abancay, gorge of Matara, Hacienda Soccospampa, 2,400-2,800 m, C. I argas C. 2231 (holotype, CUZ not seen; isotype, MO). On the basis of a photo of the plant and a sketch of the flower, Ravenna (1988) rejected Traub’s transfer of Stenomesson morrisonii to Hymenocallis, thereby refuting its affinity with Ismene (but without mentioning Velarde’s transfer of S. morrisonii to Pseudostenomes¬ son). Examination of the isotype at MO clearly reveals two basal, globose ovules in each locule of the ovary. Furthermore, the relatively large globose seed (fleshy when fresh) can be ob¬ served in a developing capsule on the isotype. The locules of all species of Stenomesson have numerous ovules, which are flattened and de¬ velop into flat, dry, winged seeds. Ismene ringens (Ruiz & Pavon) Gereau & Mee¬ row, comb. nov. Basionym: Pancratium rin¬ gens Ruiz & Pavon, FI. Peruv. 3: 53. 1802. Liriope ringens (Ruiz & Pavon) Herbert, Ap¬ pendix 42. 1821. Elisene ringens (Ruiz & Pavon) Herbert, Amaryllidaceae 201. 1837. Hymenocallis ringens (Ruiz & Pavon) J. F. Macbride, Field Mus. Nat. Hist., Bot. Ser. 13(1): 673. 1936. TYPE: Ruiz & Pavon, FI. Peruv. 3: t. 283, fig. b. 1 802 (lectotype, selected here). Ismene sublimis (Herbert) Gereau & Meerow, comb. nov. Basionym: Elisena sublimis Her¬ bert, Bot. Mag. 67: sub pi. 3873. 1841. Hy¬ menocallis sublimis (Herbert) J. F. Macbride, Field Mus. Nat. Hist., Bot. Ser. 11: 11. 1931. TYPE: Peru. La Libertad: Cajamarquilla, Ma¬ clean s.n. (holotype, K). 3. Leptochiton Sfai.y Sealy (1937) described the genus Leptochiton based on a single species, L. quitoensis (Herbert) Sealy, an ephemeral geophyte of southwestern Ec¬ uador and adjacent northwestern Peru. Meerow7 (1990) maintained this genus (as Lepidochiton, orth. var.) largely on the basis of ovary and seed mor¬ phology. Although we have not yet been able to examine type material, there can be no doubt from the illustration in Ravenna (1980) that the following new7 combination is needed: Leptochiton helianthus (Ravenna) Gereau N Meerow7, comb. nov. Basionym: Hymenocallis heliantha Ravenna, Bot. Not. 133: 97. 1980. TYPE: Peru. Cajamarca: grassy slopes in re¬ gion of Magdalena, between Pacasmayo and Cajamarca, cult, in Santiago de Chile, P. Ra¬ venna 2059 (holotype. Herb. Ravenna not seen; isotype, USM not seen). Literature Cited Baker, J. G. 1888. Handbook of the Amaryllideae. George Bell & Sons, London. Brummitt, R. K. 1987. Report of the committee for Spermatophyta. Taxon 36: 734-739. 30 Novon Dandy, J. E. & F. R. Fosberg. 1954. The type of Amaryllis belladonna L. Taxon 3: 231-232. Goldblatt, P. 1984. (748) Proposal to conserve 1176 Amaryllis and typification of A. belladonna (Amar- yllidaceae). Taxon 33: 511-516. Greuter, W., H. M. Burdet, W. G. Chaloner, V. Demoulin, R. Grolle, D. L. Hawksworth, D. H. Nicolson, P. C. Silva, F. A. Stafleu & E. G. Voss. 1988. Inter¬ national Code of Botanical Nomenclature. Regnum Veg. 118. Herbert, W. 1837. Amaryllidaceae. James Ridgway & Sons, London. Hitchcock, A. S. 1929. Genera 1-500. Pp. 114-155 in A. S. Hitchcock & M. L. Green, Standard-species of Linnean genera of Phanerogamae (1753-54), pp. 111-199 in J. Ramsbottom et al. (editors). Inter¬ national Botanical Congress, Cambridge (England), 1930. Nomenclature, Proposals by British Botanists, His Majesty’s Stationery Office, Wyman & Sons, London. Macbride, J. F. 1936. Amaryllidaceae Lindl. In: J. F. Macbride, Flora of Peru, Field Mus. Nat. Hist., Bot. Ser. 13(1): 631-690. Meerow, A. W. 1988. Evolutionary relationships in Hymenocallis Salisb. sensu lato (Amaryllidaceae). Amer. J. Bot. 75: 193. [Abstract.] - . 1990. Amaryllidaceae. In: G. Harling & L. Andersson (editors), Flora of Ecuador 41: 1-53, Berlings, Arlov, Sweden. Ravenna, P. 1980. Hymenocallis heliantha. Bot. Not. 133: 97. - . 1988. Studies in the genus Stenomesson (Amaryllidaceae). Onira 1: 17-21. Sealy, J. R. 1937. Leptochiton quitoensis. Bot. Mag. 160: tab. 9491. - . 1939. Amaryllis and Hippeastrum. Bull. Misc. Inform. 1939: 49-68. A New Ocotea (Lauraceae) from the High Mountains of Costa Rica and Panama Jorge Gomez-Laurito Escuela de Biologia, Universidad de Costa Rica, Ciudad Universitaria 2060, San Jose, Costa Rica & Herbario Nacional de Costa Rica ABSTRACT. Ocotea pharomachrosorum (Laura¬ ceae), of the Ocotea helecterifolia .-group from Costa Rica and Panama, is described here. Within the large genus Ocotea Auhlet (Laura¬ ceae), this new species belongs to the Ocotea he- lecterifolia- group, this group contains numerous borderline cases between the genera Ocotea. \ec- tandra, and Phoebe (Rohwer, 1991; 1992 in litt.). However, its outer tepals are glabrous inside, the inner ones have only a few papillae and the anthers are fleshy and smooth, not papillate, nor hood-shaped; these are all Ocotea- like characters. Although this species was collected for the first time several years ago, the material was insufficient to describe it. This collection was treated by Rohwer (1991) as Ocotea sp. A and by Burger & van der Werff (1991) as “a species of uncertain position.” Ocotea pharomachrosorum Gomez-Laurito, sp. nov. TYPE: Costa Rica. San Jose: Dota, San Gerardo, Finca de Efrain Chacon, 10°32'20"N, 83°49'05"W, 2,100 2,200 m, 10 feb. 1992, J. Gomez-Laurito , J. 1. Lopez, A. Mora & U . Barillas 12160 (holotype, CR; isotypes, E, MO, K, USJ). Figure 1. ' A speciebus quas Rohwer ad species e turma helec¬ terifolia ascripsit cornbinatione induinenti densissimi cris- pati cum basibus foliorum inaequilateralibus et nervis la- teralibus utroque 3-5 distinguenda. frees 8 12 m tall, 30-40 cm D.B.H., trunk straight, terete; hark grayish with transverse lenti- cels; inner bark yellowish; wood hard. Leafy bran- chlets densely grayish tornentulous with curled and matted hairs. Leaves alternate, distant; petioles 2.5 3.5 cm long; lamina 9- l 9 cm long, 5-7.5 cm broad, narrowly ovate to ovate-elliptic or ovate-ohlong, ta¬ pering to a short-acuminate apex, obtuse to rounded at the base, the sides of lamina unequal at the base with the sides 2-6 mm distant on the petiole, drying stiffly chartaceous or subcoriaceous, the upper sur¬ face glabrous and lustrous but with hairs above the slightly elevated proximal major veins, tertiary ve¬ nation slightly elevated, lower surface densely yel¬ lowish gray or whitish gray tornentulous, the hairs minute (0.1 -0.3 mm) and curved, with 3-5 major secondary veins on each side, the basal secondaries often strongly ascending, central secondaries arising at angles of 35°-50°. Young leaves very dense brownish tomentose. Inflorescences solitary and ax¬ illary to distal leaves or undeveloped leaves near the shoot tip, paniculate with short lateral branches sub¬ tended by conspicuous (4-7 mm) oblong bracts, peduncle, rachis and bracts densely brownish gray tornentulous. Flowers white, delicately scented; 6 tepals 4-5 mm long, 3 mm wide, externally pubes¬ cent, internally scarcely papillose; 9 fertile stamens, 6 outer, 3 inner, subsessile, thick, fleshy, smooth with hairs at base of filaments; glands sessile, 0.6 mm long. Ovary ovoid, glabrous. Style 1 mm long; stigma capitate. Cup 8-13 mm long, obconic, red- tinged. Berry ellipsoid, 3.5 cm long, up to 2 cm wide, green to purple at maturity. This new species and other Lauraceae are an important food source of one of the most beautiful birds in Central America: the resplendent quetzal ( Pharomachros rnocinno), hence the name. Ocotea pharomachrosorum is easily distinguished by its dense gray tomentum of branchlets and undersides of leaves; unusual long petioles to 3.5 cm long; subcoriaceous leaves with only 3 4 pairs of major veins and sides of the lamina unequal at the base. M oreover, the young leaves and shoots are con¬ spicuously dense brownish tomentose. It is presently known from the Pacific slope of the Cordillera de Talamanca in Costa Rica, and adjacent Chiriqui highlands in Panama, from 1.600 to ca. 2,300 m elevation. Paratypes. Same locality as type: Julio Sanchez s.n. (CR, USJ). Panama, chiriqui. Vic. of Boquete, Finca Collins, El Velo, 12 Mar. 1963 (US) Steam et al. 1985. Acknowledgments. I am gratelul to William C. Burger (Field Museum of Natural History, Chicago, IJ.S.A.) and Jens G. Rohwer (Institut fur Syste- rnatische Botanik mid Pflanzengeografie, Heidel¬ berg, Germany) for critical readings of the manu- Novon 3: 31-33. 1993. 32 Novon Figure 1. Ocotea pharomachrosorum Gomez-Laurito. — A. Flowering branch. — B. Leaf. — C. Leaf base. — D. Fruit. Drawing by lleana Ling. Volume 3, Number 1 1993 Gomez-Laurito Ocotea pharomachrosorum 33 script, and Julio Sanchez (Museo Nacional de Costa Rica) who brought the type locality to my attention. Literature Cited Burger, W. C. & H. van der Werff. 1990. Flora Costaricensis. Family # 80, Lauraceae. Fieldiana, But. New Ser. 23: 1-129. Rohwer, J. 1991. Borderline cases between Ocotea, Nectandra, and I’hoebe (Lauraceae): The “margin¬ al" species of the Ocotea helecterifolia- group, in¬ cluding the 0. heydeana-group. Bot. Jahrb. Syst. 112(3): 365-397. A Synopsis of the Panamanian Species of Ilex (Aquifoliaceae) William James Hahn Department of Botany, University of Wisconsin, Madison, Wisconsin 53706, U.S.A. ABSTRACT. The species of Ilex (Aquifoliaceae) na¬ tive to Panama are treated in a synoptic fashion. Three new species are described: Ilex fortunensis, Ilex maxima, and Ilex stellata. The Flora of Panama (Edwin, 1968) listed four species of Ilex for the country, based on only 17 specimens. One-half of these specimens were of the widespread Ilex guianensis (Aublet) Kuntze col¬ lected in the Pearl Islands. An increase in collecting activity over the past 20 years, especially in the moist uplands of Chiriqui, Code, and Bocas del Toro. Panama, has produced a current collection base of over 200 specimens representing 10 species of Ilex, including three previously undescribed. Similar changes in estimates of local species di¬ versity for the genus based on new collections occur elsewhere in the Neotropics. For example, extensive collecting in the Guayana Highlands since the 1950s has increased recorded diversity from 6 species of Ilex (Wurdack, 1961) to 55 species (Edwin, 1965) to 78 species (Steyermark, 1988). The typological species concepts employed by the latter two authors probably overstate diversity for the Guayana High¬ lands. The pattern of increase in recorded diversity is indisputable, however, and strongly suggests the need for additional collecting. In Panama, three main groupings of species can be made based on the nature of the inflorescence¬ hearing shoots. One group, composed of Ilex cos- taricensis J. D. Smith and /. pallida Standley, hears solitary, axillary inflorescences in both flower and fruit. This is equivalent to the solitary flowering unit composed of secondary dichasia sensu Loizeau (1992: figs. 2h, 13). These two species differ from each other in a number of characters and are not closely related. In the second group the inflorescence-hearing shoots are short while in flower (= brachyblasts), hut usually expand and develop normal leaves to¬ ward the apex as the fruits mature. In the termi¬ nology of Loizeau (1992: figs. 2g, 11, 12), these are proliferating thyrses. This group consists of four species in Panama: /. stellata W. ,1. Hahn, /. for¬ tunensis W. J. Hahn, Ilex lamprophylla Standley, and /. guianensis, which may be loosely related to each other. The first two taxa are endemic to Pan¬ ama and are known from only a few collections. Ilex lamprophylla is largely restricted to Panama and Costa Kica, with a few isolated populations in Nic¬ aragua. Related species are found in northern South America. Ilex guianensis is the most widespread member of a large species group centered in north¬ ern South America (Hahn, 1988). A third group of species includes /. vulcanicola Standley, I. ehiriquensis Standley, I. yuruman- guinis Cuatrecasas, and I. maxima W. J. Hahn. These species are characterized by abaxial leaf punc- tations and inflorescences borne on brachyblasts in both flower and fruit. The basic inflorescence unit is still a cyme, but the stem that bears the cymes fails to elongate as in the previous group and never bears leaves. The overall appearance of the cluster of cymes is umbellate or paniculate which corre¬ sponds to the non-proliferating thyrse of Loizeau (1992: fig. 10). The species form part of a natural group probably referable to Loesener’s series Pal- toria. This group of closely related species forms several grades that show broad continua of character states in leaf size and shape, inflorescence size and branching, and fruit size. Several collections show intermediate combinations of character states, mak¬ ing species identification difficult (see notes under /. ehiriquensis and I. yurumanguinis). As most spe¬ cies in this group are South American in distribution, a revision of the group in its main range will be necessary before the identities and relationships of the Panamanian taxa are completely resolved. Edwin (1968) reported I. maefaydenii (Walper) Rehder as possibly occurring in Panama, but this species is restricted to the Greater and Lesser An¬ tilles and possibly Mexico. It is distinguished by thin leaves with long-acuminate apices, distinctly serrate leaf margins, and generally solitary inflorescences. Key to the Species of Ilex in Panama la. Leaves epunctate abaxially; individual cymes solitary in the leaf axils when in fruit. 2a. Leaf margin entire or essentially so. Novon 3: 34-45. 1993. Volume 3, Number 1 1993 Hahn Panamanian Species of Ilex 35 3a. Leaves coriaceous, moderately to faintly discolorous, glaucous, glabrous or with a few scattered hairs, the margins revolute, secondary veins obscure to apparent; pistillate cyme branched or unbranched, peduncles 1-2 mm diain.; fruits 3-7 mm long, 2.5-7 mm diam. 4a. Leaves elliptic, drying dark brown or black abaxially, the margin revolute, strongly so at the base, venation brochidodromous, secondary veins apparent; cymes solitary in the leaf axils when in flower and in fruit; pistillate flowers 3-7 per dichasium (sometimes solitary at high elevations); fruits 3-7 mm long, 2. 5-5. 5 mm diam . 1. Ilex costaricensis J. D. Smith 4b. Leaves usually spatulate or obovate, drying olive green or brown abaxially, the margin revolute but not at the base, venation semicraspedodromous, secondary veins obscure; cymes clustered on brachyblasts when in flower, usually solitary when in fruit; pistillate flowers solitary; fruits 4.5-6 mm long, 4-7 mm diam . 2. Ilex guianensis (Aublet) Kuntze 3b. Leaves chartaceous to thin coriaceous, strongly discolorous, puberulent, the margin flat, secondary veins distinct; pistillate inflorescence branched, peduncles 2-3 mm diam.; fruits 0.9 1. 1 cm long, 0.7-1 cm diam . 3. Ilex fortunensis W. J. Hahn 2b. Leaf margin toothed or crenulate, sometimes minutely so, the teeth bearing small marginal spines. 5a. Leaves planar, elliptic to ovate or obovate, 3-6 cm long, 1.5-3 cm wide, glabrous or pubescent with simple hairs. 6a. Leaves drying pale green or olive, margin serrate-dentate, petiole 2-5 mm long; cymes solitary in flower and fruit; fruits 6-10 mm long, 4-8 mm diam . 4. Ilex pallida Standley 6b. Leaves frequently drying dark brown or black, margin crenulate or serrate to almost entire, petiole 3-45 mm long; cymes clustered or solitary in flower, usually solitary in iruit; fruits 3-6 mm long, 3-4 mm diam. 7a. Leaf margin usually distinctly crenulate, flat; petiole 3-8 mm long; cymes clustered on brachyblasts in flower; pistillate flowers solitary; pericarp thin, fruit usually deeply ribbed when dry . 5. Ilex lamprophylla Standley 7b. Leaf margin mostly entire, a few spines occasionally present, never crenulate, revolute, strongly so at the base and abaxially along the petiole; petiole 8-45 mm long; cymes solitary in the leaf axils when in flower; pistillate flowers solitary or more frequently 3 7 per dichasium; pericarp leathery, fruit surface smooth when dry . . 1. lie: v costaricensis J. L). Smith 5b. Leaves deeply to shallowly bullate, broad elliptic to rotund, 5-10 cm long, 3-8 cm wide, bearing deciduous stellate hairs . 6. Ilex stellata ^ . J. Hahn lb. Leaves with conspicuous glandular punctations abaxially; cymes in flower and fruit borne clustered on brachyblasts. 8a. Leaves coriaceous, 1 .5-10 cm long, orbicular to elliptic or obovate, secondary venation usually obscure; staminate cymes unbranched. 9a. Leaves 1.5-3 cm long; 5-10 cymes per fascicle; fruits 2.5-4 mm long ..7.1. vulcanicola Standley 9b. Leaves 3-10 cm long; 5-20 cymes per fascicle; fruits 4-7 mm long . . . 8. I. chiriquensis Standley 8b. Leaves chartaceous to coriaceous, (6-)8-40 cm long, elliptic, secondary venation distinct; staminate cymes branched. 10a. Leaves (6 )8 13 cm long, 4 7.5 cm wide with 6-8 secondary veins per side; fruit 0.3 0.6 cm long . 9. I. yurumanguinis Cuatrecasas 10b. Leaves 15-40 cm long, 10-13 cm wide, with 10-15 secondary veins per side; fruit 0.8 1 cm long . 10. /. maxima W. J. Hahn I. Ilex costaricensis .1. I). Smith, Bot. Gaz. (Graw- fordsville) 57; 416. 1914. TYPE: Costa Rica. Puntarenas: Cuesta de la Cara, hand procul ab El Paramo, Pittier 10183 (erroneously cited as Pittier 10813 in original publication) (ho- lotype, US 1380596). Ilex valerii Standley, J. Wash. Acad. Sci. 16: 483. 1926. TYPE: Costa Rica. Heredia: Cerros de Zurqui, NE of San Isidro de Las Vueltas, Standley & l a/erio 50582 (holotype, LIS 1251513; isotype, K). Ilex brenesii Standley, Field Mus. Nat. Hist., Bot. Ser. 18: 629. 1937. TYPE: Costa Rica. Entre La Palma y El Socorro de San Ramon, Brenes 6220 (holotype, F). Ilex ramonensis Standley, Field Mus. Nat. Hist., Bot. Ser. 18: 630. 1937. TYPE: Costa Rica. Entre Pata de Gallo y Santiago de San Ramon, Brenes 6639 (holotype, F). Ilex costaricensis is known throughout t Lie Tala- rnancas of western Panama and eastern Costa Rica through the Cordillera Central of Costa Rica and on Volcan Mombacho in Nicaragua. In Panama, the species occurs at elevations of 1,200 1,700 m; flowering mostly in August. Distinctive characters of Ilex costaricensis in¬ clude epunctate leaves, attenuate leaf bases, leaf margins entire and strongly revolute at the base and along the petiole, the blade drying black or dark brown, lenticellate stems drying dark brown to black, inflorescences solitary and axillary, and capitate stig¬ mas prominent in flower and fruit. Edwin (1968) treated Hex valerii from Costa Rica as a distinct species and included I. ramonensis Standley as a variety. Ilex costaricensis was ap¬ parently overlooked by Edwin, as he failed to men¬ tion the taxon in his treatment. The similarities be¬ tween the types of these three taxa are numerous 36 Novon and clearly support recognition of a single species. The type specimen of Hex brenesii (Brenes 6 266) is extreme in petiole, peduncle, and pedicel lengths, which are 3-10 times that found on other specimens of Ilex costaricensis. However, all other character¬ istics of the specimen agree with the type of Ilex costaricensis and fall within the range of variation accepted for that species. Smith (1914) placed Ilex costaricensis close to I. mexicana Hemsley based on apparent similarities in the inflorescences. Collections from Cerro Colorado, Chiriqui, on the Continental Divide {Folsom et al. 1755, McPherson 8807, and Mori <£• Dressier 7895) have smaller, thinner, narrowly elliptic to lanceolate leaves with the margin revolute at the base and along the portion attenuate to the petioles. Occasionally a few spines are present on the leaves. These differences are consistent among all collections from Cerro Colorado and are perhaps related to local environmental con¬ ditions, such as the high copper concentrations of the substrate. The specimen from Cerro Hornito ( Croat 67961) is similar to the type, however, and to most Costa Rican material. Additional specimens examined. Panama. BOCAS DEL TORO: Cerro Colorado, McPherson 8807 (MO, PMA). CHIRIQUI: Cerro Hornito, Croat 67961 (G, MO); Cerro Colorado, Folsom et al. 4755 (K, MO, PMA); Cerro Colorado, Mori & Dressier 7895 (F, MO, NY, PMA, US, NY). 2. Ilex guianensis (Aublet) Kuntze, Rev. Gen. PI. 1: 113. 1 891 . Macoucoua guianensis Aublet, Hist. PI. Gui. Fr. 88, t. 34. 1775. Ilex acu¬ minata Willdenow, Sp. PI. 1: 711. 1798. Ilex macoucoua Persoon, Syn. PI. 1: 152. 1805. TYPE: French Guiana, Aublet s.n. (holotype, P in herb. Rousseau, vol. 3, folio 100). Fig¬ ure 1 . Ilex panamensis Standley, Field Mus. Nat. Hist., Bot. Ser. 4: 221. 1952. TYPE: Panama. Bocas del Toro: Cooper 469 (holotype, F 579403; isotype, K). Ilex gentlei Lundell, Field & Laboratory 13: 5. 1945. TYPE: Belize. Toledo, Punta Gorda-San Antonio Road, Gentle 4807 (holotype, SMU not seen; photo, MO; isotype, K). Although somewhat variable, this species is char¬ acterized by its epunctate, coriaceous, spatulate to obovate leaves with entire, revolute margins and obscure secondary venation. While in flower, the cymes are borne on brachyblasts, which later expand as fruits mature resulting in clearly lateral or axillary cymes. Of the species native to Panama, Ilex guianensis has the widest distribution, ranging from the Guianas through Venezuela and Colombia along the Carib¬ bean coast of Central America to Tobasco, Mexico, and apparently throughout the Greater Antilles (Bornstein, 1989). It is also a major component of the vegetation of the Pearl Islands off the Pacific Coast of Panama (Johnston, 1949), illustrative of the numerous Caribbean species found on these is¬ lands (Bennet, 1968; D’Arcy, 1977). Flowering in Panama occurs during the major dry season of De¬ cember to April, with scattered occurrence at other times of the year, particularly for upland popula¬ tions. Closely related congeners forming Loesener’s sec¬ tion Micranthae subsect. Epunctatae are among the most variable and widespread within the genus (Bornstein, 1989). Previous studies of this group include a discussion of the relationships between /. guianensis and the widespread Caribbean taxon Ilex sideroxyloides (Swartz) Grisebach (Bornstein, 1989), and a limited study of the group as found in northern Central America (Hahn, 1988). Further study of the limits of variation in this group of species, par¬ ticularly in the Greater Antilles and in northern South America, would help in establishing species delimitations and relationships. In the present work, the species is treated as known from Central America and the Guianas; synonymy is limited to Central America. The presumably natural habitat for I. guianensis is sclerophyllous forest on seasonally dry, coastal or inland savannas typical of the type localities of Cay¬ enne and Sinamarry, French Guiana, cited by Au¬ blet. A similar sclerophyllous habitat with strong Caribbean affinities is also seen in the Pearl Islands, where most collections of the species in Panama have been made. The greatest degree of intraspecific variation, however, typically arises in recently dis¬ turbed areas at higher elevations Such middle-el¬ evation populations in Panama are known from Cer¬ ro Jefe, Cerro Campana, Cerro Azul, Boquete, and El Valle de Anton, all zones of deforestation within the last few hundred years (Bennet, 1968; D’Arcy, 1977). A number of other species closely related to Ilex guianensis have been described from Colombia (e.g., /. uniflora Bentham and /. danielis Killip & Cua- trecasas), which may represent high-elevation morphs as found in Panama. Ilex guianensis is apparently replaced in the Amazon Basin by the related /. inundata Poeppig. Reports of Ilex occidentalis Macfayden from An¬ con Hill (Hemsley, 1880; see also Standley, 1929), based on Seemann 554, and Ilex bumelioides Kunth (Hemsley, 1880), based on Duchassaing s.n., were founded on erroneous species concepts held by these authors (see Triana, 1872; Loesener, 1901), as Volume 3, Number 1 1993 Hahn Panamanian Species of Ilex 37 Figure 1. Ilex guianensis (Aublet) Kuntze. — A. Habit in fruit (Johnston 224). — B. Habit in pistillate flower (Gentle 1244). — C. Habit in staminate flower (Erlanson 411). 38 Novon these collections clearly represent Hex guianensis. The type of Ilex bumelioides Kunth is from Ec¬ uador; the species is more closely related to I. kun- thiana , while Ilex occidentalis Macfayden of Ja¬ maica is probably synonymous with I. guianensis. Additional specimens examined. Panama. BOCAS del TORO: Almirante, Bocas Island, Cooper 469 (F, G, K, NY); Chiriqui Lagoon, von IT edel 1144 (GH, MO). CHI¬ RIQUI: Llanos Francia, Dwyer & Lallathin 8698 (MO, NY); El Boquete, Holcomb’s Trail, Killip 3562a (US); Boquete, Llanos Francia, Stern et al. 1197 (GH, MO). COCLE: El Valle, Allen 752 (GH, MO); hills S of El Valle de Anton, Allen 2505 (MO, NY, US), Allen 3570 (F, G, GH, K, MO, NY, P, US); Penonome, Dwyer 2017 (MO); between El Gope and sawmill on Continental Divide, Hammel 2645 (MO, NY). COLON: along ocean trail be¬ tween Rio Indio and Miquel de la Borda, Croat 36916 (MO), Croat 36920 (MO); Salud, Lao & Holdridge 229 (MO); trail above Rio Indios, Sullivan 113 (MO). HERERRA: 10 km W of Las Minas on road to El Toro, D'Arcy & Sytsma 14319 (MO), Sytsma & D'Arcy 3245 (MO); Las Minas, Stern et al. 1807 (MO). LOS santos: road to El Cortezo, D'Arcy & Sytsma 14356 (MO, NY), panama: Cerro Jefe, Churchill 4284 (BM, G, MEXU, MO, PMA), D'Arcy et al. 15500 (G, MEXU, MO, PMA) D'Arcy et al. 15506 (G, MEXU, MO, PMA); ibique in monte Lan- con iuxta urbem ipsem, Duchassaing s.n. (A); Cerro Campana, Duke 10724(3) (MO, OSU); Isla del Rey, Duke 95 1 9 (MO); Cerro Azul, Dwyer 4087 (MO), Dwyer 4700 (MO); San Jose Island, Erlanson 77 (G, GH, NY, US), Erlanson 260 (G, P, US), Erlanson 348 (G, GH, NY, US), Erlanson 411 ((ill, LIS), Erlanson 513 (GH, US); Cerro Jefe, Folsom et al. 2516 (MO, PMA); Cerro Azul, Gentry & Dwyer 3408 (DUKE, F, NY); San Jose Island, Harlow 95 (GH, US), Johnston 224 (DLIKE, GH, US), Johnston 338 (GH), Johnston 341 (BM, GH, MO, US), Johnston 426 (GH, MO, US), Johnston 428 (DUKE, GII); Cerro Jefe, Mori 6529 (MO), Mori 6531 (MO, NY), Pipoly 7018 (MO); without locality (cited in Hemsley (1879) as Mount Lancon (Ancon Hill), near the city of Panama) Seemann 554 (K); Cerro Campana, van der If erf & Herrera 6153 (DUKE, G, K, MEXU, MO, NY, PMA), van der If erf & van Hardeveld 6916 (MEXU, MO, PMA); Plava Corona, l argas , Jr. 15 (MO), ver- ACUAS: Escuela Agricola Alto de Piedra, Croat & Eolsom 33978 (MO). 3. Ilex fortunensis W. J. Hahn, sp. nov. TYPE: Panama. Chiriqui: La Fortuna hydroelectric project, in cloud forest, along trail uphill behind camp, 1,200-1,400 m, Hammel 2164 (holo- type, MO 3488625; isotypes, PMA, US). Frutex; caules lenticellis praediti; folia integra, tenui- coriacea, glabra, valde bicoloria, epunctata; inflorescen- tiae pistillatae ramosae 2 ordene dispositae; stigma quad- rilobatum; fructus globosi, 9-10 mm x 7-10 mm. Shrub, 4 m tall, moderately branched. Stems terete, somewhat ridged when young, covered with fine white pubescence, young stems drying dark brown or black; lenticels small and sparse or absent; bark dark gray-brown, smooth to somewhat rough with numerous round or oval lenticels, 1 -2 mm diam.; stipules broad-triangular, 0.4 0.8 mm long, 0.8-1. 4 mm wide, deciduous. Leaves thin coria¬ ceous, glabrous, drying dark olive green adaxially, pale yellowish tan abaxially, epunctate, elliptic to obovate-elliptic, (5 )7— 9( 11) cm long, 3.5-5 cm wide, apex caudate to acute, base shortly attenuate, margins entire, sometimes very slightly revolute; venation brochidodromous, somewhat eucampto- dromous at the base, the secondary veins 5-7 per side, the tertiary veins reticulate, apparent; petioles 0.5-1 cm long, 0.8- 1.2 mm diam., round in cross section, caniculate adaxially, drying black. Stami- nate inflorescences not known. Pistillate inflores¬ cences not known in flower, only in fruit where they are axillary or lateral simple cymes on new wood, branched to 2 orders with 3 fruits per inflorescence; bracts ovate-triangular, ca. 2 mm long, thick cori¬ aceous; peduncles 0.5-1 cm long, 1-1.5 mm wide, flat in cross section, tapering toward the apex; pe¬ duncular bracts triangular, 0.5-0. 8 mm long, 0.8- 1 mm wide, paired laterally at the first furcation; pedicels 2-5 mm long, square or angled in cross section; bracteole absent. Fruit globose, 9-1 1 mm long, 7-10 mm diam.; sepals 4-5, oblong or round¬ ed; pericarp coriaceous, 0.3-0. 5 mm thick, deep red to purple drying black-purple, the stigma four- lobed, 3. 5-4. 5 mm diam., spreading, not prominent in profile; mesocarp copious; immature pyrenes 4, trigonal. Ilex fortunensis is known only from the type collection from the La Fortuna hydroelectric site in Chiriqui at an elevation of 1,200-1,400 m. This collection bears immature fruits collected in March. The species is distinguished by the inflorescence type and the distinctly discolorous, thin coriaceous leaves with entire margins. The infructescences are borne on relatively young stems, suggesting that the inflorescences were clustered on an abbreviated stem while flowering. 4. Ilex pallida Standley, J. Wash. Acad. Sci. 16(18): 482. 1926. TYPE: Costa Rica. He¬ redia: Cerros de Zurqui, NE of San Isidro, 2,300 m, Standley & Valerio 50608 (holo- type, IIS 1251651; isotype, F). Figure 2. Ilex pallida is found throughout the Talamancas of Panama and Costa Rica and on nearby mountains at 2,000-3,000 m, and on Volcan Mombacho in Nicaragua above 1,100 m. The species flowers late in the Panamanian dry season, February to April, and again during the brief intermittent dry spell of June to August. Fruits are borne in March and September in Panama and Costa Rica, more spo¬ radically in Nicaragua. The species is characterized by the epunctate leaves, consistently spinose margins with small but Volume 3, Number 1 1993 Hahn Panamanian Species of Ilex 39 evident serrations or crenulations, the blade drying green or olive, pistillate inflorescences with solitary flowers and fruits, and the Iruits to 1 cm long and 0.8 cm in diameter. Most collections from the Tala- maticas have relatively thin leaves. Additional specimens examined. PANAMA. BOCAS DEL TOKO: Valle de Silencio, Antonio 15 99 (G, K, MO, PMA); 6 km NW of Cerro Enchandi, Davidse et al. 25237 (CR, G, MO). CHIRIQUI: El Volcan, D'Arcy & Hammel 12496 (MO, PMA, US); Bajo Chorro, Davidson 254 (F, MO); Volcan Baru, Hammel et al. 6446 (PMA); Aguacata, Hammel et al. 70 19 (A, F, MO, NY, PMA); Bajo Chorro, Wilbur et al. 17276 (DUKE). 5. Ilex lamprophylla Standley, J. Wash. Acad. Sci. 15(21): 476. 1925. Ilex discolor Hemsley var. lamprophylla (Standley) Edwin, Anri. Mis¬ souri Bot. Card. 53; 376. 1966. TV PE: Costa Rica. Cartago: La Estrella, Standley 30 1 It) (holotype, US 1228657; isotypes. A, K). fig¬ ure 3. Ilex carpinterae Standley, J. Wash. Acad. Sci. 15(21): 477. 1925. TYPE: Costa Rica. Cartago: Cerro de La Carpintera, Standley 34491 (holotype, US 1226682). Ilex tristis Standley, J. Wash. Acad. Sci. 16(18): 482. 1926. TYPE: Costa Rica. San Jose: Cerro de las Vueltas, Standley & I alerio 43670 (holotype, LIS 1251406). Ilex davidsoniae Standley, Field Mus. Nat. Hist., Bot. Ser. 22: 88. 1940. TYPE: Panama. Chiriqui: Bajo Chorro, Boquete District, Davidson 166 (holotype, F 915592; isotypes. A, MO, US). 40 Novon Figure 3. Ilex lamprophylla Standley. — A. Habit in fruit ( Standley 39440). — B. Fruits ( Standley 39440). Usually found above 1,500 m elevation, this spe¬ cies is particularly abundant in the Cordillera Central of Costa Rica, with extensions into the Talamancas of Costa Rica and Panama and on scattered peaks in Nicaragua and El Salvador. Flowering in Panama occurs at the end of the wet season through the dry season, September to March. Ilex lamprophylla is readily recognized by its Volume 3, Number 1 1993 Hahn Panamanian Species of Ilex 41 normally rounded leaf crenulations, epunctate leaves, which frequently dry a dark brown color, lighter brown abaxially, cymes on a reduced stem, which later expands in fruit, and the deeply grooved py¬ renes, which give a ribbed appearance to the mature fruits. Ilex davidsoniae and Ilex carpinterae were con¬ sidered by Standley to be possibly synonymous with I. lamprophylla. Study of a large array of specimens supports this view. Standley (1926, 1937) also ex¬ pressed doubts about the validity of his Hex tristis, which he cited as known only from the type locality of Cerro de las Vueltas, San Jose Province, Costa Rica. A search for this taxon in the region proved fruitless, and I consider it to be merely an immature specimen of I. lamprophylla. Edwin (1968) recognized I. lamprophylla as a variety of the Mexican /. discolor, citing I. carpin¬ terae and I. davidsoniae as synonyms. In addition, a number of herbarium specimens from Central America have been annotated by Edwin with /. dis¬ color Hemsley var. tristis (Standley) Edwin, and /. discolor Hemsley var. hondurensis (Standley) Ed¬ win. No reasons were given for this transfer, al¬ though there is some gross similarity in leal shape and inflorescence arrangement. Ilex lamprophylla differs from /. discolor in its thinner, broader, epunc¬ tate leaves that dry a dark brown or black, the nature of the brachyhlast, the branching and size dimensions of the inflorescence, and in the texture of the fruits and pyrenes. Two collections from Chiriqui, McPherson 8946 and Mori & Kallunki 6000, are unusual in that the leaves, apparently immature, are relatively thin with much reduced crenulations and a few small marginal spines. Additional specimens examined. Panama. CHIRIQUI: Cerro Punta, Lao 332 (MO), Luteyn 3085 (DUKE); Bajo Chorro, Davidson 166 (A, F, MO, US,); N of San Felix, Mori & Kallunki 6000 (MFXU, MO, PMA); Fortuna Dam, McPherson 9095 (MEXU, MO, PMA); Cerro Col¬ orado, McPherson 8946 (MEXU, MO, PMA). cocle: El Valle, Sytsma 3824 (MO, PMA). 6. Ilex stellata W. J. Hahn, sp. nov. TYPE: Pan¬ ama. Veraguas: vicinity of Cerro Arizona-Tute, above Sante Fe and Altos de Piedra, McPher¬ son 12806 (holotype, MO 3821597; isotypes: G, K, PMA, WIS). Arbor; folia bullata tomentosa pilis tomentosis, epunc- tatis, 5-9 cm longa, 3-5 cm lata; petioli 1-2 cm longi; inflorescentiae pistillatae ramosae; stigma inanifeste ele- vato-capitatum. Tree to 8 m tall; moderately ramified. Stems slightly ridged on new growth, equally so on older growth, lightly beset with fine tan or white stellate and simple hairs, drying yellowish tan or greenish brown; lenticels scattered, raised, oval, ca. 1 mm long; bark dark grayish to grayish brown, moderate, ridged, densely covered with raised oval lenticels to 1 mm long; stipules triangular, ca. 1 mm long, extensions of the bark. Leaves coriaceous, strongly and deeply bullate, densely pubescent with short, white, stellate hairs, becoming glabrous with age except along the veins abaxially and on the petiole, drying brown to dark brown, epunctate, ovate to obovate, 5-9 cm long, 3-5 cm wide, apex rounded to obtuse, slightly acuminate, base obtuse, short attenuate, margin strongly revolute, essentially en¬ tire but with numerous minute teeth along the mar¬ gin, scarcely visible; venation brochidodromous, the secondary veins 5-7 per side, deeply sunken adax- ially, prominent and raised abaxially, the tertiary veins reticulate and distinct; petioles 1-2 cm long, 1-2 mm wide, densely covered with short, white, stellate hairs. Staminate inflorescence not seen. Pis¬ tillate inflorescence not seen in flower, in fruit of solitary axillary cymes borne on new wood, branched to 2 or 3 orders with 3-7 flowers per dichasium; bracts ca. 1 mm long, triangular, deciduous; pe¬ duncle 5-10 mm long, 1-2 mm wide, flat in cross section; peduncular bract triangular, awl-shaped, ca. 1 mm wide and long, laterally paired or slightly offset at the first furcation; rachis 0.2-1 cm long, 1-1.5 mm wide, flattened or angular; bracteoles 0.5 mm long; pedicels 2-8 mm long, ca. 1 mm wide, angular, tapering toward the base; floral bracteoles apparently absent. Immature fruits depressed glob¬ ular; sepals 5, explanate in fruit, rounded to ovate, 1-1.5 mm long; pericarp leathery, green, I he stigma prominent, raised; mature pyrenes unknown. The species is endemic to Panama and known only from elfin or summit forest at 1 ,000-1,200 m elevation on Cerros Caracoral and Caital in Code and Cerros Arizona and l ute in Veraguas Province. Ilex stellata is readily recognized by the deeply bullate leaves, which bear a moderately thick cov¬ ering of stellate hairs. Ilex bullata Cuatrecasas, described from Valle, Colombia, resembles I. stellata with its deeply bullate leaves, but differs in possessing simple hairs, smaller leaves (3-6 cm long, 1.5—3 cm wide) with abaxial punctations, shorter petioles (3-5 mm long), unbranched pistillate inflorescences, and an adpressed capitate stigma. The inlructesc- ences are borne solitary and axillary on young stems, suggesting that the cymes are clustered in flower as with /. lamprophylla. Paratypes. Panama, cocle: Cerro Caracoral, Kirk- 42 Novon bride, Jr. 1110 (MO); above El Valle, on trail to top of Cerro Caital, McPherson 11958(¥, K, MO, PMA, US). 7. Ilex vulcanicola Standley, J. Wash. Acad. Sci. 15(21): 477. 1925. TYPE: Costa Rica. San Jose: Las Nubes, Standley 38729 (holotype, US 1228373). Ilex vulcanicola is restricted to the highlands of western Panama and Costa Rica at elevations above 1 ,800 m. This species is related to the widespread, polymorphic Ilex kunthiana Triana of Colombia, which differs in having solitary cymes as opposed to the consistently clustered cymes found in I. vul¬ canicola. Other species described from northern South America, such as /. Jarrolensis Cuatrecasas and I. caniensis MacBride, are also closely related and together would probably be referable to series Paltoria sensu Loesener (1901, 1908). Individuals with extremely large leaves might be confused with /. chiriquensis, which differs in its generally larger leaves and fruits and in having more cymes per fascicle. In Panama, flowering of /. vulcanicola is sporadic from February to August. Additional specimens examined. Panama. BOCAS DEL TORO: Fortuna Dam, McPherson 1 1079 (MO, PMA, US). CHIRIQUI: between Gualaca and Fortuna Dam, Croat 49956 (MO, NY, PMA, US); La Fortuna, Hammel 6 243 (F, MO, NY, US); between Los Pianos de Hornito and For¬ tuna Dam, Hampshire & Whitefoord 251 (BM), Hamp¬ shire & Whitefoord 258 (BM). COCLE: Cerro Pilon, Dwyer & Lallithin 8646 (F, MO, NY); Cerro Caital, Knapp et al. 5990 (G, MO, PMA, MEXU); La Mesa, McPherson 11253 (MO, PMA, US); Cerro Caracoral, Sytsma 4049 (A, MO, NY, PMA, US), panama: Cerro Campana, Croat 35973 (F, MO); Cerro Jefe, Mon & Kallunki 6088 ( MO); Cerro Campana, Mori <& Bolten 7694 (F, MO, NY, OSU, US); Cerro Jefe, Tyson et al. 4362 (F, MO). VERaguaS: Cerro Tute, Mori et al. 7594 (A, F, MO, NY, PMA), Sytsma & Andersson 4691 (A, F, MO, US). 8. Ilex chiriquensis Standley, Pub. Field Mus. Nat. His., Bot. Ser. 22: 88. 1940. TYPE: Panama. Chiriqui: Bajo Chorro, Boquete Dis¬ trict, 1,800 m, Davidson 243 (holotype, F 915550; isotypes, MO, US). Ilex chiriquensis is largely confined to western Panama and Costa Rica, with outlying populations known at the Monteverde area of western Costa Rica. Populations in Panama are typically found at elevations of 1,500-2,000 m. Collections from the Fortuna Dam area are known from ca. 1,100 m and from Monteverde, Costa Rica, from 2,000 to 2,500 m. The flowering period of /. chiriquensis is in March and April, with fruiting from May to Au- gust. This species is distinguished by its medium-sized. thickly coriaceous, orbicular to elliptic leaves, un¬ branched staminate inflorescences, and depressed capitate fruit. It is sometimes difficult to separate from Ilex yurumanguinis but that species typically occurs at lower elevations and has thinner, larger, elliptic leaves with distinct secondary venation, branched staminate inflorescences, and ovoid-capi¬ tate fruits. Populations of I. chiriquensis from Mon¬ teverde, Costa Rica, have relatively distinct sec¬ ondary venation and more pronounced marginal teeth. An undescribed epiphytic taxon from Costa Rica appears to be closely related to I. chiriquensis, but differs in its consistently narrower, obovate leaves. Additional specimens examined. Panama. BOCAS DEL TORO: Rio Culebre, Gomez et al. 22387 (CR, G, MO); Chiriquicito- Calderas Trail, Kirkbride & Duke 993 (MO). CHIRIQUI: between Fortuna Lake and Chiriqui Grande, Croat & Grayum 59994 (MO, PMA); Bajo Chorro, Bo¬ quete, Davidson 243 (F, MO, US); Palo Alto NE of Boquete, Hammel 6066 (MO); Cerro Hornito, Hammel 6219 (MO, NY, PMA); Cerro Punta, Hammel et al. 6532 (MO); Palo Alto, Hammel 7449 (MO); Cerro Hor- queta, Luteyn <£ W ilbur 4609 (DUKE); Cerro Pate Ma¬ cho, McPherson 8058 (MEXU, MO, PMA); Cerro Hor¬ nito, Mori 7504 (BM, MEXU, MO, PMA); Cerro Punta, van der Werff & Herrera 6311 (G, MO); Cerro Horqueta, U ilbur & Luteyn 19332 (DUKE). VERAGUAS: Cerro Tute, Mori et al. 7599 (BM, MO). 9. Ilex yurumanguinis Cuatrecasas, Lloydia 1 1(3): 210. 1948. TYPE: Colombia. Valle: Rio Yurumangui, Veneral, bosques en la Quebrada del Zancudo, 10-50 m, 10 Feb. 1944, Cua¬ trecasas 16156 (holotype, US 1951538; iso¬ type, F). This species occurs in wet forests from sea level to 1,500 m in Panama through northwest Colombia into Ecuador, with an outlying population on the Cocos Islands. In western Panama, the species flow¬ ers at the end of the dry season from (December) February to April and fruits in July and August. Populations in central and eastern Panama flower slightly earlier (November to February). Collections from eastern Panama (Colon, Darien, Panama, and San Bias) from below 500 m elevation are virtually identical to material from the type locality of the Rio Yurumangui in Valle, Colombia, with thick leaves, distinct rounded crenulations, a rounded, obtuse apex, and revolute margins. Populations from elevations of 700-1,000 m in Code and 1,000-1,500 m in Chiriqui and Bocas del Toro are characterized by thinner leaves that dry strongly bicolored, the leaf apex long acuminate, the marginal crenulations less rounded or laterally flattened and almost entire with only a few scattered spines, thinner pedicels and peduncles, and more Volume 3, Number 1 1993 Hahn Panamanian Species of Ilex 43 delicate cymes. These specimens approach I. max¬ ima in leaf shape but have much smaller leaves with fewer secondary veins, smaller fruits, and different flowering and fruiting times. These populations per¬ haps deserve specific status, but further study of the Colombian taxa is still needed. One unusual collection from the ridge of Cerro Caracoral (Sytsrna 3824) shows strongly reduced, spatulate leaves and denser ramification, possibly due to the windswept habitat. Two collections from Hocas del Toro near the Fortuna Dam site (McPherson 9050B and Croat 59994) show such features as broader and thicker leaves and more grayish bark, which approach /. chiriquensis. These two gatherings were made in the vicinity of McPherson 9(450 A and Croat 59995, which fall well within the morphological range of the upland I. yurumanguinis form. These may be hy¬ brids, as both species arc found in the area. Cuatrecasas placed I. yurumanguinis near I. /au¬ reola Triana and I. affinis Gardner, two species relegated hv Loesener (1901, 1908) to series Thyr¬ si prinus sect. Thyrsi florae. Additional specimens examined. Panama, chiriqui: road from Fortuna Lake to Chiriqui Grande, Hampshire <£: W hitefoord 425 (BM); 1 km N of Fortuna Lake, Hampshire & U hitefoord 940 (BN1); Fortuna Dam, Mc¬ Pherson 8731 (MEXU, MO, PM A), McPherson 9125 (MEXU, MO, PMA); Boquete, Fortuna Dam site, van der II erf) N van Hardeveld 6815 (G, MO, PMA). COCl.K: Cerro Caracoral, Duke X- Dwyer 15078 (MO, NY); above El Valle, McPherson 12151 (BM); La Mesa, Nee & Dwyer 9157 (DUKE, F, MO, OSU, PMA, US); Cerro Caracoral, Sytsma 3824 (MO). COLON: East Ridge, Duke 15270 (MO). [IOC AS DEI, TORO: Finca Serrano, NE of Boquete, Hammel 9 168 (DUKE, MO, NY, PMA, US); Fortuna Dam Forest, McPherson 9050 (MEXU, MO, PMA). DARIEN: S of Garachineon W slope of Serrania Sapo, McPherson et al. 15379 (MO). PANAMA: Rancho Chorro, Torti Arriba, Folsom et al. 6 712 (MO, NY); El Llano Carti Road, km 16 18.5, Ace & Tyson 10957 (MO). SAN III AS: trail along Continental Divide, Mc- Donagh et al. 302 (BM, MO). 10. Ilex maxima W. J. Hahn, sp. nov. TYPE: Panama. Code: La Mesa, 4 km N of El Valle, 875 m, Nee & Hale 9626 (holotype, MO 38211595; isotypes, BM, F, G, MEXU, PMA, US). Figure 4. Arbor; caules arcuati; folia tenui-coriacea, punctata, 15-40 cm longa, 10-15 cm lata; inflorescentiae dichasiis fasciculatis; inflorescentia staminata ramosa; inflorescen- tia pistillata non ramosa. Free to 10 m tall, laxly ramified, the branches often arching. Stems slightly angular-ridged when dry, glabrous; lenticels scattered on young stems, abundant on older stems, oval, split, 0.6- 1 .3 mm long, 0.3-0. 7 mm wide; bark thin, tannish gray; stipules to 2 mm long, appearing as extensions of bark. Leaves thick chartaceous to coriaceous, drying dark brown-green adaxially, pale tan abaxially, mostly glabrous, punctate abaxially, elliptic, 1 5- 40 cm long, 10-15 cm wide, the apex acute, the base attenuate, the margin laterally flat, subentire with occasional spines, very slightly crenulate toward the apex; ve¬ nation strongly brochidodromous, the secondary veins 10-20 per side, the tertiary veins reticulate, distinct; petioles thickened, 1.5-2 cm long, drying black or dark brown. Staminate inflorescence of compound cymes borne on a much reduced stem resembling an axillary panicle; cymes branched to 2 orders, 1 — 3 flowers per inflorescence, bracts coriaceous, scale¬ like, ovate-rounded, 12 mm long, 1 1.5 mm wide; peduncle 0.5-1 cm long, 0.4- 0.6 mm wide, laterally flattened; peduncular bracts triangular, ca. 1 mm long, 0.5 mm wide, induplicately folded, oppositely paired or distinctly offset; pedicels 0.3 1 cm long, 0.3-0. 4 mm wide, laterally flattened; floral brac- teoles paired at the base of the pedicels, small. Pistillate inflorescences of unbranched cymes clus¬ tered on a much reduced stem with the complete ensemble resembling a cyme, the stem not expanding in fruit; 1 flower per dichasium; bracts coriaceous, deltoid, 1-2 mm long, 1.5-2. 5 mm wide; peduncle 0.8—1 cm long, 0.3-0. 6 mm wide, flattened in cross section, tapering; peduncular bracts, paired or es¬ sentially so, membranaceous, deltoid, 0.8— 1.2 mm long, 0.3-0. 7 mm wide; pedicels 1-2 mm long, 0.1 -0.2 mm wide, round in cross section; floral bracteoles minute. Staminate flowers 4-merous; ca¬ lyx broad cupuliform, sometimes urceolate, the se¬ pals glabrous, ovate-deltoid, 1.2 1.5 mm long, 1.2- 1.5 mm wide, the margin mostly regular, indupli¬ cately folded; corolla rotate, the petals white, gla¬ brous, 2-2.5 mm long, 1 .5-2 mm wide, the margin entire; filaments 2-2.5 mm long, flattened, tapering, the anthers yellow7, ovoid, tapering, 1 1 .5 mm long, 0.3— 0.6 mm wide; pistillodia spherical-conical, 0.4— 0.6 mm tall, 0.8- 1.2 mm diam., the stigma cylin¬ drical, 0.1 -0.3 mm tall. Pistillate flowers unknown. F ruit spherical to ellipsoid, 0.8 I cm long, 0.5- 0.7 mm diam.; pericarp leathery, moderately thick, dark purple at maturity, stigma persistent, spreading to slightly flattened capitate. 2- 2.5 mm diam., 1 mm high, 4-lobed; mesocarp moderately thick, fleshy; pyrenes 4, trigonal 4 mm long, 1-1.5 mm wide, shallowly channeled, endocarp bony. Ilex maxima occurs in wet forest in the Cordillera Central of Costa Rica, the Lalamancas of Costa Rica and Panama through central Panama to the western slopes of the Andes in Colombia. It generally occurs 44 Novon Figure 4. Ilex maxima W. J. Hahn. A. Habit (AW & Hale 96 26). B. Fruits (Stevens 18321). 1cm 1cm Volume 3, Number 1 1993 Hahn Panamanian Species of Ilex 45 at elevations of 700-1,000 m, except to 1,500 m in the Talamancas and lower elevations in Colombia. Flowering is at the end of the wet season into the onset of the dry season, (rom September to Novem¬ ber, and fruiting is from January to April. Ilex maxima is characterized by its lax habit, cymes borne clustered on reduced stems in flower and fruit often appearing cauliferous, the large spherical fruits, and the very large, elliptic leaves. It is closely related to I. latci of the Guayana High¬ land region of Venezuela and I. yurumanguinis Cuatrecasas of northwestern Colombia and Panama, but differs in its larger and relatively thicker leaves with more secondary veins, the larger, more robust staminate inflorescences, and the larger fruits. Other apparently related taxa include a number of Colom¬ bian species such as I. caliana Cuatrecasas and /. laurina Kunth, which have been referred to section Daphnophyllae (Loesener, 1901, 1908). Paratype. Panama, CHIRIQUI: confluence of the Rio Hornito and Rio Chiriqui, Stevens 18321 (MO, UTD). Acknowledgments. I thank the many individuals who have provided years of encouragement and advice, including W. G. D’Arcy, J. Miller, \\ . Ha¬ ber, S. Andrews, and P.-A. Loizeau. Special thanks are extended to Karen Krager for the illustrations and John Dwyer for the Latin diagnoses. Courtesies extended by the curators of A, BM, CR, 1)1 KE, F, C, GH, K. MO, NY. P, PM A. and US are gratefully recognized. Literature Cited Bennet, C. F. 1968. Human influences on the zooge¬ ography of Panama. Ibero-Americana 51: 1112. Bornstein, A. J. 1989. Aquifoliaceae. Pp. 107 1 18 in R. A. Howard (editor), Flora of the Lesser Antilles, vol. 5. Arnold Arboretum, Harvard Univ., Jamaica Plain, Massachusetts. D’Arcy, W. G. 1977. Endangered landscapes in Pan¬ ama and Central America: The threat to plant species extinction. Pp. 89-104 in G. T. Prance & T. S. Elias (editors), Extinction is Forever. The New York Botanical Garden, New York. Edwin, G. 1965. Aquifoliaceae. In: B. Maguire & Col¬ laborators, The Botany of the Guayana Highland part VI. Mem. New York Bot. Gard. 12(3): 121 150. - . 1967(1968). Aquifoliaceae. In: Flora of Pan¬ ama. VI. Ann. Missouri Bot. Gard. 54: 381-387. Hahn, W. J. 1988. A new species of Ilex (Aquifoliaceae) from Central America. Ann. Missouri Bot. Gard. 75: 73-735. Hemsley, W. B. 1880. Botany. In: F. D. Godman & 0. Salvin (editors), Biologia Centrali-Americana I: 186 187. R. H. Porter, London. Johnston, I. M. 1949. The Botany of San Jose Island (Gulf of Panama). Sargentia 8: 1-306. Loesener, T. 1901. Monographia Aquifoliacearum, Nova Acta Acad. Caes. Leop. -Carol. German. Nat. Cur. 78: 1-567. - . 1908. Monographia Aquifoliacearum, Nova Acta Acad. Caes. Leop. -Carol. German. Nat. Cur. 89: 5-314. Loizeau, P.-A. 1992. Proposition d’une classification des inflorescences d 'Ilex L. (Aquifoliaceae). Candollea 47: 99-112. Smith, J. D. 1914. Undescribed plants from Guatemala and other Central American Republics. Bot. Gaz. (Crawfordsville) 57: 415-427. Standley, P. 1926. The Costa Rican species of Ilex. J. Wash. Acad. Sci. 16(8): 481 484. - . 1929. Studies of American plants. Publ. Field Mus. Nat. Hist., Bot. Ser. 4(8): 221-222. - . 1937. Aquifoliaceae. In: Flora of Costa Rica. Part II. Publ. Field Mus. Nat. Hist., Bot. Ser. 8: 628-631. Steyermark, J. A. 1988. Flora of the Venezuelan Gua¬ yana. IV. Ann. Missouri Bot. Gard. 75: 320 333. Triana, J. 1872. Ilicinae. In: K. S. Kunth, Prodromus Flora Nova-Granatensis. Ann. Sci. Nat. Bot., ser. 5, vol. 16: 375-378. Wurdack, J. J. 1961. Aquifoliaceae. In: B. Maguire & J. Wurdack (editors), The Botany of the Guayana Highland, Part IV (2). Mem. New York Bot. Gard. 10(4): 1-6. A New Species and a New Combination in Salpichroa (Solanaceae) Shirley //. Kuo Keel The Nature Conservancy, Latin America Science Program, 1815 N Lynn Street, Arlington, Virginia 22209, LJ.S.A. ABSTRACT. A monographic study of the genus Sal¬ pichroa revealed a new species, S. rnicroloba, and one taxon with a new status, S. glandulosa subsp. weddellii. These taxa are discussed and information on their probable relationships is given. Salpichroa glandulosa (Hooker) Miers subsp. weddellii (Benoist) Keel, stat. et comb. nov. Basionym: Salpichroa weddellii Benoist, Bull. Soc. Bot. France 85: 408. 1938. TYPE: Bo¬ livia. Cochabamba: between Llave and Moro- chata, Dec. 1846 (fl). If eddell 41 16 (bolotype, P; isotypes, P — 2 sheets). A study of the genus Salpichroa (Keel, 1984) gave clear indication that S. glandulosa consists of two subspecies, Salpichroa glandulosa subsp. wed¬ dellii and S. glandulosa subsp. glandulosa, that are similar in growth habit, habitat preference, and many aspects of morphology. The morphological characters that distinguish subspecies weddellii from subspecies glandulosa include (1) absence of teeth between the sinuses of the corolla lobes and (2) long pedicels with calyx bases that do not enlarge after fruiting. The two taxa, representing populations sep¬ arated geographically by tbe Cordillera I res Cruces in Bolivia, are best treated as vicariant subspecies. Salpichroa rnicroloba Keel, sp. nov. TYPE: Peru. Lima: Arquircancha, near Lachaqui, 3,658 m, 2 Feb. 1979 (fl, fr), heel & Vilcapoma 397 (holotype, NY; isotypes, MO, USM). Figure 1. Frutices penduli vel effusi ad 1 m alti. Rami et ramuli interdum flexuosi. Folia ovata vel elliptica; petioli subfi- liformes vel complanati. Flores flavi vel viridi-flavescentes. Pedicelli filiformes. Calyx profunde 5-partitus, lobis li- nearibus, apice acutis vel attenuatis, connatis solum basi. Corolla 5-lobata, tubulosa, tubo recto vel apicem versus leviter dilatato, sed constricto ad faucem, lobis aestivatione valvatis, triangulatis vel ovato-triangulatis, apice acutis, ciliatis, sub anthesi per 180° reflexis. Stamina prope tubi apicem inserta. Stylus glaber, leviter exsertus vel tubum aequans, stamina excedens, stigmate subcapitato vel cap- itato. Bacca cyanea vel purpurea. Pendent or straggling shrubs to 1 m tall, the young stems, leaves, and pedicels puberulous, pilose or glabrous, sometimes with gland-tipped trichomes. Branches and twigs terete, flexuous, sometimes winged on 1 to 3 sides, each node occasionally with 2 to 4 semicircular or circular scales. Leaf blades ovate or elliptic, (1— )1 .5— 2.8(— 3.6) x (0.6-)l- 2.1(-3) cm; petioles subfiliform to flattened, 0.6- 1.8 cm long. Pedicels filiform, 0.5-1. 5 cm long; calyx deeply divided, the lobes linear, subequal, 5- 13 x 0.5-2. 5 mm, ciliate, pilose, puberulous or glabrous, connate at the base, apex acute or atten¬ uate; corolla tubular, yellow or yellowish green, the tube straight, slightly inflated on upper part, con¬ stricted at the throat, 22-34 x 3-7 mm, glabrous or puberulous externally, glabrous internally, the lobes triangular or ovate-triangular, equal to sub¬ equal, valvate, 2-3 x 1.5-3 mm, ciliate, apex acute, reflexed by 180° at anthesis; stamens in¬ cluded, equal or subequal, inserted at top !4 of the tube, the filaments 0.5- 1.5 mm, glabrous, the an¬ thers 2.5-4 mm; style nearly equal to the throat of corolla tube or slightly exserted, exceeding sta¬ mens, 2-3 cm long, glabrous, the stigma capitate or subcapitate, glabrous. Berry dark blue or purple, 3 cm long. Distribution. Sub-puna or montane cloud forests, near creeks, in agricultural fields, hedgerows, on roadsides, grass steppes, or on stone walls, rock crevices; 2,400-3,800 m; in the Department of Cotopaxi, central Ecuador (one collection only) and west-central Peru. Flowering December-May, fruits found in February. Salpichroa rnicroloba shows slight variation in degree of pubescence and size of leaves. Cerrate & Tovar 1909 is exceptional for its large, glabrous leaves and wide corolla tubes. Salpichroa rnicroloba and S. weberbaueri Daminer have nearly the same corolla length, but the shapes of corolla tubes and lobes, which are best seen in living plants, are very different. At anthesis, the corolla tube of ,S. micro- loba is constricted at its throat and the triangular or ovate-triangular corolla lobes are reflexed by 1 80°. The corolla tube of S. weberbaueri widens at the upper third of the tube; the lobes, from horizontally spreading to reflexed by 135° at anthesis, are tri¬ angular-lanceolate. In the field, one can easily sep- Novon 3: 46-48. 1993. Volume 3, Number 1 1993 Keel Salpichroa 47 arate them by the shape of corolla and the degree of reflection of the corolla lobes at anthesis. The specific epithet, microloba, refers to the relatively small size of the corolla lobes compared to the tube. Salpichroa microloba is probably most closely related to S. ramosissima Miers. Although the length of corolla tube of these two species is quite different (2. 2-3. 4 cm long in S. microloba, and 0.9- 1 .5 cm 48 Novon in S. ramosissima ), the shape of corolla and the reflection angle of lobes at anthesis are similar. Salpichroa microloba has been collected in cen¬ tral Ecuador, Department of Cotopaxi, and west- central Peru. Salpichroa ramosissima grows in the western part of central and southern Peru, and in southern Bolivia and northwestern Argentina. The ranges of the two species overlap in the Department of Lima, Peru. Local names. Peru: Ayanata ( Keel & Vilcapoma 397, Vilcapoma 129), Callalluma (Vilcapoma 130), Shuculumpa ( Cerrate & Tovar 1909). Paratypes. Ecuador, cotopaxi: road Quevedo Lata- cunga, above Pilalo, ca. 2,900 m, 3 May 1968 (fl), Hurling et al. 8993 (GB, MO). PERU. LIMA: Banos, on the way to Cerro de Pasco, Wilkes, 21-22 May 1839 (collecting date based on Wilkes, 1845) (fl) (US); Lacha¬ qui, 3,800 m, 28 Dec. 1972 (fl), Vilcapoma 129 (US), 3,400 m, 29 Dec. 1972 (fl), Vilcapoma 130 (US); Pi- rocancha, on the way to Lachaqui, 3,440 in, 1 Feb. 1979 (fl, fr), Keel <£- Vilcapoma 390 (MO, NY, US, USM); Achaca, few km beyond Lachaqui, 3,658 in, 2 Feb. 1979 (fl, fr), Keel tfc Vilcapoma 396 (NY, USM); Matucana, 2,438 m, 14-18 Mar. 1923 (fl), Macbride 2946 (F); vie. of Huarochiri, western cordillera, 2,100-3,000 m, Hrdlicka, Feb. 1913 (fl) (US); vie. of Santiago, 3,600- 3,700 m, 15 May 1953 (fl), Cerrate & Tovar 1909 (US). Acknowledgments. This paper is part of results of my doctoral dissertation. I am grateful to Ghillean T. Prance for useful advice, Rupert C. Barneby for help with the Latin description, Bobbi Angell for her plant illustrations, and William G. D’Arcy for re¬ viewing the manuscript. The curators of herbaria who made specimens available for study are thank¬ fully acknowledged. Literature Cited Wilkes, C. 1845. Narrative of the United States ex¬ ploring expedition during the years 1838, 1839, 1840, 1841, 1842. 1: 261-266. Lea & Blanchard, Philadelphia. Keel, S. H. K. 1984. A revision of the genus Salpichroa (Solanaceae). Ph.D. Dissertation, City University of New York, New York. Calathea liesneri, a New Species of Marantaceae from Venezuela Helen Kennedy Herbarium, Botany Department, University of British Columbia, Vancouver, B.C. V6T 1Z4, Canada ABSTRACT. Calathea liesneri. from Territorio Federal Amazonas, Venezuela, is unusual in having the upper side of the lamina pilose throughout. It ddTers from other species sharing ihis feature by leaf shape, and bract shape and number. It is herein described as new for inclusion in the Flora of the Venezuelan Guyana treatment. Calathea liesneri Kennedy, sp. nov. TYPE: Ven¬ ezuela. Territorio Federal Amazonas: Depto. Atahapo, in saddle between Cerro Duida and Cerro Marahuaca near base ol Cerro Duida, medium height forest, 1,000 m, 03°34'N, 65°32'W, 25 Oct. 1088, Liesner 25333 (ho- lotype, MO; isotypes, K, IIBC, \ EN). A Calathea cyclophora lamina superior utrinque pilosa et bracteolis membranaceis diftert; a Calathea squarrosa bracteis 7-10, transverse ellipticis ad rotundatis et 2- 2.9 cm longis differt. Acaulescent herb, 35-80 cm high, bearing 2-4 basal leaves. Leaf blade elliptic, apex acuminate, base obtuse to rounded, very shortly abruptly at¬ tenuate, 12-40 x 9 14 cm. Leaf blade green above, pilose throughout; leal surface gray-green, velvety tomentose below. Pulvinus elliptic in cross section, tomentose throughout, 0.5 2.3 cm long. Petiole 0 29 cm long, tomentose, hairs with cushion of cells at base. Leaf sheath not auriculate, tomentose to subhispid toward base, margins subglabrous, 14.5— 30 cm long. Inflorescence terminal, 1 per shoot, imbricate, ellipsoid, 5-6.5 cm. Peduncle pilose, hairs with cushion of cells at base, 40 64 cm long. Bracts 7-10, spirally arranged, transverse elliptic to ro¬ tund, apex rounded with an acumen to obtuse, outer margin and apex patent, margin slightly undulate in upper bracts, 2-2.9 x 2-2.8 cm, each sub¬ tending 4 or more flower pairs. Outer surface of bracts “lavenderish white” (fide Liesner 25651), glabrous; inner surface glabrous. Bicarinate prophyll membranaceous, elliptic, apex obtuse, glabrous, 1.7 2.5 x 1.2- 1.4 cm, 0.6 0.7 cm wide, earina to carina. Secondary bract membranaceous, elliptic, apex acute to obtuse, glabrous, 1 .72.2 x 1 1.2 cm. Bracteoles 1 per flower pair, channeled, mem¬ branaceous, medial, glabrous, ca. I .2 cm long. Mow¬ ers “lavenderish white” (fide Liesner 2565 / ). Sepals ovate to elliptic, acute, glabrous, 6. 5-9. 5 x 2.5- 3 mm. Corolla glabrous, ca. 50 mm, tube ca. 30 mm long; corolla lobes subequal, elliptic, obtuse, ca. 15-19 X 5. 5-8. 5 mm. Outer staminode obovate, ca. 15 mm long. Callose staminode petaloid. ca. 13 x 4 mm. Cucullate staminode ca. 8 mm long. Sta¬ men with lateral petaloid appendage; anther 2.5 mm long. Ovary glabrous, ca. 2 mm long. Capsule smooth, obovoid, glabrous, ca. 10 x 7 mm; crowned by a persistent calyx. Seeds usually 3 per capsule, tri¬ gonal, ca. 6-7 x 4 x 3 mm. Calathea liesneri belongs to Calathea sect. Free- iscapus Benthain, by virtue of its all basal leaves and inflorescence with spiral bracts borne on the leafy shoot. Paratype. Venezuela, territorio federal amazonas: Depto. Atabapo, Cerro Huachaniacari, E slope, forested, near waterfall, 03°49'N, 65°42'W, 600-700 in, 2 Nov. 1988, Liesner 25654 (MO, UBC, VEN). Acknowledgments. I thank the curators at Mis¬ souri Botanical Garden for the loan of material and Paul Berry for help with the Flora of the I enezuelan Guyana project. I thank Fred Ganders for support and Tom Baumann for the use of his computer. Novon 3: 49. 1993. New Taxa in Maerua (Capparaceae) proposed for the Flora of Ethiopia Lars E. Kers Bergius Botanic Garden, P.0. Box 50017, Stockholm, Sweden Abstract. During studies of Capparaceae carried out for the Flora of Ethiopia, t he lollowing new taxa in Maerua were discovered. Maerua intricata Kers, sp. nov. TYPE: Ethiopia. Ogaden: Mustahil to Ferfer, 5 Dec. 1954, Peg¬ gy E. Ellis 357 (holotype, K). A speciebus quas A. Brongniart et alii auctores ad Courhoniam ascripserunt habitu intricato foliis minutis costa plerumque immersa ideo utrinque occulta distin- guenda, ab eis etiam in combinatione petioli brevissimi (ut in M. pseudopetalosa (Gilg & C. Benedict) De Wolf) cum fructibus ellipsoidalibus basi regulariter per quattuor val- vas dehiscentibus (ut in M. subcordata (Gilg) de Wolf) differt. Virgate shrub 0.5-2 m tall, intricately branched, sterns erect or ascending, glabrous, ± minutely sca¬ brous, branches and branchlets stiffly divaricate, somewhat striate. Leaves simple, minute, 5-15 mm long, l-5(-8) mm wide, stiff, coriaceous, narrowly elliptic, oblong or lanceolate, seemingly often nerve¬ less (midrib imbedded), petiole very short, 0.5- 1 .5(-2) mm long. Flowers without petals, solitary and axillary; pedicel short, 7-11 (-13) mm long; receptacle cylindric, (2-)3-4.5 mm long; sepals three, (6 )810 mm long; androgynophore Anally exceeding the receptacle by 1-2.5 mm; stamens ca. 20-25; gynophore 12-16 mm long, longer than the pedicel; ovary fusiform, 4-nerved; placenta 2, ovules two on each placenta. Fruit broadly ellipsoid, with one or rarely two seeds, pericarp finally break¬ ing up into four valves at the very base of tbe fruit, otherwise indehiscent. Seed (9-) 14- 15 mm long, narrowly ovate to oblong, cotyledons pale yellowish. Maerua intricata is clearly allied to the species of Maerua that were formerly grouped in the genus Courbonia Brongniart. It differs in the intricately branched growth, the minute leaves with a generally completely immersed and consequently hidden mid¬ rib. The petioles are as short as in M. pseudope¬ talosa, whereas in shape and mode of dehiscence the fruits are comparable with those of M. subcor¬ data. Distribution. Known from the area around the River Schebelli in eastern Ethiopia and central So¬ malia and from there north into the African Horn of the northeastern Somali Republic; Irom 100 m near the coast to 350 m inland. Habitat. In Acacia Commiphora bushland or in scattered low bush. Growing on rocky slopes, in stony soil at foot of hills, in sandy areas with lime¬ stone ridges or in dry water courses. Indigenous names. Cagat, cagah (Bari region), Dugal erigiren (Hiraan region). Paratypes. ETHIOPIA. OGADEN: Scivelli to Beletuen, June 1943, Hummel 151 (K); Webi (= Schebelli River), July-Aug. 1891, Robecchi-Brichetti 228 (FT). Somalia Republic, sanaac; region: below Einad, 1 1°55'N, 48°55'E, 1957, Newbould 1022 (K). BARI REGION: Alula, without date, Anonymous s.n. (FT fragments); along road from Alula to Seyn Wayn, 9 Oct. 1982, Barbier 939 (FT, K); near Bender Merhagno, 1 1°41'N, 50°27'E, 15 Oct. 1982, Barbier 946 (K); Mt. Bolimock W of Alula, 29 Oct. 1959, Hemming 1786 (K); 40 km SW of Alula, 1 1°55'N, 50°30'E, 14 Nov. 1980, Hemming & Udtson 3087 (K); near Bosaso, to Cassim Bender, 7 July 1924, Puccioni & Stefanini 1034 (= 1 136) (FT); Wadi Merero, Scortecci, 1957 (FT); mountain pass W' of Bargal, ca. 1 1°15'N, 51°00'E, 25 Nov. 1985, Thulin & Uarfa 5532 (UPS); mountains S of Bender Murrayha, valley above Tayeega, 29 Oct. 1959, Ihulin & Uarfa 5814 (K, UPS). NUGaal region: valley slope of Nugal River, 7 Oct. 1959, Hemming 1679 (K). miidug region: 69 km SW of Gal- kayo on main road, 06°27'N, 46°53'E, 4 June 1979, Gillett, Hemming & Watson 22347 (K). hiraan region: 03°06'N, 45°04'E, 1 Aug. 1983, Abukar Sheik 2023 (K); Bulo Burti district, 4 km N of Murkayale, 03°50'N, 45°13'E, 28 Sep. 1986, Kuchar 17082 (K); El Mocoile, on road from Bulo Burti to Galgiel, 5 Apr. 1933, Suckert 66 (FT, K). Figure 1. Maerua intricata Kers. — A. Portion of stem with flowering twigs. — B. Portion of stem with immature fruits. — C. Ovary in transversal and longitudinal sections. — D-F. Leaves. — G. Gynophore with ovary, torus with stamen scars, androgynophore. — H. Flower. — I. Mature fruit showing the basal dehiscence, seed partially exposed. Novon 3: 50-54. 1993. Volume 3, Number 1 1993 Kers Maerua 51 — J. Receptacle opened showing sepals and the minute corona. — K. Embryo in lateral view and transversal section showing the cotyledons. Fragment of the seeil coat (right). A, F, Abukar Sheik 2023 (K); B, Suckert 66 (FT); C, E, G, H, J, Bar bier 039 (FT); D, Barbier 946 (K); I, K, Thulin & Warfa 3814 (UPS). 52 Novon Figure 2. Maerua gillettii Kers. — A. Portion of stem showing reflexed phyllodes. — B. Apex of petioles with rudimentary leaflets. — C. Floral bract with stipule. — D. Portion of stem with flowering twigs and spreading phyllodes. — E. Flower. — F. Androgynophore, stamens, gynophore with ovary. — G. Receptacle opened showing sepals and the lobed corona. — H. Sepals. — I. Petals. — J. Ovary in longitudinal and transversal section. — K. Fruit with gynophore, androgynophore, receptacle, and pedicel. A j, Bally & Smith 14887 (K); K, Haugen 1915 (S). Volume 3, Number 1 1993 Kers Maerua 53 Maerua gilletlii Kers, sp. nov. TYPE: Kenya. Northern Frontier Province: Moyale, 15 kin out Waijir road, 03°25'N, 39°07'E, 5 Nov. 1952, ./. B. Gillett 11118 (holotype, K; iso¬ type, FT). “ Maerua spec. A” in Flora of Trop¬ ical East Africa, Capparaceae, 49-50, 1964. Haec species M. glaucae Chiovenda verosimiliter af- finis sed phyllodia habet ideo inter omnes Maeruas afri- canas bene distincta. Shrub, glabrous, to 1 .5 m tall, stems green, striate and slender hut generally rigid, branches and hranchlets divaricate. Leaves well spaced, initially 3-foliolate but readily forming phyllodes; leaflets very reduced, rudimentary, scarious and usually dropping off early; petioles transformed into phyllodes. Phyl¬ lodes similar to the stems, becoming reflexed, grad¬ ually elongating, finally up to 19 cm long. Inflores¬ cences few-flowered and rather loose, racemose or corymbose; pedicel short, 4-6(-9) mm long; flowers small; sepals 4, 4-5 mm long; petals elliptic, 3- 4(-5) mm long; receptacle shortly cylindric to nar¬ rowly campanulate, 2.5-3 mm long; corona 1 1.75 mm long, cleft to base into usually four segments; stamens 8—1 0(— 12), 0.8-1. 1 cm long, borne in one series; anthers 1 mm long. Androgynophore 4.5- 5.5(-6) mm long, exceeding the receptacle by (2-)4- 5 mm. Gynophore thin. ca. 1.2 cm long. Ovary 1 1.5 mm long, ellipsoid at first; placenta 2; ovules 8; stigma sessile to subsessile. Fruit globose to sub- globose, ca. 0.5 cm diam., smooth, generally 1 -seeded. Seed subspherical, filling the interior of the fruit, mature seeds unknown. Maerua gilletlii is well characterized by its stem¬ like phyllodes, which are not found in other African Maeruas. The phyllodes are usually reflexed, with early dropped rudimentary leaflets. The flowers and fruits are similar to those of M. glauca Chiovenda, a species that differs in having ordinary leaves and a longer pedicel (to 1.8 cm long). Maerua filiform is Drake from Madagascar de¬ velops phyllodes, but they are not reflexed; this species differs also in its narrowly cylindrical recep¬ tacle 3-4 mm long, the shorter androgynophore exceeding the receptacle by only 1 mm, the incon¬ spicuous or practically wanting corona, the subcor- date, 2.5-3-mm-long petals, and the ovate to subglo- bose shape and blunt apex of the ovary. Distribution . Only known from northernmost Kenya and the adjacent parts of southern Ethiopia; from 600 m in Kenya, to 1,400 m in Ethiopia. Reported to be locally common below 1,300 m in southern Ethiopia. Habitat. In Acacia-Commiphora scrub or in Acacia mellifera bushland. Growing on sandy or loamy soil near exposed bedrock of granite, gneiss, or schist. Average rainfall in northern Kenya 635 mm/year with maximum rainfall in April and Oc¬ tober. This species is named after Jan B. Gillett (K), formerly of the East African Herbarium in Nairobi, an acknowledged expert on the flora of northeastern Africa and an important collector. Paratypes. Kenya, northern frontier province: S end of Huri Hills, 25 Feb. 1963, Bally 12531 (K); 16 km SE of Sololo on road to Moyale, ca. 03°26’N, 38°42'E, 18 Jan. 1972, Hally & Smith II 14887 (K): Huri Hills, ca. 03°25'N, 37°46'E, 4 Nov. 1976, Herlodier 17 9(K). Ethiopia, sidamo province: Hadessa, ca. 70 km S of Neghelle, 11 Nov. 1991, Haugen 1943 (S); ca. 60 km from Neghelle, on road to Dawa River, ca. 90 km from Dawa, 9 Feb. 1991, Haugen 1915 (S). The Maerua angolensis Complex of Northeast Africa Two species are involved in the Maerua ango¬ lensis complex of northeast Africa: M. angolensis DC. (sens, str.), which is widely distributed in Africa and also extends into Arabia; and M. socotrana (Schweinlurth ex 1. B. Baltour) Gilg, based on spec¬ imens from Socotra. The Socotran population and M. angolensis are easily separated by the length of the receptacle and the size of flowers and leaf blades. They also differ in habit, M. socotrana being a much more slender-stemmed shrub or low tree. These clearly distinct taxa were found to almost grade into each other by means of numerous, more or less isolated populations growing in the Somali Republic and eastern Ethiopia. When previously determined, these atypical specimens had generally been labeled M. socotrana. Sometimes the name \I. thomsonii T. Anderson had been applied to them. Maerua angolensis and M. socotrana cannot be kept separate specifically. The broad concept of M. angolensis results in a complex species. The mor¬ phological variation within this complex is not con¬ tinuous and is subdivided into the taxonomic entities presented below. These differ in a number ol fea¬ tures, of which only the more easily observed and measured are mentioned here. A more detailed treat¬ ment of them will be found in the Flora of Ethiopia. The geographical areas of the allied taxa seldom meet or overlap. Further studies ol the complex will be of great biogeographical interest. The Socotran population (M. socotrana sens, str.) is fairly uniform and can be separated taxonomically on a subspecific level. To some extent, its distinctness may be due to geographical isolation within a small area. Maerua thomsonii T. Anderson was described in 54 Novon 1860 from Aden. This species has previously been regarded as identical to M. socotrana (Elffers et al., 1964). Maerua thomsonii is not, however, a close ally and does not belong to this complex. Key to the Infraspecific Entities of Maerua angolensis la. Receptacle (7— )9— 1 5(— 1 9) mm long, cylindri¬ cal, androgynophore (9-)l 1 18(-20) mm long, exceeding the receptacle by 2 5 mm; leaf blades commonly 2-5.5 cm long; low trees with a rounded crown or shrubs, much branched . . . . subsp. angolensis lb. Receptacle 2-6( 9) mm long, cylindrical, in¬ fundibular or campanulate, androgynophore (3.5 — )4— 7( — 1 1 ) mm long, equal or exceeding the receptacle by 15 mm; leaf blades 1 — 2(— 3) cm long; low, often very slender trees or shrubs, few-branched . subsp. socotrana 2a. Sepals 0.5-1 cm long, stamens (0.5-)0.7- 1(— 1.5) cm long, receptacle 2-3(-4) mm long . subsp. socotrana var. socotrana 2b. Sepals 1.2-2. 2 cm long, stamens 1.5-2. 4 cm long, receptacle 2-6 mm long . . subsp. socotrana var. africana 1. Maerua angolensis DC. subsp. angolensis. TYPE: Angola. Benguela: Jose da Silva s.n. (holotype, P). I bis subspecies is characterized by the long re¬ ceptacle, commonly 9 1 5 mm long, and by the long androgynophore, commonly 11-18 mm long. It var¬ ies in shape of leaf blades, direction of sepals (spread¬ ing or reflexed), and hairiness. Widely distributed in Africa; also found in Arabia. 2. Maerua angolensis DC. subsp. socotrana (Schweinfurth ex I. B. Balfour) Kers, stat. nov. Basionym: Maerua angolensis DC. var. so¬ cotrana Schweinfurth ex I. B. Balfour, Proc. Roy. Bot. Card. Edinburgh 12: 402, 1883. Maerua socotrana (Schweinfurth ex I. B. Bal¬ four) Gilg, Engler Bot. Jahrb. 33: 228, 1903. SYNTYPES: Socotra, Hay ley Balfour 193 (BM), 588 (B, BM, P); Schweinfurth 251 (not seen), 457 (B, P), 603 (P). The flowers are smaller than in subspecies an¬ golensis, with a receptacle commonly 2-6 mm long and with an androgynophore commonly 4-7 mm long and sometimes not exceeding the receptacle. Distribution. Ethiopia, Somalia, and Socotra. 2A. Maerua angolensis DC. (subsp. socotrana) var. socotrana. TYPE: as for M. angolensis DC. subsp. socotrana. Sepals 0.5-1 cm long, 2.75-4 mm wide, glabrous within, stamens (0.5— )0.7— 1(— 1 .5) cm long, anthers 1 - 1 .5(- 1 .75) mm long, receptacle 2-3(-4) mm long. Distribution. Socotra. 2B. Maerua angolensis DC. (subsp. socotrana ) var. africana Kers, var. nov. TYPE: Ethiopia. Ogaden: Lamaloye, 7 July 1988, C. F. Hem¬ ming 1517 (holotype, K). Sepalis 1.2-2. 2 cm longis, 3-6 mm latis, intus glabris vel puberulis, staminibus 1 .5-2.4 cm longis, antheris 1 .5- 2.5 mm longis, receptaculo 2-6 rnm longo. Haec varietas plantas pius-minusque intermedias inter subsp. angolen- sem et var. socotram includit. Maerua angolensis var. africana differs from variety socotrana in the somewhat bigger flowers with sepals 1.2-2. 2 cm long and 3-6 mm wide, glabrous or puberulous within. Stamens 1.5-2. 4 cm long and with anthers 1.5-2. 5 mm long. The re¬ ceptacle is 2-6 mm long. The occurrence of variety africana is split into a number of more or less isolated local forms, in some localities rather intermediate between subspe¬ cies angolensis and subspecies socotrana var. so¬ cotrana. Distribution. Eastern Ethiopia and Somalia. Literature Cited Elffers, J., R. A. Graham & G. P. De Wolf. 1964. Capparidaceae. In: C. E. Hubbard & E. Milne-Red- head (editors), Flora of Tropical East Africa. Crown Agents, London. A New Species of Nematopoid Xyris (Xyridaceae) from the Araracuara Area of Colombia Robert Krai Herbarium, Department of Biology, Vanderbilt University, Box 1705, Station B, Nashville, Tennessee 37235, U.S.A. Joost l)ui veil voordert Tropenbos Program, Apartado Aereo 034174, Bogota, I). E., Colombia ABSTRACT. A new species of Xyris (sect. Nema- topus) has been discovered in the Caqueta basin east of Araracuara, Dept. Caqueta in the Colombian Amazon area. Xyris trachysperma is described, illustrated, and discussed as to its relationships with other taxa of the section. The affinities of A. tra¬ chysperma appear to be more with species of plan- altan Brazil than with those of the Guayana High¬ lands. During integrated surveys by personnel of the Tropenbos Program of the forest and soils of the Caqueta basin east ol Araracuara, Dept. Caqueta, in the Colombian Amazon area, J. Duivenvoorden and associates visited and collected plants in the savanna complexes of a sandstone rneseta (350 in a.s.l.) near Araracuara in 1988 and 1990. 1'his region of Colombia has long been known to be rich in Xyridaceae: its lowland and upland savannas pro¬ duce a wealth of species shared with the Orinoco and Rio Negro systems of Colombia and contiguous Venezuela and also some unshared endemics. The Araracuara area has had a fairly long history of plant exploration, most ol it originating with R. E. Schultes, H. Garcia-Barriga, and associates in the 1950s, and subsequently by B. Maguire and asso¬ ciates in the 1950s and 1960s, resulting in the discovery of several novelties, three from Rio Ca¬ queta savannas. Yet these systems are so large and so complex that the potential lor further discoveries continues. Therefore, when a large set of collections from this place was sent from Tropenbos to VDB for definitive study, one Xyris attracted particular attention. A full description of it is provided to em¬ phasize that its affinities are, surprisingly, more with species of planaltan Brazil rather than with its Co¬ lombian or Venezuelan cohabitants. Xyris trachysperma Krai & Duivenvoorden, sp. nov. TYPE: Colombia. Dept. Caqueta: Arara¬ cuara (0°37'S, 72°24'W), sobre la mesa de areniscas de la base militar, 18 Oct. 1990, Joost Duivenvoorden & I. C.leeJ 300 (liolo- type, Herbario Amazonico; isotype, \ DB). fig¬ ure la — n. Xyris trachysperma, species nova: a A. bialata Malme foliis et scapis ciliatis, bracteis fertilibus minute rufociliatis, a X. melanovaginata Krai caulibus brevioribus, scapis bialatis, areis dorsalibus distinctibus, et ab ambobus semi- nibus paucis (2-4), cupiformis, 1 .3- 1 .4 mm longis, grosse papillatis bitruncatis, bi-umbilicatis, apicibus anguste ap- iculatis, placentatio basalis differt. Sturdy caespitose perennial to 75 cm high, roots fibrous, somewhat coarse. Stems short, to 4 cm. Leaves flabellately spreading, the principal ones 40- 60 cm long, longer than the scape sheaths; sheaths entire, ecarinate at base, carinate above it. lustrous brown or dark red-brown, the keel a deep shining red-brown, often rusty-ciliolate, the sides abruptly, then gradually narrowing to a short-ligulate apex; blades elliptic-linear, 3-4 times longer than the sheaths, 5-10.1 mm wide, flat, olivaceous, with several broad, low nerves, the surfaces finely pa¬ pillose; tips abruptly incurved-acute, densely red- ciliate; margins with a narrow, deep, red-brown, inc.rassate, lustrous, entire to intermittently rusty- ciliolate border. Scapes gradually widening from base upward, broadly bi-alate, distally to 6 mm wide, green, slightly thickened and rusty-ciliolate at mar¬ gin, the sheaths angulate, short-bladed. Spikes nar¬ rowly ellipsoid or cylindric, 1 .5-2.5 cm long, obtuse, many-flowered; sterile bracts 2 pairs, decussate, the lower pair triangular, strongly carinate, 3-5 mm long, the carinas ciliolate, the inner pair obovate, ca. 3-3.5 mm long; fertile bracts broadly ovate to broadly elliptic, 6-7 mm long, convex, ecarinate, at anthesis with a light brown matrix, reddish brown, with margin rusty fimbriolate and with the dorsal areas ovate or broadly elliptic, green, with costa indistinctly antrorsely arcuately branched. Flowers with lateral sepals free, curved, inequilateral, ca. 6 mm long, acute to acuminate; keel broad, rusty- Novon 3: 55-57. 1993. 56 Novon Figure 1. Xyris trachysperma Krai & Duivenvoorden. — a. Habit sketch. — b. Leaf apex. — c. Leaf blade, midsector. — d. Leaf blade-sheath junction. — e. Leaf base. — f. Spike. — g. Scape sector. — h. Fertile bract. — i. Lateral sepal. — j. Petal blade, stamen. — k. Staminode (left) and enlarged beard hair (right). — 1. Stylar apex, m. Dehisced capsule showing one seed on elongate funiculus. — n. Seed. Volume 3, Number 1 1993 Krai & Duivenvoorden Xyris trachysperma 57 ciliate. Petals with blades orbicular, light yellow, ca. 6 mm long, apically lacerate. Staminodia 2-branched, the branches densely long-penicillate. Anthers ob¬ long, 2.5 mm long, deeply bifid, slightly sagittate; filaments terete, 1.5-2 mm long. Capsule obovoid, 3-4 mm long, acuminate; placentation basal with elongate-clavate funicles. Seeds few (2-4), barrel¬ shaped, 1.3- 1.4 mm long, dark dull brown, coarsely papillate or tuberculate, obscurely coarsely ribbed, the ends truncate-umbilicate, the apex narrowly apiculate. This species is one ol the most distinctive of American A yris, primarily because ol its unique seed form and sculpture. An attempt to align it with some previous large treatments oi South American Xyris, such as those by Idrobo (1954), Maguire & L. B. Smith (1964), L. B. Smith & R. J. Downs (1968), or myself (1988), would place it. as to general char¬ acters, nearest A. melanovnginata Krai & Lyman B. Smith of the high tepuis of Bolivar, Venezuela, or V. biala t a Malme far to the southeast in the Brazilian planalto. Both of these are likewise sturdy, with similarly pigmented foliage, leaves in fans, with strongly broadened scapes, and with similar spikes, fertile bracts, and sepals. However, \. melanova- ginata, while displaying similar leaves anil remark¬ ably similar spike outlines, has much longer stems; its broad scapes, while compressed, are not bi-alate; its spike bracts lack dorsal areas. Also, and perhaps most importantly, placentation in A. melanovagin- ata is distinctly axile. On the other hand, the Bra¬ zilian A. bialata comes nearest overall, even though it occurs far to the southeast. It combines a short stem, broadly bi-alate scapes, and iertile bracts with distinct dorsal areas; its placentation is oi the central type. Such features thus, in a general treatment, align it most closely with \. trachysperma , which is nonetheless separated by its ciliate leaf blades and scapes, its narrower spikes, its ciliolate bract edges, its basal (rather than free-central) placentation, and particularly by its fewer, larger, and distinctive seeds. Literature Cited Idrobo, J. M. 1954. Xiridaceas de Colombia. Caldasia 6(29): 184-260. Krai, R. 1988. The genus Ayres (Xyridaceae) in \ en- ezuela and contiguous northern South America. Ann. Missouri Bot. Card. 75: 522-722. Maguire, B. & L. B. Smith. 1964. Xyridaceae. In: B. Maguire, J. J. Wurdack & Collaborators, The Botany of the Guayana Highland — part \ . Mem. New A ork Bot. Card. 10(5): 7-37. Smith, L. B. & R. J. Downs. 1968. Xyridaceae. In: 1. C. Hoehne, FI. Brasilica 9(2): 1-215. A New Name for an Indian Memecylon (Melastomataceae) P. Lakshminarasimhan and Sam P. Mathew Botanical Survey of India, A & N Circle, Port Blair-744 102, India Abstract. A new name, Memecylon balakrish- nanii, is proposed for a species of Memecylon de¬ scribed by Balakrishnan & Nair. Balakrishnan & Nair (1983) described the new species Memecylon collinum from Saddle Peak, North Andaman Island, India (holotype, N. P. Ba- lakrishnan & N. G. Nair 4764 .4, CAP; isotypes, 4764 B D, PBL). The name is occupied by M. collinum Jacques-Felix (1979) from Gabon and Cameroon (holotype, N. Halle 3587, P). Thus, M. collinum N. P. Balakrishnan & N. C. Nair is a later homonym and illegitimate. A new name is required for the species described by Balakrishnan and Nair (1C BN, Art. 64. 1 ; Greuter et al., 1 988). This species is renamed after N. P. Balakrishnan, for his out¬ standing contributions to plant systematics in India. Memecylon balakrishnanii P. Lakshminarasim¬ han & S. P. Mathew, nom. nov. Memecylon collinum N. P. Balakrishnan & N. G. Nair, Bull. Bot. Surv. India 24: 30. 1982 [1983], not H. Jacques-Felix 1979: 420. Acknowledgment. We thank the Director, Bo¬ tanical Survey of India, for facilities and encour¬ agement. Literature Cited Balakrishnan, N. P. & N. G. Nair. 1982 [1983], New taxa and record from Saddle Peak, Andaman Islands. Bull. Bot. Surv. India 24: 28-36. Greuter, W., H. M. Burdet, W. G. Chaloner, V. Demoulin, R. Grolle, 1). L. Hawksworth, D. H. Nicolson, P. C. Silva, F. A. Stafleu & E. G. Voss. 1988. Inter¬ national Code of Botanical Nomenclature. Regnum Veg. 118. Jacques-Felix, H. 1979. Especes nouvelles et peu con- nues du genre Memecylon (Melastomataceae) en Af- rique. Adansonia, ser. 2, 18: 409-432. Novon 3: 58. 1993. New Species of Ma nett in (Rubiaceae) from Mesoamerica David //. Lorence National Tropical Botanical Carden, P.O. Box 340, Lawai, Kauai, Hawaii 96765, U.S.A. John D. Dwyer Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A. Abstract. Three new species of Manettia (Ru¬ biaceae) are described irom Panama: M. arboricola , M. microphylla , and M. longicalycina. Manettia Mutis ex L. (Rubiaceae) is a genus ol herbaceous or somewhat woody, usually twining vines distributed in the West Indies, Mexico, Central America, and South America south to subtropical Argentina and Paraguay. Comprising an estimated 80-130 species, this genus was revised by Wern- harn (1918, 1919) and treated for Central America and Mexico in the \orth American Flora (Standley, 1921), Flora of Guatemala (Standley & Williams, 1975), and Flora of Panama (Dwyer, 1980). Al¬ though two papers on the classification of Manettia were published by Chung (1967, 1968), no recent comprehensive accounts of the genus exist. Traditionally, Manettia has been placed in the (. inchoneae (de Candolle, 1830; Schumann, 1889; Steyermark, 1974; Robbrecht, 1988), on account of its dry fruits with numerous ovules per locule and winged seeds. Because Manettia contains raphides, Bremekamp (1966) and Verdcourt (1958) removed it from Cinchoneae and placed it in Hedyotideae, an option also mentioned by Robbrecht (1988). In a recent cladistic study ot Cinchoneae incor¬ porating cpDNA data and other inhumation, An- dersson & Persson (1991) suggested that the pres¬ ence or absence of raphides in Rubiaceae is not as important a character as previously thought. An- dersson and Persson placed Manettia in Hedyoti¬ deae in the same clade as Bouvardia, Hedyotis, and Hindsia, based on shared characters of distinct placentas attached centrally by a stipe, testal out¬ growth forming a concentric wing, wood having fiber tracheids and predominantly solitary vessels, and loculicidal capsules dehiscent only at the apex. During the course ol preparing a treatment ol Manettia for Flora Mesoamerieana, three new spe¬ cies — all from Panama — came to light and are de¬ scribed below. Most interesting are the two hemiepi- phy tic species, M. arboricola Lorence and M. microphylla Lorence & Dwyer, which climb up tree trunks by means of adventitious roots, an unusual feature previously unreported in the genus. These species are otherwise characteristic of Manettia in terms of their flower, capsule, and seed morphology. 1. Manettia arboricola Lorence, sp. nov. T\ PK: Panama. Comarca de San Bias: El Llano Card Road, km 16.7, 350 m, 9°19'N, 78°55'W, It) June 1985, G. de \evers & S. Gharnley .5907 (holotype, MO 3623655, photo, PI BO; iso¬ types, F, SCZ). Figure 1 . Species non volubili hemiepiphyto habitu, radicibus ad- ventitiis, inflorescentia terminali in ramis frondosis dis- posita, floribus 3-5, corolla caerulea, tubo gracili 0.5- 0.6 mm in diametro distinguibilis. Hemiepiphytic vine ascending tree trunk, the main stem woody, 1.5-2 mm diarn., the bark brownish white, peeling, adventitious roots produced at nodes, the primary branches 10-23 cm long, arising at nodes, sparsely branching again, the secondary branches 4-10 cm long, 1 mm diam., usually leafy and terminated by a single inflorescence, the inter¬ nodes 1-3 mm long, each side with a longitudinal ridge flanked by 2 furrows, smooth, glabrous, whit¬ ish; leaves petiolate, the petioles 2-6 x 0.5 mm, glabrous; lamina ovate, 2.5-5 x 1-2.5 cm, the base cuneate or rounded, the apex acute, glabrous, drying dark brownish green, chartaceous or mem¬ branaceous, the 2° veins 4 6 pairs, arcuate, weakly brochidodromous; stipules narrowly deltoid, 0.5 mm long, thickened. Inflorescence terminal, glabrous, cymose, 3-5-flowered, the peduncle 10-13 mm long; flowers 4-merous, the pedicels 4-10 x 0.2 0.3 mm, bracteolate, the bracteoles subulate, 1 mm long, the hypanthium obovoid-turbinate, 1.5-2 x 1 1.2 mm, glabrous; corolla blue when fresh, salverform, the tube 8-9 mm long, 0.5 0.6 mm diam., glabrous externally and internally, the lobes narrowly lan¬ ceolate-elliptic, 7-8 x 1.5-2 mm, glabrous; sta¬ mens included, affixed 2 mm below throat, the an¬ thers 1.5 mm long, linear-ellipsoid; style 1.5 mm long, the stigma bilobed, the lobes linear, 0.5 mm Novon 3: 59-62. 1993. 60 Novon Figure 1 . Manettia arboricola Lorence. De Nevers & Figure 2. Manettia microphylla Lorence & Dwyer. Charnley 5907 (holotype, MO). McPherson 12189 (holotype, MO). long. Capsules obovoid-subglobose, 5-7 X 5-7 mm, weakly 8-10-ribbed, glabrous, brown; seeds 1.4- 1.6 mm diam., subcircular, the center dark brown, 0.6-0. 7 mm diam., the wing scarious, the testa cells radiate, elongate, the margin irregularly erose-fim- briate. Distribution. Panama, known from the Cerro Tacarcuna in the Darien Province and from the type locality in the Comarca de San Bias. Habitat. Manettia arboricola occurs in tropical wet forest and premontane wet forest from about 350 to 1,300 m. Flowering and fruiting specimens were collected in January, February, and June. Manettia arboricola is most closely allied to M. microphylla (described below), a species with much smaller leaves and fruits and solitary flowers. Both species display the same specialized hemiepiphytic habit, are characterized by nontwining stems with adventitious roots, and have corollas that are gla¬ brous in the throat. The blue corollas of M. arbor¬ icola are also unusual within the genus. Paratypes. Panama. DARIEN: Cerro Tacarcuna, trail from Tacarcuna Village on Rio Tacarcuna to Cerro Mali, 800-1,300 m, A. Gentry & S. Mori 1361 6 (MO); trail from Rio Pucuro base camp up W ridge of Cerro Mali, 640-1,000 m, A. Gentry & S. Mori 14167 (MO). 2. Manettia microphylla Lorence & Dwyer, sp. nov. TYPE: Panama. Darien: Pirre Massif, Al- turas de Nique, above Cana mine, 7°45'N, 77°40'W, 850-1,150 m, 2 Mar. 1988, Mc¬ Pherson 12189 (holotype, MO 3648261 , pho¬ to, PTBG; isotypes, F, PTBG). Figure 2. Species non volubili hemiepiphyto habitu, radicibus ad- ventitiis, foliis microphyllis, lamina elliptica vel ovato- elliptica, 5-10 mm longa, 3-5.5 mm lata, petiolis 1-2 mm longis, capsulis parvis solitariis terminalibusque dis- tinguibilis. Hemiepiphytic vine ascending tree trunk, the main stem woody, 3-4 mm diam., the bark brownish white, peeling, adventitious roots produced at nodes, the primary branches produced irregularly from nodes, (5-)10-40 cm long, 0.5-1 mm diam., sparsely to profusely branched again, the secondary branches 2-8 cm long, usually leafy and terminated by a single fruit, the internodes 4-7 mm long, whit¬ ish, each side with a longitudinal ridge flanked by 2 furrows, moderately hirtellous, the hairs 0.1 0.2 mm long, white, antrorse, unicellular; leaves ap¬ pearing distichous (actually decussate), petiolate; petioles 1-2 x 0.2 mm, sparsely hirtellous; lamina elliptic or ovate-elliptic, 6 10 x 3 5.5 mm, the Volume 3, Number 1 1993 Lorence & Dwyer Manettia 61 base cuneate or rarely attenuate, the apex acute or rounded, drying dark greenish or blackish brown (in alcohol-treated specimens), chartaceous, adaxially glabrous, abaxially strigi I lose along costa and with numerous prominent white raphides oriented parallel to veins, the 2° veins 3(-4) pairs, arcuate, weakly brochidodromous; stipules narrowly deltoid, 0.2-0. 3 mm long, glabrous, compressed, the tip thickened. Mature flowers unknown; immature flower in bud on pedicel 0.3 mm long, the hypanthium obovoid, 1 x 0.7 mm, compressed, glabrous, the calyx cup 0.15 mm deep, the lobes 4, narrowly deltoid or subulate, 0.8 1.1 x 0.2 0.3 mm; immature corolla in bud ca. 1 mm long, glabrous externally and in¬ ternally. Fruits solitary, terminal, pedicellate, the pedicel 2-4 mm long, slender, bracteolate, the brac- teoles narrowly subulate, ca. I mm long; capsule subglobose, 3-5 x 3—5 mm, slightly bisulcate, slightly compressed, weakly 6-ribbed, crowned by a low, bilobed disc, and 4 narrowly deltoid or subulate calyx lobes 1.2 I .5 x 0.4-0. 5 mm, glabrous, raph¬ ides conspicuous; seeds subcircular, 1 1.3 mm diarn., the center dark brown, 0.4-0. 5 mm diarn., the wing light brown, the testa cells long, radiating, the margin irregularly erose- fimbriate. Distribution . Known only from the type locality, approximately 14 15 km north of the Alturas de Nique on the Panamanian Colombian border. Habitat. Tropical wet forest, at 850 1,150 m altitude. The type collection was fruiting in March. Manettia microphylla is undoubtedly most close¬ ly related to \1. arboricola , as both species share the nontwining, hemiepiphytic habit, have adventi¬ tious roots, similar stipule and seed morphology, terminal inflorescences on leafy shoots, and inter¬ nally glabrous corollas. The latter species differs in having much larger leaves (25-50 x 10-25 mm), larger fruits 5-7 mm long and wide, and 3-5-fiow- ered cymes. Manettia microphylla represents an extremely reduced and specialized species adapted to hemiepiphytism, growing closely appressed to tree trunks. 3. Manettia longicalycina Dwyer & I.orence, sp. nov. TYPE: Panama. Code: Alto Calvario, cloud forest, 800-900 m, 20 Apr. 1977, J. P. Folsom A' I. Jaslon 2f)()5 (holotype, MO 3608777, photo, PTBC). Figure 3. Species Manettiae reclinatae L. affinis, floribus in ped- icellis brevibus ( 1 - )3 — 5 mm longis, calycis lobis longior- ibus lineari-lanceolatis vel lineari-subulatis, 5-12 mm lon¬ gis, corolla alba vel rosea difFert. Dextrorsely twining herbaceous vine, the young stems quadrangular, the mature sterns cylindrical, Figure 3. Manettia longicalycina Dwyer & Lorence. balsam & Jaslon 2665 (holotype, MO). 2-3 mm diam., with 4 low wings 0.2-0. 3 mm wide, densely retrorsely hirtellous especially on wings. Leaves petiolate; petioles 2- 1 0( — 1 7) x 0.5 1 mm, sparsely villosulous or strigillose; lamina ovate or ovate-elliptic, ( L .5— )2 .5 9.5 x (1 )2 0 cm, ihe base rounded or obtuse, the apex acuminate, the acumen (2 )5-15 mm long, chartaceous or sub- coriaceous, drying green or blackish green, the 2° veins 4 5 pairs, arcuate, the venation visible to 2° or 3° on both surfaces; stipules broadly obtuse or semicircular, 0.5- 1 x 3-4 mm, spreading or re¬ flexed, glabrous, the margins bearing a fringe of thick, conical, pale brown colleters 0.3 mm long. Flowers axillary, solitary or in cymes of 2 5, or in axillary leafy inflorescences 2-2.5 cm long, the ped¬ icels and peduncles with a pair of subulate braeetoles 2-3 mm long, the peduncle 1 2 mm long; flowers on pedicels ( 1 — )3— 5 mm long, 0.5 mm diam., stri- gillose-hirtellous, the hypanthium obconical, 2-3 x 2 mm, compressed, hirtellous, the calyx cup 0.5— 0.6 mm deep, the lobes 8 or occasionally 6, linear- lanceolate to linear-subulate, 5-9(-12) x 0.5 1 mm, strigillose-hirtellous or glabrate, erect-spread¬ ing, each sinus with 3-4 small, brown colleters; corolla white or pink when fresh, salverform, the tube 11-16 mm long, 2-2.5 mm wide medially, 62 Novon externally sparsely hirtellous, internally with a white villous ring just above middle or glabrous, the throat barbate with villous yellow trichomes, the lobes el¬ liptic, 4-7 x 2-2.5 mm, the apex acute, usually spreading 45°, occasionally with hirtellous or fringed margins; stamens affixed 1-2 mm (thrum) or 6-7 mm (pin) below throat, anthers linear, 2.5-3 mm long, included in tube or tips exserted; style glabrous, 6-9 mm (thrum) or 13 mm (pin) long. Capsules broadly obovoid or obcordiform, strongly com¬ pressed, bisulcate, 5-7 mm long and wide, glabrate or hirtellous, venose, the pedicels 5-7 mm long; seeds subcircular, 2-3 mm diam., the dark brown center 1 mm diam., the wing 0.6-1 mm wide, brown, the testa cells radiate, elongate, the margin erose- laciniate. Distribution . Manettia longicalycina is known only from Panama, where it has been collected on the Atlantic slope in the Provinces of Code (Alto Calvario, La Mesa, El Valle Mesa, El Cope, and between Cerro Pilon and El Valle de Anton), Panama (Cerro Campana, and between Cerro Azul and Cerro Jefe), and Veraguas (Escuela Agricola Alto Piedra). Habitat. This new' species occurs in tropical wet forest and cloud forest from about 700 to 1,300 m elevation, where it grows in disturbed sites or clear¬ ings. Flowering specimens were collected in April July, September, and October, and fruiting speci¬ mens in January, April-Oetober, and December. Manettia longicalycina appears to be most closely allied to M. reclinata L., from which it differs by the characters given in the diagnosis. This new spe¬ cies is also closely related to an undescribed species from Costa Rica (C. M. Taylor, pers. comm.) char¬ acterized by its much longer peduncles 10-25 mm long and pedicels 10-25 mm long, (6-)8-merous flowers, and corollas with a shorter tube 6 10 mm long and lobes 3-4 mm long. Manettia longica¬ lycina typically has white or less frequently [link corollas. However, a single collection from the Canal Zone [Fort Sherman Site, U.S. Army Tropic Test Center, J. D. Dwyer 8595, MO] was said to have dull purple-red flowers. In addition to the anomalous flower color, this collection occurs in the wrong climatic zone and at an atypically low altitude for the species, a little above sea level. Its taxonomic status is therefore doubtful. Paratypes. Panama. COCLE: between Cerro Pilon and El Valle de Anton, 700-900 m, J. A. Duke & J. D. Dwyer (MO); La Mesa, 2 km W of Cerro Pilon, 900 m, G. A. Sullivan 509 (MO, US); 7 km N of El Cope, near Rivera sawmill, 700-850 m, J. P. Folsom 5230 (MO); 2.2 km beyond sawmill in forest along lumber road above El Cope, 900 m, B. Hammel 992 (MO); El Cope on Pacific side, '/(.-hour walk from sawmill, 2,600 ft., T. Antonio 2060 (MO); above El Potroso sawmill at Con¬ tinental Divide, N of El Cope, K. Sytsma & L. Andersson 4567 (MO); sawmill above El Cope, along stream E of sawmill, 2,300 ft., B. Hammel 4080 (MO); above sawmill above El Cope, 1,000 m, J. S. Miller et al. 824A (MO); 7 km N of El Cope, Alto Calvario, 700-850 in, J. P. Folsom et al. 5732 (MO); El Valle Mesa, 5.6 km along Mesa road from main road in El Valle, 700-800 m, J. P. Folsom 3891. Z (MO); La Mesa, above El Valle de Anton, ca. 2 km W of Cerro Pilon, 900-930 m, T. B. Croat 37483 (MO), panama: Cerro Campana, K. L. Dressier 3524 (MO), 780-875 m, T. B. Croat 25190 (MO), 2,600-2,800 ft., J. D. Dwyer 1922 (MO), 850 m, J. S. Miller et al. 769 (MO); between Cerro Azul and Cerro Jefe, R. L. Dressier 3248 (MO), veraguas: 0.2 mi. beyond fork in road at Escuela Agricola Alto Piedra on road to Rio Calovebora, 750 rn, T. B. Croat & J. P. Folsom 33899 (MO). Acknowledgments. We are grateful to Charlotte M. Taylor and an anonymous reviewer for their critical reading of the manuscript and helpful sug¬ gestions. Literature Cited Andersson, L. & C. Persson. 1991. Circumscription of the tribe Cinchoneae (Rubiaceae) — A cladistic ap¬ proach. PI. Syst. Evol. 178: 65-94. Bremekamp, C. E. B. 1966. Remarks on the position, the delimitation and subdivision of the Rubiaceae. Acta Bot. Neerl. 15: 1-33. Candolle, A. P. de. 1830. Prodromus Systematis Na- turalis Regni Vegetabilis, IV. Paris, Truettel & Wiirtz. Chung, I. 1967. Studies in Manettia (Rubiaceae) sec¬ tion Heterochlora Schum. Phytologia 15: 272 288. - . 1968. Studies in Manettia (Rubiaceae) section Pyrrhanthos Schum. Phytologia 17: 353-366. Dwyer, J. D. 1980. Manettia. In: Flora of Panama. Rubiaceae. Ann. Missouri Bot. Card. 67: 277-282. Robbrecht, E. 1988. Tropical woody Rubiaceae. Opera Bot. Belgica 1: 1-271. Schumann, K. 1889. Rubiaceae. In: C. F. P. von Mar- tius, A. G. Eichler & I. Urban (editors), Flora Brasili- ensis 6(6). Fleischer, Leipzig. Standley, P. C. 1921. Rubiaceae. In: North American Flora 32: 96-100. - & L. (). Williams. 1975. Manettia. In: Flora of Guatemala. Rubiaceae. Fieldiana, Bot. 24(11): 121-123. Steyermark, J. A. 1974. Rubiaceae. In: Flora de Ven¬ ezuela 9(1): 121-158. Verdcourt, B. 1958. Remarks on the classification of the Rubiaceae. Bull. Rijksplantentuin Bruss. 28: 209 281. Wernham, H. F. 1918. The genus Manettia. J. Bot. 57 (Suppl.): 1-16. - . 1919. The genus Manettia. J. Bot. 57 (Suppl.): 17-44. A New Species of Ardisia (Myrsinaceae) from Madagascar James S. Miller and John J. Pipoly III Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. Abstract. Recent exploration in the Marojejy Natural Reserve has resulted in the discovery of a new species, Ardisia (subg. Akosmos) marojejyen- sis. The species is described and illustrated, and its phylogenetic relationships are discussed. A key to distinguish the three known Madagascan species of Ardisia is provided, and subgenus Madardisia is relegated to synonymy under subgenus Akosmos. Ihe Reserve Naturelle Integrale de Marojejy (number 12) comprises 60,150 hectares of wet, tropical forests that range from ca. 100 to 2,137 m in elevation (Jenkins, 1987). Henri Humbert bot¬ anized extensively in the region and made approx¬ imately 3,000 collections, many of which were spe¬ cies new to science (Humbert, 1955). Humbert's efforts were undoubtedly one of the factors drawing interest to the rich, largely endemic flora and leading to the declaration of the area as a reserve in 1952. Despite the efforts of Humbert and botanists who have since continued collecting in the reserve, the flora remains poorly known and continues to yield novelties. Recent estimates (Miller, ined.) indicate that the flora of the reserve contains about 2,000 species, perhaps 20% of the total flora of the coun¬ try. Collecting efforts in the southern part of the reserve in February 1989 have yielded a previously undescribed species of Ardisia Swartz (Myrsina¬ ceae), only the third reported for Madagascar. Ardisia (suhg. Akosmos) marojejyensis James S. Miller & Ripoly, sp. nov. TV PE: Madagascar. Antsiranana: Reserve Naturelle Integrale de Marojejy, along the trail to the summit of Ma¬ rojejy Est, NW of Mandena, wet, evergreen forest above the second camp, 700 850 m, 14°26'S, 49°16'E, 10 Feb. 1989 (fl), James S. Miller A- P. P. Lowry 3936 (holoytpe, MO 4064879; isotypes, K, P, TAN, US). Figure 1. Arbor ad 15 m alta. Folia persistentia; lamina obovata, 18-28 cm longa, 8-12 cm lata, apice late obtuso ad rotundato, basi cuneata ad obtusa; petiolo 3 5 cm longo. Flores pedicellis 1 -2 mm longis inserti, sepalis 1.7-2 mm longis, petalis 4-5 mm longis, antheris 2.5-3 mm longis, stylo ca. 3 mm longo. Fructus nobis non visus. Free 10-15 m tall; branchlets terete, 1 1.5 cm diam., densely furfuraceous-lepidote, the scales so densely packed as to form an apparent sheet, later breaking up and somewhat glabrescent. Leaves per¬ sistent; lamina carnose when fresh, drying coria¬ ceous, obovate, 18-28 cm long, 8-12 cm wide, the apex widely obtuse to rounded, the base cuneate to obtuse, decurrent on the petiole, midrib depressed above, prominently raised and black punctate-lin- eate below, the secondary veins 35-40 pairs, 0.5- 1 cm apart, inconspicuously pellucid punctate, gla¬ brous above and below; petiole marginate, 3 5 cm long, densely furfuraceous-lepidote, glabrescent. In¬ florescences clustered near the apices of branches, internodal, erect, paniculate, the rachis 15-30 cm long, the lateral branches 5-10 cm long, glabrous or nearly so. Flowers bisexual, borne on pedicels 1 2 mm long; buds ovoid to conical; sepals 5, quin- cuncial, greenish white, ovate, 1 .7-2 mm long, gla¬ brous, sparsely pellucid punctate, the margin scar- ious, hyaline, glabrous; petals 5, quincuncial, bright pink, basally connate, ovate to narrowly ovate, 4 5 nun long, 1 .5-2 nun wide, apex acute, symmetric, prominently pellucid punctate, the margin irregular, entire, glabrous; stamens 5. the filaments 0.2-0. 4 mm long, basally adnate to the petals, free from each other, the anthers bright yellow, lanceoloid, 2.5-3 mm long, apex long-apiculate, base cordate, dehiscing first by apical pores, then narrow, longi¬ tudinal slits, the connective conspicuously brown punctate dorsaily; ovary ovoid, 0.5 1 mm long, the placenta depressed-globose, 0.4 0.6 mm long, 0.6 0.8 mm diam., apex apiculate, the ovules 4, biseri- ate, the style ca. 3 mm long, the stigma punctiform. Fruit unknown. Local names: “Talandoha” (near Mandena); “Maimbola” (Ambatosoratra). Ardisia marojejyensis is a spectacular, appar¬ ently mass-flowering tree, covered with pink flowers, making it easily visible from a distance. It is un¬ common in the reserve at middle elevations, and few individuals were seen even though all in a given valley can easily be counted. Because this species appears to mass flower (all individuals flower syn¬ chronously in a short period of time), it is probable that it has been missed by previous collectors purely Novon 3: 63-65. 1993. 64 Novon Figure 1. Ardesia marojejyensis James S. Miller & Pipoly. — A. Flowering branch, showing erect, pyramidal panicles with racemose branches. — B. Section of the inflorescence showing conical buds and open flowers. — C. Anthers showing apical pores and longitudinal slits. — D. Longisection of ovary showing placenta and biseriate ovules. Drawn from Miller & Lowry 3936. by chance. However, when it was recently collected, the senior author noted it was a conspicuous element ot the forest, and many individuals were readily visible Irom most vantage points. Paratype. MADAGASCAR, antsiranana: Reserve Na- turelle Integrate de Marojejy, N slopes of Ainbatosoratra, wet montane forest on steep slopes, 700-900 m, 14°32’S, 49°41'E, 24 Feb. 1989 (fl), James S. Miller 4196 (MO, TAN). Volume 3, Number 1 1993 Miller & Pipoly Ardisia 65 Key to the Species of Ardisia in Madagascar la. Leaves 6- 16 cm long; inflorescences umbelli- form, long-pedunculate; anthers dehiscent by terminal pores; plants of western forests .... . A. didymopora (H. Perrier) Taton lb. Leaves 18-40 cm long; inflorescences race¬ mose or corymbose, in pyramidal panicles; an¬ thers dehiscent by terminal pores, then narrow, longitudinal slits; plants of eastern forests. 2a. Leaves borne on petioles 3-5 cm long; panicles erect, pyramidal, the branches racemose; pedicels 1-2 mm long; petals 4-5 mm long, the apices acute . . . A. marojejyensis James S. Miller & Pipoly 2b. Leaves sessile or nearly so; panicles pen¬ dent, columnar, the branches corymbose; pedicels 10-20 mm long; petals 10-11 mm long, the apices abruptly acuminate- caudate . A. procera Capuron Ardisia marojejyensis appears closely related to A. procera Capuron, another species from the wet forests of northeastern Madagascar. The paniculate inflorescences subequal to the leaves with racemose branches and long peduncles, the slightly curved style subequal to the petals, punctiform stigma, and anthers with first poricidal, then longitudinal dehis¬ cence, clearly indicate that both species are mem¬ bers of subgenus Akosmos Mez. Subgenus Akosmos was thought to have its center ol diversity in sub¬ tropical central Asia and eastern Asia (Mez, 1902), but recent studies by Stone (1989, 1990) indicate its center of diversity lies in the Malesian region. Capuron (1963) placed Ardisia procera in his new subgenus Madardisia, which we consider a taxo¬ nomic synonym of subgenus Akosmos. In comparing the descriptions of the subgenera, we have found that the principal difference was the fact that Ca¬ puron considered Madardisia to have strictly porici¬ dal anther dehiscence, while Mez (1902) indicated that Akosmos had longitudinal anther dehiscence. Our examination of representative species of Ardisia subg. Akosmos from Madagascar, central tropical Asia, eastern subtropical Asia, and Malesia indicates that the anthers in many species open first by con¬ spicuous or inconspicuous pores (terminal or sub¬ terminal), then by longitudinal slits. Failure to un¬ derstand this morphogenetic process led Mez and Capuron to have incomplete concepts of the group. Therefore, the fact that two of the three Ardisia species thus far known from Madagascar belong to subgenus Akosmos is phytogeographically interest¬ ing. We hope that further collection in Madagascar and in the Malesian region will help the current dearth of knowledge on the population biology of these species. Ardisia procera and A. marojejyensis are similar in general aspect and occur in similar forest types. However, Ardisia marojejyensis differs from A. procera in having leaves borne on marginate petioles 3-5 cm long, erect pyramidal panicles with race¬ mose branches, and smaller flowers on short (1-2 mm) pedicels. Capuron (1963) described A. procera as deciduous, but this appears to be either the result of his describing a western dry forest race (or eco¬ type) formation, or an error. Ardisia marojejyensis certainly appears to have persistent foliage, and A. procera, also from a high-rainfall region, is most probably evergreen as well, at least in the rainforest zone. On the other hand, the other Madagascan species of the genus, Ardisia didymopora (H. Per¬ rier) Taton, from dry regions in the west, is clearly deciduous. Acknowledgments. We thank Hoy E. Gereau for helpful comments on the manuscript and Latin de¬ scription, and John Myers for the illustration. Field¬ work at Marojejy was funded by the National Geo¬ graphic Society (3647 87), the World Wildlife Fund-U.S., and World Wide f und for Nature (pro¬ ject #6502); their support is gratefully acknowl¬ edged. Literature Cited Capuron, R. 1963. Contributions a l’etude de la flore de Madagascar. XIV. Le Genre Ardisia Swartz (Myr- sinacees) a Madagascar. Adansonia, ser. 2, 3: 380- 385. Humbert, H. 1955. Une rnerveille de la nature du Madagascar: Premiere exploration botanique du mas¬ sif du Marojejy et des ses satellites. Mem. Inst. Sci. Madagascar, ser. B, Biol. Veg. 6: 1 272. Jenkins, M. D. 1987. Madagascar. An Environmental Profile. IUCN, Gland, Switzerland. Mez, C. 1902. Myrsinaceae. In: A. Engler (editor). Das Pflanzenreich. IV. 236: 1-437. Wilhelm Engel- mann, Leipzig, Germany. Stone, B. C. 1989. New and noteworthy Malesian Myr¬ sinaceae, III. On the genus Ardisia Sw. in Borneo. Proc. Acad. Nat. Sci. Philadelphia 141: 263-306. - . 1990. Studies in Malesian Myrsinaceae: 5. Additional new species of Ardisia Sw. Proc. Acad. Nat. Sci. Philadelphia. 142: 21-58. Tapirira obtusa comb. nov. (Anacardiaceae) John I). Mitchell The New York Botanical Garden, Bronx, New York 10458-5126, IJ.S.A. ABSTRACT. The discovery that one of the syntypes cited hy Engler under the name Tapirira mar- chandii is also the type for an earlier, validly pub¬ lished name, Mauria obtusa Bentham, necessitates the transfer of Mauria obtusa to Tapirira. The new combination Tapirira obtusa takes priority over Tapirira marchandii Engler. A review of the literature and specimens of Tap¬ irira (Anacardiaceae) has revealed the existence of an earlier name, which has priority over Tapirira marchandii Engler. The specimen “ Schomburgk 892" [= Robert Schomburgk series II 892 ] is included in the list of syntypes cited by Engler (1876) under the original description of Tapirira marchandii. This is the same specimen as one of the two syntypes mentioned by Bentham (1852) for his species, Mauria obtusa. Bentham described two other species of Mauria in the same article. Two of the species, Mauria mul¬ tiflora C. Martius ex Bentham and Mauria subbi- juga C. Martius ex Bentham, were later treated as synonyms of Tapirira guianensis Aublet by Engler (1876). However, Engler overlooked the third spe¬ cies of Mauria, Mauria obtusa, described by Ben¬ tham (1852: 16) in the same article. Later authors were also apparently unaware ol Bentham’s third Mauria species. The name Mauria obtusa is not listed in any of the editions of Index Kewensis or the Gray Card Index. In conclusion, the basionym Mauria obtusa has priority over the name Tapirira marchandii. The syntypes of Mauria obtusa, Robert Schom¬ burgk II 892 and Richard Schomburgk 1442, clearly agree morphologically with the other syn¬ types listed by Engler under the name Tapirira marchandii. These syntypes evidently belong in the genus Tapirira, not Mauria, because they are uni¬ sexual, the petals are imbricate, and the staminate pistillode and the pistil are both crowned by five styles. This is the typical condition of the flowers of Tapirira. Mauria, on the other hand, has bisexual flowers with a valvate corolla and a single style crowned by three sessile stigmas. Bentham (1852) wrote that he was aware of the affinity of his three taxa of Mauria to Tapirira guianensis. (He actually published M. obtusa as “ Mauria (Tapirioides) obtusa") He probably mis¬ placed these three species in Mauria instead of in Tapirira because of the incorrect fruit description and illustration in Aublet (1775). Aublet included a fruit from a different plant family in his description and illustrations of Tapirira guianensis. The following is my proposed new combination and synonymy. Tapirira obtusa (Bentham) Mitchell, comb. nov. Basionym: Mauria obtusa Bentham, Hookers J. Bot. Kew Card. Misc. 4: 16. 1852. TYPE: Guyana, 1842-1843, Robert Schomburgk II 892 (lectotype, designated here, K; isolecto- type, NY). \Richard Schomburgk 1442 was probably collected from the same plant.] Tapirira pao-pombo Marchand var. major Marchand in Warming, Vidensk. Meddel. Dansk. Naturhist. For- en. Kjobenhavn 15: 59. 1873. TYPE: Brazil. Minas Gerais: Lagoa Santa, Warming s.n. (syntypes, C). Tapirira marchandii Engler in C. Martius, FI. Bras. 12(2): 379. 1876. Based on Tapirira pao-pombo Marchand var. major Marchand. Tapirira peckoltiana Engler in C. Martius, FI. Bras. 12(2): 379. 1876. TYPE: Brazil. Rio de Janeiro: Canta Gallo, Peckolt 348 (holotype, BR). Tapirira pao-pombo var. major was placed in synonymy by Engler (1876) in his description of Tapirira marchandii. 1 agree with Barfod’s (1987) taxonomic decision that T. peckoltiana and 7. mar¬ chandii are the same species. Acknowledgments. I thank Scott Mori for helpful comments and Rosemary Lawlor for typing the manuscript. Literature Cited Aublet, F. 1775. Histoire des Plantes de la Guiane Fran^oise, 4 vols. Didot, Paris. Barfod, A. 1987. Anacardiaceae. Flora of Ecuador 30(104): 11-49. Bentham, G. 1852. Second report on Mr. Spruce’s collections of dried plants from North Brazil. Hook¬ er’s J. Bot. Kew Gard. Misc. 4: 8-18; 12(2): 367- 418. Engler, A. 1876. Anacardiaceae. In: C. F. P. von Martius, FI. Brasiliensis 12(2): 367-418. Novon 3: 66. 1993. Two New Mesoamerican Species and a New Combination in Huperzia (Lycopodiaceae) Benjamin 0llgaard Department of Systematic Botany, Institute of Biological Sciences, Aarhus University, 68 Nordlandsvej, DK 8240 Kisskov, Denmark Abstract. Two new species ol Lycopodiaceae, Huperzia mesoamericana and //. talamancana, are described as a result of revision of material for the pteridophyte volume of Flora Mesoamericana. The relationship of H. talamancana to //. crassa is discussed. A new combination, Huperzia curvi - folia var. parvifolia, is proposed. Huperzia mesoamericana B. 011gaard, sp. nov. TYPE: Panama. Chiriqui: Cerro Hornito, epi¬ phyte, in cloud forest, 6,200 It., 8 May 1978, Hammel 3061 (holotype, MO). Figure 1. Species Huperziae taxifoliae (Swartz) Trevisan vero- similiter affinis, a qua differt foliis surculorum basalium 12-15 x 4-5 mm, ellipticis usque ad late lanceolatis, obtusis usque ad late acutis, et foliis surculorum dense sporangiferorum 6-8 x 2.5-3 mm, late lanceolatis, planis vel fere planis. Pendent, up to 25 cm long; shoots slightly and gradually tapering from ca. 20-30 mm diam. in¬ cluding leaves at the base to 10-15 mm in distal divisions; stems (excluding leaves) 2-3 mm thick at the base; leaves gradually modified, borne in irreg¬ ular, alternating whorls of 3, covering the stem; leaves of basal divisions spreading to perpendicular, elliptic to broadly lanceolate, obtuse to broadly acute, 12-15 x 4-5 mm, almost flat, firmly herbaceous to subcoriaceous, somewhat shining, with slightly revolute, entire margins; leaves of middle and distal divisions gradually smaller, more acute, more ar- cuately ascending, flat, with flat or slightly revolute margins; leaves of fully sporangiate divisions 6-8 x 2.5-3 mm; sporangia 1.5 2 mm wide. Huperzia mesoamericana grows as an epiphyte in cloud forests, from 1 ,300 to 2,050 in, and is endemic to Costa Rica and Panama. It is a very distinct species of uncertain taxonomic affinity. Be¬ cause it is a pendent epiphyte with gradually het¬ erophyllous shoots and rather wide leaves of firm texture, this species is referred tentatively to the Huperzia taxifolia group of 011gaard (1987, 1989). It differs from //. taxifolia by the shorter growth and more compact aspect, by the elliptic to widely lanceolate leaves, 12-15 x 4-5 mm, with obtuse to broadly acute apices, in the basal divisions of the plant, and by the leaves of the densely sporangiate distal divisions 6 8 x 2.5-3 mm, widely lanceolate and flat, with fiat or slightly revolute margins. In contrast, H. taxifolia has linear-lanceolate to lan¬ ceolate leaves, 14-23 x 2-3 mm, in the basal divisions, and the leaves of the densely sporangiate distal divisions 3-8 x 1-1 ,5(~2) mm, lanceolate or narrower, usually with a widened and somewhat clasping base, and a long, narrow, involute apex. Paratypes. Costa Rica, cartago: San Mateo et le chemin de Guacimo au bord de la mer du Pacifique, Brenes, A. M. s.n. (NY), alajuela-puntarenas-cuana- CASTE: Cordillera de Tilaran, Monteverde, 1,300-1,400 m, Aug. 1978, Dryer 1618 ( F). puntarenas: Montever¬ de, Veracruz river valley S of Reserve, Pacific slope wet forest, 10°15'N, 84°46'W, 1,300-1,500 m, 22 Aug. 1986, Haber ex Bello <£■ Clagget 5367 (MO). Panama. CHlRiyiJl: Cerro Colorado, 1,600 m, 9 Aug. 1984, Schmalzel 1997 (MO); Cerro Colorado, 11.2 kin along ridge road from main road to Escopeta, 1,700 m, 16 Aug. 1977, Folsom 4873 (MO). Huperzia talamancana B. 011gaard, sp. nov. TYPE: Costa Rica. San Jose: Cordillera de la- lamanca, Cerro de la Muerte, Pan American Highway at km 91, Ericaceae-Chusquea scrub, 3,000-3,300 m, 25 Aug. 1983, Hickey 898 (holotype, AAU). Figure 2. Species Huperziae crassae (Humboldt & Bonpland ex Willdenow) Rothmaler affinis, a qua differt surculis bas- alibus homoblasticis, foliis (6— )8 1 1 mm longis, (1.5-)2- 3 mm latis, ascendentibus usque ad laxe imbricatis, abax- ialiter epidermide laevi et saepe nitida, vel interdum cel- lulis paucis protrusis pustuliformibus, non pruinosis. Erect from an ascending base, forming large clumps, sometimes up to 60 cm tall; shoots homo- blastic, homophyllous, equally thick throughout or slightly tapering, 7—1 5( -25) mm diam. including leaves; stem (excluding leaves) 3-5 mm thick; leaves borne in crowded, irregular, alternating whorls of (4-)5-7, spreading to arcuately ascending or loosely imbricate, lanceolate, (6-)8-l 1 x 2 3 mm (leaves of distal divisions sometimes 1 .5-2 mm wide), with an inconspicuous to prominent, often prominently decurrent basal swelling (air cavity), abaxially fiat Novon 3: 67-72. 1993. 68 Novon Figure 1. Huperzia mesoamericana B. 011gaard — a. Entire plant. — b. Portion of basal division with large leaves. — c. Portion of distal, sporangiate division. ( Hammel 3061, MO.) Volume 3, Number 1 1993 Ollgaard Huperzia 69 5 Figure 2. Huperzia talamancana B. 011gaard — a. Holotype. — b. Portions of two adjacent middle divisions of same. — c. Portion of middle division of plant from shaded habitat, (a and b: Hickey 898, AAU; c: Burger & Liesner 6349, GH.) to rounded or with a shallow central ridge, smooth and often shining, sometimes with a few inconspic¬ uous, slightly protruding, blisterlike epidermal cells, with irregularly protruding, clear, marginal cells, green or yellowish tinged, not pruinous; sporangia 2-3 mm wide. Huperzia talamancana grows as a terrestrial plant and under open scrub in paramo, in marshes and high swamps, often associated with Sphagnum. It appears to be rather frequent at high altitudes, mainly above 3,000 m in Costa Rica, and is known from two localities in western Panama. It belongs 70 Novon Figure 3. Huperzia crassa (Humboldt & Bonpland ex Willdenow) Rothmaler sens. lat. — a. Part of young plant with strongly heteroblastic basal divisions. — b. Portions of several middle divisions, (a: Evans & Lellinger 143, MO; b; Davidse el at. 25941 , AAU.) to the Huperzia saururus group (011gaard, 1987, 1989) and has commonly been included in a wide concept of Huperzia saururus (as Lycopodium or Hrostachys ); see, e.g., Lellinger (1989) and Rolleri (1977a, b, 1981). These authors emphasize similar features of specialized epidermal cells at the leaf margins. However, true II. saururus, as represented in the Andes from Peru to Argentina, besides Africa, differs from the present species in the strongly het¬ eroblastic development of basal divisions, and the usually very tightly appressed leaves in the erect shoots. The erect shoots of H. saururus are densely aggregated and squeezed together at the base so that the lower leaves appear etiolated, and the hor¬ izontal divisions may have strongly reduced leaf development. In herbaria this species has been referred to ( ro- stachys erikssonii Nessel (type from Colombia), and l . orion is Herter (type from Venezuela) (Herter, 1958). The taxonomic identity of the types of these names is not well understood at present, but they do not correspond to any material 1 have seen from Central America. Huperzia talamancana is apparently related to //. crassa, with which it often coexists in Costa Rica, and with which it has often been confused. Huperzia crassa (Fig. 3) differs from //. talamancana by the heteroblastic shoots (Fig. 3a), by the smaller and usually closely imbricate, somewhat pruinose leaves, (4 )5 8 x 1-1.5(-1.8) mm (Fig. 3b). The abaxial leaf epidermis is usually rugose due to protruding, blisterlike cells, while the margins have few or no specialized, protruding cells. In contrast, II. tala¬ mancana has homoblastic shoots (Fig. 2a), and leaves (6-)8-l 1 x (1.5-)2-3 nun, ascending to loosely imbricate (Fig. 2b, c), abaxially smooth and often shining, not pruinous, or sometimes with few pro¬ truding, blisterlike epidermal cells, while the margin Volume 3, Number 1 1993 0llgaard Huperzia 71 cells are specialized and usually individually pro¬ truding. It is a variable species, especially with re¬ gard to the direction of the leaves, responding strongly to the light conditions ol the habitats, as shown by the specimens in Figure 2b, from a light-exposed habitat, and Figure 2c, from a somewhat shaded habitat. Paratypes. Costa Rica, san jose: Cerro de las Vueltas, 2,700-3,000 m, 1 Jan. 1926, Standley & Valerio 43611 (US), 44004 (US), 44006 (F, US); Road Cartago-S. Isidro del General, km 88.5, Asuncion, 3,335 m, 29 Jan. 1986, A. Smith 2008 (MO, UC); Cerro de la Muerte, paramo de Buenavista, 1-3 km S of Interamerican High¬ way, 3,350 m, 9 Aug. 1967, Lellinger 869 (US), 870 (MO, US); Cerro de la Muerte, N slopes, 11,000 ft., 25 May 1971, Proctor 32070 (MO), 3,300-3,400 m, Al- verson 1853 (MO); Cerro Asuncion, near Cerro de la Muerte, 3,335 m, 4 Aug. 1965, Croat 507, 509 (MO), Stone 2035 (UC); Cerro de la Muerte, 25 July 1983, Hickey & Regan 890 (AAU); Asuncion summit, 3,335 in, 22 July 1966, Davidse & Pohl 101 18 (MO); Hotel La Georgina-Cerro Frio, Cerro de la Muerte, 3,100- 3,400 m, 9°33'N, 83°43-46'W, Davidse et al. 25008 (AAU, UC); Cerro de la Muerte, km 91 of Pan American Highway in Cordillera de Talamanca, 3,300 m, W hitmore 41 (F, GH); Cerro de la Muerte, 3,200 m, Burch 4731 (MO); Cerro de la Muerte, 3,200 m, Lorence 1751 (MO), 1752 (MO); Cerro de la Muerte, 3,400 m, Taylor 1 1730 (MO); Cerro de la Muerte, paramo Buena Vista, 1-3 km 5 of Interamerican Highway, 3,350 m, Mickel 3249 (UC, US); paramo de Buenavista, 3,400 m, 14 June 1967, de la Sola 5047 (US); Cerro de la Muerte, 3,300 m, 7 Aug. 1976, Solomon 2532 (MO), san jose-cartaco: near Asuncion, 9°34'N, 83°45'W, 3,300-3,400 m, Burger 6 Stolze 5982 (F, MO), 5977 (F, GH), 5985 (F, MO); Interamerican Highway near La Asuncion, 3,200-3,300 m, 9°34'N, 83°45'W, 21 Nov. 1969, Burger & Liesner 6349 (F, GH, MO); Cerros Cuerici, 9°35'N, 83°38'W, 3,200 m, Davidse 24762 (AAU); paramo at La Asuncion, 3,335 m, Mickel 2117 (US), 2118 (US), 2120 (NY). CARTACO: Cerro de la Muerte, La Virgen de los Angeles paramo, 3,000 in, Brown CR-88 (US); Talamanca range, near Pan American Highway, 3,400 m, 25 Feb. 1965, Lent 392 (F); San Isidro del General-Cartago, 3,400 m, 11 Jan. 1964, McKee 11203 (P, UC, US); Cordillera Talamanca, 3,300 m, 25 Aug. 1961, C. Weber 6067 (US); Cerro de la Muerte, 3,000 m, 19 Feb. 1957, M. Carlson 3512 (GH); Cerro de la Muerte, La Virgen de los Angeles paramo, 3,470 m, Godfrey 66717 (GH); Cerro Sakina, paramo Buena Vista, 3,000 m, Churchill 3367 (AAU); Cerro de la Muerte, paramo de Buena Vista, E side, 3,000 m, 24 Jan. 1981, Churchill 3711 (AAU); Cerro Buenavista, 3,000-3,300 in, 12 May 1982, Huft et al. 2141 (AAU); Cordillera Talamanca, Mt. Cerro de la Muerte, El Alto de Asuncion, 3,335 m, 4 Mar. 1966, Molina R. et al. 18338 (F, US); Cerro de la Muerte, 3,200 m, 17 May 1956, L. O. Williams 20052 (US), 3,300 m, 30 Mar! 1949, L. O. W illiams 16087 (F, MO, US), 3,200 rn, 9°35'N, 83°45'W, 1 Feb. 1965, L. 0. Williams et al. 28833 (F, GH, US); Cerro de la Muerte, 11,000 ft., 11 July 1961, Brown C. A. CR-88 (US). LIMON: SW foot of Cerro Kamuk, 9°16'N, 83°02'W, 3,200-3,350 m, Davidse et al. 25978 (MO). HEREDIA: Buenavista Massive, 10,900 ft., 33 mi. NW of San Isidro del General, 31 July 1964, Woodruff s.n. (US). Panama. CHIRIQUI: Volcan de Chiriqui, Loma Larga to summit, 2,500-3,380 m, 4-6 July 1938, Woodson et al. 1079 (GH, MO, US), bocas DEL TORO: between Itamut and Bine peaks, Fabrega Massif, 3,200 m, 5-9 Mar. 1984, Gomez et al. 22259 (AAU, UC). Huperzia curvifolia (Kunze) Holub var. parvi- folia (Nessel) B. 011gaanl, comb. nov. Basio- nym: Urostachys verticillatus (L. f.) Herter var. parvifolius Nessel, Revista Sudamer. Bot. 6: 163, f. 43. 1940. TYPE: Costa Rica. Volcan Barba, Brade & Brade 283 (I IB not seen, BONN-Herb. Nessel no. 247). Lycopodium verticillatum L. f. var. parvifolium (Nessel) Lellinger, Proc. Biol. Soc. Wash. 89: 719. 1977. The material treated under ibis name is varia ble and referred here tentatively. Most specimens are somewhat intermediate between Huperzia acerosa and //. curvifolia var. curvifolia (Andes). Some specimens approach slender forms of //. filiformis. In general, the distinction of species in this group is problematic. Huperzia curvifolia var. curvifolia is an extremely thin form, with almost capillary stems, strongly upward curved leaves in proximal divisions, and closely appressed and reduced non- sporangiate leaves in the narrow, distal divisions. Variety parvifolia has thicker stems and is larger, with slightly less closely appressed nonsporangiate leaves in distal divisions, but shares the upward curved leaves in proximal divisions with the type variety. The epithet parvifolia is unfortunate in the pres¬ ent context because it does not describe the relation to the type variety, but it is the only legitimate choice. Acknowledgments. I thank Kirsten Tind for preparation of Figure 1, and Anni Sloth lor Figures 2 and 3. Literature Cited Herter, W. 1958. Some critical and new Central Amer¬ ican species of Urostachys. Amer. Fern J. 48: 81 84. Lellinger, D. B. 1989. The Ferns and Fern-allies of Costa Rica, Panama, and the Choco (Part 1: Psilo- taceae through Dicksoniaceae). Pteridologia 2 A: 1- 364. 011gaard, B. 1987. A revised classification of the Ly- copodiaceae s. lat. Opera Bot. 92: 153-178. - . 1989. Index of the Lycopodiaceae. Biol. Skr. Dan. Vid. Selsk. 34: 1-135. Rolleri, C. 1977a. The correlation of morphology and geographical distribution in Lycopodium saururus. Amer. Fern J. 67: 109-120. 72 Novon - . 1977b. Estudios inorfologicos y sistematicos en la seccion Crassistachys Herter del genero Ly¬ copodium L.: Lycopodium crassum H. B. ex Willd- enow y Lycopodium saururus Lam. Obra Centenario Mus. La Plata 3: 97-110. - . 1981. Sinopsis de las especies de Lycopodium L. (Lycopodiaceae Pteridophyta) de la seccion Cras¬ sistachys Herter. Revista Mus. La Plata (n. s.) Bot. 13 (71): 61-113. The Identification of Hieracium kaimii (Asteraceae) James L. Reveal Department of Botany, University ot Maryland, College Park, Maryland 20742-5815, U.S.A. ABSTRACT. The types of Hieracium kaimii and II. canadense are taken to represent two extremes within a species complex. A new combination, II. kaimii var. canadense, is proposed. Hieracium kaimii L. (Asteraceae) was established by Linnaeus (1753: 804) based on a Pehr Kahn collection ( 954.43 , holotype, LINN) obtained in Pennsylvania. The unusual fruit prompted Monnier (1829: 81) to place the species in its own genus, Sclerolepis, which proved to be a later homonym of Sclerolepis Cassini, and was renamed Pachylepis by Lessing (1832: 139). Gray examined the Kalin sheet and suggested (Torrey & Cray, 1843: 479) that the plant was probably not native to North America. Gray (1884: 424) later concluded that //. kaimii was not a species of Hieracium. Zahn (1923: 1 563), the last monographer ol Hieracium, retained H. kaimii in Pachylepis without comment. Fernald (1943) could not satisfactorily dispose of //. kaimii, and while he felt it was similar to II. canadense Michaux (1803: 86), and especially the southern variety fasciculatum (Pursh) Fernald, he regarded the Kalin sheet as representing an introduced exotic. Since 1943, H. kaimii has been essentially excluded by American authors from regional floras except for Lepage (1960), who took up II. kaimii for H. can¬ adense in part. Lepage (1960) considered the species represented by the Kalin specimen to be a plant of southeastern Canada and New England, and to be most closely related to H. canadense and H. scabriusculum Schweinitz. His conclusion was not accepted by Glea¬ son & Cronquist (1962) nor by Scoggan (1979), and the name has not been taken up by subsequent authors. My own examination of the Kahn sheet initially suggested that the plant belonged to the genus Cre- pis, specifically nearest the European C. vesica ri a L. subsp. taraxacifolia (Thuillier) Thellung. The short, plumpish achenes were unlike any I found in herbarium material of this now widespread and weedy subspecies found in Europe, Africa, and North America. Like Gray, I could not match the achenes with any species of Hieracium. When the late Arthur Cronquist (NY) visited I'he Natural History Museum (BM) in August 1991, I outlined the problem and produced the Kalin sheet for his examination. He concluded that the specimen was indeed a member of Hieracium, and the achenes were merely immature. We concluded that the Kahn specimen represented what Cronquist (Gleason & Cronquist, 1991) was (in August 1991) about to name II. kaimii. The variation within Hieracium kaimii is com¬ plex and no doubt subject to many interpretations. Lepage (1960) recognized both II. canadense and H. kaimii at the species rank, and recognized va¬ rieties in each. He associated the Kalm specimen with the smaller, less branched, more northern form of the species with an open capitulescence. The larger, more southern expression with an umbelli- forin capitulescence he named II. kaimii var. fas- ciculatum (Pursh) Lepage. My evaluation of the Kalm sheet, like Fernald s (1943), is that it is a depauperate form of variety fasciculatum. Linnaeus stated that the Kalm spec¬ imen was found in Pennsylvania, and while one cannot discount the possibility that Kalm gathered it in John Bartram’s garden near Philadelphia, it probably came from the southern part of the species’ range. As Hieracium kaimii (1753) has priority over //. canadense (1803), the following combination is proposed: Hieracium kaimii L. var. canadense (Michaux) Reveal, comb. nov. Based on H. canadense Michaux, FI. Boreali-Amer. 2: 86. 1803 var. canadense established by II. canadense var. latifolium Torrey & A. Gray, FI. N. Amer. 2: 476. 1843. The variety kaimii is the taller (5-15 dm) and more robust expression of the species with numerous (25-50) firm leaves that seldom have long hairs on the lower surface, and umbelliform capitulescences with short, stout peduncles 2-4 cm long. The variety ranges from Quebec to Minnesota south to New Jersey, Ohio, Illinois, and Missouri. The variety can¬ adense is a shorter (1.5-10 dm) and more slender expression with few (5-30) thinner leaves typically Novon 3: 73-74. 1993. 74 Novon with long hairs on the lower surfaces, and more open capitulescences with longer and more lax peduncles 2-10 cm long. This variety occurs from Labrador and New England to Manitoba and Montana, and is disjunct lrom British Columbia to Oregon. Acknowledgments. I thank Fred R. Barrie for comments, the late Arthur Cronquist for review of the Kalm sheet, and John L. Strother for helpful review. Work on the typification of temperate east¬ ern North American plants is supported by National Science Foundation Grant BSR-8812816 to the Missouri Botanical Garden. This is Scientific Article A-6271, Contribution No. 8440, of the Maryland Agricultural Experiment Station and Cooperative Extension Service. Literature Cited Cronquist, A. 1980. Asteraceae. Vase. FI. SE U.S. 1: 1-261. Chapel Hill. Fernald, M. L. 1943. VIII. Notes on Hieracium. Rho- dora 45: 317-325. Gleason, H. & A. Cronquist. 1963. Manual of Vascular Plants of Northeastern United States and Adjacent Canada. D. van Nostrand, New York. - & - . 1991. Manual of Vascular Plants of Northeastern United States and Adjacent Canada, 2nd ed. New York Botanical Garden, New York. Gray, A. 1884. Hieracium. Syn. FI. N. Amer. 1(2): 424-432. Lepage, E. 1960. Hieracium canadense Michx. et ses alliees en Amerique du Nord. Naturaliste Canad. 87: 59-84, 85-107. Lessing, C. F. 1832. Synopsis Generum Compositarum. Black, Young & Young, London. Linnaeus, C. 1753. Species Plantarum. L. Salvius, Stockholm. Michaux, A. 1803. Flora Boreali-Americana, Vol. 2. C. Craplet, Paris. Monnier, A. 1829. Essai Monographique sur les Hier¬ acium. C.-J. Hissette, Nancy. Scoggan, H. J. 1979. Compositae. Natl. Mus. Natural Sci. Publ. Bot. 7(4): 1442- 1626. Torrey, J. & A. Gray. 1843. Hieracium. FI. N. Amer. 2: 474-480. Zahn, K. H. 1923. Hieracium. In: Engler, Das Pflan- zenreich IV(280), 82: 1532 1574. New Species of Ericentrodea from Bolivia and Colombia (Asteraceae, Coreopsidinae, Heliantheae) Harold Robinson Department of Botany, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. ABSTRACT. Ericentrodea davulsmithii from Boliv¬ ia and E. ramirezii from Colombia are described. A key is provided for the six known species of the Andean genus Ericentrodea. The genus Ericentrodea Blake & Sherff (Blake, 1923) was established for a group of Coreopsidinae in Ecuador and Colombia that had strongly obcom- pressed achenes with contracted apices, 6 15 pap¬ pus bristles mostly in two groups over the angles of the achenes, and a marginal wing bearing a dense fringe of spreading setulae. A species with more highly dissected leaves later described from northern Peru, E. decomposita Blake & Sherff (Sherff, 1931), was the southernmost known record for the genus for over 60 years. Now, material collected in 1989 by the late David Smith in Bolivia is described, representing a considerable southeastward extension of the known generic range. The Bolivian species is similar to the Peruvian species in its highly dissected leaves. The species is named here after the collector. A second new species is also described from the northern part of the generic range in Colombia. The Colombian species is coauthored by Santiago Diaz- Piedrahita of the Universidad Nacional de Colombia, who brought the material for study, and is named for the collector B. B. Ramirez P. of the Herbario Universidad de Narino. Ericentrodea davidsinithii II. Robinson, sp. nov. TYPE: Bolivia. Santa Cruz: Provincia Manuel Maria Caballero, 50 km al norte de Mataral (por la carretera Santa Cruz-Comarapa) pas- arido por San Juan del Potrero y hajando a la cuenca del alto Rio Ichilo, 2,000 2,100 m, 26 May 1989, Smith, Quintana & Garcia 13419 (holotype, US; isotype, MO), f igure 1A, D, E. Ad E. decompositam simila sed in caulibus glabris in capitulis subsolitariis et majoribus in bracteis involucri interiores ad 9 mm longis et 3.5 mm latis in acheniis superne in collo prologatis et breviter bilobatis et in setis pappi majoribus utrinque ca. 8 differt. Flexuous vines with internodes ca. 8-9 cm long, stems essentially terete, multistriated with 6 stronger striae, glabrous, narrowly fistulose. Leaves opposite. petioles 4. 5-6. 5 cm long, petioles and petiolules glabrous; blade (Fig. IE) broadly triangular, mostly 10-15 cm long and wide, tri- to quadri-ternate, 30- 40 ultimate pinnae deeply lohed to pinnatifid, gen¬ erally ovate, 15-30 mm long, 12-17 mm wide, apices obtuse to shortly acute and minutely mucro- nate, surfaces glabrous with slightly prominulous veinlets. Inflorescence of ca. 3 heads on small lateral branches, holotype specimen with only 1 head fully developed at end ol branch; peduncle ca. 3.5 cm long, glabrous. Head (Fig. 1A) homogamous, ca. 13 mm high, when mature with spreading bracts ca. 20 mm wide; outer involucral bracts herbaceous, linear, 7-8 mm long, ca. 1.5 mm wide; inner in¬ volucral bracts reddish yellow, oblong, 8 9 mm long, 2. 5-3. 5 mm wide, apex shortly acute, outer surface glabrous, with numerous longitudinal reddish lines; pales flat, similar to inner involucre in size, shape, texture, and lack of pubescence. Rays lacking. Flo¬ rets 20-25, all bisexual; corolla yellow, ca. 8 mm long, tube ca. 2.5 mm long, glabrous, throat abruptly and narrowly campanulate, ca. 5 mm long, glabrous, lobes triangular, ca. 0.4 mm long and 0.3 mm wide, with scattered, narrow, monoseriate, multicellular hairs outside; anther thecae and appendages black, thecae ca. 3.5 mm long, appendages ovate, ca. 0.8 mm long, with median reddish duct reaching near tip; style branches flat, with narrow median groove and median reddish duct. Achenes (Fig. ID) im¬ mature, strongly obcornpressed, ca. 5.5 mm long, with narrow marginal wing, glabrous inside and out¬ side, densely fringed by biseriate- and triseriate- celled setulae along margins, constricted above into a glabrous, broad neck ca. 1 mm long, apex shal¬ lowly but distinctly bilobed; pappus with 6-8 stout, retrorsely barbed bristles on each lobe, 1 .5-2.0 mm long, with single smaller, smooth setulae at inner ends of each cluster. Pollen grains ca. 28 gm diam. Ericentrodea davidsmithii is known only from the type. The habitat given for the collection is “bosque primario, bosque pluvial montano; laderas con abundante Prumnopitys y pastizales anthro- pogenicos." The new species appears closest in relationship to the geographically closest Ericentrodea decorn- Novon 3: 75-78. 1993. 76 Novon Figure 1. Ericentrodea. A, D, E. E. davidsmithii H. Robinson. B, C, F. E. decomposita Blake & Sherff. — A, B. Heads. — C, D. Achenes. — E, F. Portions of leaves. Volume 3, Number 1 1993 Robinson Ericentrodea 77 posita of northern Peru. The latter is the only other member of the genus with as highly dissected, tri- to quadri-ternate leaf blades, having 20-40 leaflets. The four northern species in Ecuador and Colombia also have divided leaves, but the blades are only ternate or biternate. The most dissected leaves of E. mirabilis (Sherff) Blake & Sherff and the follow¬ ing new species, E. ramirezii, have 15-20 leaflets at most. Ericentrodea davidsmithii differs from E. decomposita by its glabrous stems, petioles, petio- lules, and peduncles, more ovate leaflets, nearly solitary and much larger heads with broader and larger inner involucral bracts, achenes with a prom¬ inent neck, slight apical sinus, and its ca. 8 retrorsely barbed bristles in each of the two clusters. In con¬ trast, E. decomposita has puberulous stems, peti¬ oles, and peduncles, rather elliptical leaflets, more numerous and smaller heads 7-8 mm high, achenes without a glabrous neck or bilobed tip, only 3 or 4 retrorsely barbed bristles in each of the clusters, and with ca. 2 smaller setulae on the inner ends of the clusters. Each species is known from a single collection, and achenes of both are somewhat im¬ mature, but the degree of immaturity seems nearly the same, and the differences in achene apices are regarded as reliable species differences. On the basis of the limited material, the stem of the new species also seems more terete. The stems of the holotype (F) and isotype (US) of E. decomposita examined are strongly hexagonal. Illustrations are provided for a head, an achene, and a portion of a leaf for the new species and for the previously unillustrated E. decomposita. Ericentrodea ramirezii H. Robinson & S. Diaz- Piedrahita, sp. nov. TYPE: Colombia. Narino: Mpio. de Pasto, 2-3 km E de la poblacion de Dolores, 2,900-3,000 m, 8 Feb. 1992, li. H. Ramirez P. & J. A. Cuayal M 4584 (holotype, COL; isotype, US). In foliis biternatis in foliolis 15-20 apice rotundatis vel breviter obtusis in capitulis homogamis et in setis pappi erectis solum in fasciculis binis dispositis a congeneribus differt. Vines with internodes 3. 5-5.0 cm long, stems weakly hexagonal, sparsely puberulous, not fistulose. Leaves opposite, petioles 2.5 3.0 cm long, petioles and petiolules sparsely puberulous; blade broadly triangular, up to 8 cm long, 8-10 cm wide, biternate or with basal triternate lobules, ca. 20 broadly ovate leaflets mostly 15-25 mm long, 9-17 mm wide, apices rounded to shortly obtuse, margins crenate- serrate, upper surface minutely puberulous on veins, lower surface sparsely pilosulous, secondary veins pinnate, ascending, slightly prominulous. Inflores¬ cences on short lateral branches bearing reduced bi - 1 r i - foliol a te leaves, pyramidally thyrsoid; pedun¬ cles 7-15 mm long, glabrous. Heads homogamous, ca. 8 mm high, submature 7-8 mm wide; outer involucral bracts herbaceous, narrowly oblong, 2.5- 4.0 mm long, ca. 0.8 mm wide; inner involucral bracts reddish yellow, oblong, ca. 6 mm long, 1.5- 2.0 mm wide, apex shortly acute, outer surface glabrous, with numerous longitudinal reddish lines; pales flat, similar to inner involucre in size, shape, texture, and lack of pubescence. Rays lacking. Flo¬ rets 20-25, all bisexual; corolla yellow, submature, ca. 5 mm long, mostly glabrous, basal tube ca. 1 mm long, throat narrowly campanulate, ca. 3 mm long, lobes triangular, 1 mm long, ca. 0.5 mm wide, pilosulous with a few monoseriate multicellular hairs outside; anther thecae and appendages black, thecae ca. 2 mm long, appendages ovate, ca. 0.5 mm long; style branches flat, with depressed median groove inside, shortly apiculate. Achenes strongly obcom- pressed, immature, ca. 1 mm long, glabrous inside and outside, densely fringed by setulae along mar¬ gins, not obviously constricted above; pappus of two widely separated clusters of 4-5 retrorsely barbed bristles, 1 .5-2.5 mm long, each group on short raised lip of apical callus. Pollen grains ca. 30 gm diam. Ericentrodea ramirezii is known only from the type. Detailed habitat data is lacking, but the cited elevation of 2,900-3,000 m is common for the genus in the northern Andes. The area around Pasto is associated with the eastern Cordillera in Narino. Ericentrodea ramirezii is closest to E. homo¬ gama and E. mirabilis, but it differs from both by the rounded or shortly obtuse tips on the leaflets. The species differs further from E. mirabilis by the lack of extra pappus bristles between the two pri¬ mary clusters. The achenes are too immature to see whether a lateral wing will develop or whether the pappus will ultimately be raised on a prominent lobe and be directed laterally as in E. homogama. In spite of the lack of certainty about the achene, the new species can be distinguished from E. homogama vegetatively by having leaves with ca. 20 blunt- tipped leaflets rather than having less dissected leaves with about nine pointed leaflets in three sets of three. The six known species of Ericentrodea can be distinguished by the following key: Key to the Species of Ericentrodea la. Leaf blades tri- to quadri-ternate, highly dis¬ sected into 20-40 leaflets; species of Peru and Bolivia. 78 Novon 2a. Stems hexagonal; stems, petioles, and pe¬ duncles puberulous; inflorescences with 5- 10 heads; heads 7-8 mm high; inner in- volucral bracts and pales 5-6 mm long and to 2 mm wide; achenes constricted but without a short glabrous neck above, not apically bilobed; pappus with 3-4 retrorse- ly barbed bristles in each cluster; northern Peru . E. decomposita Blake & Sherff 2b. Stems terete; stems, petioles, and pedun¬ cles glabrous; inflorescences with 1-3 heads, often only 1 developed; heads 12- 14 mm high; inner involucral bracts and pales to 9 mm long and 3.5 mm wide; achenes with a distinct glabrous neck at upper end, slightly bilobed apically; pappus with 6 8 retrorsely barbed bristles in each cluster; central Bolivia . . E. davidsmithii H. Robinson lb. Leaf blades undivided to ternate or biternate, rarely with more than 12 leaflets; species of Colombia and Ecuador. 3a. Heads with rays; in fruit ca. 9 mm high and to 1 7 mm wide; upper leaves often simple; Colombia and Ecuador . . E. corazonensis (Hieronymus) Blake & Sherff 3b. Heads without rays, homogamous; in fruit 6-8 mm high and 10 mm wide; upper leaves rarely simple. 4a. Achenes with pappus erect, often with 2-3 shorter bristles on each side be¬ tween the main clusters; heads with ca. 12 florets; peduncles ca. 1 cm long; Ecuador . . ... E. mirabilis (Sherff) Blake & Sherff 4b. Achenes with pappus separated at apex into two distinct, often laterally spreading clusters; heads with 20 or more florets; peduncles up to 3 or more cm long. 5a. Leaves biternate with usually nine leaflets; leaflets with apices acute; Ecuador . . E. homogama (Hieronymus) Blake & Sherff 5b. Leaves biternate with 15-20 leaf¬ lets; leaflets with apices rounded or shortly obtuse; Colombia . . . . E. ramirezii H. Robinson & Diaz-Piedrahita Acknowledgments. The Field Museum of Natural History, Chicago, Illinois, is thanked for the loan of the holotype of Ericentrodea decomposita. The il¬ lustration was prepared by Alice Tangerini of the Department of Botany, Smithsonian Institution. Literature Cited Blake, S. F. 1923. Two new genera related to I\lar- valina. J. Wash. Acad. Sci. 13: 102-105. Sherff, E. E. 1931. New or otherwise noteworthy Com- positae. VII. Bot. Gaz. (Crawfordsville) 92: 202-209. Reappraisal of Subtribe Vargasiellinae (Maxillarieae, Orchidaceae) Gustavo A. Romero Oakes Ames Orchid Herbarium, Harvard University Herbaria, 22 Divinity Avenue, Cambridge, Massachusetts 02138, U.S.A. German Carnevali Missouri Botanical Garden, B.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. Abstract. Subtribe Vargasiellinae, proposed by C. Schweinfurth in 1952, is here validated. It is placed in tribe Maxillarieae, rather than in the Malaxideae, as originally proposed by Schweinfurth, based on new findings on the pollinarium structure of l a r- gasiella peruviana C. Schweinfurth. Schweinfurth described I argasiella in 1952 from two Peruvian specimens collected by C. Vargas. The morphology of the inflorescence and pollinarium puzzled the author; the absence of complete polli- naria in the specimens he examined led him to believe l argasiella was close to subtribe Liparideae (in Schlechter's old system, referred to tribe Ma¬ laxideae Lindley in Dressier, 1990: 217). However, based on leaf and flower characters, Schweinfurth (1952: 152) suggested subtribal status for Varga- siella, but failed to validly publish the taxon. Schweinfurth (1958: 219) described a second species from the Venezuelan Guayana; the flowers he had available did not have complete pollinaria. Vargasiella venezuelana was collected a second time by G. C. K. Dunsterville (Dunsterville & Garay, 1965: 326; Dunsterville & Dunsterville, 1982: 37), but this material and his subsequently published drawing (Dunsterville & Garay, 1965: 326-327; 1979: 1037) did not present critical details of the pollinarium. Dressier (1990: 249) suggested the placement of Vargasiella in subtribe Zygopetalinae: “The An¬ dean Vargasiella has been described as having na¬ ked pollinia, but the pollinia appear to be superposed, and I suspect that it is closer to this group” (i.e., subtribe Zygopetalinae). However, examination of recently collected specimens of Vargasiella peru¬ viana indicate the pollinarium has four plano-convex pollinia, a short stipe, and a hooked viscidium (Fig. 1). The pollinia do not appear to be superposed and flattened as in Zygopetalum and other allied genera. These features, combined with the fact that the labellum does not have any ridges or keels, a di¬ agnostic feature of subtribe Zygopetalinae, indicate that Vargasiella, although closely allied to the Zygo¬ petalinae, merits subtribal status. Vargasiellinae C. Schweinfurth ex G. Romero & Carnevali, subtrib. nov. TYPE: Vargasiella C. Schweinfurth, Hot. Mus. Leafl. 15: 150. 1952. Herba epiphytica vel terrestris; caulis elongatus, sine pseudobulbis; folia disticha, plicata, articulata; inflores- centia laxe racemosa; labellum simplex, ecallosum; pol¬ linarium viscidio unico, stipite unico, polliniis quatro, plano- convexis, non-superimpositis. The authors would like to request live and/ or pickled specimens of Vargasiella from orchid col¬ lectors and growers from Peru and Venezuela. This material is urgently needed to undertake further studies, particularly of cytology, growth habits, and the development of the inflorescence. We hope the examination of additional specimens will shed some light on the placement of this interesting genus within the Maxillarieae. Specimens examined. Vargasiella peruviana C. Schweinfurth. Peru, cuzco: Prov. Paucartambo, Pilla- huata, 3,400 m, epiphyte in rainforest, floral segments white lined with pink, 1 1 Dec. 1942, C. I argas 3010 (AMES); Convencion Prov., hills of Pintobamba, in humus forest, 2,400 m, perianth white with pinkish lip, 3-4 Mar. 1943, C. Vargas 3288 (holotype, AMES), pasco: Oxa- pampa Prov., Cordillera Yanachaga, road over shoulder of Cerro Pajonal to Villa Rica drainage, 12 km SE of Oxapampa, cloud forest, 2,300-2,500 m, 10°35'S, 75°20'W, “Terrestrial 1 m; stem zig-zag; petioles grooved; bracts green with purple rim; two lower sepals purple outside, whitish inside; upper sepal white with purplish stripe down middle; petals white, lip and undulate margin maroon tongue; stem purple,” 9 Oct. 1982, K. 11. Foster & D. Smith 9 094 (MO); same locality, 14 km E of main Oxapampa-Villa Rica road, lumber road at top of cor¬ dillera; stunted cloud forest, 2,450 m, 10°35'S, 75°15'W, terrestrial, flower white, maroon outside and on lip, 2 Mar. 1982, A. Gentry & D. Smith 35984 (MO); border Provincias Oxapampa and Pasco, San Cotardo, in dwarf forest, rich in Ericaceae, bamboo and Blechnum, with sphagnum layer below, in swampy places, 2,700 m, ter¬ restrial, flowers white with red, 7 Mar. 1986, //. van der Her l fetal . 8508 (MO). Novon 3: 79-80. 1993. 80 Novon Figure 1. Vargasiella peruviana C. Schweinfurth (Foster & Smith 9094). — A, B. Column. — C-E. Anther. F. Pollinarium. (Scale bars equal 2 mm.) Vargasiella venezuelana C. Schweinfurth. VENEZUELA. BOLIVAR: Chimanta Massif, NW part of summit of Aba- capa-tepui, Bonnetia forest, 2,125-2,300 m, terrestrial, inflorescence erect, all parts of flower rich purple, 3 sepals projecting backwards, 2 lateral petals erect, bracts green, peduncle dull lavender, leaves firmly membranous, deep green above, dull green below, 13 Apr. 1953, J. A. Steyermark 74914 (holotype, AMES; isotype, AMES); Auyan-tepui, above El Pehon, growing almost completely hidden in low thick brush and grass on steep slope, ele¬ vation ca. 6,500 ft., G. C. K. & E. Dunsterville 861 (line drawing, AMES). Acknowledgments. We thank Cirilo Nelson for comments on the Latin description and Bruno Man- ara lor the line drawings. Literature Cited Dressier, R. L. 1990. The Orchids. Harvard Univ. Press, Cambridge, Massachusetts. Dunsterville, G. C. K. & L. A. Garay. 1965. Venezuelan Orchids Illustrated III. Andre Deutsch, London. - & - . 1979. Venezuelan Orchids, an Illustrated Field Guide. Harvard Univ. Press, Cam¬ bridge, Massachusetts. - & E. Dunsterville. 1982. Auyan-tepui: Rem¬ iniscences of an orchid search. Pp. 19-38 in J. Arditti (editor). Orchid Biology and Perspectives II. Cornell Univ. Press, Ithaca, New York. Schweinfurth, C. 1952. Vargasiella. Bot. Mus. Leaf!. 15: 150-152. - . 1958. Vargasiella venezuelana. Bot. Mus. Lead. 18: 219-223. New Species of Hedyosmum (Chloranthaceae) from Northern South America Carol A. Todzia Plant Resources Center, Department of Botany, University of Texas, Austin, Texas 78713, U.S.A. ABSTRACT. Four new species of Hedyosmum, all belonging to subgenus Tnfalla sect. Microcarpa, are herein proposed. Hedyosmum uniflorum occurs in mid-elevation montane regions in central Ecuador and differs from its closest relative, //. bonplan- dianum, by its basally rounded, sometimes oblique leaves, flocculose midveins, slightly fimbriate stipular appendages, and long, linear, dissected stigmas. Hedyosmum narinoense, which inhabits the western slopes of the western cordillera in southern Colom¬ bia, also has single- flowered cymules but is unique among the solitary-flowered cymule species by hav¬ ing shorter pistillate inflorescences and scabrous, scurfy leaf sheaths. The remaining two new species, II. tepuiense and II. intermedium, are found in the Venezuelan Guayana. Hedyosmum tepuiense is rec¬ ognized as distinct from the broadly ranging II. racemosum by virtue of its long stipular appendages, verrucose leaf sheaths, floccose mid veins, and longer inflorescences. Morphologically intermediate be¬ tween H. gentryi and II. neblinae, Hedyosmum intermedium is distinguished by its glabrous, elliptic leaves and 2-3-flowered purple- fruited cymules. Recent collections from South America have clar¬ ified the status of several populations of Hedyosmum that were previously incorporated into the variation of broadly ranging species (Todzia, 1988). All of these newly described species fall into subgenus Taf- alla sect. Microcarpa, the largest section in Hedy¬ osmum, now with 28 species. This section is rec¬ ognized by racemose or paniculate pistillate inflorescences, cymules of 1 -8 flowers, fleshy floral bracts that are fused in fruit, and white, rarely purple, fruiting cymules. Hedyosmum uniflorum Todzia, sp. nov. TYPE: Ecuador. Pichincha: old road Quito-Santo Do¬ mingo, 12-15 km NE of old road from junction with new road near Alluriquin, 1,430-1,540 m, ca. 0°20'S, 78°55'W, 3 Feb. 1982, Luteyn, Pipoly, Lebron- Luteyn & Kallunki 8754 (ho- lotype, TEX; isotypes, NY, QCA not seen). Figure 1 . Hedyosmo bonplandiano foliis laevibus et cymulis flore singulo instructis similis sed differt appendicibus fimbriatis stipularum, foliis ad basim rotundatis aliquando obliqu- isque trichomatibus floccosis in nervi centrali paginis in- feris, et stiginatibus longis linearibusque. Dioecious (?), aromatic trees or shrubs 4-8 m tall; young stems quadrate; older stems terete, with tubular leaf bases persisting, ultimately disintegrat¬ ing and leaving circular scars; internodes 3-8 cm long. Leaves elliptic, 7.5-15.5 cm long, 3-6.5 cm broad, with long-acuminate tips 0.5-1 cm long, rounded, often oblique at base, at margins sharply serrate with teeth 3-4 mm distant, drying subcor- iaceous, smooth to slightly scabrous; midveins im¬ pressed above, raised beneath, with sparse floccose multibranched trichomes; larger lateral veins 6-9, 0.6- 1.8 cm distant, arcuate, raised beneath, gla¬ brous or occasionally with sparse strigose to floc¬ culose hairs; free portion of petioles 0.6- 1.4 cm long, glabrous except for occasional flocculose hairs running down midrib onto petiole; petiolar sheaths 1.3- 1.8 cm long, 0.5-1 cm broad at apex, inflated, slightly flared at apex, with two strigose lines ex¬ tending down from the free portion of petioles, per¬ sistent but disintegrating with age, each distal margin with two slightly fimbriate, stipular appendages 1 2 mm long, extending above free portion of petioles ca. 0.5 mm. Staminate inflorescences not seen. Pis¬ tillate inflorescences axillary or terminal, racemes or sparsely branched panicles, 3.5 8 cm long with 16-46 cymules; subtending bracts similar in size and shape to the leaves or more frequently smaller; cymules l(-3)-flowered, alternately or oppositely ar¬ ranged on the rachis, 3-10 mm distant, sessile or on short peduncles 1-3 mm long; subtending floral bracts enclosing one-fourth to three-quarters of flow¬ er, with acuminate tips 0.5-1 mm long, margin entire or with occasional strigose hairs. Pistillate flowers slightly trigonous, ca. 3 mm long, ca. 2 mm broad, with or without a small pore on each face of the ovary; perianth lobes small, deltoid; stigmas usu¬ ally highly dissected, linear, 2-3 mm long, papillose. Fruiting cymules green (?). Distribution, habitat, and phenology. Known Novon 3: 81-85. 1993. 82 Novon Figure 1 . Distribution of the new species of Hedyosmum. Each symbol represents the presence of that respective species in a degree square. only from central Ecuador (Prov. Pichincha) on the western slopes of the western cordillera, where it occurs between 1,300 and 2,200 m in disturbed cloud forest and montane wet forest. Flowering spec¬ imens have been collected in February, July, Sep¬ tember, and December. Hedyosmum uniflorum appears to be most close¬ ly allied to //. bonplandianum and //. racemosum. This new species is similar to //. bonplandianum in having usually single-flowered cymules and leaves smooth upon drying. It differs from //. bonplan¬ dianum in having long, linear stigmas (vs. stigmas shorter and more clavate), leaves rounded and some¬ times oblique at base (vs. attenuate), flocculose hairs on midvein beneath (vs. entirely glabrous), and stip- ular appendages slightly fimbriate (vs. usually line¬ ar). If its fruiting cymules are green at maturity as indicated by label data, this would also be a character at variance with H. bonplandianum. Fruiting cy¬ mules of all previously known species of subgenus Tafalla sect. Microcarpa are either white or purple. The southernmost populations of //. bonplandian¬ um occur in southern Colombia (Depto. Cauca). Although this population was originally placed into H. racemosum, it was noted that its solitary- flowered cymules and long (up to 3 mm), highly dissected stigmas were anomalous within the //. ra¬ cemosum species concept (Todzia, 1988: 75). Mor¬ phological characters shared with //. racemosum include slightly fimbriate stipular appendages and leaves smooth upon drying. As noted above, //. uniflorum differs from //. racemosum in having usually single-flowered cymules, highly dissected stigmas, as well as rounded often oblique leaf bases (vs. attenuate and not oblique) and fruiting cymules green not white. Paratypes. Ecuador. PICHINCHA: Carretera Quito- Aloag-Santo Domingo de los Colorados, km 94, a 10 km al S de la carretera, estribaciones occidentales del Volcan Corazon, 0°21'30"S, 78°51'15"W, 1,300-1,500 m, 24 Dec. 1986, Zak 1492 (MO, TEX); Reserva Floristica- Ecologica “Rio Gualjalito,” km 59 de la carretera antigua Quito-Santo Domingo de los Colorados, estribaciones oc¬ cidentales del Volcan Pichincha, 78o48'10"W, 0°13'53"S, 1,800-2,200 m, 9 July 1985, Jaramillo & Zak 7735 (MO, TEX); Reserva Floristica-Ecologica “Rio Gualjali¬ to,” km 59, carretera antigua Quito-Santo Domingo, Volume 3, Number 1 1993 Todzia Hedyosmum 83 faldas al oeste del Volcan Pichincha, 0°13'S, 78°48'W, 1,800-2,200 m, 24 Sep. 1988, Zak & Jaramillo 3827 (TEX). Hedyosmum narinoense Todzia, sp. nov. TYPE: Colombia. Narino: 2-8 km E of Junin on Tu- maco-Tuquerres road, 1°15'N, 78°09'W , ca. 1,100 m, 26 July 1986, Gentry et al. 55235 (holotype, MO; isotype, TEX — 2 sheets). Fig¬ ure 1 . Hedyosmo bonplandiano ac H. unifloro cymulis flore singulo instructis similis sed ab utroque differt inflores- centiis pistillatis brevioribus et vaginis foliorum furfuraceis scabris aetate provectis laciniatisque. Dioecious (?) trees ca. 8 m tall; young stems quadrate, scurfy; older stems terete, becoming gla¬ brous, with tubular leaf sheaths persisting and ul¬ timately leaving circular scars; internodes 2.5-6 cm long. Leaves narrowly elliptic to elliptic, 8.5-12 cm long, 2.5-5 cm broad, with acuminate tips 0.5-1 cm long, rounded to attenuate at base, at margins sharply serrate with teeth 2.5-5 mm distant, drying coriaceous, smooth above, scabrous beneath; mid¬ veins impressed above, raised beneath, with sparse, floccose, multibranched trichomes; larger lateral veins 6-7, 11-20 mm distant, arcuate, raised and gla¬ brous beneath; free portion of petioles 0.8-1 .1 cm long, scabrous, rough; petiolar sheaths 1 .6-2 cm long, 0.7- 1.3 cm broad at apex, slightly inflated to somewhat closely appressed to stern, scurfy, becom¬ ing laciniate at upper margin with age, with slightly raised longitudinal lines extending down from distal margin, stipular appendages absent or early cadu¬ cous. Staminate inflorescences not seen. Pistillate inflorescences axillary or terminal, racemes or sparsely branched panicles, 2. 5-5. 8 cm long with ca. 50 flowers, alternately arranged 1-8 mm distant on the rachis, subsessile or on short pedicles 1-3 mm long; lower inflorescence bracts similar to leaves but much smaller. Flowers usually solitary on rachis, rarely aggregated into clusters of 2, subtended by a fleshy cupular bract, on one side with an acuminate free tip ca. 1 mm long, enclosing lower half of flower. Pistillate flowers slightly trigonous, ca. 2 nun long, ca. 1 mm broad; perianth lobes united at base; stigmas clavate, dissected distally, ca. 2 mm long. Fruits white, ellipsoidal, 3-4 mm long, ca. 3 mm broad. Distribution , habitat, and phenology. Known only from southern Colombia (Prov. Narino) on the western slopes of the western cordillera where it occurs in pluvial forests and disturbed roadsides along cloud forest at elevations between 900 and 1 ,000 m. The three known collections were collected in flower and fruit in July. Hedyosmum narinoense appears to share dis¬ tinctive characters with several other species of He¬ dyosmum. It has the solitary flowers of II. bon- plandianum, which ranges from Nicaragua to Colombia, and //. uniflorum of central Ecuador, yet differs from both by its shorter pistillate inflores¬ cences, and scabrous and scurfy leaf sheaths that become laciniate with age. These laciniate leaf sheaths, along with its scabrous leaves, show a pos¬ sible relationship to //. scaberrimum, which also occurs in southern Colombia. It differs from the latter species by its flowers, which are solitary on the inflorescence rachis (vs. in cymules of 2-4 flow¬ ers). Thus it is on the following suite of characters that the above collections are described as new: flowers borne singly on the inflorescence axis, not clustered into cymules; scabrous leaves with sparsely dentate teeth 2.5-5 mm distant; and scurfy, sca¬ brous leaf sheaths becoming laciniate along the distal margin. Paratypes. Colombia. NAKifio: Junin-Barbacoas road, 2-10 km N of Junin, 1°30'N, 78°10'W. 900 1 ,000 m, 26 July 1986, Gentry et al. 55329 (MO, TEX). Hedyosmum tepuiense Todzia, sp. nov. TYPE: Venezuela. Amazonas: Depto. Rio Negro, Cerro de la Neblina, Expedition Camp VII, 0°50'N, 65°58'W, 1,850 m, 29 Nov. 1984, Anderson 13389 (holotype, TEX; isotypes, MICH not seen, NY not seen, VEN not seen). Figure 1 . Hedyosmo racemoso inflorescentibus racemosis pani- culatisque cymulis floribus aliquot instructis similis sed differt appendicibus stipularum maxime longis et vaginis foliorum verrucosis. Dioecious, aromatic trees or shrubs 4 10 m tall; young stems quadrate; older stems terete with tu¬ bular leaf bases persisting, ultimately disintegrating and leaving circular scars; internodes 3-6 cm long. Leaves elliptic, 7.5-19 cm long, 2.3-10 cm broad, with long-acuminate tips 0.5- 1 .5 cm long, attenuate at base, at margins coarsely serrate with teeth 3- 6 mm distant, drying subcoriaceous, scabrous; mid¬ veins impressed above, raised beneath, with floccose or strigose hairs; free portion of petioles 0.7-2 cm long, glabrous; petiolar sheaths 1.5-2. 5 cm long, 1-1.4 cm broad at the apex, inflated, slightly flared at apex, smooth to verrucose especially along distal margin, extending above free portion of petioles ca. 0.5 mm, persistent but disintegrating with age, each distal margin with two linear to slightly fimbriate stipular appendages ca. 3 mm long. Staminate in¬ florescences composed of a straight rachis 2. 5-6. 5 cm long, with 2 3 nodes; nodes usually alternate, sometimes opposite, with 1 spike per node; spikes 84 Novon sessile or on short peduncles 1 -4 mm long; mature spikes 0.6- 1.5 cm long, ca. 4 mm broad; stamens 50-100; anthers 1-1.5 mm long, connectives ex¬ tended into a pad ca. 0.5 mm long with an apicule at summit. Pistillate inflorescences axillary or ter¬ minal, racemes or sparsely branched panicles, 3- 10 cm long with ca. 20 cymules; cymules sessile or on short peduncles 1-5 mm long, oppositely or alternately arranged on rachis, 0.2- 1.4 cm distant, very irregularly globose, 4-6 mm diam., subtending floral bracts connate in lower one- to two-thirds, enclosing one- to three-fourths of flower, the margin usually entire, occasionally with a few sparse strigose hairs. Pistillate flowers slightly trigonous, 2 3 mm long, 1-2 mm broad, with a small pore on each face of the ovary; perianth lobes small, ca. 0.3 mm long, rounded at apex; stigmas irregularly clavate, 1-2 mm long, 3-angled, papillose. Fruiting cymules white, irregularly globose, 4 9 mm diam. Seeds ca. 3 mm long, ellipsoidal, brown, smooth. Distribution , habitat, and phenology. Occurs throughout a large region of the Venezuelan Gua- yana at elevations of 1,200-2,660 m in wet forest and riparian habitats. Flowering specimens have been collected in February, April, and November and fruiting specimens in March, October, November, and December. The sandstone plateaus or tepuis of the Vene¬ zuelan Guayana are not a region of great Hedy os- mum diversity as they are for some other groups of plants (Steyermark, 1979). Three species were pre¬ viously reported from that area: //. racemosum (Ruiz & Pavon) G. Don, //. gentryi D’Arcy & Liesner, and II. neblinae Todzia (Todzia, 1988). This new species, //. tepuiense, was previously treated as //. racemosum in Todzia (1988). Another new species described below, //. intermedium, was previously treated as the southernmost populations of II. gen¬ tryi. Considering these taxonomic reevaluations, the three species found in the Venzuelan Guayana are now called //. tepuiense, H. intermedium, and II. neblinae. Hedyosmum tepuiense is obviously closely allied to Hedyosmum racemosum, with both having race¬ mose and paniculate inflorescences with several- flowered cymules. This new species differs from //. racemosum by its extremely long stipular append¬ ages (4 mm long vs. 12 mm long for II. race¬ mosum) and verrucose leaf sheaths (vs. glabrous in II. racemosum). Hedyosmum tepuiense has con¬ sistently larger leaves than populations of II. ra¬ cemosum in the Andes, as well as floccose trichomes on the lower side of the midvein and longer inflo¬ rescences. Paratypes. Guyana. Upper Potaro River region, upper slopes of Mt. Wokomung, 05°05'N, 59°50'W, ca. 1,500 in, 4 July 1989, Boom <£~ Samuels 9063 (NY, TEX), 1,530 m, 13 July 1989, Boom ct Samuels 9200 (NY, TEX). Venezuela, amazonas: Depto. Rio Negro, Cerro de la Neblina, Expedition Camp VII, 00°50'N, 65°58'W, l, 850 m, 29 Nov. 1984, Anderson 13410 (TEX), 1 Dec. 1984, Anderson 1 3440 (TEX); Cerro Neblina, camp #7, S slopes of Canon Grande, 1,800 m, along river below camp, 0°55'N, 66°0'W, 29 Nov. 1984, Croat 59449 (MO, TEX), Croat 59463 (MO, TEX); trail S from Cerro Neblina Camp #5, 1,200 1,300 m, 0°49'N, 66°0'W, 12 Apr. 1984, Gentry & Stein 46570 (TEX); Depto. Rio Negro, E of and below Neblina Camp 7, 0°55'N, 66°0'W, S slopes Canyon Grande, ca. 1,700 m, 29 Nov. 1984, Krai 71905 (TEX); Depto. Atabapo, Cerro Marahuaca, summit SW side of center, sides and bottom of 120 m deep sinkhole, 3°39'N, 65°26'W, 2,600 m, 22 Oct. 1988, Liesner et al. 25171 (TEX); Depto. Rio Negro, Cerro de La Neblina, ridge at divide between Brazil and Venezuela, 26 km ENE of Neblina Base Camp, approx. 0°53'N, 65°56'W, 2,000 m, 15 Apr. 1984, Plowman & Thomas 13629 (NY, TEX); Depto. Rio Negro, Camp 7, 00°52'N, 65°58'W, 1,730-1,850 m, 1 Feb. 1985, Renner 2058 (US). BOLIVAR: Cerro Apacara, Rio Caroni, 1,850 m, 11 Nov. 1946, Cardona 1956 (NY, US); Cerro Roraima, ca. 5°12'N, 60°40'W, 2,280- 2,600 m, Luteyn & Aymard 9769 (TEX); Ilu-tepui, Gran Sabana, 7-8,000 ft., 17 Mar. 1952, Maguire 33470 (TEX); Mount Roraima, SW slopes, 2,400 m, 10 Jan. 1939, Pinkus 130 (BR, F, G, GH, MO, NY, S, US); Cerro Uananapan, S of Uei-tepui, between Luepa and Cerro Venamo, 1,450 in, 25 Apr. 1960, Steyermark & Nilsson 746 (NY, VEN); Chimanta Massif, SW edge of Apacara-tepui, 1,800-2,000 in, 14 Apr. 1953, Steyer¬ mark 74972 (F, NY); Auyan-tepui, entre la escarpa su¬ perior, este del paso de acceso a la cuinbre del sur y “El Penon,” 1,800 m, 17 May 1964, Steyermark 94083 (NY, VEN); Massif Chimanta, along Rio Asaporko, 1,300 m, 7 Jan. 1953, Wurdack 34033A (NY, US). Hedyosmum intermedium Todzia, sp. nov. TYPE: Venezuela. Amazonas: Depto. Atures, Sierra Maigualida, NW sector, small valley along an upper tributary of Cano Iguana, 5°30'N, 65°1 5'W, 2,000 m, 28 Feb.-3 Mar. 1991, Berry, Huber & Rosales 4913 (holotype, TEX; isotypes, MO, MYF not seen, VEN not seen). Figure 1 . Inter Hedyosmum neblinae ac //. gentryi interme¬ dium; H. neblinae similis dispositionibus florum ac cy- mularum sed foliis ellipticis et vaginis foliorum glabris differt; H. gentryi similis foliis ellipticis ac vaginis foliorum glabris sed cymulis floribus 2 3 instructis differt. Dioecious, aromatic trees or shrubs 1.5-8 m tall; young stems quadrate, sometimes dull red; older stems terete with tubular leaf bases persisting, ul¬ timately disintegrating and leaving circular scars; internodes 17 cm long. Leaves elliptic, 6-14 cm long, 2. 3-6. 4 cm wide with abruptly acuminate apices 0.5-1. 2 cm long, attenuate at base, at mar¬ gins coarsely serrate with teeth 5 8 mm distant. Volume 3, Number 1 1993 Todzia Hedyosmum 85 glabrous, drying subcoriaceous and smooth; midvein glabrous beneath; larger lateral veins 10-14, 7-10 mm distant, obscure, leaving midvein at ca. 90° angle, arcuate toward margin, free portion of peti¬ oles 0.4- 1.8 cm long, glabrous; petiolar sheaths 0.8-1 . 1 cm long, 0.6-0. 9 cm broad at apex, slightly flared at apex, glabrous, smooth; stipular append¬ ages linear, ca. 2 mm long, caducous. Staminate inflorescences 5.5 1 1 cm long, composed ol ra- chises with 1-4 nodes with 1-3 spikes per node; mature spikes 1 -2 cm long, subtended by 2-4 ovate bracts; stamens ca. 130-200, congested on axis, ca. 1.5 mm long, ca. 1 mm broad at apex, con¬ nective ca. 0.3 mm long, flattened distally. Pistillate inflorescences axillary or terminal, simple or basally branched racemes 3-4 cm long; cymules with 2-3 flowers, opposite on inflorescence axis, 4-10 mm distant, sessile; subtending floral bracts 2.5-4 mm long, free almost to base or connate in lower one- fourth, acute at apex, margin ciliate. Pistillate flow¬ ers strongly trigonous, 3-4 mm long, 1-2 mm broad on each face, with a large pore on each face of the ovary; perianth lobes minute, deltoid, rounded at apex. Fruiting cymules red to purple; seeds ca. 4 mm long, strongly trigonous, minutely papillose. Distribution, habitat, and phenology. Wide¬ spread in the Venezuelan Guayana, occurring in semi-open and medium-height forest, savannas, and stream and river edges between 1,200 and 2,000 m. Flowering specimens have been collected in Feb¬ ruary, March, and October, while fruiting is reported in March and October. Numerous recent expeditions in the Venezuelan Guayana have provided sufficient material to assess these populations that were originally placed in He¬ dyosmum gentryi (Todzia, 1988). Along with H. gentryi, H. neblinae, and H. pseudoandromeda, H. intermedium is one of the few purple-fruited species in subgenus Tafalla sect. Microcarpa. He¬ dyosmum intermedium is morphologically inter¬ mediate between Hedyosmum gentryi, which rang¬ es from eastern Panama (Darien) and northern Colombia (Antioquia) to northern Venezuela (Ara- gua, Falcon, Merida, Miranda, Monagas, Portu- guesa, and Yaracuy), and //. neblinae (known only from Cerro Neblina, which is further south than the localities presently known for //. intermedium). Hedyosmum intermedium has a flower and cymule arrangement similar to //. neblinae, but differs from it by glabrous (vs. verrucose) leaf sheaths and elliptic (vs. ovate) leaves. It shares with H. gentryi elliptic leaves and glabrous leaf sheaths, but differs in having 2-3 flowers per cymule (vs. 1 flower per cymule). Paratypes. Venezuela. AMAZONAS: Serrania Paru, Cano Asisa, Rio Ventuari, 2,000 m, 7 Feb. 1951, Cowan & Wurdack 31336 (NY); Depto. Atabapo, slope of Cerro Marahuaca, Rio Yameduaka arriba, 3°38’N, 65°28'\\ , 1,225 m, 19 Feb. 1985, Liesner 17665 A (MO. TEX, VEN); Depto. Atabapo, below Salto Los Monos on trib¬ utary of headwaters ot Rio Iguapo, 3°35'N, 65°23'W, 1,500-1,650 m, 12 Mar. 1985, Liesner 18582 (MO, TEX, VEN); Depto. Rio Negro, Cerro Aracamuni, sum¬ mit, Proa camp, 1°32'N 65°49'W, 1,400 in, 27 Oct. 1987, Liesner Carnevali 22517 (MO, TEX, VEN), Liesner & Carneva l, 22535 (MO, TEX, VEN); Depto. Atabapo, Cerro Marahuaca, slopes, “Sima” area, 03°43'N 65°30'W, 1,200 m, 16 Oct. 1988, Liesner 24961 (MO, TEX, VEN), 18 Oct. 1988, Liesner 25068 (MO, TEX, VEN); Depto. Rio Negro, Cerro Aracamuni, summit, Proa camp, 1°32'N 65°49'W, 1 ,400 m, 25 Oct. 1 987, Liesner & Carnevali 22438 (MO, TEX, VEN), 26 Oct. 1987, Liesner & Carnevali 22472 (MO, TEX, VEN). bolivar: Meseta del Jaua, Cerro Jaua, 1,820-1,880 m, 28 Feb. 5 Mar. 1974, Steyermark et al. 109677 (A, K, NY); Distr. Cedefio, Meseta de Jaua, sector centro meridional, cabeceras del Rio Marajano, afluente del Rio Cacaro, 04°48'N, 64°32'W, 1,750-1,800 m, 20 Nov. 1989, Huber 13048 (TEX). Acknowledgments. I thank the curators at MO, NY, US, and VEN for sending Hedyosmum material for identification. 1 also thank Guy Nesom for the Latin diagnoses and Paul Berry for review ol the manuscript. Literature Cited Steyermark, J. A. 1979. Plant refuge and dispersal centres in Venezuela: Their relict and endemic ele¬ ment. Pp. 185-221 in K. Larsen & L. B. Holm- Nielsen (editors), Tropical Botany. Academic Press, London. Todzia, C. A. 1988. Chloranthaceae: Hedyosmum. FI. Neotropica Monogr. 48: 1 139. Jaimehintonia (Amaryllidaceae: Allieae), a New Germs from Northeastern Mexico H. L. Turner Department of Botany, University of Texas, Austin, Texas 78713, U.S.A. ABSTRACT. A new genus and species, Jaimehin¬ tonia gypsophila, from gypseous soils of Nuevo Leon, Mexico, is described and illustrated. It belongs to the tribe Allieae of the lamily Amaryllidaceae, where it is most closely related to Androstephium and associated genera. It differs from most of these in having, in combination, coarsely tunicate bulbs, purple to pink perianths, and a well-developed hy- panthium to which the gynoecium is not fused. Continuing identification and curation of gypso- philic plants from Nuevo Leon, Mexico, has revealed the following novelty. Jaimehintonia gypsophila B. Turner, gen. et sp. nov. TYPE: Mexico. Nuevo Leon: along highway 68, 15 mi. S of Ascension and 3.5 mi. NW of La Escondida, ca. 97 mi. N of Matehuala, steep exposed gypsum hill adjacent to the road, “associated with ferns, Dalea and Agave," 24 Sep. 1973, James L. Reveal 3426, with N. D. Atwood (holotype, LEX). Figure 1. Herba bulbi tunicis grosse fibrosis. Folia basalia paene teretia 12-15 cm longa. Scapus 25-30 cm longus, um- bellam apicalem florum 3-6 ferens, umbella bracteis 3 6 scariosis lineari-lanceolatis 3-5 mm longisque subtenta. Pedicelli 3-4 cm longi ad summum articulati. Hypanthium roseum vel purpurascens anguste tubulare in 10 mm basali, versus apicem gradatim dilatatum; tepala 6 ex- pansa elliptica vel elliptici-oblanceolata 1-nervata 10 12 mm longa 3-4 mm lataque. Ovarium glabrum stipitatum hypanthio non adnatum. Bulb turbinate with very coarse fibrous coat, ca. 3 cm long x 2 cm wide, not deeply buried, perhaps only 2-3 cm below the soil surface. Leaves 12- 15 cm long, 1.0— 1.5 mm wide, terete or somewhat 4-sided in cross section (dried), glabrous, the angles adorned with minute clear pustulate cells, these ar¬ ranged in lines. Scapes naked, terete, 25-30 cm long, glabrous, except for very scattered minute recurved callosities, f lowers 3 6 to an inflorescence, arranged in an apical umbel, the base of the umbel subtended by 3 6 linear-lanceolate scarious bracts 3-5 mm long. Pedicels 3-4 cm long. Hypanthium reportedly “purple to pink or rarely white,” tubular below for about 10 mm, then gradually flaring to the apex, the hypanthium or fused portion 16-20 mm long, 4-6 mm wide at the top, the tepals elliptic to elliptic-oblanceolate, somewhat spreading, 10-12 mm long, 3-4 mm wide, 1 -nervate, the apices obtuse to rounded. Stamens 6, the filaments ca. 8 mm long, united below into a short scarious tube ca. 1 mm long; anthers basifixed, yellow, ca. 2.1 mm long. Ovary glabrous, elliptic, ca. 4 mm long, 2 mm wide, borne on a free stipe 8-12 mm long; style somewhat longer than the filaments, the stigma ± capitate. Capsule ellipsoid, 8-10 mm high, 3-4 mm wide, 3-valvate, each carpel containing ca. 16 seeds. Seeds irregularly quadrate, black, the surfaces finely mu- ricate (40 x ). This remarkable plant was first called to my at¬ tention by the Hinton collection cited below. I strug¬ gled for an identification of the specimen, which lacked below-ground parts. In a search through sun¬ dry genera housed in the collection at LL, TEX, I came upon another collection of the taxon by James Reveal among the unidentified materials of the Lil- iaceae, one that had a tunica-covered bulb and well- preserved flowers. Superficially, the two specimens are similar to Androstephium coeruleurn (Scheele) Torrey, having pedicels jointed at the summit, the perianth segments united into a distinct hypanthial tube, and the stip- itate ovary not fused with the hypanthium. They lack, however, the prominent corolla tube or corona of that species. They appear closely related to An¬ drostephium and cohorts, keying to this group in Moore’s (1953) synoptical key to the American gen¬ era of the Allieae; among these ( Androstephium , Rloomeria, Brodiaea, Dichelastemma, Muilla, Tri- teleia, Triteleiopsis), 1 could find no satisfactory genus in which to position the present novelty. The species of this group appear to be apportioned among genera (as currently accepted) distinguished by rath¬ er small differences. Such also appears to be true for the Allieae in general. Without a comprehensive study of the tribe, however, and the identification of groups that perhaps are more natural, the new species described here cannot be placed to genus. Its recognition as a monotypic genus is in keeping with current classificatory trends. Novon 3: 86-88. 1993. Volume 3, Number 1 1993 Turner Jaimehintonia 87 Figure 1. Jaimehintonia gypsophila, from the holotype. 88 Novon Jaimehintonia is similar to the recently described Milla mortoniana II. E. Moore (1953), having the blue (or purplish on drying) perianth and remarkably similar umbellate inflorescence, hut it lacks the an- droecial and gynoecial features of that species (cf. Moore’s illustration, 1 10 B). I take pleasure in naming the genus for Jaime Hinton, son of the late George B. Hinton, whose early collections from the Pacific slopes of Mexico are well known (Hinton & Rzedowski, 1972). Jaime is a Mexican citizen and, along with his wife, son George, and grandson, now resides on the western lower slopes ol Cerro Potosi near the village of San Rafael on his “Rancho Aquillilla.’’ The most re¬ markable attribute of Jaime, in my opinion (as op¬ posed to most nonacademic people of my acquain¬ tance), is his exceptional dedication to things botanical. He revealed to me once (following a ques¬ tion put to him regarding this matter) that one of the happiest periods of his life had to be the several years he spent enduring the rugged terrain and difficult conditions collecting plants with his father, mostly in western Mexico and Guerrero. Proof of that has been his continued zeal in pursuing similar fieldwork in the mountainous terrain of northeastern Mexico with his son George; more recently, both have been joined by the latter’s son, making this a four-generation operation, all of their collection numbers having been extended from those of George the elder, hence the collector’s notation, G. B. Hin¬ ton et al. Bara type. Mexico. NUEVO leon: Mpio. Aramberri, gypsum hillsides, Aramberri, 1,145 m, 1 Sep. 1990, G. B. Hinton et al. 20560 (TEX). Acknowledgments. I am grateful to Guy Nesom for the Latin description and to him and T. I*. Ramamoorthy for reviewing the manuscript. Nancy Webber provided the illustration. Literature Cited Hinton, J. & J. Rzedowski. 1972. George 15. Hinton, collector of plants in southwestern Mexico. J. Arnold Arbor. 53: 141-181. Moore, H. E., Jr. 1953. The genus Milla and its allies. Gentes Herb. 8: 263-294. A New Species of Oplonia (Acanthaceae) from Southern South America Dieter Wasshausen Department of Botany, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Cecilia Ezcurra Universidad Naeional del Comahue, Centro Regional Universitario Bariloche, Casilla de Correo 1336, 8400-San Carlos de Bariloche, Prov. de Rio Negro, Argentina ABSTRACT. A new species of Oplonia from the province of Jujuy, Argentina, 0. jujuyensis, is de¬ scribed, illustrated, and compared with its closest relative. It differs markedly from O. hutehisonii by the corolla, which is weakly zygomorphic, and by the erect limb, with the posterior lip deeply two- lobed. This is the first report of this genus from Argentina. I he present study of t fie Acanthaceae for the flora of the Province of Jujuy, Argentina, has led to the recognition of a new species of Oplonia from that region. The genus belongs to the subfamily Acanthoideae tribe Odontonemeae in accordance with Lindau's (1895) infrafamilial classification of the Acanthaceae, or in the tribe Justicieae subtribe Odontoneminae if one prefers to follow Breme- kamp’s (1965) system of subfamilial classification. It is characterized by flowers that are either solitary in the leaf axils or borne in comparatively few- flowered axillary fascicles with minute bracts. This much reduced inflorescence distinguishes Oplonia from Odontonema Nees, the latter of which has flowers borne in terminal racemes or panicles. The corolla is zygomorphic, occasionally only weakly so, with a slender tube and a two-lipped limb, the pos¬ terior lip is two-lobed and the anterior lip is three- lobed. There are two stamens, which may be ex- serted or included, and two staminodes. Presently, two species are known from northern Peru, eight from the West Indies, and five from Madagascar. This is the first time that the genus is reported from Argentina. Although the genus Oplonia is easily recognized and defined, the delimitation and distribution of the taxa are more complex and somewhat unusual. Most species grow in dry open habitats as part of scrub forest. The new taxon is found in thicket of tran¬ sitional forest between the Chaco and Yungas (sensu Cabrera, 1976). The evolution of the genus (Stearn, 1971) would appear to be from a mesopbytic spine¬ less South American ancestor, resembling the small- flowered species of Odontonema, by progressive re¬ duction in leaf size, by the contraction of the inflo¬ rescence from a narrow panicle or raceme into a few-flowered axillary fascicle or solitary flower, and by the conversion of lateral branches into spines, by which changes they became xerophytic. We be¬ lieve that Oplon ia jujuyensis, with its reduced leaves, an inflorescence of few flowers in axillary fascicles, small, Odontonema- like flowers, and habitat rep¬ resents an isolated transition between the genera Odontonema and Oplonia. Oplonia jujuyensis Wasshausen & Ezcurra, sp. nov. TYPE: Argentina. Jujuy: Depto. Ledesma, Parque Naeional Calilegua, near the road to Mesada de las Colmenas, 850 m, 9 Aug. 1985, ludica & Ramadori 91 (holotype, SI). Figure 1 A-E. Frutex vel suffrutex, 1.5 m altus, ramosus, inermis, caulibus teretibus cupraceis vel pallidis nitidis cortice ex- foliato; foliorum laminae ovato-ellipticae 4-10 cm longae et 3-4 cm latae utrinque attenuatae glabrae; floribus axillaribus fasciculatis, geminis ternisve brevissime pedi- cellatis; bracteolis 1-2 mm longis subulatis brevibus; co¬ rolla alba late tubulosa 1 5 mm longa, labis subaequalibus, superiore profunde bilobato, lobis ovatis vel obovatis, in- feriore prope basim trilobato, lobis superioribus similibus leviter latioribus; staminibus sterilibus brevibus; capsula ignota. Branching shrub or subshrub to 1.5 m high; stem erect, cylindrical, the bark pale or copper-colored, exfoliating, glabrous, with prominent leaf scars; spines absent; leaves opposite, petiolate, the lower leaves often fascicled and on short axillary branches, re¬ duced, deciduous toward base, the blades elliptic - ovate, 4-10 cm long, 3-4 cm wide, acute and acuminate, narrowed at the base and decurrent on the petiole, membranous, entire or undulate, gla¬ brous, the cystoliths prominent under a lens; petiole 10-13 mm long; flowers 2-3 in sessile dichasia. Novon 3: 89-91. 1993. 90 Novon Figure 1. A-E. Oploriia jujuyensis ( Iudica & Ramadori 91). — A. Habit. — B. Bracteoles, calyx segments, and corolla. — C. Corolla expanded, stamens, and staminodes. — D. Bracteoles, calyx segments, ovary, and style. — E. Portion of style and stigma. occasionally reduced to 1 flower subtended by 2 bracteoles, borne in the axils of the upper leaf blades; bracteoles triangular, small, 12 mm long, glabrous or puberulous; calyx 3-4 mm long, the segments linear, ca. 3 mm long, 1 rnm wide, subulate, pu¬ berulous or glabrous; corolla white, tubular, puber- ulent within at the base of the tube, glabrous without, 15 mm long, the tube 12 mm long, 1 mm wide at the base, gradually expanded to 3.5-4 mm diam. at the throat, the limb erect, the posterior lip deeply 2-lobed with the lobes ovate to obovate, 3 mm long, 1 .6-2.2 mm wide, obtuse or rounded at apex, ciliate along the margins, the anterior lip similar only the lobes slightly larger, 2. 2-2. 5 mm wide; stamens included, inserted near the middle of the corolla tube; filaments 2.5 mm long, glabrous; anthers 1.5 mm long, the thecae subequal, muticous; pollen grains spheroid, 36 x 33 /urn, 3-colporate, each colpus flanked by 2 pseudocolpi, pitted with minute lumina (Fig. 2A-B); staminodes 2, minute, 1-2 mm long; ovary glabrous, 2 mm long; stigma slightly bifid. Capsule wanting. Distribution. Growing in thicket of foothills of transitional forest between the Chaco and Yungas. Oplonia jujuyensis is not closely allied to the other known species of Oplonia. It is perhaps near¬ est in relationship to O. hutchisonii Wasshausen from the Department of Amazonas, Peru; however, this species is unique in that the corollas are much smaller (8-8.5 mm long), strongly zygomorphic with the posterior lip shallowly two-lobed and the lobes Volume 3, Number 1 1993 Wasshausen & Ezcurra Oplonia 91 Figure 2. A, B. Scanning electron (SEM) photomicrograph of Oplonia jujuyensis (Iudica & Ramadori 91). — A. Equatorial view, x 1,700. — B. Polar view, x 1,500. of the anterior lip spreading, 7-15 flowers are borne in sessile axillary fascicles, and the pollen grains are prolate, 66 x 36 pm, not distinctly pitted with minute lumina. In contrast, O. jujuyensis has co¬ rollas 15 mm long, these weakly zygomorphic with the posterior lip deeply two-lobed and the lobes of the anterior lip erect, 2-3 flowers are borne in sessile axillary fascicles, and the pollen grains are sphe¬ roidal, 36 x 33 pm, distinctly pitted with minute lumina. Paratype. Argentina, salta: Depto. Grab San Martin, Quebrada de Capiazuti, Schulz & l arela 5378 (CTES). Literature Cited Bremekamp, C. E. B. 1965. Delimitation and subdi¬ vision of the Acanthaceae. Bull. Bot. Surv. India 7: 21-30. Cabrera, A. L. 1976. Regiones Eitogeograficas Argen- tinas. In: Enciclopedia Argentina de Agricultura y Ganaderia, 2da. edic., Tomo 2(1), Editorial Acme, Buenos Aires. Lindau, G. 1895. Acanthaceae. Nat. Pflanzenfain. IV. 3b: 274-354. Stearn, W. T. 1971. A survey of the tropical genera Oplonia and Psilanthele (Acanthaceae). Bull. Brit. Mus. (Nat. Hist.) But. 4(7): 261-323. A New Combination in Pottia (Musci) Richard Zander Buffalo Museum of Science, 1020 Humboldt Parkway, Buffalo, New York 14211, U.S.A. Abstract. Pottia fosbergii, of the southwestern United States and Mexico, is recognized at the va¬ rietal level: P. starkeana var. fosbergii. While working on t lie genus Pottia for the forth¬ coming Moss Flora of Mexico (Sharp et al., in press), I found that plants referable to Pottia fosbergii E. B. Bartram usually occur in mixture with P. starke¬ ana (Hedwig) C. Muller subsp. starkeana and may actually be hybrids between P. starkeana and an¬ other species. The operculum is usually undiffer¬ entiated and the peristome absent, hut a Californian specimen that is clearly P. fosbergii (Ikenberry 369, CANM), called to my attention by Terry McIntosh, has a weakly differentiated operculum and rudi¬ mentary peristome teeth. The gametophytes of both the above taxa are essentially identical. Pottia fos¬ bergii should henceforth be recognized at the va¬ rietal level, as it differs from P. starkeana only in the operculum weakly differentiated or not differ¬ entiated, and the peristome absent or rudimentary. Pottia starkeana var. fosbergii (E. B. Bartram) R. H. Zander, comb, et stat. nov. Basionym: Pottia fosbergii E. B. Bartram, Bryologist 33: 18. 1930. TYPE: U.S.A. California: Los An¬ geles Co., Claremont, Fosberg 1)24 (holotype, FH). Literature Cited Sharp, A., H. Crum & P. Eckel (editors). Moss Flora of Mexico. New York Bot. Gard., Bronx, New York (in press). Volume 3, Number 1, pp. 1-92 of NOVON was published on 19 March 1993. Volume 3 Number 2 1993 NOVON Nj||^ Lepidium tayloriae (Brassicaceae), a New Species from Chile Ihsan A. Al-Shehbaz Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. ABSTRACT. Lepidium tayloriae from Atacama (Region III), Chile, is described and illustrated. Its relationship to L. pseudodidymum and /.. strictum is discussed, and its distinguishing characters from these and other South American species are given. Several South American species of Lepidium have been discovered since Hitchcock’s (1945) revision, and the total number, including this new species and L. peruvianum Chacon (1990), bring the total es¬ timate of Al-Shehbaz (1989) to 52 species. The South American species of Lepidium belong to sev¬ eral complexes (e.g., L. abortanifolium Turczani- now, L. aletes Macbride, L. chichicara Desvaux, L. depressum Thellung, and L. meyenii Walpers), the taxonomy of which requires critical study. The limits of component species ol a given complex are sometimes obscure. As shown below, however, the new species and its relatives are readily distinguished from all the other South American lepidiums. Lepidium tayloriae Al-Shehbaz, sp. nov. TYPE: Chile. Atacama (Region III): Copiapo Province, between Huasco and Copiapo, ca. 20-25 km W of Totoral on the road to the coast, dry plains and hillside, 20°00'S, 70°50'W, 8 Oct. 1991, C. M. Taylor, C. von Bohlen & A. Marticorena 10804 (holotype, MO; isotype, CONC). Figure 1 . Herba annua hirsuta 17-24 cm alta. Folia caulina infima pinnatisecta 2-4 cm longa, segmentis 0.2-0. 5 mm latis; folia superma anguste linearia integra, 1-4 cm longa, 0.3- 1.5 mm lata. Flos sepalis ovatis 0.6-1 mm longis; petalis filiformibus usque ad 0.2 mm longis; pedicellis fructiferis erectis 1.2-2 mm longis, anguste alatis. Fructus suborbicularis, medio secus replum subconstrictus et sub- didymus, (3.4-)3.7-4 mm longus et latus, valvis alatis, manifeste reticulato-venosis. Herbs annual, erect, 1 7-24 cm tall; branches many, ascending, moderately to densely hirsute with spreading trichomes to 0.5 mm long. Basal leaves not seen; lowermost cauline leaves pinnatisect, 2- 4 cm long, hirsute; petiole winged, 0.5- 1.5 cm long, to 1 mm wide; lateral lobes 3-6, narrowly linear, 2-6 x 0.2-0. 5 mm; rachis as wide as petiole; uppermost leaves narrowly linear, entire, 1-4 cm long, 0.3- 1.5 mm wide. Inflorescences ebracteate racemes, greatly elongated in fruit. Sepals ovate, 0.6-1 x 0.3-0. 5 mm, glabrous or rarely sparsely hirsute, purplish, scarious at margin, usually falling ofF after fruit maturation; petals filamentous, to 0.2 mm long; stamens 2, the filaments white, ca. 0.6 mm long, the anthers ca. 0.2 mm long; nectar glands 4, toothlike, ca. 0.1 mm long. Fruiting pedicels 1.2- 2 mm long, erect, subappressed to rachis, narrowly winged, hirsute. Fruit suborbicular, subdidymous, (3.4-)3.7-4 mm long and wide, prominently retic¬ ulate, winged, glabrous, emarginate at apex, trun¬ cate-rounded at base; wing encircling all fruit, ca. 0.6 mm wide at fruit apex; apical notch ca. 0.3 mm deep, ca. 0.6 mm wide; style absent; stigma entire. Seeds ovate, brown, ca. 2 x 1 mm; cotyledons incumbent. Lepidium tayloriae, which is named after Char¬ lotte M. Taylor, one of the collectors of the holotype, is most closely related to the Argentinian L. pseu¬ dodidymum and the Chilean L. strictum (S. Wat¬ son) Rattan. These three species are readily distin¬ guished from all of the other South American lepidiums by the prominently reticulate-veined fruits. The new species differs from L. pseudodidymum Novon 3: 93-95. 1993. 94 Novon Volume 3, Number 2 1993 Al-Shehbaz Lepidium tayloriae 95 by its winged fruits (3.4-)3.7-4 mm long and wide, erect fruiting pedicels, entire upper cauline leaves, and sessile stigma. By contrast, L. pseudodidymum has wingless fruits 2-2.6 x 2.2-3 mm, arcuate- spreading fruiting pedicels, 2- or 3-pinnatifid or - pinnatisect upper cauline leaves, and styles ca. 0.17 mm long. It is separated from L. strictum by its larger fruits with glabrous keel, toothlike nectar glands, entire upper cauline leaves, trichomes to 0.5 mm, and constricted fruit replum distinctly lower than the valve surface. Lepidium strictum has smaller fruits (2.5 X 2 mm) with puberulent keel, linear nectar glands, pinnatisect upper leaves, tri¬ chomes to 0.3 mm, and raised fruit replum higher than the valve surface. Acknowledgment. I am grateful to Roy Gereau for checking the Latin description. Literature Cited Al-Shehbaz, I. A. 1989. Lepidium boelckei and L. jujuyanum (Brassicaceae), new species from Jujuy, Argentina. Ann. Missouri Bot. Card. 76: 1 189-1 192. Chacon, G. 1990. La maca (Lepidium peruvianum Chacon sp. nov.) y su habitat. Rev. Peru. Biol. 3: 169-272. Hitchcock, C. L. 1945. The South American species of Lepidium. Lilloa 11: 75-134. Romanschulzia mexicana (Brassicaceae), a Remarkable New Species from Guerrero, Mexico Ihsan A. Al-Shehbaz Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. Hugh H. litis Department of Botany, University of Wisconsin-Madison, Birge Hall, 430 Lincoln Drive, Madison, Wisconsin 53706-1381, U.S.A. ABSTRACT. Romanschulzia mexicana is described as new and illustrated. It is readily distinguished from all other 12 species of Romanschulzia by its fruiting pedicels and gynophores that are both very long and rather slender. Romanschulzia 0. E. Schulz is one of two genera of the Brassicaceae (Cruciferae) that are distributed exclusively at high allitudes in the tropics (Al-Sheh¬ baz, 1984). The other genus is the African Oreo- phyton O. E. Schulz. Romanschulzia consists of 1 2 species distributed from central Mexico south into Panama at 1,800-3,300 m (Hollins, 1942, 1956, 1984). The genus belongs to the Thelypo- dieae Prantl, a tribe considered by some to include genera that appear to be “primitive” among the Brassicaceae (Cruciferae) (Al-Shehbaz, 1973, 1985; Rollins, 1956). Because of its exceptionally long gynophores and shrubby habit, the new species was thought to be a member of the Capparaceae and thus was sent to one of us (H. H. litis). litis identified it as an unknown Romanschulzia because of its long gynophore, as well as the presence of a complete septum and incumbent cotyledons not invaginated between the radicle and cotyledons; the latter both are characters typical of Brassicaceae and not Cap- paraceae-Cleomoideae. Although the single collec¬ tion on which R. mexicana is based contains no flowers, the species is described because of its unique fruiting material. Romanschulzia mexicana litis & Al-Shehbaz, sp. nov. TYPE: Mexico. Guerrero: Mun. Le¬ onardo Bravo, Pedregal, 28 km by road WSW of Kilo de Caballo, 10 June 1985, W . Thomas <£• J. L. Contreras 3788 (holotype, NY; isotype, WIS; fragments, MEXU, UC). Figure 1. Frutex scandens usque ad 2.5 m; folia petiolata lan- ceolata vel oblongo-lanceolata obscure denticulata, folia proxime infra racemos auriculata vel ainplexicauli; ped- icelli fructiferi tenuississimi, arcuati vel divaricati, 4-8 cm longi; gynophoro tenuississimo, 2-3.2 cm longo; fruc- tus cylindricus, 2.5-4. 1 cm longus; semina oblonga, un- iseriata, 2. 6-3. 6 x 1-1.3 mm; cotyledones incumbentes. Scandent shrubs to 2.5 m, glabrous throughout. Stems terete, green when young, becoming straw colored with age; lateral branches 3-10 cm long; pith solid. Lowermost leaves of lateral branches pet- iolate, lanceolate to oblong-lanceolate, 2.5-6 x 0.4- 2.2 cm, cuneate at base, obscurely denticulate, acute at apex; leaves just below racemes sessile, auriculate to amplexicaul, with rounded to subacute basal lobes. Flowers not seen. Infructescences lax racemes, 3- 7 cm long, terminating lateral branches; fruiting pedicels very slender, divaricate at base, strongly arcuate to straight, 4-8 cm long, 0.3 mm diam. Receptacle 1.2-2 mm diam.; nectar glands contin¬ uous, subtending bases of all filaments. Gynophore very slender, 2-3.2 cm long, ca. 0.3 mm wide, obscurely and longitudinally 4-striate or 4-winged. Fruit narrowly cylindrical, 2.5-4. 1 cm long, 2-3 mm wide; valves acute at both ends, with a prom¬ inent midvein and conspicuously reticulate lateral veins; septum complete, thin, opaque, pushed to valves by seeds on other side; style absent; stigma entire, minute. Seeds narrowly oblong, brown, 2.6- 3.6 x 1-1.3 mm, plump, uniseriately arranged in each locule, wingless, faintly and minutely reticulate, with a tiny appendage distally; cotyledons incum¬ bent, that facing radicle smaller. Distribution. Known only from the type material, collected in forests and pasture on karstic limestone at 1 ,800-1,900 m. The relationship of Romanschulzia mexicana is not entirely clear. The shrubby habit in Roman¬ schulzia is known only in another species, R. ape- tala Rollins, which is a narrow endemic of the Cor¬ dillera Talamanca, Prov. Cartago, Costa Rica, once collected by R. W. Holm and H. H. litis in 1949 (Holm A' litis 536, MO). II owever, the latter, only Novon 3: 96-98. 1993. Volume 3, Number 2 1993 Al-Shehbaz & litis Romanschulzia mexicana 97 Figure 1. Romanschulzia mexicana litis & Al-Shehbaz (Ihomas & Contreras 3788). — A. Lateral vegetative branch. — B. Infructescences. — C. Fruit. — D. Receptacle. — E. Seed. Scales A C = 1 cm; D, E = 1 mm. 98 Novon remotely related to R. mexicana, is readily distin¬ guished from the new species by its long cauline leaves to 7 cm, flattened fruits, slender styles 4-5 mm, shorter gynophores 5-8 mm, biseriate seeds, and stout pedicels 1.5-2. 5 cm. Romanschulzia mexicana is easily separated from all other species of Romanschulzia by its very slen¬ der, arcuate to spreading fruiting pedicels 4-8 cm long, slender gynophores 2-3.2 cm long, promi¬ nently veined fruit valves, and sessile stigmas. In fact, the combination above of the very long and slender pedicels and gynophores is rather unique in the Brassicaceae, and the only other genus with such long gynophores, but with much shorter (0.4- 2 cm) fruiting pedicels, is the western North Amer¬ ican Stanleya Nuttall, another member of the tribe Thelypodieae. Literature Cited Al-Shehbaz, I. A. 1973. The biosystematics of the genus Thelypodium (Cruciferae). Contr. Gray Herb. 204: 3-148. - . 1984. The genera of Cruciferae (Brassica¬ ceae) in the southeastern United States. J. Arnold Arbor. 65: 343-373. - . 1985. The genera of Thelypodieae (Crucif¬ erae: Brassicaceae) in the southeastern United States. J. Arnold Arbor. 66: 95-111. Rollins, R. C. 1942. A tentative revision of Roman¬ schulzia. Contr. Dudley Herb. 3: 217-226. - . 1956. Some new primitive Mexican Crucif¬ erae. Rhodora 58: 148-157. - . 1984. Studies on Mexican Cruciferae II. Contr. Gray Herb. 214: 19-27. A New Species of Xylopia (Annonaceae) from Southern Venezuela Paul E. Berry Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, IJ.S.A. David M. Johnson Ohio Wesleyan LTniversity, Department of Botany-Microbiology, Delaware, Ohio 43105, U.S.A. Abstract. Xylopia plowmanii is newly described from plants from Estado Apure and Estado Ama¬ zonas, Venezuela. It most closely resembles a group of emarginate-leaved species that includes X. emar- ginata, X. spruceana, and X. venezuelana. During the preparation of the Annonaceae treat¬ ment for the upcoming Flora of the Venezuelan Guayana, the following new species of Xylopia was recognized and is described below. Xylopia plowmanii P. E. Berry & D. Johnson, sp. nov. TYPE: Venezuela. Amazonas: Tobo- gan de la Selva, along Rio Coromoto, 35 km S of Puerto Ayacucho, approximately 5°22'N, 67°33'W, T. Plowman & F. Guanchez 13523 (holotype, MO; isotypes, F, NY, U). Figure 1. Species Xylopiae emarginatae Martius, X. sprucean- ae Bentham et X. venezuelanae R. E. Fries proxirna, differt vero: caulibus juvenilibus sparse adpresseque pub- erulentibus deinde subglabratis, foliis latioribus atque el- liptico-oblongis, pagina inferiore glabris, excepta costa adpresse strigosa, calyce glabro leviter lobato, 2 mm lon- go, 3. 5-4. 5 mm lato, petalis exterioribus 14.5-18 mm longis, dorso inconspicue adpresso-pubescentibus, petalis interioribus 1 4 20( 24) mm longis. tree 12-16(-20) m tall. Leaf-bearing twigs 0.8 2.8 mm thick, brown to grayish brown, at first hnely appressed-puberulent, later becoming glabrate, len- ticellate. Blades of mature leaves 5-8 cm long, 2- 3 cm wide, subcoriaceous, discolorous, oblong to elliptic-oblong, rounded and conspicuously emargin- ate at the apex, narrowed toward the base but angled sharply 2-4 mm from the petiole, making the base obliquely truncate, glabrous except for midrib, which is strigose abaxially with hairs 1-2 mm long; midrib impressed adaxially, raised abaxially; secondary veins faint, 7-9 per side, at 60-70° angle to midrib, slightly raised on both surfaces of the blade, anas¬ tomosing ca. 3 mm from the margin; petioles 3 4 mm long, 1.1 -1.3 mm wide, terete, shallowly can¬ aliculate and pubescent adaxially, glabrous abaxi¬ ally. Flowers axillary, commonly two (sometimes three) per axil; inflorescence axis below articulation with pedicel 1-2.2 mm long, 1.1 -1.3 mm thick; pedicel 2.8-4 mm long, 0.7-0. 8 mm thick at mid¬ point, with an amplexicaul bract attached 1.2 1.5 mm above the articulation; bract 0.9-1 mm long, 1-1.3 mm wide, quadrate to semicircular. Calyx cuplike, 3. 5-4. 5 mm diam., divided to about the midpoint into broadly triangular lobes 1.5-2 mm long, glabrous. Petals pale yellow adaxially, orange except for the yellowish base abaxially; outer petals 14.5-18 mm long at maturity, 2 mm wide at mid¬ point, strap-shaped, widening to 3 mm at base, base concave, apex obtuse, densely puberulent adaxially, very sparsely appressed-puberulent abaxially; inner petals subequal in length to outer ones and similar in shape or exceptionally to 24 mm long, 1 .2 mm wide at midpoint, 2 mm wide at base, puberulent along the midvein adaxially, uniformly puberulent abaxially. Stamens numerous, 1-1.4 mm long, 0.2- 0.4 mm wide, the outermost ones converted into staminodes 0.9- 1.2 mm long, 0.4-0. 5 mm wide; connectives dome-shaped, short conic or truncate, papillate; filaments 0.4 mm long. Carpels 6-10, linear; ovaries 0.9-1 mm long, pubescent; style/stigma 4. 2-4. 8 mm long, linear, with a few sparse hairs along edges; ovules 4 in a single row. Torus in flower 2.1 mm diam., the stamens leaving raised scars on the torus surface, the center of the torus excavated, with the ovaries concealed within the cavity with only the styles exserted. Pedicel in fruit 10 mm long, 2 mm wide, glabrate. Monocarps 4-6, stipitate, the seed-containing portion 1.9-2. 8 cm long, 1-1.8 cm wide, irregularly ovoid, densely covered by a matted, ferruginous indument, turning brown and glabrescent at maturity; stipes 4-7 mm long, 1.5-2 mm wide. Seeds usually 3 per mono¬ carp, at 60° angle to long axis of monocarp, 6.5- 7.5 mm long, 4-5 mm wide, ellipsoid, smooth; aril bilobed. Distribution. Southern Estado Apure and north¬ western Estado (formerly Territorio Federal) Ama- Novon 3: 99-101. 1993. 100 Novon Figure 1. Xylopia plowmanii (from Davidse <£ Gonzalez 15875, except for floral details from Plowman & Gudnchez 13523). — a. Fruiting branch. — b. Flowering branch. — c. Floral bud showing cuplike calyx. — d. Single carpel, in lateral view, showing the elongated style. — e. Staminode, abaxial view. — f. Stamen, abaxial view. — g. Lower (abaxial) surface of a leaf showing the strigose indument along the midrib and the angular base of the blade. zonas, Venezuela, 75-550 m elevation, in gallery forests and on forested slopes along granite outcrops. This species resembles the species group of Xy¬ lopia emarginata Martius, X. spruceana Bentham, and X. venezuelana R. E. Fries, which are char¬ acterized by relatively small flowers and somewhat leathery, emarginate leaves. It differs from them mainly in the broader, chartaceous (vs. coriaceous) leaves and the relatively glabrous outer petals. Al¬ though occasionally found in other members of the Volume 3, Number 2 1993 Berry & Johnson Xylopia plowmanii 101 “emarginata” group, the leaves consistently have a prominent, angular projection toward the base and a strigose abaxial surface of the midrib. 1 his species is dedicated to the late Timothy Plowman, who was an extremely knowledgeable and enthusiastic neo¬ tropical botanist. Paratypes. VENEZUELA. Amazonas: Serrania Ba¬ tata, 2 km al NE del Salto Colorado, 55 km SE de Puerto Ayacucho, 5°33'N, 67°08'W, 550 m, Sanoja et al. 3327 (MO, PORT). Apure: Distrito Pedro Camejo, Cano (or Morichal) El Caballo, 16 airline km NW of Mata de Guanabana, between the Rio Meta and the Rio Cinaruco, 6°19'N, 68°19'W, elev. 75 m, 27 Feb. 1979 (fr), G. Davidse & A. Gonzalez 15875 (MO, U, VEN). New or Noteworthy Orchids for the Venezuelan Flora IX: New Taxa, New Records, and Nomenclatural Changes, Mainly from the Guayana Shield and Northern Amazonas German Carnevali and Ivon Ramirez de Carnevali Fundacion Instituto Botanico de Venezuela, Herbario Nacional de Venezuela, Aptdo. 2156, Caracas 1010-A, Venezuela; present address: Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299. U.S.A. Abstract. Research on the Orchidaceae of the Venezuelan Guayana and neighboring areas has yielded several novelties, new records, and nomen- clatural changes. Nine species (one Dichaea, two Lepanthes, one Maxillaria, three Pleurothallis, one Sarcoglottis, and one Uleiorchis ), a natural hybrid in Maxillaria, and two subspecies (one Myoxanthus and one Stelis) are newly described. New synony¬ mies are proposed in Campy locentrum, Habenaria, and Octomeria. In addition, several miscellaneous country records are reported. Fifteen of the taxa presented are illustrated. Continued studies related to the orchid treatment for Steyermark et al.’s Flora of the Venezuelan Guayana have yielded several novelties and no¬ menclatural changes for the orchid flora of the area and from other parts of the Neotropics. These nov¬ elties have resulted from the study of material re¬ ceived from several recent expeditions or from the study of critical material mainly at the Orchid Her¬ barium of Oakes Ames (AMES), the Herbario Na¬ cional de Venezuela (VEN), and the Missouri Bo¬ tanical Garden (MO). The taxa are arranged alphabetically by subfamilies, subtribes, and genera, following Dressier (1990). Subfamily Cypripedioideae Selenipedium steyermarkii Foldats, Bol. Soc. Venez. Gi. Nat. 21: 254, fig. 1. 1961. This species was recently collected for the first time outside Venezuela, extending its range into Guyana. Specimens examined. GUYANA. Potaro— Siparuni Region: summit of Mt. Kopinang, 5°00'N, 59°55'W, 1,700-1,800 m, 8 Apr. 1988, Hahn, Judziewicz & Gopaul 4392 (MO, US). Subfamily Epidendroideaf. SUBTRIBE ANGRAEC.INAE Campylocentrum hondurense Ames, Sched. Orchid. 5: 37. 1923. TYPE: Honduras. Lan- cet ilia Valley near Tela, 250 ft. altitude, 16 Mar. 1923, Ames II 210 (holotype, AMES). Figure 1 . Campylocentrum steyermarkii Foldats, Act. Bot. Venez. 3: 316, f. 4. 1968, syn. nov. TYPE: Venezuela. Tachira: El Pinal, Rio Frio, 250-300 in, 27 Aug. 1966, Steyermark & Habe 96715 (holotype, VEN). Campylocentrum steyermarkii was based on a specimen in young buds with not fully developed spurs; otherwise it is identical to the type of C. hondurense. We have seen living material collected close to the original locality that confirms this idea. Recently C. hondurense has been collected in the Amazon Basin in Venezuela and Peru; this is the first record of this species for Peru. The species is now known from Honduras, Belize, Venezuela, and Peru. It is apparently always cleistogamous. Specimens examined. PERU. Madre de Dios: Tam- bopata, Cuzco Amazonico, Lodge Camp site 1, Plot E, 200 m, 14 June 1989, Nunez et al. 1 1029 (MO). VEN¬ EZUELA. Territorio Federal Amazonas: Rio Ma- vaca, 1°59'N, 65°6'W, ca. 200 m, 23 Mar. 1988, Ra¬ mirez & Laskowski 246 (VEN). SUBTRIBE DICHAEINAE Dichaea venezuelensis Carnevali & I. Ramirez, sp. nov. TYPE: Venezuela. Bolivar: Distrito Piar, Amarway-tepui, S side, forest 5-25 m tall, 700-910 m, 28 Apr. 1986, Liesner <£• Holst 20477 (holotype, VEN; isotype, MO). Figure 2. (Species) D. riopalenquensi Dodson verosimiliter affinis sed floribus minoribus, sepalis 5-6 mm longis (vs. 8 mm), petalis 4.5 mm longis, 2 mm latis (vs. 8x3 mm) et labello proportione latiore, 4.5 mm longo, 5.5 mm lato (vs. 9x7 mm) differt. Dichaeam cleistogamam Dodson in mentem revocans sed foliis sepalisque angustioribus recedit. Novon 3: 102-125. 1993. Volume 3, Number 2 1993 Carnevali & Ramirez Venezuelan Orchids IX 103 2 cm Figure 1. Campylocentrum hondurense. — A. Fruiting plant. — B. Detail of habit. — C. Flower, frontal view (based on Ramirez & Laskowski 246). 104 Novon Figure 2. Dichaea venezuelensis. — A. Flowering habit. — B. Lateral view of flower with developing capsule. — C. Perianth segments, flattened (based on Carnevali & Ramirez 2030). Epiphytic herbs, small to medium sized, suberect to pendulous. Stems 7-15 cm long, 3-4 mm thick, usually simple, if branching, then only near base, laterally compressed. Leaves 25-31 mm long, 3- 4 mm wide, articulate with sheaths, linear-oblong or linear-elliptic, acute, aristate-acuminate, margins apically minutely ciliolate. Inflorescences uniflorous, solitary in leaf axils; peduncle filiform, terete, 7-10 mm long, with two basal sheaths 3 mm long, 2.5 mm wide; the sheaths imbricate, oblong-elliptic, acute dorsally carinate. Floral bract 2.8-3. 1 mm long, ca. 2 mm wide, broadly ovate, acuminate, dorsally carinate. Bracteole 2.8-3 mm long, 0.3 mm wide, linear, acuminate. Ovary 0.8-1 mm long, densely inuricate with white hairs (when living). Flowers small, not opening widely and frequently cleistoga- mous, with greenish white, basally red-speckled se¬ pals and petals; labellum white; column green with red margins. Sepals and petals fleshy, 5-nerved. Dorsal sepal 4. 5-5. 2 mm long, 2. 2-2. 4 mm wide, ovate-elliptic, acute. Lateral sepals 5. 3-6. 2 mm long. 2. 4-2. 8 mm wide, obliquely ovate-elliptic, acumi¬ nate. Petals 4. 1-4.5 mm long, 1. 9-2.1 mm wide, oblong-elliptic to oblong, acute and apiculate or shortly acuminate. Labellum 4-4.2 mm long, flesh¬ ier than other perianth segments, very broadly ob- ovate, anchorlike from a broadly cuneate-unguicu- late claw, the claw 2. 5-2. 7 mm long, the blade 5- 5.5 mm wide between the apices of the extended lateral lobes, dilated into an abruptly crescentiform anterior portion, apically very broadly rounded or subtruncate, shortly and obtusely apiculate in the center, the apical lateral lobes retrorse, 1.5-2 mm long. Column 1.5 mm long on dorsal face, 2.8-3 mm long on ventral face, ca. 1.5 mm thick, with a pubescent, triangular infrastigmatic ligule 0.5 mm wide. Paratypes. VENEZUELA. Bolivar: 78 km S tie Uri- man-tepui, bosques en alrededores de pequena cima de arenisca, 04°39'N, 62°36'W, 450 m, Sep. 1986, Fer¬ nandez 3424 (MYF, VEN); Gran Sabana, Amarway- tepui, 600-900 in, floracion en cultivo de una planta Volume 3, Number 2 1993 Carnevali & Ramirez Venezuelan Orchids IX 105 originalmente colectada por R. Liesner & B. Holst, 30 Sep. 1986, Carnevali & Ramirez 203(7(VEN); cercanias de Wonken, Rio Akaruai, vegetacion riparina, 800-900 m, 1 1 Sep. 1983, Morillo et al. 9445 (VEN). Miranda: Parque Nacional Guatopo, bosque nublado, 600-800 m, 10°03'N, 66°27'W, 27 May 1987, Capote 140 (VEN). Tachira: ca. 35 km SSE de San Cristobal, La Buenana, 6-12 km from Quebrada Colorado, primary forests, 600- l, 000 m, 07°28'N, 72°09'W, 20-21 Mar. 1981, Liesner & Gonzalez 10848 (MO, VEN). Territorio Federal Amazonas: Rio Arari, afluente del Matapire, 3-6 km aguas arriba de la desembocadura del primero, 530-550 m, 01°30'N, 65°13'W, 14 Nov. 1982, Gudnchez 2250 (TFAV, VEN). This new Dichaea is one of about 15 species in the genus with articulated leaves and echinate ova¬ ries. It appears to be related to the Ecuadorean D. riopalenquensis, which has larger flowers and a differently proportioned labellum. Another related species is D. cleistogama Dodson, also from Ec¬ uador, with which it shares the tendency toward cleistogamous flowers, but D. venezuelensis is dif¬ ferent in its narrower leaves and perianth segments. Specimens of D. venezuelensis have previously been confused with I). brachypoda Reichenbach. f., a similar but larger species. In D. brachypoda the perianth segments are brownish violet within and are 7. 5-8. 5 mm long. Dichaea venezuelensis grows as an epiphyte at altitudes of 450- 1 ,000 m, in wet forests. It is a widespread, though uncommon species in Venezuela. SUBTRIBE GASTRODIINAE Uleiorchis liesneri Carnevali & I. Ramirez, sp. nov. TYPE: Venezuela. Territorio Federal Amazonas: Departamento Atures, lower for¬ ested E slope of unnamed 1,760 m peak, 8 km NW of Yutaje, 1,050-1,200 m, 5°41'N, 66°9'W, 13 Mar. 1987, Liesner & Holst 21893 (holotype, VEN; isotype, MO). Fig¬ ure 3. Species haec Uleiorchis ulaei (Cogniaux) Handro af- finis sed differt labello longiore, proportione angustiore, 15-16 mm longo, 3-3.2 mm lato vs. 11 mm longo, 4.5 mm lato, i.e., 5-plo longiore quam latiore (vs. 3-plo long¬ iore quam latiore), anguste oblongo-elliptico (vs. ovato- elliptico), longiore vel subaequante quam copa perianthina (vs. breviore quam copa perianthina). Saprophytic, erect herbs, 15-33 cm high, shade- loving. Basal corms 3-10 mm thick, horizontal, fusiform. Stems 1-2 mm thick, terete, straight or arched, simple or bifurcate, mostly naked except for membranous sheaths at nodes; sheaths 4-5 mm long, 3 mm wide, ovate, acute to subacuminate, 5- 6-nerved. Flowers solitary at stem apex, resupinate, horizontal, not widely opening. Floral bracts 5-6 mm long, 3-4 mm wide, similar to cauline sheaths. loosely fitting the pedicellate ovary. Pedicellate ova¬ ry obconic, subtrigonous; pedicel 3-4 mm long, terete, ovary 4-4.5 mm long. Sepals and petals fused into a cup or sac longitudinally opened at the ventral face, this opening entirely filled in natural position by the labellum. Perianth cup dorsally 14- lb mm long, 17-18 mm long ventrally from base of columnar foot, 5 6 mm wide, subtriangular in lateral view; free portions of sepals and petals 0.5- 1.5 mm long, triangular elliptic, acute, obtuse or truncate, the sepaline tails somewhat longer; ex¬ panded perianth cup 13-15 mm wide at base. La¬ bellum 15-16 mm long, 3-3.2 mm wide, simple, narrowly oblong-elliptic, apically subrounded or broadly obtuse, basally subrounded-truncate with a claw 1 mm long by which the blade is hinged to column-foot; blade concave, margins undulate-cre- nate, ventral face laterally finely and densely black- verruculose, with an intense grayish hue, medially smooth and cream-colored. Column 8-10 mm long; basal zone subcylindric, 4-4.5 mm long; apical, stigmatic zone 4.5-5 mm long, obovate, truncate, pubescent; column-foot 4. 5-5. 5 mm long, forming an angle of ca. 60° with the ovary; rostellurn 2 mm long, flat, horizontal. Capsules obovoid, 10 12 mm long with pedicel up to 12 mm long. This is the second known species in this sapro¬ phytic genus. It differs from U. ulaei by its solitary flower and longer and narrower labellum, which is longer or nearly equal in length to the perianth cup. Uleiorchis liesneri seems to inhabit altitudes above 1,000 m, while U. ulaei is only known from below 500 m. The species is named after Ronald Liesner of the Missouri Botanical Garden, one of the most important plant collectors in the Venezuelan Gua- yana. Key to the Species of Uleiorchis la. Labellum narrowly oblong-elliptic, 5 times or more as long as wide, 15-16 mm long, longer than or subequaling the perianth cup . . U. liesneri lb. Labellum ovate-elliptic or ovate, 2.5-3 times as long as wide, up to 1 1 mm long, shorter than the perianth cup . U. ulaei Notes on Uleiorchis ulaei Uleiorchis ulaei was previously known from southeastern and Amazonian Brazil and southern Venezuela, but it is now known to occur in Panama and Peru as well. In Venezuela, it has recently been collected in the northwestern state of Zulia, in the foothills of the Sierra de Perija. Perija is known to have floristic relationships both with the Amazonian Basin (Steyermark, 1982), as shown by such orchids as Epidendrum huebneri Schlechter and E. smar- agdinum Lindley, and with the Atlantic Coast of 106 Novon ] . 5 cm 1.5 cm Figure 3. Uleiorchis liesneri. — A. Flowering plant. — B. Labellum, flattened (based on Liesner & Holst 21893). Panama and Central America, as shown by such orchids as Leucohyle subulata (Swartz) Schlechter, Trigonidium egertonianum Bateman, Trichocen- trum capistrntum Reichenbach f., and Encyclia hunteriana Schlechter. Specimens examined. PANAMA. Canal Zone: Mil¬ itary reserve Fort Sherman, between Gatun and Pina, 0- 170 m, 2 Apr. 1973, Liesner 1367 (MO). San Bias: El Llano-Carti Road km 26.5, 9°19'N, 78°55'W, 200 m, 8 Apr. 1985, de Nevers el al. 5248 (MO). PERU. Pachitea: Puerto Inca, 250-300 m, 09°18'S, 74°58'W, 15 Sep. 1982, Foster 8825 (MO). VENEZUELA. Bo¬ livar: Sierra Maigualida, 300-500 m, 26 Apr. 1966, 6°N, 65°W, Steyermark 95775 (VEN). Zulia: 55 km SW of Machiques by air, Aricuaisa, 100-250 m, 9°36'N, 72°5'W, 24-25 Mar. 1982, Liesner tS: Gonzalez 13177 (VEN). Territorio Federal Amazonas: Cerro de La Neblina, 150 500 m, 0°50'N, 66°10'W, 20 Feb. 1984, Liesner 15934-a (MO, VEN), 20 Feb. 1984, Liesner 16129 (MO, VEN). SUBTRIBE MAXILLARIINAE Maxillaria x dunstervillei Carnevali & I. Ra¬ mirez, nothosp. nov. TYPE: Venezuela. Ter¬ ritorio Federal Amazonas: Cerro de la Neblina, Camp II, 2. 5-3. 5 km of Pico Phelps, wet mead¬ ow with Heliamphora, grasses and sphagnum surrounded by low forest of predominantly Eu¬ terpe and Ronettia, “Flowers yellow, growing Volume 3, Number 2 1993 Carnevali & Ramirez Venezuelan Orchids IX 107 Figure 4. Maxillaria x dunstervillei. — A. Flowering habit. — B. Pseudobulb, bare. — C. Apex of leaf. — D. Perianth segments flattened. — E. Detail of flattened labellum. — F. Transversal section through midlobe of labellum. — G. Labellum and column, other perianth segments removed. — H. Apex of labellum in natural position (based on Dunsterville 1155; illustration by G. C. K. Dunsterville, courtesy of AMES). on tree in moss,” 0°50'N, 65°48'W, 2,000 m, 1 7-22 Feb. 1 984, Funk 6258 (holotype, VEN; isotype, US). Figure 4. Planta et flos inter Maxillariam meridensem Lindley et M. quelchii Rolfe quasi intermedia, vel variabilis. A Maxillaria quelchii similis sed rhizomatis vaginis sca- briusculis longioribus; foliis pseudobulbisque angustioribus, sepalis petalisque longioribus, flaveobrunnescentibus con- coloribus vel parum bicoloribus, labelli lobulo centrali ova- to, subacuto vel rotundato, parum verruculoso, callo elon- gatiore differt. This natural hybrid was first noted by Dunsterville (1972) but he failed to provide a Latin diagnosis, probably because he was not sure of its identity. Material collected in the same general area indicates that these plants probably represent the product of introgression between Maxillaria meridensis and M. quelchii. Several lines of evidence point to the hybrid origin of these populations restricted to the highlands of the Neblina Range in southern Vene¬ zuela and northern Brazil. First, these plants show 108 Novon intermediate stages between the putative parents in most morphological characters. Second, the distri¬ bution of the character states is by no means even, some plants being almost exactly intermediate, oth¬ ers closer to one parent, others to the alternative parent. As Hurst (1902) pointed out, “in orchid hybrids of the first generation single specific char¬ acters are inherited in all degrees of blending, form¬ ing as a whole, a perfect series between the re¬ spective characters of the two parents.” Third, these plants grow in mixed populations with both parents, being apparently (based on abundant collections of this complex in the area) rarer than any of them. Fourth, Maxillaria meridensis and M. quelchii are the only Camaridium-Yike species in the genus pres¬ ent at such high elevations in the area where these natural hybrids occur. The only other species of the Camaridium group to be found in the Neblina Range, but usually at lower altitudes, is M. mapiriensis (Kranzlin) L. 0. Williams, which has different pol- linaria, much smaller flowers and leaves, and is unlikely to have the same pollinator as the two putative parents. Maxillaria meridensis is characterized by long rhizomes with small (1.5-3. 5 cm long, 1-1.5 cm wide), narrowly fusiform pseudobulbs set distantly apart. Both pseudobulbs and rhizome are concealed by scabrous-pustulose sheaths. Leaves are linear to linear-lanceolate, 9-28 cm long, 1-1.5 cm wide and acute. The perianth segments are yellowish or yellow-brown to dull orange, with sepals 17-22 mm long. The apical lobe of the labellum is very fleshy and ovate to triangular-ovate, and its surface is smooth to lightly scabrous. Maxillaria quelchii, on the other hand, has a more compact growth habit with pseudobulbs closely set on the rhizome, al¬ though it can be definitely creeping; the pseudohulbs and rhizome are concealed by striate sheaths. The pseudobulbs are 2.5-3 cm long, 1 .5-2 cm wide. Leaves are oblong-lanceolate to oblong-oblanceolate and obtuse to broadly acute. The perianth segments are white or yellowish basally, grading abruptly to a very dark maroon that is shiny in the outer surface; sepals are 31-37 mm long. The apical lobe of the labellum is transversely elliptic to obovate-suborbi- cular and coarsely verruculose. Depending on the individual, Maxillaria x dunstervillei shows vari¬ able combinations of all these character states. In all the plants collected, the abrupt change in color that occurs in the sepals and petals of M. quelchii has been obliterated or somewhat “washed out” due to the influence of M. meridensis. The holotype is a plant closer to M. quelchii, but with narrower leaves and pseudobulbs and an almost smooth central lobe approaching M. meridensis; the sepals in these specimens are up to 48 mm long, apparently showing some degree of hybrid vigor. A paratype (Stein & Gentry 1571) is much closer to M. meridensis vegetatively, but the flowers are large and similar to those of M. quelchii. The other two paratypes (Dunsterville 1154, 1155) are more intermediate between the putative parents. The pollinaria of the hybrids are intermediate among those of the putative parents. All the collections made of this hybrid come from a small area in the Neblina Range, although both putative parents are rather common and wide¬ spread in all the Guayana Highlands. Maxillaria schlechteri Foldats (syn. M. rugosa Schlechter), described from Roraima, may represent this hybrid, although the description suggests only a narrowly central-lobed form of M. meridensis. This taxon has never been collected again, and the type seems to have been destroyed in the bombard¬ ment of the Berlin herbarium during World War II. Paratypes VENEZUELA. Territorio Federal Amazonas: Cerro La Neblina, Camp. II, 2.8 km NE of Pico Phelps, 2,100 m, 0°49'40"N, 65°59'W, open bog, 15 Apr. 1984, Stein & Gentry 1571 (MO, YEN). BRA¬ ZIL. Amazonas: Cerro La Neblina, Pico Zuloaga, close to NE lateral cliff fall, 2,400 m, Oct. 1972, Dunsterville 1154 (AMES, HB), Dunsterville 1155 (AMES, HB). Maxillaria foldatsiana Carnevali & I. Ramirez, Ann. Missouri Bot. Gard. 76: 376. 1989. T his is the first collection of this species for Guy¬ ana. It was previously known only from the type collection in southern Venezuela. Specimen examined. GUYANA. Mazaruni— Potaro: Membaru Creek, upper Mazaruni River, 13 Feb. 1939, Pinkus 209 (AMES). Maxillaria grobyoides Garay & Dunsterville, Ven. Orch. Ill. 5: 186, 1972. This taxon seems to be widespread, though rare or only locally common in the northern Amazon Basin. The following are the first collections reported for Colombia and Peru. It was previously known from Brazil and Venezuela. Specimens examined. COLOMBIA. Vaupes: Rio Ap- aporis, Raudal del Jirijirimo, 7 ago. 1951, Schultes Cabrera 13439 (AMES). PERU. Loreto: Alto Amazo¬ nas, carretera Oleoducto secundario entre Andoas y Ca- pahuari Sur, 2°50'S, 76°28'W, 210 m, 13 Sep. 1979, Diaz <& Jaramillo 1374 (MO). Maxillaria maleolens Schlechter., Feddes Rep. Sp. Nov. Beih. 19: 233. 1923. Maxillaria maleolens is closely related to M. violaceo punctata Reichenbach f. from the Guianas Volume 3, Number 2 1993 Carnevali & Ramirez Venezuelan Orchids IX 109 and the Amazon Basin, from which it differs mainly by details of labellum shape and coloration, by its broader sepals, and by its fragrance, which despite the specific epithet is not disagreeable. This species is apparently widespread but uncommon in Mexico, Belize, Costa Rica, and Panama. It is now recognized from Colombia, close to the Panamanian border. Specimen examined. COLOMBIA. Choco: Bahia So¬ lano, 0-100 rn, 25 July 1973, Warner & White 37 (MO). Maxillaria nuriensis Carnevali & I. Ramirez, sp. nov. TYPE: Venezuela. Bolivar: Altiplanicie de Nuria, cloud forest on summit of SE-facing escarpment, E of Cerro El Picacho, N of Las Nieves and Las Chicharras, 45 km N of Tum- eremo, vicinity of Deborah, 600-650 m, 5 8 Feb. 1967, Steyermark 89128 (holotype, VEN; isotype, NY). Figure 5. (Species) Maxillariae melinae Lindley similis sed la- bello cuarto apicali trilobato, petalis angustioribus recedit. Epiphytic herbs, medium sized, caespitose, erect. Pseudobulbs 2.5-3 cm long, 1 1.3 cm wide when dry, obliquely oblong, apically unifoliate, clothed by 2-4 sheaths, the innermost 1-2 bearing leaves. Leaves 9.5-25 cm long, 2.1-3 cm wide, the blades submembranous, oblong-elliptic, acuminate, basally attenuate into a pseudopetiole 4-6.5 cm long, ca. 2 mm thick, 2 3 times longer in the apical leaf. Inflorescences 5-8 cm long, uniflorous, much longer than pseudobulb but shorter than leaves (nearly equaling petioles), entirely clothed by tubular, scar- ious, subimbricate sheaths. Flowers medium sized for the genus, resupinate, with subparallel, creamy- colored perianth segments. Pedicellate ovary 2.3 2.8 cm long, subterete. Floral bract 1.8 cm long, 0.7 cm wide, elliptic, acute. Perianth segments sub- membranous, strongly nerved in rehydrated mate¬ rial. Dorsal sepal 2-2.2 cm long, 0.5-0. 6 cm wide, narrowly triangular to triangular, acuminate, 9- nerved. Lateral sepals 2.2 cm long, ca. 0.5 cm wide, similar to dorsal but somewhat oblique. Petals 1.6 1.8 cm long, 0.3 0.32 cm wide, narrowly trian¬ gular-oblong, acuminate, 7 8-nerved. Labellum 1.5- 1.7 cm long, about 0.5 cm wide, narrowly elliptic- obovate, slightly 3-lobed above apical fourth, the lateral margins erect in natural position; lateral lobes small, subtruncate to rounded; central lobe 4 mm long, 2.7 mm wide, fleshy, ovate, obtuse to subacute, somewhat verruculose; disk with an oblong, acute callus slightly above middle of blade of labellum. Column ca. 8 mm long, apically 3 mm thick, hem- icylindric, somewhat laterally flattened, basally pro¬ duced into a 5-mm-long, almost perpendicular foot. This new species is related to the Maxillaria melina Lindley complex. Among the species of this group it can be recognized by the following character combination: labellum 3-lobed above apical !4; co¬ lumnar foot 6 mm long, slightly shorter than column (8 mm long), sepals 2 2.2 cm long, and narrow petals (1.6- 1.3 cm long, 0.3-0.32 cm wide). Max¬ illaria nuriensis can be readily distinguished from the closely related M. melina by its apically 3-lobed (not pandurate) labellum and its narrower petals. Maxillaria santanae Carnevali & I. Ramirez, Aim. Missouri Bot. Card. 76: 377. 1989. The following collection is the first of this species for Guyana. All known localities in Venezuela and Guyana are very close to the borders ol Brazil, so it is expected that this species will be found there also. It is also known from Ecuador. Specimen examined. GUYANA -BRAZIL BORDER. Akarai Mountains, between Rio Mapuera (Trombetas trib¬ utary) and Shodikar Creek (Essequibo tributary), 600- 800 m, 19 Jan. 1938, Smith 2987 (AMES, NY). Maxillaria tenuis C. Schweinfurth, Bot. Mus. Leaf!. 11: 289, t. 18. 1945. The study of previously undetermined material allows us to report the first collections from Colombia of this typically Amazonian species. It was previously known from Venezuela, Brazil, and Peru. Specimens examined. COLOMBIA. Vaupes: Rio Kanarari, Cerro Isibukuri, 250-700 m, 4 Aug. 1951, Schultes & Cabrera 13371 (AMES); Rio Kuduyari, 300 m, 23-25 June 1958, Garcia-Barriga et al. 15775 (AMES, MO). SllBTRIBE ORNITHOCEPHALINAE Caluera Dodson & Determann Caluera surinamensis Dodson & Determann, Amer. Orchid Soc. Bull. 52: 377. 1983. Material belonging to this species was collected more than 20 years ago in Venezuela, but remained unidentified until now. Besides the herbarium spec¬ imen mentioned here, the authors have seen living material collected in Parque Nacional Guatopo, Es- tado Miranda and cultivated by Halina Mendez, in Caracas, Venezuela. This is the first record of the genus Caluera for Venezuela. This species is also known from Surinam and French Guiana. Specimen examined. VENEZUELA. Rarinas: Re- serva Forestal Caparo, bosque tropofilo, Unidad I, parte S de la pica 7 y 9, 16-18 km SE del Campamento Cachicamo, E de El Canton, 100 m, 9 abr. 1968, Stey¬ ermark et al. 101989 (MY, VEN). 110 Novon Volume 3, Number 2 1993 Carnevali & Ramirez Venezuelan Orchids IX 111 Figure 6. Eloyella panamensis. — A. Flowering habit. — B. Detail of the flower. — C. Labellum, flattened (illustration by J. Myers, based on Stergios & Taphorn 1341 6). Eloyella Ortiz Eloyella panamensis (l)ressler) Dodson, Icon. Plant. Trop., ser. 2: pi. 0455. 1989. Figure 6. This diminutive twig-epiphyte was recently col¬ lected for the first time in Venezuela, growing in a high tree at 15-20 m in riparine forests. This is the first record of the genus Eloyella for Venezuela. Figure 5. Maxillaria nuriensis. — A. Flowering habit. — B. Perianth segments, flattened (based on Steyermark 89128). 112 Novon Figure 7. A-C. Brachionidium parvum. — A. Flowering plant. — B. Flower. — C. Labellum (based on Liesner & Holst 21724). D H. Lepanthes unitrinervis. — D. Flowering plant. — E. Flower. — F. Perianth segments flattened. — G. Labellum. — H. Lateral view of labellum and column (based on Liesner 23433). Eloyella panamensis was previously known from Panama and Ecuador. Specimen examined. VENEZUELA. Bolivar: Distri¬ to Sucre, Cano Icutu, Rio Nichare, afluente del Rio Caura, 5°53'N, 69°51'W, 150-200 m, 24 mar. 1989, Stergios & Taphorn 13416 (PORT, YEN). SUBTRIBE PLEUROTHALL1DINAE Brachionidium Lindley Hrachionidium parvum Cogniaux, Repert. Spec. Nov. Regni Veg. 6: 307, 1909. Figure 7 A-C. Volume 3, Number 2 1993 Carnevali & Ramirez Venezuelan Orchids IX 113 This species was previously known only from Gre¬ nada, Dominica, and Haiti, but recently it has been collected several times in the Venezuelan Guayana. It had been erroneously included in the Venezuelan Flora by Garay (1956) when he reduced B. lon- gicaudatum Ames & C. Schweinfurth to its syn¬ onymy. This latter taxon is clearly a different, much larger species that grows at higher elevations on tepui summits. Foldats (1970a), in bis treatment of the genus, followed Garay’s interpretation of B. par¬ vum, and the description he provided was a mixture of B. parvum and B. longicaudatum. Here we offer the first complete description based on all the ma¬ terial available of B. parvum. Data on populations from the West Indies were kindly provided by C. A. Luer. Epiphytic or subterrestrial, humicolous, erect herbs, 4-8 cm high including inflorescences. Roots few, thick, relatively short. Rhizome to 12 mm long, erect, subterete, concealed by scarious sheaths. Ramicauls 3 4 mm long and ca. 1 mm diam., spaced 1 -4 mm on the rhizome, subterete to subtrigonous, concealed by two sheaths, these 2-3 mm long, in¬ flated, costate, mucronate. Leaves 8-16 mm long, 4-5.5 mm wide, 7-nerved, fleshy, erect to suberect, elliptic, acute with a 3-denticulate apex, basally at¬ tenuate into a pseudopetiole 1-3 mm long. Inflo¬ rescences from the ramicaul apex, solitary, uniflo- rous, erect; peduncle 12 22 mm long, subterete, straight, with a median, long-mucronate sheath 2 3.5 mm long, tuhulose, inflated, subcostate. Floral bract similar to peduncle sheath, 2-2.5 mm long. Pedicel 0.5-1. 5 mm long, laterally compressed. Fil¬ ament 2.5—4 mm long, filiform, thickened toward apex. Ovary 2-2.5 mm long, subtrigonous. Flowers greenish to hyaline, nonresupinate, often fertilized. Dorsal sepal 5-7.5 mm long, 3-nerved, blade 4.5 mm long, 2.5 mm wide, ovate, acute, caudate, the margins finely ciliate, tail 1.5-3. 5 mm long; lateral sepals connate into a 4-nerved synsepal, dimensions as for dorsal sepal, tail minutely bifid. Petals 5-7.5 mm long, 3-nerved, elliptic to oblong-elliptic, acu¬ minate, blade 3.5-5 mm long, 1.5-2 mm wide, the tail 1.5-2. 5 mm long, the margins finely ciliate. Labellum 1.2- 1.5 mm long, 1.5-1.75 mm wide, fleshier than other perianth segments, transversely rhombic or subtrapeziform, apically rounded or ob¬ tuse and with a triangular apiculum 0.4 mm long, basally cuneate, the anterior margin erose; disk fine¬ ly papillose, apically concave, basally with an ovoid, cushionlike, microscopically pubescent callus. Col¬ umn 0.5-0. 7 mm long, ca. 1 mm broad. Pollinia 6. Capsule 8 mm long, 3-4 mm diam., oblong- ellipsoid. Specimens examined. VENEZUELA. Bolivar: 17 km E of El Pauji, Rio Las Ahallas, 850 m, 4°30'N, 61°30'W, 1 Nov. 1985, Liesner 19270 (MO, VEN). Territorio Federal Amazonas: Departarnento Atures, along stream on plateau N of unnamed 1,760-m peak, 9 km N W of settlement of Yutaje, 4 km W of Rio Corocoro, W of Serrania de Yutaje, forested area, 5°41'N, 66°10'W, 1,050 1,300 m, 7 Mar. 1987, Liesner & Holst 21724 (AMES, MO, VEN); Departarnento Rio Negro, Cerro Ta- macuari, Sierra Tapirapeco, 1°N, 65°W, 1,400 m, Feb. 1989, Santana & Gutierrez 43 (VEN). This small species is apparently restricted in Ven¬ ezuela to relatively low elevations in the Guayana. It is known from three stations 500-700 km apart and probably occurs in intervening places. Among other Brachionidium species it can be recognized by the following character combination: plant small; rhizome short, erect; leaves elliptic; inflorescences much longer than the leaves; sepals and petals mi¬ nutely ciliate, long-caudate, and labellum trans¬ versely rhombic with a triangular apiculum. Lepanthes Sw. Phis is a genus of perhaps 600 or more mainly Andean species (Luer, 1 986). The Amazon Basin and the Guayana Highlands are surprisingly species- poor, with less than 20 known species. To date, we know of nine species of Lepanthes in all of the Venezuelan Guayana. Most of the species in this area are either endemic or show an interesting close relationship with the Lepanthes flora of the Southern Antilles. Two species of the Venezuelan Guayana have proved to be undescribed, and a third one has recently had its known range extended into Brazil. Lepanthes marahuacensis Carnevali & I. Ra¬ mirez, sp. nov. TYPE: Venezuela. Territorio Federal Amazonas: Departarnento Atabapo, Cerro Marahuaca, summit on SE corner, open rocky plateau and ravines, 3°37'N, 65°21'W, 2,700 m, 13 14 Oct. 1988, “at base of bluff. Fruit greenish brown,” Liesner 24798 (holo- type, VEN; isotype, MO). Figure 8. Lepanthi dussii Urban similis sed ramicaulibus proli- feris, scandentibus; rachidibus laxioribus; sepalis lateral- ibus in quintam partem basalem tantum connatis; columna valde elongata discrepat. Plants small, epiphytic near ground on shrubs or subhumicolous, scandent or erect to prostrate. Roots relatively thick, 1-2 originating from bases or apices of ramicauls. Ramicauls 14-35 mm long, relatively stout, straight or arcuate, prolific, concealed in 4- 7 close-fitting, imbricate, lepanthiform sheaths with scabrous to microscopically pilose costae and ostia. 114 Novon Figure 8. Lepanthes marahuacensis. — A. Flowering plant. — B. Detail of lepanthiform sheaths. — C. Flower, front view. — D. Petals. — E. Blades of labellum. — F. Column. — G. Lateral view of column, labellum, and petals (based on Liesner 24798; flowering plant by Bruno Manara, rest of the figure by Carlos Reynel). Leaves 10-18 mm long, 5-8 mm wide, fleshy, per¬ pendicular to ramicaul, 9-nerved, elliptic, obtuse to broadly obtuse at apex, subapically mucronulate, basally attenuate into a broadly channeled, 3-mm- long pseudopetiole. Inflorescences 2-4-flowered ra¬ cemes, 30-60 mm long, 3-4 times as long as the subtending leaf, erect; peduncle 25-45 mm long, 4-8-articulate, terete, with bracts to 2.5 mm long, tubulose, much shorter than internodes, oblong-el¬ liptic, abruptly acute, the margins scabrous; rachis to 20 mm long, lengthening with each successive flower, fractiflex, terete, the internodes 5-8 mm long. Floral bracts 1-2 mm long, similar to peduncle bracts. Pedicel 1.5-2. 5 mm long, terete. Ovary 1.5-2 mm long, conspicuously ridged. Flowers, ap¬ parently nodding, hyaline to brownish purple. Sepals dorsally carinate, especially toward base, micro¬ scopically ciliolate within, glabrous without. Dorsal sepal 5 mm long, 3-3.5 mm wide, 3-nerved, ovate, acuminate; lateral sepals 5 mm long, 2.5-3 mm wide, 1 -nerved, ovate-elliptic, acuminate, slightly oblique, connate in the basal fifth. Petals 0.6-0. 8 Volume 3, Number 2 1993 Carnevali & Ramirez Venezuelan Orchids IX 115 mm long, ca. 2 mm wide, microscopically ciliolate on both sides, transversely elliptic-reniform; upper lobe 0.8-0. 9 mm long, obtuse-rounded; lower lobe 1.1 - 1.2 mm long, obtuse. Labellum with the blades 2. 2- 2. 5 mm long, ca. 1 mm wide toward the base, minutely pubescent, triangular-semilunate with obliquely acute upper end and acute-acuminate low¬ er end; the blades overlapping at their lower ends when expanded; connectives and body dorsally pu¬ bescent, broadly cuneate, connate above basal third of the column; sinus acute; appendix very small, subtriangular, densely long pubescent. Column 2.5 mm long, elongate, cylindric, minutely pubescent. Anther dorsal. Stigma apical. Paratype. VENEZUELA. Territorio Federal Amazonas: Departamento Atabapo, Cerro Marahuaca, type locality, 15 Oct. 1988, Liesner 24828 (MO, VEN). This species is closely related to Lepanthes dussii Urban and L. unitrinervis Carnevali & I. Ramirez, both occurring in the highlands of the Venezuelan Guayana. Lepanthes marahuacensis is easy to dis¬ tinguish from these two species by its proliferous ramicauls and a much more elongate column. Le¬ panthes unitrinervis is known only from llu-tepui and Cerro Roraima in Estado Bolivar, while L. mar¬ ahuacensis is known from the summit of Cerro Marahuaca in Territorio Federal Amazonas; both species grow at elevations from 2,500 to 2,700 m. Lepanthes dussii was described from Guadeloupe in the Lesser Antilles; elsewhere it is known only from two very distant stations in the Venezuelan Guayana (Gran Sabana and Cerro Neblina) at 1,200- 1,500 m. Several other species of Lepanthes show similar patterns of distribution. Lepanthes cercion Luer, L. pariaensis Foldats, and L. pectinata Luer occur only in the Peninsula de Paria, an area known to have a floristic relationship with the Antilles, and in the Guayana Highlands of Venezuela and Brazil. Lepanthes duidensis Ames & C. Schweinfurth is known to occur outside the Guayana area only in the Isla de Margarita, also known for its floristic relationship with the Antilles and located very close to Paria. Brachionidium parvum Cogniaux also shows the same basic distributional pattern. Lepanthes pariaensis Foldats Acta Bot. Venez. 3; 339, fig. 11. 1968. This species is common and widespread in the Venezuelan Guayana in cloud forests at elevations of 750-1,500 m. The populations from this area have more pubescent petals than those of the typical population. Recently, it was collected for the first time in Brazil. Specimens examined. BRAZIL. Amazonas: plateau of N Massif of Serra Araca, 0°51-57'N, 63°21-22'W, 1,400 m, cloud forest, 17 Feb. 1984, Prance et al. 29125 (INPA, NY, VEN); same locality, 21 Eeb. 1984, Prance et al. 29201 (INPA, NY, VEN). Lepanthes unitrinervis Carnevali & I. Ramirez, sp. nov. TYPE: Venezuela. Bolivar: llu-tepui, lower plateau with varied habitats, rocky, bog¬ gy, shrubby and short forest to 5 m tall, 5°25'36"N, 60°29'W, 2,500 m, 16 Apr. 1988, “in crack in rock. Flower buds reddish purple,” Liesner 23433 (holotype, VEN; isotype, MO). Figure 7D-H. Species Lepanthi dussii Urban affinis sed ramicaulium vaginis scabrosis; rhachidis internodiis pedicellos 3-4-plo excedentibus; sepalis lateralibus in quartam partem bas- alem tantum connatis; petalorum lobis angustioribus, lon- gioribusque; labello bilobato differt. Epiphytic herbs, small to minute, erect, musci- colous, caespitose. Roots proportionately thick and long. Ramicauls 5-8 mm long, enveloped by 3 tight¬ ly fitting, minutely scabrous, lepanthiform sheaths. Leaves 9-10 mm long, 6-7 mm wide, fleshy, broad¬ ly elliptic, apically obtuse to subrotund, basally at¬ tenuated into a twisted, 2-2.5 mm long, broadly channeled pseudopetiole. Inflorescence 25-30 mm long, a long peduncled, laxly 1-3-flowered raceme. Peduncle 20 mm long, 3-bracted. Floral bracts 1.2 mm long. Pedicels 0.8 mm long, 3-4 times shorter than internodes of rachis. Flowers apparently nod¬ ding. Fruiting ovary 1.8 mm long. Dorsal sepal 4.2 mm long, 2.4 mm wide, 3-nerved, dorsally carinate, ovate-elliptic, acuminate. Lateral sepals 5 mm long, 2.3 mm wide, 1 -nerved, connate in basal fourth, ovate elliptic, acuminate, carinate. Petals 2.2 mm long, 0.5 mm wide, transversely narrowly elliptic, inconspicuously bilobed with both lobes somewhat falcate, the upper obtuse, the lower truncate; mar¬ gins ciliolate. Labellum bilaminate, the blades 2 mm long, minutely pubescent, elliptic-lunate with obtuse upper end and acute lower end, overlapping when expanded; connectives and body broadly cuneate, connate to very base of column, dorsally pubescent; sinus elliptical with a very small, subtriangular ap¬ pendix. Column 1.8 mm long, elongate-subterete, dorsally carinate, minutely pubescent. Anther dor¬ sal. Stigma apical. Paratype. VENEZUELA. Bolivar: Cerro Roraima, valley of Rio Arabapo, 2,600 m, Sep. 1976, Dunsterville 1356 (AMES). This taxon is closely related to Lepanthes mar¬ ahuacensis , from which it differs chiefly by its non- prolific ramicauls, the more obtuse basal lobes of its 116 Novon 5 cm Volume 3, Number 2 1993 Carnevali & Ramirez Venezuelan Orchids IX 117 labellum, and its shorter column. It is distinguished from L. dussii by its scabrous, nonpilose, cauline sheaths, clearly bipartite labellum, more transverse petals with narrower lobes, and a less crowded ra- chis. The name refers to the 1 -nerved lateral sepals and the 3-nerved dorsal sepal. Myoxanthus Myoxanthus aspasicensis suhsp. arenicola Car¬ nevali & I. Ramirez, suhsp. nov. TYPE: Ven¬ ezuela. Territorio Federal Amazonas: Depar- tamento Atabapo, Cerro Huachamacari, E slope, rainforest, 03°49'N, 65°42'W, 600-700 m, 3 Nov. 1988, “terrestrial, flowers red-purple in¬ side,’’ Liesner 25776 (holotype, VEN; isotvpe, MO). Figure 9E-H. Myoxanthus aspasicensis subsp. aspasicensis similis sed habitu arenicola, rhizomate valde elongato, foliis ram- icaule brevioribus vel raro hunc subaequantibus, basi ob- tusa rotundatave vel raro subacuta (vs. cuneata vel acuta), 7.5-13 cm longis, 1.5-3. 3 cm latis (vs. 16-25 x (3.5 — )5-6.5 cm), sepalo postico 11-13 mm longo, 2. 1-2.7 mm lato (vs. 17 x 6 mm), petalis 3-4 mm longis, 1 1.3 mm latis (vs. 6x2 mm), labello synsepalo 2-plo breviore (vs. 3-plo breviore). Erect herbs, medium-sized, psammicolous to lit li- ophytic on sandstone, very rarely epiphytic, growing in sun or shade. Rhizome horizontal to suberect, short to long, relatively thick, bearing ramicauls at intervals of 6-11 mm, entirely concealed by scar- ious, defibrating sheaths. Roots thick. Ramicauls 8 20 cm long, terete, apically unifoliate, 4-articulate; each articulation entirely clothed by a tubular sheath; sheaths papery, apiculate eventually defibrating and deciduous, brown-pustulose, especially the lower ones. Leaves 7.5-12.5 cm long, (1 .5— )1 .7— 3.0(— 3.3) cm wide, thickly coriaceous, perpendicular to ramicaul or nearly so, narrowly oblong-elliptic or elliptic to oblong-elliptic, apex obtuse or subacute to (rarely) almost rounded, base obtuse or rounded to (rarely) subacute. Inflorescences uniflorous, fasciculate, originating from a conspicuous spathe about 1-2 cm long. Peduncle 1.4 2.2 cm long, terete, hispid, basally clothed by 1-2 tubular, imbricate, hispid sheaths. Floral bract 3-4.5 mm long, tubular, hispid, apiculate, completely enclosing the pedicellate ova¬ ry. Pedicel 1.5-3 mm long, stout, terete, subglab- rous. Ovary 3-3.5 mm long, obconic, stout, hispid. Flowers medium sized, 1 5-20 mm long, open, fleshy, resupinate. Sepals with pubescence in small mounds on the outer face and at margins. Dorsal sepal 11- 13 mm long, 2. 1-2.7 mm wide, 5-nerved, oblong- elliptic, obtuse or acute, somewhat concave. Lateral sepals 10-12 mm long, 3. 5-4. 5 mm wide, 3-nerved, coherent into an elliptic synsepal; synsepal 7 8 mm wide, rounded or broadly obtuse, concave, free in apical third. Petals 3-4 mm long, 1-1.3 mm wide, 3-nerved, narrowly elliptic to oblong, obtuse or sub¬ acute to somewhat subulate-attenuate. Labellum 4.5- 5 mm long, 2-2.8 mm wide, fleshier than other perianth segments, articulate with column-foot, somewhat concave, elliptic in overall shape, apically rounded, the ventral face verruculose, with a pair of linear, uncinate lobes 0.5-0. 9 mm long in basal fourth, and a pair of suboblong low lateral calli at middle, the disk with an inverted U-shaped callus near base and extending forward between lateral lobes as a pair of verruculose keels. Column some¬ what arcuate, winged in upper third, 3.8 4.2 mm long. Capsule 3 4 cm long, oblongoid, hairy. Paratypes. BRAZIL. Amazonas: Plato da Serra Ar- aca, 1,150-1,250 m, 21 Feb. 1984, Tavares et al. 112 (NY). GUYANA. Cuyuni— IVlazaruni Region : Timehri rock paintings, 2 km SSW of Maipuri Falls, 700-850 m, 5°40'N, 60° 1 7'W, 23 Dec. 1989, Gillespie A Smart 2865 (MO, US). VENEZUELA. Bolivar: El Pauji, 900 m, 4°30'N, 61°36'W, 8 Nov. 1985, Liesner 10745 (MO, VEN); Cerro Jaua, 1,850 1,920 m, 4°48'50"N, 64°34T0"W, 28 Feb. 1974, Steyermark et al. 100455 (VEN), Steyermark et al. 100811 (VEN); La Escalera, 20 Aug. 1979, Stergios & Rodriguez 6646 (PORT, VEN); Auyan-tepui, 1,200 m, 17 Sep. 1968, holdats 7 174 (VEN); 201 km S of El Dorado, 1,200 1,400 m, 22 Eeb. 1972, Steyermark et al 105408 (VEN); Rio Paragua, Oct. 1974, Steyermark s.n. (VEN); medio Rio Paragua, minas de Maraima, 300 in, 6°7'N, 63°45'\\ , 13 June 1987, Stergios 10258 (PORT, VEN); Cerro Guaiquinima, 750 in, 5°44'4”N, 63°4U8"\\, 24 Jan. 1977, Steyermark & Dunsterrille 1 1 8460 (\ EN); Lue- pa. Gran Sabana, 1,250 m, 21 Oct. 1987, Ramirez A Carnevali 147 (TFAV, VEN); Rio Outrun, Dunsterrille 206 (AMES, VEN). Territorio Federal Amazonas: Cerro Duida, 1,350 m, 3°10'N, 65°31'W, I I Feb. 1975, Tillett 752-161 (MYF, VEN); Cerro Duida, Sep. 1977, Tillett et al. 752-200 (MYF. VEN); Cerro Neblina, 15 Apr. 1984, Gentry A Stein 4666 1 (MO, VEN), 15 18 Mar. 1984, Liesner 76690 (MO); Cerro Yapacana, 825- 1,000 m, 3°45'N, 66°45'W, 7 May 1970, Steyermark 6 Bunting 108147 (MY. VEN). Figure 9. A-D. Pleurothallis elvirana. —A. Flowering plant. — B. Flower. — C. Perianth segments, flattened. — D. Labellum (based on Liesner & Stannard 16808). E H. Myoxanthus aspasicensis subsp. arenicola. — E. Old plant, showing creeping habit. — F. Flowering habit. G. Perianth segments, showing detail of externa pu escence. — H. Labellum, flattened (based on Liesner 25776). 118 Novon This new subspecies is apparently restricted to the sandstone outcrops and sandy soils derived from the Roraima Formation in southern Venezuela, Guy¬ ana, and northern Brazil. Myoxanthus aspasicensis subsp. arenicola usually grows in open situations, in full sun or under light shrubs. It has been re¬ peatedly cited in the literature of Venezuelan orchids as Pleurothallis uncinata Fawcett & Rendle [/i. uncinata (Fawcett & Rendle) Luer] (Dunsterville & Garay, 1959; Foldats, 1970), a taxon occurring from Belize to Panama and in Jamaica. The closest relative of this new taxon is M. aspasicensis subsp. aspasicensis (Reichenbach f.) Luer, which it resem¬ bles in its creeping rhizome, the relatively broad leaves for the species complex, and the broad, ver- ruculose, apical lobe of its labellum. Myoxanthus aspasicensis subsp. arenicola is easily distinguished from the typical subspecies by its smaller stature, sand-loving habit, more clearly creeping rhizome, smaller leaves that are almost always shorter than the ramicaul with a rounded or obtuse to very rarely subacute base, shorter spathe (1-2 cm long vs. 3- 4 cm long), smaller flowers (dorsal sepal 11-13 mm long, 2. 1-2.7 mm wide vs. 17 mm long, 6 mm wide), and labellum, which is about 2 times shorter than the synsepal (in the typical subspecies it is about 3 times shorter). The typical subspecies is mainly an Andean taxon known from Costa Rica, Colombia, Ecuador, and Bolivia. Myoxanthus as¬ pasicensis subsp. arenicola was illustrated and de¬ scribed by Dunsterville & Garay (1959: 354-355) as Pleurothallis uncinata Fawcett & Rendle; the drawing is somewhat misleading because it does not show the verrucae on the ventral face of the label¬ lum. Octomeria R. Brown Octomeria exigua C. Schweinfurth, Bot. Mus. Leafl. 3: 86. 1935. TYPE: Guyana. Arakaka: Hoodsline, Feb. 1896, Im Thurn 1 15b (holo- type, AMES; isotype, K). Octomeria exigua var. elata C. Schweinfurth, Bull. Tor- rey Bot. Club 75: 216. 1948. TYPE: Guyana. Kai- eteur Plateau, 5 May 1944, Maguire & Fanshawe 2325b (holotype, AMES; isotype, NY). Octomeria deltoglossa Garay, Bot. Mus. Leafl. 18: 201. 1958. TYPE: Venezuela. Bolivar: Rio Carrao, July 1955, Dunsterville 289 (AMES). Octomeria kestrochila Garay & Dunsterville, Ven. Orch. Ill. 6: 288. 1976. TYPE: Venezuela. Territorio Fed¬ eral Amazonas: Cerro Yapacana, 1,100 m, 5 May 1970, Steyermark 103130 (holotype, AMES; iso¬ type, VEN). The study of numerous herbarium specimens, including the types and abundant living material has shown that these four concepts belong to one vari¬ able species, widespread in the Guayana Highlands in Guyana and Venezuela, at elevations of 450- 1,500 m. The labellum varies in shape from elliptic (as in 0. kestrochila) to oblanceolate-obtrullate (as in the types of the other three taxa). The surface of the labellum varies from verrucose to almost smooth, with all intermediate stages, sometimes even in the same population. The plants are also rather variable in stature, ranging from 5 to 20 cm in height; the leaves can be shorter or longer than the ramicaul. Selected specimens examined. GUYANA. Arakaka: Hoodsline, Feb. 1896, Im Thurn 115 (AMES, K); Kai- eteur Plateau, 5 May 1 944, Maguire & Fanshawe 2325b (AMES, NY). VENEZUELA. Bolivar: 12 km N of Apar- aman-tepui, 950 m, Sep. 1986, A. Fernandez 352b (MYF, VEN), Chimanta tepui, Rio Tirica, 1,000 m, May, Steyermark 75571 (VEN); La Escalera, 500-1,200 m, Jan. 1962, Foldats 28b 1, 2883 (both VEN); Amarway- tepui, 600-900 m, Dec. 1986, cultivated from material collected by R. Liesner & B. K. Holst, Carnevali 20b5 (VEN); Cerro Sarisarihaina, 700-1,350 m, Feb. 1974, Steyermark et al. 1092b0 (VEN), 109091 (VEN). Ter¬ ritorio Federal Amazonas: Cerro Sipapo, 1,400 m, Maguire & Politi 27727 (NY, VEN); Cerro Yapacana, 1,000-1,200 m, 5 May 1970, Steyermark & Bunting 103130 (AMES, VEN); Cerro Duida, 1,440 m, 1931, Tate 554 (NY); Serrania de Yutaje, 5°41'N, 66°09'W, 1,000-1,050 m, 3 Mar. 1987, Liesner & Holst 215b3 (MO, VEN). Octomeria gemmula Carnevali & I. Ramirez, Ernstia 39: 15. 1986. We here report this species for the first time from Colombia. Specimen examined. COLOMBIA. Vaupes: Rio Pir- aparana (tributary of Apaporis), between 0°15'S, 70°30'\V and 0°25'N, 70°30'W, 18 Sep. 1952, Schultes & Ca¬ brera 1751b (AMES, MO). Pleurothallis R. Brown Pleurothallis elvirana Carnevali & I. Ramirez, sp. nov. TYPE; Venezuela. Territorio Federal Amazonas: Departamento Rio Negro, Cerro de La Neblina, Camp 5, valley N of base of Pico Cardona, 1,250 m, 0°49'N, 66°0'W, 21-24 Mar. 1984, Liesner & Stannard 16898 (ho¬ lotype, VEN; isotype, MO). Figure 9A-D. Species haec in Pleurothallis lindeniae Lindley com- plexu his concurrentibus notis distinguitur: planta elongata (50 cm attinenti); foliis ellipticis; inflorescentiis pendulis 5-8 floris; floribus luteis parvis, haud estriatis; petalis ellipticis; labello 3-nervato, ecalloso vel subecalloso, apice truncato. (Subgenus Pleurothallis, sect. Pleurothallis, subsect. Macrophyllae-Racemosae. ) Volume 3, Number 2 1993 Carnevali & Ramirez Venezuelan Orchids IX 119 Epiphytic herbs, erect, densely caespitose, 30- 50 cm high. Hoots many, relatively thick. Ramicauls 20-35 cm long, terete, clothed by 2 tubulose, scar- ious sheaths, the lower 1.5-2 cm long, the upper 3.5- 4 cm long. Leaves 14-17 cm long, 2. 1-3.5 cm wide, erect, coriaceous, narrowly elliptic to nar¬ rowly obovate-elliptic, acute, the apex acuminate, tridenticulate, the base subcuneate. Inflorescences 5-8-flowered pendulous racemes to 9 cm long, 1- 4 simultaneously from a fibrous sheath 4-5 mm long, ramicauls producing up to 8 racemes; peduncle subterete; rachis straight. Floral bracts 3. 5-4. 5 mm long, tubulose, basally appressed to rachis, apically expanded and cupuliform with attenuate apex. Ped¬ icel and ovary subterete; pedicel 4-5 mm long; ovary 2.5 mm long. Flowers not resupinate; perianth segments expanded, subfleshy, yellow, glabrous. Dorsal sepal 6. 5-7. 5 mm long, 3.5-4 mm wide, 3- nerved with shorter lateral nerves extending only to apical third, elliptic, obtuse to subacute, abruptly pointed apically. Lateral sepals 6-6.5 mm long, 4. 5- 5. 5 mm wide when expanded, 4-nerved, con¬ nate into a broadly elliptic, concave synsepal. Petals 5-5.5 mm long, 1.4- 1.6 mm wide, 3-nerved, flesh¬ ier than sepals, apically thickened, narrowly ovate- elliptic, acute, slightly oblique. Labellum 1 .8-2 mm long, 2. 2-2. 5 mm wide when expanded, fleshy, broadly ovate or transversely oblong, apically round¬ ed or subtruncate, with a small, rounded mucro, with three prominently raised nerves in dry material, these inconspicuous when rehydrated; disk ecallose or with 2 inconspicuous pulvinate calli 0.1 mm high. Column 1.5- 1.7 mm long, subconic or cylindric, footless; anther apical; stigmatic surfaces 2, apical. This new species belongs to the Pleurothallis lindenii complex, a group composed of many in¬ terrelated species that differ only in subtle ways. Many of these species are only known from one mountain or mountain range. Pleurothallis elvirana is the first species in this complex to be collected in the Guayana area; all the other species are from the Andean Range or the Coastal Cordillera of Ven¬ ezuela. Pleurothallis elvirana can be distinguished from most of its relatives by its 3-nerved labellum, which is ecallose or with two inconspicuous, pulvi¬ nate calli, and has a truncate apex provided with a small, rounded, apical mucro. There are two other species in this complex with ecallose labella: P. apo- rosis Luer and P. decora Luer, from Ecuador and Colombia respectively, but in these two species the labellum is longer than wide, while it is wider than long in the new species. The species is named after Elvira Cotton, from Venezuela, who distributed the material collected during the Neblina Project. Pleurothallis morilloi Carnevali & I. Ramirez, sp. nov. TYPE: Venezuela. Tachira: on Rio San Buena, 10 km E of La Fundacion, primary areas around Represa Dorada, forest ever¬ green, soils tending to be sandy, derived from metamorphic rock (schist or gneiss), 700-1,000 m, 7°47-48'N, 71°46-47'W. 13-14 Mar. 1980, Liesner , Gonzalez & Smith 9535 (ho- lotype, VEN; isotype, MO). Figure 10. Pleurothallis chamensi Lindley affinis, statura vege- tativa manifeste minore differt; rhizoma elongata pros¬ trate; foliis basaliter cuneatis, quam caulibus secundariis longioribus vel subaequalibus; inflorescentiis tantum 1-3- floris; floribus minoribus; sepalo dorsali extus carinato trianguliter oblongo; petalis vix inconspicue serrulatis; la- belo leviter carnoso. (Subgenus Acianthera sect. Sicariae subsect. Sicariae.) Figure 6. Epiphytic herbs, shortly creeping, shade-loving, 4-5.5 cm high. Roots filiform. Rhizome 4-8 cm long, terete, with 2-4 articulations between rami¬ cauls. Ramicauls 1.1 -2. 2 cm long, 0.4-1. 5 cm wide, erect to horizontal or subpendulous, basally ancipital, apically triquetrous. Leaves 2. 3-3. 2 cm long, 1-1.7 cm wide, erect to horizontal, fleshy- coriaceous, elliptic to ovate, obtuse to acute, apically minutely bidenticulate, basally cuneate or rounded to (rarely) subcordate, the midrib abaxially promi¬ nent. Inflorescences in fascicles of 1-3 but only 1 fertile at any given time, 1-3-flowered, with 1-2 flowers in simultaneous anthesis; peduncle 0.2-0. 4 cm long. Pedicellate ovary ca. 2.3 mm long, 6- ribbed, two-winged, hence triquetrous, 5 mm long after pollination. Floral bracts 0.5 mm long, green, broadly triangular, obtuse to acute. Flowers erect to suberect, not opening completely, sometimes cleistogamous; perianth segments rigid, subfleshy, greenish yellow; labellum greenish white. Sepals 3- nerved, concave, dorsally carinate, the carinae con¬ tinuous over ovary. Dorsal sepal 4 mm long, 0.9- 1.1 mm wide, 3-nerved, oblong to oblong-elliptic, acute. Lateral sepals 3.5 mm long, 1.3 mm wide, 3-nerved, triangular-ovate, acute, connate in basal V* to Vi into an oblong-elliptic synsepal. Petals 2.3 mm long, 0.4 mm wide, hyaline, 1 -nerved, linear oblong, obtuse to acute, the apical margins finely and irregularly serrulate. Labellum 2.5 mm long, 1.3- 1.5 mm wide, membranous, 3-nerved, ovate- elliptic, the apex rounded to broadly obtuse, mi¬ nutely erose, simple, concave, ecallose, the basal margins erect. Column 2 mm long, slightly recurved, with two small apical wings. Anther apple-green or yellow, papillose. Pollinia 2. Paratype. VENEZUELA. Bolivar: Rio Aonda (Au- yantepui), ca. 600 m. Mar. 1969, Dunsterville 1108 (AMES, line drawing AMES). 120 Novon B \ _ C - ^ — D H Figure 10. Pleurothallis morilloi. — A. Fruiting habit. — B. Cross section of ramicaul. — C, D. Two lateral views of flower at anthesis. — E. Sepals and petals, flattened. — F. Labellum, flattened. — G. Column. — H. Anther (based on Dunsterville 1108; illustration by G. C. K. Dunsterville, courtesy of AMES). This small creeping species, named after Gilberto Morillo, former curator at VEN, is closely related to P. chamensis Findley. Pleurothallis morilloi dif¬ fers from Lindley’s species by its smaller vegetative and floral dimensions, its long creeping rhizome, its cuneate leaves subequal to the length of the rami¬ caul, its 1-3-flowered inflorescence, and its dorsal sepal which is dorsally carinate and triangular-ob¬ long (linear-oblong to linear spathulate in l1. cha¬ mensis ); furthermore, the petals are only minutely serrulate and the labellum is thinly membranous in P. morilloi. Pleurothallis chamensis is a species of medium to high elevations, while P. morilloi is found at lower elevations ( to 1,000 m). Living material of this species was collected hy the first author close to the town of Wonken, Akaruai River, Estado Bo¬ livar in 1984, but no voucher of it was made. Pleu¬ rothallis chamensis, widely distributed and very common in the Venezuelan Coastal Range and in the Andes, is absent from the Guayana. Volume 3, Number 2 1993 Carnevali & Ramirez Venezuelan Orchids IX 121 Pleurothallis tepuiensis Carnevali & 1. Ramirez, sp. nov. TYPE: Venezuela. Bolivar: Auyan- tepui, region del Oso, 2,200 m, 10 sep. 1958, Pannier & Schwabe 1819 (VEN). Figure 11A-C. (Species) P. batillaceae Luer affinis sed statura minore, foliis multo angustioribus (6-10-plo longioribus quam la- tioribus vs. 2-3-plo longioribus quam latioribus), inflores- centiis solitariis succesivis, synsepalo sepalum dorsale su- baequante (vs. angustiore) differt. (Subgenus Crocodeilanthe.) Epiphytic herbs, caespitose to shortly creeping, to 30 cm high. Rhizome short-creeping to ascending, clothed by scarious sheaths. Ramicauls 9 17 cm long, terete, basally enclosed in a series of large scarious sheaths, the lowermost enclosing 13 ram¬ icauls, the ramicauls with a tubular median sheath. Leaves 6- 1 2(- 1 4) cm long, 1 .4- 1 .8(-2.5) cm wide, coriaceous, linear-elliptic to narrowly oblong-elliptic, apically obtuse, minutely tridenticulate, basally at¬ tenuate into a pseudopetiole 1.5-2 cm long. Inflo¬ rescences 4-12 cm long, solitary in anthesis but produced in succession, racemose, laxly to sub- densely 1 0-25-flowered, bearing flowers from very base, subtended by a conduplicate spathe 1.3-2 cm long. Floral bracts 1.5 2.3 mm long, cupuliform. Pedicel ca. 2 mm long; ovary 1-2 mm long, ridged, arcuate. Flowers campanulate, hyaline with purplish nerves, nodding, glabrous. Sepals subfleshy, dorsally carinate, concave; dorsal sepal 3.5-4 mm long, 2.5- 3 cm wide, 3-nerved, elliptic, broadly acute; lateral sepals connate about half their length into a synsepal 3.5 4.5 mm long, 2 3 mm wide, the free portions 2- nerved, triangular-elliptic, acute. Petals 2. 7-3. 2 mm long, 0.8-0. 9 mm wide, 1- to obscurely 3- nerved, narrowly oblong to narrowly oblong-oblan- ceolate, apically rounded to subtruncate. Labellum 1.8 2.2 mm long, 1.7-1. 9 mm wide, articulate to column-foot, fleshier than other perianth segments, 3- nerved, concave in natural position, broadly ob- ovate in outline when expanded, basally attenuate into an elliptic to oblong, concave claw, this separate from blade by a transverse bar; blade 3-lobed in the apical third, the lateral lobes 1 mm long, 0.4-0. 5 mm wide, callose-thickened, triangular-semilunate, rounded, the central lobe 0.5-0. 6 mm long, 0.8 mm wide, very broadly ovate, rounded to subtrun¬ cate, thickened, the ventral surface rough. Column 1.2- 1.3 mm long; column-foot 0.4 mm long, finely papillose. Paratypes. VENEZUELA. Bolivar: Meseta del Jaua, Cerro Jaua, curnbre, 4°48'50"N, 64°34T0"W, al 0 del tributario del Rio Marajano, 1,800 m, 24 feb. 1974, Steyermark et al. 109446 (YEN); Cerro Sarisarinama, curnbre, porcion NO, 4°41'40’'N, 64°13'20"W, 700 m. 12-15 feb. 1974, Steyermark et al. 109105 (AMES, VEN). This species belongs to the Pleurothallis galeata Lindley complex, composed of many very closely related species. Pleurothallis tepuiensis is similar to the Andean P. batillacea Luer, from which it differs by its much narrower leaves, its smaller size, by producing successive, solitary inflorescences, and by its synsepal as broad as the dorsal sepal, not narrower as in Luer’s taxon. The flowers of both species are very similar, but the vegetative char¬ acters and minor floral features make them clearly distinct. Pleurothallis tepuiensis is restricted to dwarf forests on tepui summits at elevations of 700-2,200 m. Stelis Swartz Stelis gemma Garay, Orquideologia 4: 77, fig. 1969. This unique species, easy to recognize by its short, congested inflorescences, relatively large, white flow¬ ers, 7-nerved sepals, and wide leaves, had been collected in Peru in 1959, but the material remained unidentified until now. It was previously known from Ecuador and Colombia. Specimen examined. PERU. Huanueo: Mufia, 7 Mar. 1959, If bytkowski 5241 (MO). Slelis umbelliformis Hespenheide & Dressier subsp. brevipedunculata Carnevali & I. Ra¬ mirez, subsp. nov. TYPE: Colombia. Narino: Municipio de Tumaco, km 63 carretera Tu- maco Pasto, Vereda El Carmen, 240 m, 19 feb. 1984, Benavides 4284 (holotype, PSO; isotype, MO). A subsp. umbelliformi inflorescentia quam folia 2-plo breviore (vs. quam folia longiore), sepalis petalisque atro- purpureis, petalis latioribus, margine acutis discrepat. Although the floral differences between the sub¬ species are not particularly remarkable, the much shorter inflorescences make subspecies brevipedun¬ culata distinctive. The typical subspecies is from the Province of Colon, Panama, and has inflores¬ cences ranging from a little longer than the sub¬ tending leaf to about two times as long, petals with rounded to truncate upper and lower margins, and flowers described as “verdes con bario morado.” In the new subspecies the inflorescences are half the length of the subtending leaf or shorter, the petals have acute to acuminate upper and lower margins, and the flowers are described as “color pardo os- curo.” The two known populations of this species are separated by a distance of about 900 km. 122 Novon Figure 11. A-C. Pleurothallis tepuiensis. — A. Flowering habit. — B. Lateral view of the flower. — C. Perianth segments flattened with two views of the labellum (based on Steyermark et al. 109446). D-F. Eltroplectris calcarata. — D. Flowering habit. — E. Flower. — F. Perianth segments, flattened (based on Fernandez 778). Volume 3, Number 2 1993 Carnevali & Ramirez Venezuelan Orchids IX 123 Figure 12. Sarcoglottis stergiosii. — A. Flowering habit. — B. Lateral view of the flower. — C. Perianth segments, flattened (based on Stergios & Aymard 4400). Subfamily Orchidoideae SUBTRIBE HABENARIINAE Habenaria pygmaea C. Schweinfurth & R. Schultes, Amer. Orchid Soc. Bull. 23: 822, fig., 1954. TYPE: Colombia. Comisaria del Amazonas-Vaupes: Rio Apaporis, Caehivera de Jirijirimo y alrededores, 250 m, 13 June 1951, Schultes & Cabrera (AMES). Habenaria guanchezii Carnevali & I. Ramirez, Ernstia 33: 15. 1986, syn. nov. TYPE: Venezuela. Terri- torio Federal Amazonas: Departamento Atures, Rio Guayapo, Raudal Moriche, 100- 120 m, 20 Apr. 1984, F. Gudnchez 3085 (holotvpe, YEN: isotypes, MY, TFAV). When we described //. quanchezii, we over¬ looked Schweinfurth’s species. The Venezuelan ma¬ terial is almost identical to the type collection and 124 Novon grows in the same kind of specialized psammicolous association. It is now known from Venezuela, Co¬ lombia, and Ecuador. Subfamily Spiranthoideae SUBTRIBE SPIRANTHINAF, Eltroplectris Kafinesque Eltroplectris calcarata (Swartz) Garay & Sweet, J. Arnold Arbor. 53: 390, fig. 71. 1972. Figure 11D-F. This showy species is scattered along its distri¬ butional range and may prove to be more common than the present collecting record indicates. Re¬ cently, it was collected twice in Venezuela; besides the specimen cited below we have seen living ma¬ terial from the Sierra de San Luis, in northwestern Venezuela. The Venezuelan material is intermediate between the “calcarata" form and the “setacea" form, confirming Garay & Sweet’s (1972) opinion that they were just extreme forms of a variable complex. It is now known to occur in Florida, the West Indies, Colombia, Ecuador, Venezuela, and Brazil. Specimen examined. VENEZUELA. Bolivar: Cerro Arimagua, close to Cerro Bolivar, 480 m, light semide- ciduous forest, Jan. 1985, Fernandez 778(POKT, YEN). Sarcoglotds C. Presl Sarcoglottis stergiosii Carnevali & I. Ramirez, sp. nov. TYPE: Venezuela. Territorio Federal Amazonas: Departamento Rio Negro, selva plu¬ vial, alrededores de San Carlos de Rio Negro y camino que conduce a Solano, 100-120 m, 23-29 July 1982, Stergios & Aymard 4400 (holotype, VEN; isotypes, PORT, TFAV). Fig¬ ure 12. Plantae 10-16 cm alta. Folia 3-5, pseudopetiolata; lamina usque ad 6 cm longa, 1.8 cm longa, elliptica vel late elliptica. Bracteae anguste ovato-ellipticae, usque ad 1 3 mm longae; floribus albescentibus, parvulis, extus pi¬ losis; sepalis 3-nervatis, 9.5-10 mm longis, 1.5-2 mm latis, anguste ellipticis; petabs 3-nervatis, 7.5 mm longis, 1 mm latis, lineari-ellipticis; labello 7-nervato, 13 mm longo, 2.5 mm lato, ambitu lineari-obovato, lobo apicali ovato-subrhombico, undulato, 2.7 mm longo, 2.5 mm lato, disco in triente superiore pubescente; columna usque ad 15 mm longa. Terrestrial herbs, 10-16 cm high. Roots few, short, fleshy, lanuginose. Leaves 3-5, 2-6 cm long, 1-1.8 cm wide, present at anthesis, rosulate, some¬ what oblique, elliptic to broadly elliptic, acute to rarely obtuse, basally cuneate and attenuate into a 1 - 1 .5-cm-long channeled pseudopetiole, adaxially solid purple-brown or blotched or marbled with olive- green, abaxially deep green or solid purple. Scape 9-15 cm high, erect, laxly pubescent, terete, clothed by 4-6 sheaths 1.3-2. 7 cm long, 4-4.5 mm wide, narrowly elliptic, acuminate, glabrous to glabres- cent, longer than the internodes. Inflorescence densely 3-7-flowered, the rachis 2.1-3 cm long, subterete, laxly long pubescent. Flowers small for the genus, totally white. Ovary 12-15 mm long, slightly arcuate, subdensely long-pubescent. Floral bracts 9.5 13 mm long, 2. 1-2.8 mm wide, nar¬ rowly ovate-elliptic, acuminate. Sepals externally subdensely pubescent; dorsal sepal 8 10 mm long, about 1.5 mm wide, narrowly elliptic, acute to ob¬ tuse, 3-nerved, concave; lateral sepals subparallel to column, with spreading apices, basally long-de¬ current on ovarian wall, the free portions 8. 5-9. 5 mm long, ca. 2 mm wide, narrowly oblong-elliptic, obtuse, falcate. Petals 6. 5-7. 5 mm long, ca. 1 mm wide, 3-nerved, connate with dorsal sepal for % of their length, the apices somewhat divergent, nar¬ rowly elliptic, falcate, acute to subobtuse, glandular and with ciliate margins. Labellum 12-13 mm long, 2.5 mm wide, long-unguiculate, 7-nerved, linear- obovate, basally long-sagittate; basal lobes linear, 2. 6-2. 8 mm long; blade glabrescent, gradually wid¬ ened toward apex from a narrowly oblong basal zone, slightly contracted in apical !4 and then broadened into a deflexed apical lobe; apical lobe 2. 5-2. 7 mm long, ca. 2.5 mm wide, broadly ovate-subrhombic, obtuse, finely and densely short-pubescent, coarsely undulate. Column 5.6 mm long, narrow, ventrally pubescent; rostellum 1.2 mm long, laminar, soft, truncate-emarginate apically; stigmata confluent; anther narrowly ovate. Paratype. VENEZUELA. Territorio Federal Amazonas: IVIC Study Area, 4 km E of San Carlos de Rio Negro, 120 m, 01°56'N, 67D04'W, 21 Nov. 1977, Liesner 3728 (VEN). This new species can be distinguished from other species of Sarcoglottis by the following combination of features: small size ( to 16 cm high); leaves solid purple-brown, blotched or marbled with olive-green, present al anthesis; inflorescences 3-7-flowered; flowers small, totally white; and broadly ovate- subrhombic, coarsely undulate apical lobe of the labellum. Sarcoglottis stergiosii is known only from the drainage of the Casiquiare and Negro rivers, where it is quite rare and forms small colonies in rather open, wet forests. Acknowledgments. We are indebted to the late J. A. Steyermark and the Flora of the Venezuelan Guayana Project for financial support, including a trip to AMES for the first author, and for permitting Volume 3, Number 2 1993 Carnevali & Ramirez Venezuelan Orchids IX 125 us the use of several of their unpublished plates. B. Manara prepared all the illustrations unless other¬ wise indicated and reviewed some of the Latin de¬ scriptions; J. Dwyer and R. Gereau gave advice with other Latin diagnoses; their help is gratefully ac¬ knowledged. P. A. Berry, C. H. Dodson, A. Gentry, R. Gereau. B. K. Holst, G. A. Luer, J. MacDougal, and G. A. Romero offered valuable suggestions or provided important information to improve the manuscript. The curators of the following herbaria kindly made their specimens available: AMES, MO, NY, PORT, SEL, US, VEN. The Orchid Herbarium of Oakes Ames kindly permitted us the use of several published and unpublished plates, prepared by the late G. C. K. Dunsterville; J. Myers and C. Reynel prepared illustrations for Eloyella panamensis and Lepanlhes marahuacensis, respectively. Literature Cited Dressier, R. L. 1990. The major clades of the Orchi- daceae-Epidendroideae. Lindleyana 5: 117-125. Dunsterville, G. C. K. 1972. Some orchids of Brazil's highest highlands. Bradea 1: 83-121. - & L. A. Garay. 1959. Venezuelan Orchids Illustrated, vol. 1 . Andre Deutsch, Amsterdam. Foldats, E. 1970a. Orchidaceae, Brachionidium Lind- ley. Flora de Venezuela 15(2): 5-13. - . 1970b. Orchidaceae, Pleurothallis R. Br. Flora de Venezuela 15(2): 173-453. Garay, L. 1956. The genus Brachionidium Lindley. Canad. J. Bot. 34: 721-743. - & H. Sweet. 1972. Notes on West Indian Orchids II. J. Arnold Arbor. 53: 390-398. Hurst, C. C. 1902. Mendel’s principles applied to orchid hybrids. J. Roy. Hort. Soc. 27: 614-624. Luer, C. 1986. leones Pleurothallidinarum I. System- atics of the Pleurothallidinae (Orchidaceae). Monogr. Syst. Bot. Missouri Bot. Card. 1 5. Steyermark, J. A. 1982. Relationships of some Vene¬ zuelan forest refuges with lowland tropical floras. Pp. 183-220 in G. T. Prance (editor), Biological Diver¬ sification in the Tropics. Columbia LJniv. Press, New York. Una Nueva Especie del Genero Erythroxylum (Erythroxylaceae) de Antioquia, Colombia Alvaro Cogollo P. Herbario J AUM, Fundacion Jardin Botanico, “Joaquin Antonio Uribe,” Apartado Aereo 5 1 407 , Medellin, Antioquia, Colombia John J. Pipoly III Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. RESUMEN. Erythroxylum plowmanlanum es una nueva especie de la region alrededor de la cuenca del Rio Claro, Colombia, una zona conocida como rica en especies endemicas sobre rocas calcareas. La nueva especie se describe y se ilustra, y se discute su parentezco. ABSTRACT. Erythroxylum plowmanianum is a new species from the region of the basin of the Claro River, Colombia, an area known to be rich in en¬ demic species that grow on calcareous rocks. The new species is described and illustrated, and its re¬ lationships are discussed. La region del Rio Claro forma parte del Valle Medio del Rio Magdalena, en las faldas orientales de la Cordillera Central de Colombia. Esta cuenca se conoce por sus bosques humedos tropicales re- manentes en las cuales actualmente se esta llevando a cabo un inventario intensivo de la fitodiversidad. A1 determinar especimenes para la puhlicacion del listado de la flora, encontramos una nueva especie del genero Erythroxylum , descrita a continuacion. Erythroxylum plowmanianum Cogollo & Pi¬ poly, sp. nov. TIPO: Colombia. Antioquia: Mcpio. San Luis, Canon del Rio Claro, sector occidental, margen izquierda, subida a la Do- lina, 350-475 m, 8 jul. 1983 (fl, fr), Cogollo tV Horjn 47 6 (holotipo, JAUM; isotipos, COL, HUA, MO). Figura 1. Quoad stipulas aristatas manifeste longitudinaliter stria- tasque sectioni Rhabdophyllo O. E. Schulz pertinet, et intra illam ob laminas membranaceas ellipticas vel oblon- go-obovatas, stipulas persistentes, ad E. Jimhriatum Pey- ritsch valde arete affinis, sed ab ea rainulis longitudinaliter porcatis (non angulatis), aristis stipulinis 2 (non 3), inte- gerrimis (nec fimbriatis), pedicellis fructiferis 3-3.5 (non 5) mm longis, lobis calycinis 0.5-1 (non 1-1.3) mm longis ad apices attenuatis (nec acutis) carinatis (nec planis) denique drupa oblonga vel ellipsoidea (non ovoidea) 10- 13 (nec 6 8) mm longa statim separabilis. Arbusto o arbol hasta 9 m de alto; ramitas fle- xuosas, canaliculadas a teretes, 1-1.5 mm de dia- metro, entrenudos 5-30 mm de largo; corteza ma- rron, lisa a ligeramente fisurada longitudinalmente, escasamente lenticelada; estipulas persistentes, dis- ticas, erectas, adpresas por la base del tallo, mem¬ branaceas, triangular-ovadas, 1-2 mm de largo, api- ce agudo, no fimbriado, con dos lacinias 0.3-0. 5 mm de largo, el margen entero. Hojas disticas, la¬ minas membranaceas, elipticas, u oblongo-ovadas, apice acuminado, base aguda, 4-9.5 cm de largo, 1.1 -3. 9 cm de ancho, nervadura central y secun¬ daria emergente en la haz y enves, los nervios se- cundarios de 8-11 pares, rectos, anastomosandose de 4-7.5 mm del margen, enves no lineado, pero con panel central verdoso-palido; peciolos subtere- tes, 2-3 mm de largo, ca. 1 mm de diametro, ligeramente pianos adaxialmente. Flores fascicula- das, bisexuales, 1-2 por nudo, blanquecino-cre- mosas; bracteas florales membranaceas, oblongas, concavas, 0.5-1 mm de largo, abruptamente acu- minadas en el apice; pedicelo cilindrico, estriado, en antesis 2-3 mm de largo, 0.5 mm de diametro, en fruto elongandose hasta 3.5 mm de largo; caliz 1 .5- 2 mm de largo, dividido V3-% de su longitud, los lobulos membranaceos a cartaceos, ovados, 0.5-1 mm de largo, apice atenuado, carinados, concavos, no recurvados en fruto; petalos membranaceos, ova¬ dos a elipticos, 2. 8-3. 2 mm de largo, 1.3- 1.5 mm de ancho, apice redondeado, ligeramente concavos, con un surco central prominente en la parte abaxial, el margen entero; ligula bilobada, ca. 2 mm de largo, la auricula abaxial suborbicular, 1.3- 1.5 mm de largo, 0.8-1 mm de ancho, la auricula adaxial 0.8- 1 mm de largo, 0.6-0. 8 mm de ancho; copa esta- minal 1.5-2 mm de largo; estambres dolichistiladas, los filamentos antisepalosos, 0.8-1 mm de largo, filamentos antipetalosos 1.3-1. 5 mm de largo, las anteras suborbiculares, 0.2-0. 3 mm de largo y an¬ cho; ovario elipsoide, 1.3- 1.5 mm de largo, 0.6- 0.8 mm de diametro, truncado en el apice, los estilos Novon 3: 126-128. 1993. Volume 3, Number 2 1993 Cogollo & Pipoly Erythroxylum plowmanianum 127 Figura 1. Erythroxylum plowmanianum Cogollo & Pipoly. — A. Habito. — B. Apice ramular, rnostrando las estipulas biaristadas. — C. Inflorescencia, rnostrando las ligulas y copa estaminal. — D. Petalo y ligula. — E. Androceo, rnostrando los dos niveles de estambres. — F. Pistilo. — G. Lobulo calicino, rnostrando el apice atenuado. — H. Fruto. AG, del isotipo. H, de Cogollo et al. 4275. 128 Novon 3, unidos solo en la base, ca. 1 mm de largo, el estigma deprimido-capitado, ca. 0.3 mm de largo. Fruto drupaceo, oblongoide a elipsoide, 10-13 mm de largo, 3-5 mm de diametro, rojo al madurar, agudo u obtuso en el apice, endocarpio variable- mente sulcado. Distribution. Endemica a la region del Rio Claro, y sus alrededores, en el Municipio de San Luis, Departamento de Antioquia, Colombia, 350-580 m. Ecologia. Erythroxylum plowmanianum crece en bosques humedos con afloramientos calcareos, actualmente relegados a remanentes debido a la destruccion forestal. Etimologfa. Para nosotros es muy grato dedicar esta especie a la memoria de Iimothy Plowman, anteriormente del Field Museum of Natural History, autoridad pre-eminente en la sistematica de la fa- milia Erythroxylaceae y gran estudiante de la flora colombiana. Erythroxylum plowmanianum pertenece a la seccion Rhabdophyllum Schulz debido a sus esti- pulas longitudinalmente estriadas y aristadas. La seccion contiene aproximadamente 45 especies den- tro de las cuales E. plowmanianum es muy afin a E. fimbriatum Peyritsch. Sin embargo, se separa k. plowmanianum facilmente por sus ramitas porca- tas, estipulas biaristadas, enteras, pedicelos mas cor- tos, lobulos calicinos 0.5- 1 mm de largo, atenuados, y carinados, y la drupa oblonga o elipsoidea y mas larga. Pardtipos. COLOMBIA. Antioquia: Mcpio. de San Luis, Canon del Rio Claro, sector occidental, margen izquierda, subida a la Dolina, 350-450 m, 25 oct. 1983 (fl, fr), Cogollo 814 (COL, HUA, JAUM, MO), 3 die. 1983 (fr), Cogollo 1036 (COL, JAUM, MO), 9 mar. 1984 (fr), Cogollo 1442 (COL, JAUM, MO), 30 mar. 1984 (fr), Cogollo 1470 (COL, JAUM, MO), 2 sep. 1984 (fl, fr), Cogollo 1905 (COL, HUA, MO); vereda Las Confusas, Finca Las Confusas, 6°03'N, 74°48'W, 350- 500 m, 10 abr. 1990 (fl, fr), Cardenas et al. 2677 (COL, JAUM, MO); corregimiento El Prodigio, 6°06'N, 74°48'W, 350-580 m, 24 jun. 1990 (fr), Cardenas et al. 2845 (COL, JAUM, MO), 22 sep. 1990 (fl, fr), Cardenas et al. 2920 (COL, JAUM, MO); Mcpio. Puerto Triunfo, alrededores de la gruta “El Condor,” 5°56’N, 74°50'W, 350 m, 22 oct. 1989 (fr), Cogollo et al. 4275 (COL, JAUM, MO). Agradecimientos. Deseamos agradecer la cola- boracion de Dayron Cardenas por su trabajo en el campo, la de Luz Marina Velez, por colaboracion administrativa, y el apoyo del Antonio Jose Lopez de Mesa, actual director de la Fundacion Jardin Botanico, "Joaquin Antonio LJribe.” Investigaciones en diversidad biologica en Colombia estan subven- cionadas por la John D. and Catherine T. MacArthur Foundation al segundo autor. Apreciamos el trabajo de la dibujante Linda Ellis por el dibujo aqui pre- sentado. Paspalum longiaristatum (Poaceae: Paniceae), a New Serpentine Endemic from Goias, Brazil, and the First Awned Species in the Genus Gerrit Davidse Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A. Tarciso S. Filgueiras Reserva Ecologica do IBGE, Caixa Postal 08770, 70200-200 Brasilia, DF, Brazil Abstract. Paspalum longiaristatum of subgenus Ceresia is described and illustrated. It is tbe first known awned species in the genus and the first known annual species in the subgenus. During routine determinations of grass specimens from the Brazilian state of Goias, an unusual annual grass was studied, which we consider to represent an undescribed species of Paspalum L. Paspalum longiaristatum Davidse & Filgueiras, sp. nov. TYPE: Brazil. Goias: Niquelandia, Ma- cedo, 14°18'S, 48°23'W, campo limpo, 13 abr. 1992, 7. S. Filgueiras 2277 (holotype, IBGE; isotypes, B, FLAS, ICN, ISC, K, MO, NY, P, SI, SP, R. RB, UB, UFG, US). Figure 1. Ab omnibus speciebus Paspali glurna superiore et lem- mate inferiore aristatis optime distincta. Speciebus subge- neris Ceresia (Persoon) Reichenbach simile sed habitu annuo, gluina superiore latissima super medianam partem absimile. Tufted, delicate annual. Culms 15-36 cm tall, with 4-9 elongated internodes, branching sparingly from the lowermost nodes; internodes hollow, gla¬ brous, stramineous to purplish; nodes papillose-hir¬ sute, dark. Leaves mostly cauline; sheaths rounded on the back, papillose-hirsute with hairs mostly 0.5- 1 mm long, the midrib prominent, the margins free; ligule a ciliate membrane, the membrane 0.5-0. 8 mm long, tbe cilia 0.2-0. 8 mm long; blades 4-8.2 cm long, 1-2 mm wide, flat, linear, papillose-hirsute, the marginal hairs to 3 mm long, slightly narrowed basally, acuminate apically, the midrib projected abaxially. Inflorescence of 1 — 2(— 4) racemosely ar¬ ranged, unilateral racemes, solitary from the up¬ permost leaf sheath; peduncle included to well ex- serted, glabrous to sparsely pubescent; rachis of the main axis 1.3-2 cm, glabrous to papillose, rarely extending to 3 mm beyond the base of the upper raceme as a naked bristle, with a tuft of hairs at the base of each raceme; racemes 2-6.8 cm long, arcuate, ascending at anthesis, divergent to slightly reflexed at caryopsis maturity, acuminate apically; raceme rachis extending beyond the spikelets to an acuminate apex, prominently winged, the wings 4.5- 6 mm wide, completely enclosing the spikelets, the central portion herbaceous, green, inconspicuously nerved, the middle portion membranous, purple, nerveless, the outer portion membranous, colorless, nerveless, abaxially papillose-ciliate, adaxially gla¬ brous, marginally minutely ciliolate; spikelets soli¬ tary, alternatively arranged on each side of the rachis, the pedicels 0.1 0.2 mm, puberulent. Spike¬ lets 1.8-2. 2 mm long, ca. 0.5 mm wide, dorsally compressed, ahaxial, disarticulating below the glume and falling as one unit, narrowly elliptic-lanceolate in outline, awned, with 2 florets; lower glume absent; upper glume 1.8-2. 2 mm, as long as the spikelet, hyaline, 3 -nerved, somewhat narrowed to the base, convex in the lower !/„ flat in the upper densely pubescent on the back in the lower '/j-'/j, otherwise glabrous, the margins ciliate with hairs to 1 mm, the awn 6.0-12.2 mm, minutely antrorsely sca- berulous, flexuous, not geniculate; lower floret ster¬ ile, consisting only of a lower, awned lemma 1 .6- 1 .9 mm long, slightly longer than the upper floret, hyaline, 3-nerved, flat, densely pubescent on the back in the lower %, otherwise glabrous, the margins ciliate, the awn 0.3-2 mm long, minutely antrorsely scaberulous, flexuous; upper floret 1 .5 1 .8 mm long, bisexual, slightly coriaceous, pale; upper lemma ob¬ scurely 5-nerved, acute, glabrous, smooth but mi¬ nutely scaberulous along the margins in the upper *4, the margins slightly inrolled in the upper %\ upper palea as long as the upper lemma; lodicules absent; stamens 3, the anthers 1 .4-1.9 mm long, initially yellow, becoming purple with age; styles 2, separate; stigmas plumose, initially white, becoming yellowish with age; stamens and styles terminally exserted. Novon 3: 129-132. 1993. 130 Novon Figure 1. Paspalum longiaristatum Davidse & Filgueiras. — A. Habit. — B. Ligular area of the leaf. — C. Portion of the winged rachis with the spikelets fallen, showing the minute pedicel bases. — D. Spikelet at the beginning of anthesis. — E. Upper glume. — F. Lower lemma. — G. Upper floret, lemma side. — H. Upper floret, palea side. (Based on Brooks et al. 144.) Volume 3, Number 2 1993 Davidse & Filgueiras Paspalum longiaristatum 131 Caryopsis 1-1.2 mm; embryo h/[u-Vul as long as the caryopsis; hilum punctate, basal, ca. Zw as long as the caryopsis. Paspalum longiaristatum was found growing in huge populations in the campo limpo form of cerrado on serpentine soils. Plants were sampled in two ran¬ dom plots of one square meter each and gave the following results: 93 plants of P. longiaristatum, plus 5 plants of 4 other species; and 7 1 plants of P. longiaristatum, plus 4 plants of 4 other species. The combination of abaxial, plano-convex spike- lets, a slightly coriaceous upper lemma with its mar¬ gins inrolled, broadly winged unilateral racemes, and absence of a lower glume point to a relationship with Paspalum. Paspalum is a genus of about 330 species (Clay¬ ton & Renvoize, 1986), none of which have spikelets with awns or even the suggestion of a mucro. The conspicuously awned spikelets of P. longiaristatum thus make this a most incongruous species in the genus. For this reason generic status was seriously considered. However, this has not been adopted for the following reasons. First, awn development is notoriously variable at the generic and higher taxonomic ranks in the family and may sometimes be quite variable within a spe¬ cies. Numerous instances are known of genera with a minority of awned species and a majority of awnless species or vice versa. Some examples of genera in the Paniceae with variable awn development among their species are Digitaria Haller, Eehinochloa P. Beauvois, Eriochloa Kunth, Hymenachne P. Beau- vois, Melinis P. Beauvois, Mesosetum Steudel, Pun icum L., Poecilostachys Hackel, Pseudoechino- laena Stapf, Rhynchelytrum Nees, and l rochloa P. Beauvois. Especially notable among these genera are the following two examples: (1) Panic um aristellum Doell has shortly awned glumes, the only awned species in a genus of approximately 470 species. Despite a membranous upper lemma, which might suggest a relationship to the genus Hymenachne , Zuloaga & Soderstrom (1985) maintained this species in Pan- icum. (2) Digitaria aristulata (Steudel) Stapf has an awned lower glume, but in other respects closely resembles other species of Digitaria. It is the only awned species in a genus of approximately 230 species. (We here recognize Digitariella remotig- luma De Winter, an awned species with a strongly developed callus from southern Africa, as belonging in its own monotypic genus, although some recent authors (Clayton & Renvoize, 1986; Gibbs Russell et ah, 1990) have merged it into Digitaria.) In the Chloridoideae a similar example is known in Eragrostis Wolf, where the Namibian E. aristata De Winter is the only awned species in a genus of about 350 species, although there are numerous species with acuminate lemma apices, some of which grade into small awns, such as in E. dinteri Stapf. Thus, admitting awned species to otherwise large, awnless genera is not without precedent. Emphasizing the fact of variability of awn de¬ velopment at another level is that the awn of the lower lemma of P. longiaristatum ranges from 0.3 to 2 mm in the same individual. Second, there is little doubt about the relationship of the new species to the major infrageneric groups of Paspalum. The racemosely arranged racemes, the broad, partially purple, membranous rachis wings, the conspicuously hairy, lanceolate spikelets, the upper glume convex in the lower half and flat in the upper half, and the slightly coriaceous, pale upper lemma and palea indicate a close relationship to Paspalum subg. Ceresia. Although the characters just enumerated do not leave any doubt about its overall relationship with subgenus Ceresia , para¬ doxically this species does not seem closely related to any of the described species and can be imme¬ diately distinguished by its annual habit, small, awned spikelets, and the upper glume widest above the middle. No other annual species is known in this subgenus. Although the annual habit of the new species is also distinctive, it, like awn development, is a labile character that has repeatedly and inde¬ pendently evolved in many grass genera. There is little doubt that if it were not for the prominent awns, this species would be easily accom¬ modated in subgenus Ceresia. The single difference is therefore considered to be insufficient to establish a new% monotypic genus. The broader question of whether subgenus Ceresia itself deserves generic status is beyond the scope of the present study, but seems eminently worth reconsidering. Other American genera with racemose inflores¬ cences and broadly winged rachises that were con¬ sidered in evaluating the relationship of this species include Mesosetum and Thrasya. Mesosetum can be excluded because its spikelets are adaxial with a developed lower glume, and the inflorescences are always solitary racemes (Filgueiras, 1989). Thrasya can be excluded because it is characterized by al¬ ternately adaxial and abaxial, awnless spikelets borne in one row due to the adnation of the pedicels of the spikelet pairs to the rachis (Burman, 1985). The lack of functional lodicules seems to be as¬ sociated with the terminal exsertion of the stamens and stigmas in the newT species. This has also been observed in Paspalum ceresia (Kuntze) Chase, the type species of subgenus Ceresia, although some 132 Novon specimens seem to have minute (nonfunctional?) lodicules. Other species of the suhgenus have nor¬ mally developed lodicules and terminally or laterally exserted stamens and styles. At caryopsis maturity, when the leaves are also losing their chlorophyll, each raceme becomes di¬ vergent to slightly reflexed. As part of this process, the pulvinus at the base of each raceme twists so that the wing margins point upwards and the midrib of the raceme faces downwards. In this way a nar¬ row, curved, troughlike structure is formed, which holds the spikelets, with the spikelet awns protruding beyond the raceme wings. Presumably this structure is functional in dispersal, but exactly how is not known, although T. Filgueiras observed in the field that the flexuous awns twist together, and because of this the spikelets tend to disarticulate in large groups. The species name is meant to emphasize the prominently awned upper glume of this species. Paratypes. BRAZIL. Goias: Macedo, ca. 15 kin N of Niquelandia, S-facing hill slope, stable peridotite/dunite- based scree and flat area below, ca. 500-800 m E of nickel workings, campo-cerrado, ca. 1,000 m alt., 14°18'S, 48°23'W, 21 Apr. 1988, Brooks, Reeves, Baker & Dias Ferreira BRASPEX 144 (MO, NY, SI, UFG); a 18 km de Niquelandia, local chamado Macedo, 10 June 1992, Filgueiras 2342 (BM, IBGE, ICN, ISC, K, MO, P, SI, SP, UB, US); Niquelandia, 14 Sep. 1991, Filgueiras 2003 (IBGE, SP). Acknowledgments. Filgueiras gratefully thanks CAPES (Brazilian Post-Graduation Agency), Brasi¬ lia, Brazil, for a fellowship that enabled him to work at the Missouri Botanical Garden. He also thanks the Missouri Botanical Garden for making their re¬ search facilities available to him. We thank Robert H. Brooks for a duplicate collection, which alerted us to this species. The excellent illustration was made by Vladimiro T. Dudas. Literature Cited Burman, A. G. 1985 [1987], The genus Thrasya H.B.K. (Gramineae). Acta Bot. Venez. 14: 7-93. Clayton, W. D. & S. A. Renvoize. 1986. Genera Gra- minum. Kew Bull., Addit. Ser. 13: 1-389. Filgueiras, T. S. 1989. Revisao de Mesosetum Steudel (Gramineae: Paniceae). Acta Amazon. 19: 47-114. Gibbs Russell, G. E. et al. 1990. Grasses of Southern Africa. National Botanic Gardens, South Africa. Zuloaga, F. 0. & T. R. Soderstrom. 1985. Classification of the outlying species of New World Panicum (Po- aceae: Paniceae). Smithsonian Contr. Bot. 59: 1- 63. Murdannia cryptantha (Commelinaceae), a New Species with Cleistogamous Flowers from Australia and Papua New Guinea Robert B. Faden Department of Botany, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. ABSTRACT. Murdannia cryptantha (Commelina- ceae), a new species from Australia and Papua New Guinea, is described. It is most closely related to M. graminea, and it is the first species of Murdannia reported to have cleistogamous flowers. The species of Murdannia Hoyle once were most¬ ly included in the genus Aneilema R. Brown (Faden, 1978). Thus, when I attempted to determine the correct generic placement for all epithets in Anei¬ lema, I also surveyed the species of Murdannia (Faden, 1991). In the course of that investigation I came upon slender plants of a Murdannia species from northern Queensland, Australia, that generally resembled the widespread species M. graminea (R. Brown) G. Brueckner except lor the presence of sessile, axillary flowers and capsules, in addition to terminal inflorescences and infructescences. It was not until I obtained living material (A. Faden 2/91) that I was able to confirm that such plants represent a distinct species. Murdannia cryptantha Faden, sp. nov. TYPE: Australia. Queensland. Port Curtis Dist.: road from Rockhampton to Emerald, 3 km beyond Gogango, fenced area across railway line, bri- galow bushed-grassland with frequent small temporary water holes, edge of temporary pools, 3 Mar. 1991, A. J. Faden 2/91 (holotype, US; isotypes, K, BRI). Figures 1-2. Herba perennis foliis omnibus caulinibus 1.5—7(13) cm longis, 1.5 -5. 5 mm latis, thyrso terminali et floribus axillaribus sessilibus vel subsessilibus cleistogamis, capsulis (5 )6 10(12) mm longis, ca. 2 mm crassis, loculis (4-)6- 8(-15)-seminalibus, seminibus 0.6-0.9( - 1 . 1 ) mm latis. Perennial with thickened, sometimes fusiform roots to 3 mm thick. Shoots erect to ascending, often rooting at the lower nodes, to ca. 35 cm long, unbranched or sparsely branched basally. Inter¬ nodes 2-7 cm long, glabrous to pubescent, some¬ times the pubescence confined to a longitudinal line. Leaves all cauline, distichous or in an open spiral; sheaths 0.5-2 cm long, often splitting to the base, ciliate or ciliolate at the apex and along the fused edge, sheath surface glabrous or pubescent; lamina sessile, linear-lanceolate, decreasing in length dis- tally on the flowering shoot, conduplicate or planar, often recurved at the apex, 1 .5 — 7 ( — 13) cm long, 1.5-5. 5 mm wide, with base rounded to somewhat amplexicaul and apex acuminate to acute, margins undulate or planar, ciliate or ciliolate at the base of the lamina, sometimes scabrid at the apex, otherwise glabrous, both surfaces of the lamina glabrous to sparsely pubescent, sometimes only the abaxial sur¬ face pubescent. Inflorescences axillary and terminal, glabrous, the axillary enclosed within a leaf sheath and consisting of a single sessile, bracteolate cincinnus that pro¬ duces 1 -4 sessile or subsessile cleistogamous flowers, the terminal inflorescence a lax thyrse consisting of 1- 4 cincinni, each cincinnus to 4 cm long and producing up to 8 chasmogamous flowers. Bracts of the cincinni of the terminal inflorescences linear to ovate-lanceolate, 1-9 mm long; bracteoles of the terminal inflorescences ovate to lanceolate-elliptic, 0.7-2. 5 mm long. Flowers all bisexual or an occasional chasmoga¬ mous one staminate. Chasmogamous flowers slightly zygomorphic; pedicels erect, 4-8.5 nun long in flow¬ er, to 9 mm long in fruit, glabrous. Sepals subequal, boat-shaped, elliptic to oblong-elliptic, 3-7 mm long in flower, to 8 mm long in fruit, glabrous. Petals subequal, not clawed, ovate to suborbicular, 5 9 mm long, pale purple to mauve or lavender, gla¬ brous. Filaments of stamens and staminodes free. Stamens antesepalous, equal but all bending to one side of the flower, filaments bending outwards near the middle, 2.5-4 mm long, densely bearded with moniliform hairs below the middle, anthers dorsi- fixed, elliptic or oblong-elliptic, 1-2 mm long. Stam¬ inodes antepetalous, symmetrically arranged, fila¬ ments shorter and more slender than those of the stamens, densely bearded below the middle, an- therodes basifixed, 0.5 -0.7 mm long, usually ovate with a cordate base, yellow. Ovary sessile, trigonous, 2- 2.5 mm long, trilocular, glabrous, locules up to 13-ovulate, ovules uniseriate, style bent away from Novon 3: 133-136. 1993. 134 Novon Figure 1. Murdannia cryptantha Faden. — A. Partial habit. — B. Chasmogamous flower, front view. — C. Chas- mogamous flower, front/side view. — D. Staminode. — E. Gynoecium. — F. Cleistogamous flower, slightly opened. — G. Cleistogamous flower, front sepal and two petals pulled back. — H. Cleistogamous flower, sepals removed. All from A. Faden 2/91 except inflorescence on left shoot in A added from Brass 18709 (GH). Volume 3, Number 2 1993 Faden Murdannia crypt ant ha 135 Figure 2. Murdannia cryptantha Faden. A. Mature capsule and bud in axillary inflorescence. — B. Mature capsules from cleistogamous flowers in axillary inflorescence. — C. Mature capsule from cleistogamous flower, before dehiscence. - D. Capsule from cleistogamous flower, dehisced. E. All of the seeds from a single capsule locule, showing arrangement in locule. — F. Seed, dorsal view. — G. Seed, ventral view, showing hiluin on raised ridge. — H. Seed, apical view. All from A. Faden 2/91. the stamens, straight, 1-1.5 mm long, stigma cap¬ itate. Cleistogamous flowers sessile, difficult to dif¬ ferentiate from large buds, on the one hand, and young, developing fruits, on the other hand. Sepals oblong-elliptic, 3.5-4 mm long, glabrous. Petals el¬ liptic to oblong-elliptic, 2. 5-2. 8 mm long, 0.7 I mm wide. Filaments of stamens and staminodes free. Stamens subequal, filaments ca. 2-2.5 mm long, glabrous, jointed just below a swollen summit, an¬ thers apparently dorsifixed, elliptic, ca. 0.5-0. 7 nun long, dehiscence introrse. Staminodes variable in form, filaments ca. 1.1 1.3 mm long, very slender, glabrous, antherode minute, oblong to ovate, irreg¬ ularly lobed or unlobed. Ovary sessile, ca. 2.5 mm long, trilocular, locules up to 15-ovulate, ovules uniseriate or partially biseriate, style ca. 0.5 mm long, stigma capitate. Capsules trilocular. trivalved, dehiscent at least at the apex, cylindric-trigonal, apiculate, (5 )6 10 (-12) mm long, ca. 2 mm thick, usually dark brown at maturity, glabrous, cells of the outer wall trans¬ versely elongate, locules (4— )6— 8( - 1 5)-seeded, seeds uniseriate. Seeds very variable in size, often within the same locule, rectangular to trapezoidal, rarely triangular in outline, the apical and basal seeds rounded on one end, (0.35 -)0.6 — 1 . 5( — 1 .75) mm long, 0.6—0. 9( —1.1) mm wide, testa brown to gray, rugose or verrucose and variously pitted, often far¬ inose in the depressions and around the embryotega, hilum punctiform to elliptic, often on a raised lon¬ gitudinal ridge, embryotega ± lateral. Distribution. Northern Queensland and northern Northern Territory, Australia to Papua New Guinea. 136 Novon Murdannia cryptantha is undoubtedly most closely related to the very variable M. graminea because of its geophytic habit, thyrsiform inflores¬ cence, bearded stamen and staminode filaments, and several-seeded capsule locules. Murdannia cryp¬ tantha can be distinguished by the presence of ax¬ illary, sessile or subsessile, cleistogamous flowers and capsules, and by its narrower capsules (2.5-4 mm thick in M. graminea ), more numerous seeds per locule (2-5 per locule in M. graminea), and smaller seeds (1.2-3. 9 mm long, 0.9-2 mm wide in M. graminea). It further differs by the absence of basal leaves (present in many specimens of M. graminea) and by the usually smaller leaves and terminal thyrses. In our present state of knowledge, speci¬ mens of M. cryptantha that lack fruits or cleistog¬ amous flowers might not be separable from certain small plants of M. graminea unless characters such as style length (shorter than the ovary in M. cryp¬ tantha and longer (always?) than the ovary in M. graminea) prove to be consistent and other floral differences are found. Murdannia cryptantha is the first in this genus of approximately 50 species to be recorded as having cleistogamous flowers. The best known examples of cleistogamy in the Commelinaceae are in Cotnme- lina benghalensis L. and C. forskaolii Vahl (see Faden, 1993). In the above descriptions it should be noted that, following Faden (1991: 42), seed length is the dis¬ tance between the apical and basal surfaces, i.e., the dimension parallel to the capsule length, and seed width is the distance between the lateral sur¬ faces, i.e., transverse to the locule. Paratypes. AUSTRALIA. Queensland. Cook Dist.: Douglas Creek, ca. 7 mi. SE of Mareeba, open forest on a stony ridge, 9 Apr. 1967, Brass 33523 (BRI); Lock¬ erbie, 10 mi. WSW of Somerset, alt. 30 m, wet teatree flat, 22 Apr. 1948, Brass 18361 (CANB, GH); Cape York Peninsula, Newcastle Bay, 2.5 mi. S of Somerset, alt. 50 m, wallum scrub in hollow between coastal sand dunes, 9 May 1948, Brass 18709 (CANB, GH); Mt. Molloy, moist flats, Jan. -Apr. 1941, Carr 195 [in Flecker Herbarium 7473], Carr 206 [in Flecker Herbarium 7484], Carr 207 [in Flecker Herbarium 7485] (all on one sheet) (QRS); near Mareeba, poorly drained soil with sandy surface in Eucalyptus polycarpa-E. leptophleba community, 23 Apr. 1967, Pedley 2276 (BRI); near Mareeba, layered woodland with E. alba, E. leptophleba, and Melaleuca minutifolia, 27 Feb. 1962, Webb & Tracey 5879 (BRI — 3 sheets); Cairns, [collector illegible] (BRI# 155853). Darling Downs Dist.: “Woodlands” 5 mi. SW of Westmar, gray clay soil in cleared brigalow, 14 Dec. 1961, Pedley 925 (BRI). Northern Territory: Arnhem Land Aboriginal Reserve, South Bay, Bickerton Island in the Gulf of Carpentaria, 13°45'S, 136°06'E, moist Melaleuca leucodendra stand, 11 June 1948, Specht 531 (BRI, CANB, US). PAPUA NEW GUINEA. Central Dist.: Rubulogo Creek, ca. 18 mi. N of Port Moresby, alt. ca. 140 ft., Eucalyptus savannah land, 12 Apr. 1967, Pullen 6688A (CANB; other duplicates cited but not seen: A, BRI, E, K, L, LAE). Acknowledgments. I thank the directors of the herbaria cited for the loan of specimens; an anon¬ ymous reviewer for helpful comments; Alice Tan- gerini for the illustrations; and my wife, Audrey J. Faden, for collecting the new species and maintain¬ ing my research plants for many years. Literature Cited Faden, R. B. 1978. Review of the lectotypification of Aneilema R. Br. (Commelinaceae). Taxon 27: 289- 298. - . 1991. The morphology and taxonomy of A nei- lema R. Brown (Commelinaceae). Smithsonian Contr. Bot. 76: 1-166. - . 1993. The misconstrued and rare species of Commelina (Commelinaceae) in the eastern United States. Ann. Missouri Bot. Card. 80: 208-218. Six New Species of Adenocalymna (Bignoniaceae) from Eastern South America ALwyn //. Gentry Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A. ABSTRACT. Six new species of Adenocalymna are described from Brazil and F rench Guiana. Adenocalymna is one of the genera of Bignoni¬ aceae tribe Bignonieae that is most prone to local endemism (Sutherland, 1983). Six new1 species of the genus are included in recent collections from eastern South America. Adenocalymna dichilum A. Gentry, sp. nov. TYPE: Brazil. Piauhy: Bom Jesus Sao Kai- mundo Nonato, caatinga, 11 May 1979 (II), A. Fernandes s.n. (FAC 6185), (holotype, EAC; isotype, MO). Frutex scandens, ramulis teretibus. Folia 2(-3?)-fo- liolata, interdum cirrho simplici, foliolis ovatis vel ellip- ticis, infra valde villosis. Inflorescentia floribus in racemo terminali dispositis. Calyx cupulatus, 5-dentatus, 12 14 mm longus; corolla lutea, valde bilabiata, extus puberula. Capsula ignota. Liana; branchlets terete, longitudinally striate, with scattered small inconspicuous lenticels, finely pu- berulous with small suberect whitish trichomes; pseu¬ dostipules ovate, 3-4 mm long; interpetiolar glan¬ dular fields absent. Leaves 2(-3?)-foliolate, the terminal leaflet sometimes replaced by a simple ten¬ dril, the leaflets ovate to elliptic, obtuse to emargin- ate at apex, rounded to distinctly cordate at base, 2-4 cm long, 1.5-3. 5 cm wide, above ± glabres- cently puberulous with short erect trichomes, below persistently villous, coriaceous, drying brownish above, olive below, the tertiary venation prominulous below, finer venation obscured by the indumentum; petiole 1 .5-2.5 cm long, petiolules 0.2 0.9 cm long, rather densely tomentose with multicelled trichomes. Inflorescence a terminal raceme, densely tannish tomentose, with pair of caducous obovate coriaceous bracts ca. 1.5 cm long subtending each node, each bract with several large plate-shaped glands, the pedicels 1-1.5 cm long, with pair of ca. 1-cm-long elliptic bracteoles near apex subtending calyx. Flow¬ ers with the calyx cupular, shallowly 5-dentate, 12- 14 mm long, 9-10 mm wide, tannish puberulous, with conspicuous plate-shaped glands near margin; corolla yellow, ± tubular, strongly bilabiately split almost to middle, ca. 3.5 cm long, ca. 1 cm wide at mouth of tube, the tube ca. 1.8 cm to base of split, the lower lobes narrow, ca. 1 cm long, the upper two lobes with free part ca. 3 mm long but basally fused to form 1.5-cm-long upper lip, densely puberulous outside except for base of tube; stamens subexserted, the thecae subparallel, pendulous, ca. 6 mm long; pistil and disk not examined. Fruit not seen. Distribution. Known only from the type from the caatinga of northeastern Brazil. This species is most similar to A. bracteatum (Ghamisso) DC. in its large bracts and coriaceous, densely puberulous calyx and keys out with that species in Flora Brasiliensis (Bureau & K. Schu¬ mann, 1896-1897). The most striking difference from A. bracteatum is in the deeply bilabiately split corolla that looks more like an overgrown Tynanthus flower than like that of other species of Adenoca¬ lymna. Curiously, several quite unrelated species of Bignoniaceae share this unusual floral morphology in northeastern Brazil. In addition to .1. dichilum , they are Arrabidaea tynanthoides A. Gentry, and Arrabidaea crassa Sprague (not closely related to each other), and to a lesser extent, Mansoa ven- tricosa A. Gentry. Adenocalymna fruticosum A. Gentry, sp. nov. TYPE: Brazil. Bahia: Serra do Sincora, 6 km N of Cascavel on road to Mucuge, alt. 1,200 m, 41°25'W, 1 3°6'S, 25 Mar. 1980 (fl), B. M. Ilarley, G. /,. Bromley, A. M. de Carvalho & Ci. Martinelli 26018 (holotype, GEPEC; isotypes, K not seen, MO). Suffrutex, ramulis teretibus. Folia 2-3-foliolata, ecir- rhata, foliolis ovalibus vel subrotundatis, glabris. Inflores¬ centia floribus in racemo terminali dispositis, pedicellis bracteis 1-1.5 cm longis subtends. Calyx campanulatus, minute 5-denticulatus, 8-11 mm longus; corolla lutea pallida, tubulo-campanulata, tubo extus glabro. Capsula ignota. Spindly shrub to 1 m tall; branchlets terete, lon¬ gitudinally striate-ridged, sparsely minutely puber¬ ulous; pseudostipules subulate, ca. 1 mm long, in- Novon 3: 137-141. 1993. 138 Novon terpetiolar glandular fields lacking, a ridge connecting the opposite petiole pairs. Leaves 2 3-foliolate, with¬ out tendrils, the leaflets oval to almost suborbicular, rounded to truncate or emarginate at base and apex, 3-5.5 cm long, 2.3-4 cm wide, completely glabrous, the lower surface appearing somewhat glandular lep- idote, coriaceous, drying olive, the venation raised above and below, the margin cartilaginous; petioles 1-3 cm long, the petiolules mostly very short, 0.1- 1 cm long, glabrous except for a few tiny lepidote trichomes on flattened upper surface. Inflorescence a terminal raceme, minutely puberulous with simple and thick-stellate trichomes, with a pair of olive¬ drying longitudinally ribbed foliaceous bracts 1-1.5 cm long subtending each pair of pedicels, the ped¬ icels ca. 5 mm long, densely dendroid puberulous, with a pair of ca. 1-mm-long bracteoles near base. Flowers with the calyx campanulate, truncate, mi¬ nutely 5-denticulate, 8-11 mm long, 8 11 mm wide, densely minutely tannish puberulous with den¬ droid trichomes, also with dark-drying glandular ar¬ eas; corolla (only 1 seen) creamy yellow, tubular- campanulate, 5 cm long, 1.2 cm wide at mouth of tube, the tube 4 cm long, the lobes 1 cm long, the tube glabrous, the lobes densely capitate-glandular; stamens included; pistil 3.5 cm long, the ovary ob¬ long, 2 mm long, 1 mm wide, glandular surfaced; disk thick-pulvinate, 1 .5 mm long, 3 mm wide. Fruit not seen. Distribution. Known only from the type collected in “campo geral” grassland rich in dicot herbs and shrubs and acaulous palms. This species is unusual in the genus in having an erect shrubby habit. Although there are a few erect species of Adenocalymna in the forest understory of coastal Brazil, this is the first member of its genus known to adopt the characteristic suffrutescent form so typical of the plants of the open cerrado and allied vegetations. Adenocalymna fruticosum is re¬ lated to A. apparicianum J. C. Gomes by its similar calyx, but that species differs in lacking persistent foliaceous bracts, climbing habit, and different leaves with a very different dull rough-reticulate texture and trichomes on the undersurface. Adenocalymna apparicianum occurs further north in the caatinga of Ceara and Rio Grande do Norte. Adenocalymna gracielzae A. Gentry, sp. nov. TYPE: Brazil. Para: Senador Jose Porfirio (Soz- el), Rio Xingu, 3 Dec. 1991 (fl), G. dos Santos, N. A. Rosa & M. R. dos Santos 276 (holotype, MG; isotypes, MO, MAD). Frutex scandens, ramulis teretibus, minute puberulis. Folia 2-3-foliolata, interdum cirrho simplici, foliolis ellip- ticis vel anguste ellipticis, infra sparsim puberulis saltern costa. Inflorescentia floribus in racemo terminali dispositis. Calyx cupulatus, minute 5-denticulatus, 3-6 mm longus; corolla lutea vel cremosa, anguste tubulo-campanulata, extus puberula. Capsula oblonga, haud compressa, valvis valde bialatis, puberulis, seminibus non alatis. Liana, the branchlets terete, minutely puberulous with small erect trichomes, at length glabrescent, inconspicuously striate, drying brown, with minute lenticels when older, the pseudostipules blunt-coni¬ cal, 2-3 mm long. Leaves 3-foliolate or 2-foliolate with a simple tendril or tendril scar, the leaflets elliptic to narrowly elliptic, obtuse to acuminate, the base rounded, 3-21 cm long, 1.5-8. 5 cm wide, chartaceous to subcoriaceous, main veins ± plane above, noticeably raised below, veinlets very slightly raised above and below, drying olive to olive brown above, yellowish olive to grayish olive below, usually rather shiny and varnish-surfaced, especially above, glandular-lepidote above and below, above glabrous except for short crisped trichomes on midvein, below puberulous with short crisped trichomes at least scattered along base of main veins and sometimes sparsely over surface; petiolules 0.5-2. 5 cm long, petiole 1-4 cm long, puberulous. Inflorescence a contracted terminal raceme, densely puberulous, drying tannish, a pair of small early caducous lan¬ ceolate bracts 1-2 mm long subtending each pair of pedicels in young bud, the pedicels 3-5 mm long, each with a pair of small caducous 1-2 mm long triangular bracteoles near its base, the bracteoles puberulous and sometimes glandular. Flowers with the calyx cupular, minutely 5-denticulate, 3-6 mm long, 3-5 mm wide, finely and uniformly puberulous, with a few conspicuous scattered raised hollow-cen¬ tered, pustulelike glands; corolla yellow to greenish cream, tubular-campanulate above a 2 3-cm-long narrowly tubular base, 5-7 cm long, 1-1.3 cm wide at mouth of tube, the tube 3.5-6 cm long, the lobes 0.5-1 cm long, the lobes and tube outside puber¬ ulous with thick-stellate trichomes, the tube outside also with conspicuous glands at base of lobes, inside glandular villous at level of stamen insertion; sta¬ mens didynamous, inserted 25-30 mm from base of corolla tube, the anthers included or the longest pair subexserted, the thecae divergent to subparallel, 4- 5 mm long, the staminode ca. 7 mm long; pistil 5- 5.5 cm long, the ovary linear with conspicuously raised angles, 3-4 mm long, < 1 mm wide, glabrous, becoming densely puberulous with glandular angles in young fruit; disk annular-pulvinate, 1.5 mm long, 1 .5-2 mm wide. Capsule oblong or oblong-ellipsoid, not at all compressed, the valves woody, with pair of prominent thick wings along each valve, the fruit thus somewhat tetragonal in cross section, 10-18 Volume 3, Number 2 1993 Gentry New Species of Adenocalymna 139 cm long, 5-7 cm thick and wide (including wings), drying light grayish, the midrib not visible, densely minutely puberulous, the surface slightly wrinkled- striate, the wrinkles making an acute angle with axis of Iruit; seeds thick and wingless, 2-3 cm long, 2- 3.5 cm wide. Distribution. Restricted to the margins of the Rio Xingu and its tributaries, where it mostly occurs in restingas or riverside scrub forest. This species is closely related to A. purpurascens Rushy, which is widespread in Amazonia, and to A. divaricatum Miers, which occurs in the caatinga and deciduous forest of eastern Brazil. These are the only other Adenocalymna species to have the distinctive winged fruit that characterizes the new species. Adenocalymna gracielzae differs dramat¬ ically from these species in the thick wingless seeds, presumably representing a water-dispersed deriva¬ tive of one of these widespread wind-dispersed spe¬ cies. It also differs in the unusual flower shape with a relatively long narrow corolla having the narrow basal part of the tube much elongated. It is likely that A. gracielzae represents incipient hawkmoth pollination in this otherwise mostly bee-pollinated genus. Paratypes. BRAZIL. Para: Porto de Moz, Rio Acai, 2 Dec. 1991 (fl), G. dos Santos , N. A. Rosa & M. dos Santos 260 (MG, MO), 2 Dec. 1991 (fr), G. dos Santos et al. 260 (MG, MO), 2 Dec. 1991 (st), G. dos Santos el al. 2b 1 (MG, MO); Senador Jose Porfirio (Sozel), Rio Xingu, 3 Dec. 1991 (fr), G. dos Santos et al. 280 (MG, MO), 5 Dec. 1991 (fl), G. dos Santos et al. 311 (MG, MO); Parauapebas, Reserva Biologica da Serra dos Ca- rajas, 650 m, 21 Nov. 1991 (st), G. dos Santos et al. 199 (MG, MO), 21 Nov. 1991 (fr), G. dos Santos et al. 200 (MG, MO). Adenocalymna hatschbachii A. Gentry, sp. nov. TYPE: Brazil. Parana: Morro do Arrastao, Municipio Morretes, mata pluvial encosta de morro, 200 m, 18 July 1981 (fl), Hatschbach 43936 (holotype, MO; isotype, MBM). Frutex scandens, ramulis teretibus, minute puberulis. Folia 2-3-foliolata, interdum cirrho simplici, foliolis lan- ceolatis vel elliptico-lanceolatis, membranaceis, plerumque fere glabris. Inflorescentia floribus in racemo axillari dis- positis. Calyx campanulatus, 5-dentatus, 8-12 mm lon- gus; corolla lutea, tubulo-campanulata, extus puberula; stamina subexserta. Capsula anguste oblonga, haud com- pressa, puberula, seminibus non alatis. Liana (rarely treelet?), the branchlets terete, sparsely to densely minutely puberulous with crisped trichomes, not obviously lenticellate, longitudinally minutely striate, the pseudostipules subulate, to 5 mm long. Leaves 3-foliolate or 2-foliolate with an unbranched terminal tendril or tendril scar, the leaf¬ lets lanceolate to lanceolate-elliptic, long acuminate, broadly cuneate to rounded at base, 3-17 cm long, 10.8-4.5 cm wide, membranaceous, drying gray- green with the venation light tannish below, the tertiary venation prominulously raised above and below, glabrous below or with a few trichomes at base of midvein or scattered along main veins, above puberulous along midvein; petiole 0.8 -6 cm long, puberulous, the petiolules 0.3-4. 5 cm long. Inflo¬ rescence an axillary or apparently terminal raceme, finely puberulous with whitish trichomes, the bracts caducous, linear to narrowly lanceolate, eglandular, to 1 .5 cm long, the bracteoles linear, mostly ca. 0.5 cm long. Flowers with the calyx campanulate, shal¬ lowly and broadly 5-dentate, 8-12 mm long, 5-10 mm wide, densely minutely puberulous, drying tan¬ nish to brownish, thin and eglandular or with few inconspicuous glands; corolla yellow, tubular-cam- panulate above the 1 - 1 ,.5-cm-long basal tube, 5- 6.5 cm long, 12 cm wide at mouth of tube, the lobes ca. 1 cm long, the tube 4. 5-5. 5 cm long, rather densely minutely puberulous outside and on lobes inside, villous with long multicellular trichomes at level of stamen insertion; stamens didynamous, the anthers divaricate, 4-5 mm long, at least the longest pair exserted or subexserted; pistil with the ovary narrowly oblong, strongly tetragonal with raised angles, 4-5 mm long, ca. 1 mm wide, glabrous; disk pulvinate-patelliform, 1 mm long, 2-3 mm wide. Fruit narrowly oblong, the valves woody, strongly convex, 9-17 cm long, 3. 5-4. 5 cm wide, finely tannish puberulous, the surface usually minutely longitudinally striate, also with a few irregular widely scattered warty projections; seeds thick, wingless, 1.5-2 cm long, 2-2.5 cm wide. Distribution. Restricted to the wet subtropical forests of southern coastal Brazil, occurring near sea level from Parana to southern Sao Paulo, and perhaps north to Rio de Janeiro at higher altitudes. This species is most closely related to A. dusenii Kranzlin, which has similar thin calyces and pale intricately prominulous venation on the leaf under¬ surface, but differs in larger bracts and bracteoles, usual presence of conspicuous calyx glands, thicker distinctly coriaceous leaves, and the corolla more infundibuliform-campanulate with included anthers. It is also closely related to A. comosum (Chamisso) DC., and the Neuwied collection, consisting of a single leaflet and flower, was identified as “ Adeno- calymma spec, fors comosum var. lanceolata ” (af¬ ter an original determination with an unpublished name "‘Bignonia divergens”). Adenocalymna hatschbachii differs from A. comosum in the much narrower long-acuminate membranaceous leaflets 140 Novon with a very different less prominulous-veined surface texture. The Brade collection from Rio de Janeiro was identified by Laroche as A. guilleminii (i.e., A. grandifolium var. guilleminii ), a taxon that is veg- etatively rather similar but has a completely differ¬ ent, much larger and thicker calyx. This specimen, which has thicker calyces, nonexserted anthers, and was described as a 3-m treelet, may not be conspe- cific with the material from Parana. One specimen that may be referable to this species is not included in the above description. This is Hatschbach 1925 1 (MBM, MO) from Rio Cambara, Mun. Paranagua, Parana, 50 m elev., which differs in more long-exserted anthers and more elliptic leaf¬ lets. It is possible that the corolla is also somewhat more zygomorphic than in typical A. hatschbachii. Whether this collection represents an extreme form of A. hatschbachii or a closely related, undescribed taxon cannot be determined without additional ma¬ terial. Paratypes. BRAZIL. Parana: Reserva Biologica de Sapitanduva, Mun. Antonina, 40 m, 25°30'S, 48°45'W, 24 Jan. 1985 (fr), Gentry & Zardini 49788 (MBM, MO), 4 Apr. 1982 (fl), Hatschbach 45193 (MO); Rio Canavieiras, Mun. Guaratuba, 20-50 m, 21 July 1987 (11), Hatschbach & Cordeiro 51271, 2 Oct. 1986 (fr), Silva & Cervi 193 (MBM, MO); Rio Brumado, Mun. Morretes, 4 July 1984 (fl), Hatschbach 48076 (MBM, MO). Rio de Janeiro: Angra dos Reis, Serra do Mar, 600 m, 29 June 1935 (fl), Brade 14910 (B, MO, RB); sin. loc. “Brasilia,” Neuwied 479 (P). Sao Paulo: Sete Barras, S entrance to Carlos Botelho State Park, 70 m, 46°55'W, 24°22'S, 4 Sep. 1987 (st). Gentry, Dias & Franco 59037 (MO, SPF). Adenocalymna saulense A. Gentry, sp. nov. TYPE: French Guiana. Saul, 3°37'N, 53°12'W, between Carbet and “steep, mean hill,” 250- 350 m, 27 Aug. 1988 (fl), Mori el al. (10 collectors) 19192 (holotype, MO; isotypes, CAY, NY not seen). Frutex scandens, ramulis teretibus. Folia 2-3-foliolata, interdum cirrho simplici, foliolis ellipticis vel oblongo-el- lipticis, coriaceis, infra valde aspris. Inflorescentia floribus in racemo ramifloro dispositis. Calyx campanulatus, 2-3- labiatus, 8-10 mm longus; corolla alba, tubo extus glabra, lobis puberulis. Capsula ignota. Liana, the branchlets terete, minutely puberulous, conspicuously swollen at nodes. Leaves 3-foliolate or 2-foliolate with a terminal tendril or tendril scar, the leaflets elliptic to oblong-elliptic, obtuse to acu- tisli at apex, rounded to truncate at base, 7-17 cm long, 3-7.7 cm wide, coriaceous, drying gray above, gray to gray-green below, glabrous above or with a very few minute trichomes at base of midvein, below strongly asperous from the scattered very short sub¬ erect trichomes, these occasionally in part forked; petiole 2-3 cm long, the petiolules 1-3 cm long, puberulous with minute erect trichomes, distinctly subwoody. Inflorescence a ramiflorous raceme (7- flowered in type), finely grayish puberulous with a mixture of simple and stellate trichomes, each ped¬ icel subtended by a minute bract < 1 mm long, bracteoles absent. Flowers with the calyx campan- ulate, irregularly 2-3-labiate, 8-10 mm long, 5-7 mm wide, densely minutely puberulous with mixture of stellate and simple trichomes, drying grayish with few striking, large (nearly 1 mm long), black elliptic glands near middle; corolla white, yellow in bud, tubular (?, only in bud in holotype), the tube glabrous except toward apex, tube apex and lobes puberulous outside; ovary oblong, ± tetragonal, 2.5 mm long, 1 mm wide, the surface papillate; disk destroyed in only flower dissected. Fruit unknown. Distribution. Known only from the Saiil area of French Guiana (see Mori, 1987) where it occurs in well-drained moist forest on lateritic soil. This species is well-marked vegetatively by the strikingly scabrous leaflet undersurface, and in flow¬ er by the reduced bracts, absence of bracteoles, and the unusually large plate-shaped glands on the ir¬ regularly bilabiate gray-drying calyx. Its closest rel¬ atives are A. prancei A. Gentry and A. subincanum Huber, which have nonasperous leaves with the ve¬ nation more raised-reticulate below and tan-drying, 5-denticulate calyces with much smaller glands. An¬ other noteworthy character of A. saulense is the strongly swollen woody nodes and subwoody strongly articulated petioles and petiolules. The material available to me has only buds and calyces of fallen flowers, so the corollas at anthesis cannot be de¬ scribed. Paratype. FRENCH GUIANA: Saiil, second growth near airport, 220 m, 24 June 1988 (st). Gentry 63176 (CAY, MO). Adenocalymna subspicatum A. Gentry, sp. nov. TYPE: Brazil. Ceara: Sitio Para, Vigosa do Ceara, carrasco, 14 June 1979 (fl), A. Fer¬ nandes, P. Martins & Matos s.n. ( FAC 6501 ) , (holotype, EAG; isotype, MO). Frutex scandens, ramulis teretibus, minute puberulis. Folia 2-3-foliolata, interdum cirrho simplici, foliolis lan- ceolatis vel anguste ovatis, chartaceis vel subcoriaceis, plerumque fere glabris. Inflorescentia floribus in racemo subspicato terminali dispositis, pedicellis 0.1 -0.3 cm lon- gis, bracteis 1.5-2. 5 cm longis subtentis. Calyx tubulo- campanulatus, 5-dentatus, 9-14 mm longus; corolla extus alba, intra lutea, tubulo-infundibuliformis, extus puberula. Capsula ignota. Volume 3, Number 2 1993 Gentry New Species of Adenocalymna 141 Liana, the branchlets terete, distinctly longitu¬ dinally ridged, glabrous to minutely puberulous with crisped trichomes, pseudostipules subulate, tan, 2- 4 mm long. Leaves 3-foliolate or 2-foliolate with a simple tendril, the leaflets lanceolate to narrowly ovate, acute to acuminate but with the apex ulti¬ mately minutely ernarginate, the base usually asym¬ metric, rounded to truncate or subcordate on one side, 2.5-10 cm long, 1-3.5 cm wide, cbartaceous to subcoriaceous, scattered lepidote-glandular above and below, ± inconspicuously and minutely puber¬ ulous along mid vein above and/or below, otherwise glabrous, drying olive above and below, the venation not prominently raised, the margin inconspicuously and minutely cartilaginous; petiole 0.7-2. 5 cm long, petiolules 0.3-1 .4 cm long, flattened above, sparsely lepidote and puberulous, especially above. Inflores¬ cence a very narrow terminal raceme, sparsely to conspicuously lepidote and puberulous with rather long bent trichomes, the bracts foliaceous, mem¬ branaceous, lanceolate, mostly 1.5-2. 5 cm long, 6- 8 mm wide, ± puberulous, essentially reduced leaves and occasionally ± trifid, the pedicels 0.1 0.3 cm long, usually with pair of thin 1 - 1 .5-cm-long brac- teoles near middle. Flowers with the calyx tubular- campanulate, broadly and shallowly 5-dentate, 9- 14 mm long, 5-6 mm wide, minutely and rather sparsely puberulous, the margin more densely pu¬ berulous and whitish, prominently 5-ribbed; corolla white outside, yellow inside, tubular-infundibuliform, 5-6 cm long, 1.2-1. 5 cm wide at mouth of tube, the tube 4-5 cm long, the lobes ca. 1 .5 cm long, puberulous outside with multicellular kinky tri- chomes, the lobes densely ciliate, lobes inside glan- dular-lepidote, tube mostly glabrous inside except at level of stamen insertion; anther thecae divaricate, 3-4 mm long; ovary cylindric, 2.3 mm long, ca. 0.8 mm wide, minutely lepidote; disk patelliform, I mm long, 2-3 mm wide. Fruit unknown. Distribution. Apparently endemic to the “car- rasco” vegetation of the caatinga region of northeast Brazil. The outstanding feature of this species is its long, rather thin, tubular calyx with five prominent lon¬ gitudinal ribs. Despite its large bracts, this species is probably most closely related to A. divaricatum Miers, which has similar branchlets and pseudosti¬ pules, but differs in having minute inconspicuous inflorescence bracts, a shorter, thicker, dark-drying unridged calyx, usually longer pedicels, yellow co¬ rolla, and the leaflets prominulously reticulate be¬ neath. Adenocalymna divaricatum occurs further east in the caatinga and dry habitats nearer the coast. Adenocalymna subspicatum occurs in the transition zone between the dry caatinga vegetation and easternmost Amazonian forests. Vegetatively, A. subspicatum is also reminiscent of A. flavurn in the color and texture of its leaves and bracts, but that species has a much smaller, nonribbed calyx. Paratypes. BRAZIL. Maranhao: Km 380-375, Be¬ lem- Brasilia, beira da estrada, terra firrne, 28 Aug. 1960 (H), E. Oliveira 1067 (UB, fragment MO). Ceara: Ibia- pina (?), Allemao de Cysneiros 1045 (P). Acknowledgments. I thank the National Science Foundation (BSR-86071 13) for support of my study of Bignoniaceae. Preparation of the manuscript was supported by a Conservation and the Environment Fellowship from the Pew Foundation. Literature Cited Bureau, E. & K. Schumann. 1896-1897. Bignonia¬ ceae. In: C. Martius, Flora Brasiliensis 8(2): 1-452. Mori, S. 1987. The Lecythidaceae of a lowland neo¬ tropical forest. La Fumee Mountain, French Guiana. Mem. New York Bot. Card. 44. Sutherland, C. 1983. The use of Bignoniaceae Data in Preserve Selection: A Neotropical Example. M.S. Thesis, Southern Illinois University, Edwardsville. A New Species of Styloceras (Buxaceae) from Peru Alwyn H. Gentry Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A. Gerardo Ay mar d C. UNELLEZ-Guanare, Programa de R.N.R. Herbario Universitario, Mesa de Cavacas, Portuguesa, Venezuela 3323 ABSTRACT. A new species of Buxaceae from the central Peruvian Andes is described as Styloceras penninervium. While revising collections of Styloceras during the process of identifying the first collection of this genus from Venezuela (Bautista & Aymard, in press), the junior author discovered two specimens that do not fit any of the described species. He brought them to the attention of the senior author, who has previously studied this genus (Gentry & Foster, 1 98 1 ). We concur that they represent a new species, which is described here. Styloceras penninervium A. Gentry & Aymard, sp. nov. TYPE: Peru. Junin: Provincia Tarma, sector “Agua Dulce,” 1,800 m, 23 Mar. 1948 (fl), Woytkowski 35497 (holotype, MO). Fig¬ ure 1 . Arbor parva dioecia. Folia anguste elliptica, glabra, coriacea, pinnatim nervata. Inflorescentia masculina ax- ilaris, spicata, 3-6 cm longa, floribus apetalis, subsessi- libus, bracteolis ovatis subtentis, antheris 2-2.5 mm lon- gis. Fructus globosus 2-2.5 cm diametro, stigmatum duorum elongatorum basibus disjunctis 1 cm. Dioecious shrub or small tree to 4 m tall, the branchlets glabrous, striate. Leaves alternate, 12- 18 cm long, 5-9 cm wide, elliptic to narrowly el¬ liptic, coriaceous, glabrous above and below, the margin revolute, apex acutish, sometimes minutely emarginate, cuneate at base, the midvein prominent below and impressed above, pinnately veined, the secondary veins plane above and nearly plane below, the tertiary venation barely visible, plane above and below; petioles canaliculate, glabrous, 2-3 cm long, 2-3 mm wide. Masculine inflorescence an axillary spike, in fascicles of 1-3, glabrous, striate, 3-6 cm long, the apetalous flowers subsessile on pedicel ca. 1 mm long, subtended by an acuminate deltoid-ovate bract 3-4 mm long and 2-3 mm wide; the flowers apetalous, the 12-17 naked anthers 2-2.5 mm long. Fruits pedicellate, globose, 2-2.5 cm diam., the pedicels glabrous, 1-2 cm long, with several small triangular bracts 1-1.5 mm long, the minute calyx persistent, with 5 triangular lobes, the two conspic¬ uous stigmas persistent, recurved at apex, ca. 2 cm long, the bases separated by 1 cm, irregularly de¬ hiscing. Distribution. Known only from the eastern slope of the central Peruvian Andes between 1 ,800 and 1 ,850 m. Styloceras penninervium differs from all four of the previously known Styloceras species in its pin¬ nately veined leaves, not at all triplinerved at the base. It also has the secondary veins less prominent below than in any of the known species. On the basis of the large fruits, one of the specimens of S. penninervium was originally identified (with a query) as a variant of lowland S. brokawii A. Gentry & R. Foster, the species that is also most similar in having the least strongly 3-veined leaves heretofore known in the genus. However, S. brokawii differs in the very different, more slender male inflores¬ cence with smaller (1 mm long) bracts subtending smaller flowers with anthers ca. 1 mm long. It also is distinct in the much thinner leaf texture and more broadly elliptic leaves with a distinctly 3-plinerved base. On the basis of the relatively dense inflores¬ cence, rather large anthers, and coriaceous leaves, S. penninervium is closer to S. laurifolium (Willde- now) HBK, which grows at higher altitudes (2,300- 3,800 m), and differs strongly in the more pro¬ nounced secondary veins and especially in having the basal pair of lateral veins strongly ascending. It also differs in having a smaller fruit 1.5-2 cm in diameter. Paratype. PERU. Pasco: Provincia Oxapampa, 5 km SE of Oxapampa, 1,850 m, 75°23'W, 10°36’S, 1 Feb. 1983 (fr), D. Smith 3206 (MO, USM). Key to the Five Species of SmocERAS la. Leaves pinnately veined; secondary veins below plane or barely prominulous . .... S. penninervium A. Gentry & G. Aymard C. Novon 3: 142-144. 1993. Volume 3, Number 2 1993 Gentry & Aymard Styloceras penninervium 143 Figure 1. Styloceras penninervium A. Gentry & Aymard. — A. Habit ( Woytkowski 35497 , MO). — B. Fruit (D. Smith 3206, MO). 144 Novon lb. Leaves 3-plinerved; secondary veins prominent below. 2a. Stigma bases fused in fruit; tertiary veins distinctly raised below . . S. columnare Muller Argoviensis 2b. Stigma bases separated by > 3 mm in fruit; tertiary veins plane below, the lower sur¬ face smooth between the secondary nerves. 3a. Inflorescence consistently monoe¬ cious, with apical female flowers; an¬ thers 3 mm long; Ecuador . . S. kunthianum Jussieu 3b. Plants mostly dioecious (rarely with a female flower at base of male inflo¬ rescence); anthers 1-2 mm long; Bo¬ livia to Venezuela. 4a. Leaves coriaceous, strongly 3 -veined from near base; anthers 1.5-2 mm long; fruit 1.5-2 cm diam.; southern Peru to Vene¬ zuela, above 2,200 in . . . . S. Inurifolium (Willdenow) HBK 4b. Leaves chartaceous or firmly membranaceous, weakly 3-veined from near base; anthers 1 mm long; fruit 2-3 cm diam.; lowland Bolivia and southern Peru .... S. brokaivii A. Gentry & R. Foster Acknowledgments. The junior author thanks the curators of MO and NY for hospitality and help during the visits to their herbaria. We thank Nidia Cuello for reviewing the manuscript and P. Bick for the illustration. Literature Cited Bautista, J. & G. Aymard. El genero Styloceras (Bux- aceae) en los Andes de Venezuela. Biollania 9 (in press). Gentry, A. H. & R. Foster. 1981. A new Peruvian Styloceras (Buxaceae): Discovery of a phytogeo- graphical missing link. Ann. Missouri Bot. Gard. 68: 122-124. Anomatheca laxa subsp. azurea (Iridaceae), a New Subspecies from Coastal Southern Mozambique and Natal, South Africa Peter Goldblatt B. A. Krukoff Curator of African Botany, Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A. Anne Hutchings Botany Department, University of Zululand, P. Bag XI 001, Kwadlangezwa 3886, South Africa ABSTRACT. Populations until now treated as An¬ omatheca laxa that occur along the coast of north¬ ern Natal and southern Mozambique are referred to a new subspecies azurea. these plants differ in their pale blue perianth with dark blue to violet markings and in their generally smaller flowers with a fairly long perianth tube. Subspecies azurea flow¬ ers in the cool months, June to mid-September, and is restricted to sandy soils, unlike typical red- to pink-flowered subspecies laxa, which flowers mainly from November to February, and is largely montane, favoring rocky sites. White- to pale blue-flowered plants with dark blue to violet or purple marks on the lower tepals cur¬ rently included in Anomatheca laxa (Thunberg) Goldblatt, one of five species currently admitted to this tropical and southern African genus (Goldblatt, 1972, 1991), have long seemed awkwardly placed. Except for flower color, however, there seemed little to separate the white- to pale blue-flowered individ¬ uals from typical A. laxa, which has pink to red flowers with dark red blotches at the bases of the three lower (anterior) tepals. Examination of addi¬ tional specimens of the blue-flowered plants in prep¬ aration for a treatment of Iridaceae for Flora Zam- besiaca (Goldblatt, in press) indicates a need for their taxonomic recognition, and they are described here as subspecies azurea. Typical Anomatheca laxa [Syn. Lapeirousia cruenta Lindley; Anomatheca cruenta (Lindley) Ba¬ ker], extends from the Uitenhage District in the southeastern Cape (South Africa), probably the type locality, through eastern tropical Africa to Uganda and western Kenya (Fig. 1). The range is, however, discontinuous, and A. laxa evidently does not occur in Zimbabwe, central and northern Mozambique, or southern Malawi, although it is present in central and northern Zambia, Shaba Province (Zaire), and southern Tanzania. Further north it is restricted to the mountains of eastern and northern Uganda and western Kenya. Plants generally favor rocky sites in light shade in mountain areas, but in South Africa they occasionally occur close to sea level. Related, and the only other tropical member of the genus, A. grandiflora Baker, which extends from the Transvaal (South Africa) to southern Tanzania, oc¬ curs in light woodland and open savanna, habitats generally drier than those favored by A. laxa. In contrast to the wide geographic range of sub¬ species laxa, subspecies azurea is restricted to coastal habitats in southeastern Africa from Richards Bay, Natal, in the south to Massinga, Mozambique, in the north (Fig. 1). As far as we know, subspecies azurea always grows in coarse sand, either in ex¬ posed sites such as grassy dunes or sometimes in light shade. Also, unlike subspecies laxa these coast¬ al plants only flower in the winter and early spring, June to September. Occasionally its flowering over¬ laps with that of subspecies laxa, which has been Novon 3: 145-147. 1993. 146 Novon o o o °° O ° o ° o°° Q ° O O ° A A o A A A A 18 20 22 24 26 28 30 32 34 Perianth tube mm Figure 2. Graph showing the relationship between peri¬ anth tube length and tepal length. Circles = plants with pink to red flowers with dark red markings (subspecies laxa ); triangles = plants with white to pale blue flowers with blue to violet markings (subspecies azurea ). recorded in flower from August to February, al¬ though mostly November to January. In coastal Zululand, both typical subspecies laxa and the blue- flowered subspecies azurea are known to occur in close proximity. Near Kwambonambi at a disturbed site in the Amangwe forest in Zululand, a series of hybrids occurs among the typical pale bluish sub¬ species azurea and some plants that resemble closely subspecies laxa ( Hutchings & Hutchings 301 9, 3020). Color differences alone are seldom sufficient ev¬ idence for taxonomic recognition, but in Anoma- theca laxa this is correlated with separate flowering times and habitat preferences. There also seems to be a nearly absolute difference in perianth size (Fig. 2). Healthy plants of subspecies laxa have a perianth tube (19-)21-28(-32) mm long and tepals 11-13 (-15) mm long. In contrast, flowers of subspecies azurea have a tube (25-)27-32 mm long and tepals 9-11 mm long (measurements from herbarium spec¬ imens). Flower size alone may be sufficient to sep¬ arate the subspecies, but in conjunction with peri¬ anth color, habitat, and flowering time there is little chance of confusing them. Care must be taken in making measurements to have well pressed, fully expanded flowers of healthy plants — depauperate individuals or those at the end of their flowering often have smaller flowers; also depending on care with specimen preparation, flowers may shrink to 80% of original size. We believe that subspecies rank for the white- to pale blue-flowered populations of Anomatheca laxa is appropriate in view of its generally close resemblance to typical A. laxa , and the considerably greater difference between all the populations of A. laxa and the other species of the genus. Systematic^ Anomatheca laxa subspecies azurea Goldblatt & Hutchings, subsp. nov. TYPE: Mozambique. Maputo: Inhaca Island, 20 July 1980, de Kon- ing <& Navunga 8312 (holotype, LMU; iso¬ types, BR, K, MO, SRCH). Plantae floribus pallide albescentibus ad caeruleos, basi tepalorum inferiorum atrocaerulea ad violaceam notata, tubo perianthii (25— )27— 32 mm longo, tepalis 9-11 mm longis, antheris ca. 4 mm longis. Plants with pale bluish (to nearly white) flowers, the lower three tepals each with a dark violet to indigo blotch near the base. Perianth tube (25-) 27-32 mm long; tepals 9-11 mm long, elliptic, ca. 4 mm wide. Filaments exserted for ca. 1 mm; an¬ thers ca. 4 mm long. Style dividing just above the base of the anthers, the branches ca. 2.5 mm long, forked for ca. 1 mm, and tangled in the anthers. Occurring along the coast in sandy soil, usually in sun, or in light shade; flowering mainly June to August. Differing from subspecies laxa in the white to pale blue flowers with dark blue to violet or purple markings at the base of the three lower tepals, sub¬ species azurea also generally has a longer perianth tube, usually (25-)27-32 mm long and smaller te¬ pals, 9-11 mm long. Pink- to red-flowered subspe¬ cies laxa has dark red marks at the bases of the lower tepals, a tube ( 1 9— )2 1 — 28(— 32) mm long and tepals 11 — 1 3(— 1 5) mm long. Flowers are seldom well enough preserved for us to be sure how constant other floral differences are, but there is at least a tendency for the filaments of subspecies laxa to be somewhat longer and exserted 1.5-2 mm from the tube. The style of subspecies laxa also seems to be somewhat longer than in subspecies azurea, dividing opposite the middle of the anthers. Examination of additional living populations may make it possible to determine whether these differences are consis¬ tent. Paratypes. MOZAMBIQUE. Inhambane: Massinga, rio das Pedras, 8 July 1981, de Koning & Hiemstra 8933 (BR, LMU, MO, SRGH); Massinga, Pomene, 13 July 1981, de Koning <£■ Hiemstra 9007 (LMU, MO). Gaza: between Chiconela and Gumbe, 26 May 1965, Pereira, Marques & Balsinhas 469 (LMU); Bilene, sand in dense bush, 31 July 1948, Torre 8046 (LISC); Vila de Joao Belo, Limpopo mouth, 17 June 1960, Lemos <£• Balsinhas 136 (K, LISC). Maputo: Inhaca Island, 10 July 1957, Barbosa 7626 (K, LMA); Maputo, Ponta do Ouro, 9 July 1961, Myre Macedo 4344 (LMU); Ma¬ puto, Ponta Malongane, 18 July 1971, Correia & Mar¬ ques 2194 (LMU, WAG). SOUTH AFRICA. Natal: Kosi Bay, sea level, 13 July 1947, Bayer T26 (NU); Sodwana State Forest, burned grassland, 30 m, 7 July, Nicholas Volume 3, Number 2 1993 Goldblatt & Hutchings Anomatheca laxa subsp. azurea 147 1467 (NH); St. Lucia, outside crocodile farm, 21 Aug. 1991, Hutchings <£ Hutchings 3017 (MO); grassland around Mt. Tabor, St. Lucia, Eastern Shores State Forest, 25 June 1989, Phillipson 3196 (MO, RLIH); St. Lucia Estuary, dune forest, Aug. 1984, Balkwill & Cadman 1728 (NU); Mapalane, 16 Aug. 1991, Hutchings & Hutchings 3016 (MO); Kwambonambi, Amangwe Forest, 25 Aug. 1991, Hutchings & Hutchings 3019 (MO); Monzi turnoff on road to Mtubatuba, forest track, 9 Aug. 1977, Pooley 1870 (NU); Richards Bay, 26 July 1974, Ward 5 (NU), 4 Sep. 1970, Strey 9906 (NH). Key to the Subspecies la. Flowers pink to red usually with dark red mark¬ ings on the lower tepals; perianth tube 19-28 (-32) mm long; dorsal tepal 11-15 x 5 mm . subsp. laxa lb. Flowers pale blue to whitish with dark blue to violet markings on the lower tepals; perianth tube (25 — )27— 32 mm long; dorsal tepal 9-1 1 x 4 mm . subsp. azurea Literature Cited Goldblatt, P. 1972. A revision of the genera Lapei- rousia Pourret and Anomatheca Ker in the winter rainfall region of South Africa. Contr. Bolus Herb. 4: 1-111. - . 1991. An overview of the systematics, phy- logeny and biology of the African Iridaceae. Contr. Bolus Herb. 13: 1-74. Ixia acaulis , a New Acaulescent Species of Iridaceae: Ixioideae from the Knersvlakte, Namaqualand, South Africa Peter Goldblatt B. A. Krukoff Curator of African Botany, Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A. John C. Manning Compton Herbarium, National Botanical Institute, Kirstenbosch Botanic Gardens, Claremont 7735, South Africa AUSTRACT. The generic placement of a new spe¬ cies, Ixia acaulis, from limestone outcrops in the Knersvlakte, Vanrhynsdorp District, South Africa, is not immediately clear in part due to its reduced acaulescent habit. The bright yellow flowers have a subterranean ovary and are borne above ground level on a long perianth tube. An acaulescent habit is unusual in Iridaceae, although known in several genera of the two larger subfamiles, Iridoideae and Ixioideae. In an effort to correctly assign the plant to a genus, we have examined leaf anatomy and chromosome cytology as well as macromorphology and palynology. A basal rooting corm places the species firmly in Ixioideae, as does the perforate exine sculpturing. We conclude that it belongs in the southern African genus Ixia, although an acau¬ lescent habit is otherwise unknown in this genus and other aspects of the plant do not exactly conform to this genus. The uncertain generic position of a new species of Iridaceae subfamily Ixioideae from the Knersv¬ lakte north of Vanrhynsdorp, Cape Province, South Africa, required detailed investigation of its mor¬ phology, leaf anatomy, cytology, and palynology. It is unusual in the subfamily in lacking an aerial stem. The flower has a subterranean ovary and is raised above the ground by means of a long perianth tube. Finely fibrous corm tunics seem to preclude any relationship with Hesperantha, Romulea, or Syr- ingodea, in which acaulescence is the rule or at least not uncommon, but in which the corm tunics are more or less woody. However, a combination of reduced and specialized features makes it uncertain whether the species belongs in Ixia, Tritonia, Spar- axis, Anoniatheca, or Duthieastrum, all of which have fibrous corm tunics, but only the last is nor¬ mally acaulescent. Morphology VEGETATIVE ORGANIZATION The small plants (Fig. 1A, B) grow singly or in small clumps, each individual consisting of an elon¬ gate, more or less symmetrical, round-based corm with fibrous corm tunics (Fig. IB) that accumulate in older plants to form a thick matted fibrous layer. The corms produce roots from the basal area but in no particular pattern. The 3-5 foliage leaves are basal and sheathed below the ground by 2 mem¬ branous, entirely sheathing cataphylls. The leaf blades are equitant and more or less linear. Under natural conditions the blades are more or less prostrate on the ground. Although the blades are unifacial, there is no morphological or anatomical differentiation be¬ tween the leaf surfaces facing toward or away from the surface of the ground. Grown under greenhouse conditions with more water than normally available (as were the plants illustrated), the leaves tend to be inclined rather than prostrate. The stem is short, 1-2 cm long, but does not reach ground level at anthesis and bears one or occasionally two flowers. As the capsules ripen the peduncle elongates so that the seeds are released at ground level. FLORAL STRUCTURE The flowers are sessile (i.e., lack pedicels) and are subtended by a pair of opposed membranous bracts inserted at the base of the ovary (Fig. 1C, G), thus conforming to the pattern in all Ixioideae. In the three other subfamilies of Iridaceae the flow¬ ers are normally pedicellate, rarely sessile, and al¬ ways subtended by a single bract (Goldblatt, 1 990). Also typical of Ixioideae, the outer bract is larger than the inner in Ixia acaulis, although the two bracts are more or less equal in length (Fig. 1G). The inner bract is 2-veined and apically forked. The Novon 3: 148-153. 1993. Volume 3, Number 2 1993 Goldblatt & Manning Ixia acaulis 149 1cm Figure 1. Ixia acaulis Goldblatt & J. Manning. — A. Habit. — B. Entire plant with corm. — C. Flower and paired bracts and apex of stem. — D. Longitudinal section of upper half of flower. — E. Transverse section of perianth tube at mid level showing filament flanges and central style. — F. Detail of style branches. — G. Detail of outer and inner bracts. — H. Polar view of pollen grain with operculum lying over the aperture. I. Seed. (Drawn by J. C. Manning.) ovary is subterranean and the flower is raised above the ground by a long perianth tube, 15-20 mm long, that reaches about 10 mm above ground. The flower is actinomorphic and the bright yellow tepals are subequal and spread at right angles to the tube (Fig. IB, C). The short filaments are exserted ca. 1.5 mm from the perianth (Fig. ID) and are de¬ current, extending to the middle of the tube as flanges that reach almost to the center ol the tube (Fig. ID, E). The filiform style reaches to about the mouth of the tube where it divides into three short branches, ca. 1 mm long, and each branch is adax- ially channeled above, but the margins are condu- plicate below so that the lower two-thirds are tubular 150 Novon A. Tissue distribution in half leaf (pseudomidrib vein pair to margin). — B. Cell detail of marginal area (palisade mesophyll shaded). (Drawn by J. C. Manning.) (Fig. IF). The stiginatic tips of the style branches emerge between the adjacent filaments a short dis¬ tance below the base of the anthers (Fig. 1 D). The erect and more or less parallel anthers are linear- oblong, submedianly fixed, and longitudinally dehis¬ cent (Fig. ID). FRUIT AND SEED The fruit is a globose capsule with somewhat cartilaginous walls, corresponding to the presumed plesiomorphic state for Ixioideae. The seeds are glo¬ bose, extremely hard, and lack primary or secondary sculpturing (Fig. II). The only notable feature is the white filiform vestige of the ovular vasculature lying over the raphe, thus outside the body of the mature seed. This seed type is derived in shape, absence of surface sculpturing, and in the peculiar excluded vascular trace (Manning & Goldblatt, in prep.). Sim¬ ilar hard rounded seeds without primary or second¬ ary sculpturing characterize several genera of Ixioi¬ deae tribe Ixieae, among them Sparaxis , Ixia, Dier- atna , Duthieastrum, Tritonia, its close allies Cro- cosmia, Chasmanthe, and Devin (de Vos, 1974; Goldblatt & Manning, 1990; in prep.), as well as Freesia and most species of Anomatheca. The two latter have a characteristically inflated raphe (Gold¬ blatt & Manning, in prep.) and do not have the excluded vascular trace that occurs in the other genera listed above (seeds of Duthieastrum are un¬ known in this respect). Leaf Anatomy In cross section (Fig. 2) the blades of the unifacial leaves are oblong and conform to the basic pattern for Iridaceae (Arber, 1921; Hudall & Goldblatt, 1989) in having vascular bundles in two opposed rows, with the phloem directed toward the epidermis and the xylem toward the interior of the leaf. The median pair of veins are slightly larger than the others and thus constitute a pseudomidrib, generally visible as such in surface view. The epidermal cells are slightly wider than high and papillate; the mar¬ ginal epidermal cells are similar to those of the laminar surface except in their slightly smaller size (Fig. 2B). Stomata are sunken and the guard cells are small. Immediately below the marginal epidermis there is a small strand of sclerenchyma (= phloem cap) contiguous with the phloem of a small marginal vein. There is a well-developed palisade two cell layers thick and a sclerenchyma phloem cap on all primary and secondary bundles reaching the epi¬ dermis in all bundles except the median pair. The bundle sheaths are unthickened except at the phloem pole. Presence of submarginal sclerenchyma asso¬ ciated with a marginal vein is probably plesiomorphic for Ixioideae (Rudall & Goldblatt, 1989; Goldblatt, 1990) , and a pseudomidrib is plesiomorphic for tribe Ixieae. The leaf anatomy thus corresponds with the hasal condition for Ixieae (Rudall & Goldblatt, 1989), including Ixia and Dierama among genera possibly closely related to the new species. A derived leaf margin anatomy in which the leaf margin cells are columnar and heavily thickened on the anticlinal walls and the margins lack subepider- mal sclerenchyma characterizes Tritonia (excepting T. marlothii and T. delpierrei ) (de Vos, 1980), Sparaxis (including Synnotia ), Anomatheca, and a few more genera (Rudall & Goldblatt, 1989). Duthieastrum has leaves without either subepider- mal marginal sclerenchyma or columnar marginal epidermal cells with thickened walls, but it does have a marginal vein (Goldblatt, unpublished). Pollen Grains The pollen grains (Fig. 1H), examined in the fully expanded state in 1% aceto-orcein, are monosulcate and rounded, ca. 62 x 54 jum, thus slightly longer in the plane of the sulcus. The sulcus has a con¬ spicuous, narrowly elliptic operculum lying across Volume 3, Number 2 1993 Goldblatt & Manning Ixia acaulis 151 the long axis and nearly as long as the sulcus and ca. 1 2 /am wide in the midline. The exine is perforate with the surface bearing scattered small spinulae. Both in the presence ol an operculum and in the perforate and sparsely spinulate exine the species corresponds with the plesiomorphic state for Ixioi- deae (Goldblatt, 1990). In the presumed plesiomor¬ phic condition for the subfamily, also the most com¬ mon, the operculum is double-handed (Goldblatt et ah, 1991). A derived single-handed operculum is so far known only in Ixia, some species of Anomatheca (Ixieae), and Lapeirousia and Thereianthus (Wat- sonieae) (Goldblatt et ah, 1991; Goldblatt & Man¬ ning, in prep.). Apparently only in Ixia is the oper¬ culum consistently single-handed, although only a selection of species have so far been examined. The grains are some 30% larger than recorded in other species of Ixia (Goldblatt et ah, 1991), possibly related to the long perianth tube and consequent need for larger reserves to permit growth through a longer style. Chromosome Cytology Root tips (examined by M. Takei), harvested from sprouting corms and treated as described by Gold¬ blatt et ah (1993), showed a diploid chromosome number ol 2 n = 20 at mitotic metaphase. The chromosomes are relatively small, the longest four pairs 2. 7-2. 3 fim and the smaller pairs 1 .8 1 .2 /am long, and asymmetric in arm ratio. The second lon¬ gest and one of the shorter pairs are nearly nieta- centric, and the rest are acrocentric to submeta- centric. The karyotype is weakly bimodal with three longer pairs standing out from the remaining shorter chromosomes. Satellites were not seen clearly but appear to be small and located on one of the longer chromosome pairs. Other Ixioideae with a base num¬ ber of x = 10 (Goldblatt, 1971; de Vos, 1974) include Ixia, Dierama, Sparaxis (including Syn- notia ), and Duthieastrum (all Ixieae) and Ther¬ eianthus, Micranthus, and section Fastigiata of Lapeirousia (Watsonieae) and Pillansia, only genus of Pillansieae. The ancestral base number for Ixioi¬ deae may be x = 10. In general size the chromo¬ somes of Ixia correspond most closely with those of the new species. Chromosomes of Sparaxis are some 20% smaller (total chromosome length) (Goldblatt & Takei, in prep.) and exhibit no bimodality. Discussion GENERIC POSITION A basally rooting corm, long-lasting flowers with a well-developed perianth tube, paired bracts in¬ serted below a sessile ovary, operculate pollen grains, and perforate exine, synapomorphies for Ixioideae (Goldblatt, 1991; Goldblatt et ah, 1991), are all present in Ixia acaulis and confirm its subfamilial position. Within Ixioideae it corresponds with Ixieae and Watsonieae in having leaves with a pseudomid¬ rib. Association with Watsonieae may be excluded because members of this tribe have corms that de¬ velop entirely from an axillary bud (the new- corm lies lateral to the flowering stem), and they also have deeply divided style branches, the former possibly apomorphic and the latter almost certainly so (Gold¬ blatt, 1990, 1991). There is little doubt that the new species belongs in Ixieae. Among the 20 genera of the tribe, the new species has similarities with several genera, especially Tritonia, Duthieastrum , Anomatheca, and Ixia, although the acaulescent condition and subterranean ovary are atypical of all but the monotypic Duthieastrum. Only one species of Tritonia, T. florentiae (E. Phillips) Goldblatt, lacks an aerial stem, but no acaulescent species of Ixia or Anomatheca are known. Acaulescence has ap¬ parently evolved repeatedly in Iridaceae and fre¬ quently in Ixioideae, in which at least some species of Lapeirousia and Hesperantha, most species of Romulea, and all of Syringodea and Crocus (as well as Duthieastrum) lack an aerial stem at an- th esis. Acaulescence, especially common in African Ixioideae among species of arid and semiarid areas, is obviously of direct adaptive significance, and alone is probably of little or no value in considering the relationships of species of Ixieae. More important in determining relationships among Ixioideae are the nature of the style branches (either simple or deeply divided), leaf margin anat¬ omy, bract texture (green and herbaceous or mem¬ branous and transparent or scarious), seed mor¬ phology, and possibly pollen grain morphology. The most likely generic position for this new species seems to us to be Ixia, with which it corresponds mainly in plesiomorphic features (unspecialized leaf anatomy, chromosome number, simple style branch¬ es); it also shares a notable synapomorphy with Ixia, pollen grains with a single-handed operculum. The membranous and transparent bracts may also be apomorphic for Ixia. In some species of Ixia alone of the genera of Ixioideae, the style divides at the mouth of the perianth tube and has short branches (in Hesper¬ antha the style consistently divides at or below the mouth of the tube but the branches are long and trailing). The new species does not accord with An¬ omatheca, Tritonia, Duthieastrum , or Sparaxis in leaf margin anatomy or pollen grain structure. These four genera have columnar marginal epidermal cells 152 Novon and, except for Duthieastrum, lack submarginal sclerenchyma whereas their pollen grains have dou¬ ble-banded opercula. Additionally, the style branch¬ es of Anomatheca are always deeply divided, and the seeds of this genus do not have an excluded vascular trace. Basic chromosome number in Spar- axis is also x — 10, but both Tritonia and Anom¬ atheca have x = 1 1 , excepting one derived species of Tritonia, T. fiorentiae, which has the secondary base of x — 10 (Goldblatt & Takei, in prep.). RELATIONSHIPS WITHIN IXIA Ixia subg. Dichone is derived in the genus in its peculiar, stout, and so-called subdidymous anthers (Lewis, 1 962), which appear to lack a connective and are additionally normally (possibly always) ba- sally porate. The six species of the subgenus also always have pink flowers and, according to Lewis (1962), involute style branches. Thus, I. acaulis falls within the remaining subgenus Ixia, where it accords best with section Hyalis and Morphixia. It may be most closely related to I. paucifolia of the latter (de Vos, pers. comm.), which also has a long perianth tube, and included, decurrent fila¬ ments. In one aspect, however, I. acaulis seems to differ from Ixia: it has subcentrally fixed anthers, whereas near basifixed anthers are characteristic of the genus, according to de Vos (pers. comm.). Systematics Ixia acaulis Goldblatt & J. Manning, sp. nov. TYPE: South Africa. Cape: 31.18 (Vanrhyns- dorp) Knersvlakte, farm Rooiberg, low lime¬ stone ridges (BG), 19 May 1992, Snijman & Manning 1 249 (holotype, NBG; isotypes, MO, PRE). Figure 1. Plantae 2-3 cm altae, interdum caespitosae, cormo elongato, tunicis fibrosis, foliis productis 3-5, laminis li- neato-lanceolatis 2-4 mm latis, floribus solitaris actino- morphis flavis, tubo perianthii 15-20 mm longo, tepalis aequalibus ovato-oblongis 6-7x3 mm, filamentis erectis ca. 1.5 mm longis, antheris linearo-oblongis erectis con- tiguis ca. 2 mm longis, stylo diviso prope apicem tubi, ramis styli ca. 1 mm longis. Plants acaulescent, often growing in small clumps, leaves and flowers reaching 2-3 cm above the ground. Corm elongate, ca. 3-4 mm wide near the base, tapering above, ca. 15 mm long, the tunics moderately to finely fibrous and with age accumu¬ lating as a dense covering. Foliage leaves 3-5, all basal, sheathed below by 1-2 transparent cataphylls, these just reaching ground level; leaf blades linear to narrowly lanceolate, ca. 1.5-2 mm wide, often prostrate, or inclined toward the ground, ± obtusely acuminate. Stem subterranean, 9-18 mm long, usu¬ ally unbranched or with one branch. Inflorescence a solitary flower (or the flowers solitary on each branch); bracts membranous, the outer larger, ca. 10 mm long, somewhat fringed along the upper margin, inner bract about as long but narrower, 2-veined and lightly forked apically, reaching to 5 mm above the ground. Flowers actinomorphic, bright yellow, whitish in the tube; perianth tube 15-20 mm long, arising below ground level and extending to 10 mm above the ground, cylindric, widening gradually above; tepals subequal, oblong-ovate, ob¬ tuse to emarginate, 6-7 x 3 mm, spreading at right angles to the tube. Stamens erect, the filaments arising at the top of the tube, ca. 1.5 mm long, decurrent and forming prominent ridges reaching downward to the middle of the tube, the ridges extending inward to the style but not joined to it; anthers linear-oblong, longitudinally dehiscent, sub¬ centrally fixed, erect and ± contiguous, ca. 2 mm long. Ova ry oblong, ca. 2.5 mm long; style filiform, dividing at or just beyond the mouth of the tube, the branches short, ca. 1 mm long, the apices emerg¬ ing between the filaments, terminally stigmatic, adaxially channeled in the upper part but the mar¬ gins conduplicate below and the branches thus tu¬ bular in the lower half. Capsules globose, 5-6.5 mm long, borne at ground level. Seeds few per locule (usually 3-4), globose, smooth, dark red-brown, ca. 1 .5 mm diam., the vascular trace excluded and often persisting as a white strand along the raphe. Chro¬ mosome number 2 n — 20. DISTRIBUTION AND HABITAT Ixia acaulis appears to be restricted to limestone ridges, clearly a specialized habitat, in the arid Knersvlakte of southern Namaqualand. This area of low rainfall, less than 100 mm p.a., normally falling in the winter months, is known for its diversity of habitats and extreme local endemism, particularly among Aizoaceae-Mesembryanthemoideae. lrida- ceae are not well represented here, but other Knersvlakte endemics in the family include Homeria maximiliani Schlechter, Moraea deserticola Gold¬ blatt (both Iridoideae), and Lapeirousia angusti- folia Schlechter (if this is regarded as distinct from the related T. pyramidalis (Lamarck) Goldblatt). Among these Iridaceae, /. acaulis is unusual in flowering at the beginning of the wet season in May (and possibly June) rather than in the early spring in July and August. This unusual flowering time as well as the specialized habitat together account for its late discovery in an area that has been explored botanically since the late 1 9th century. So far, /. Volume 3, Number 2 1993 Goldblatt & Manning Ixia acaulis 153 acaulis is known from two limestone ridges on the farm Hooiberg just to the north of the Saldanha- Sishen rail line. It grows in rock cracks or wedged between boulders in a virtually lithophytic situation. The few other limestone outcrops in the Knersvlakte, an area of predominantly fine clay soils often covered by a dense superficial layer of quartz pebbles, remain to be examined for the presence of the species. Paratypes. SOUTH AFRICA. Cape Province: 3118 (Vanrhynsdorp) Knersvlakte, Farm Rooiberg, low lime¬ stone ridges (BC), Perry, 1991 (NBG), 27 Aug. 1991 (fr), Goldblatt & Manning 9132 (MO, NBG, PRE). Acknowledgments. This work was supported in part by the U. S. National Science Foundation grant HSR 89-06300. We recognize the discovery ot this new species by Pauline Perry and thank her lor providing the live specimens studied here. We also express our thanks to M. P. de Vos, Stellenbosch, South Africa, for constructive comments on the manuscript. Literature Cited Arber, A. 1921. The leaf structure of the Iridaceae, considered in relation to the phyllode theory. Ann. Bot. 35: 301-336. - . 1980. The African genus Tritonia Ker-Gawler (Iridaceae): Part 1. J. S. African Bot. 48: 105-163. Goldblatt, P. 1971. Cytological and morphological stud¬ ies in the southern African Iridaceae. J. S. African Bot. 37: 317-460. - . 1990. Phylogeny and classification of the Iridaceae. Ann. Missouri Bot. Card. 77: 607-627. - . 1991. An overview of the systematics, phy¬ logeny and biology of the African Iridaceae. Contr. Bolus Herb. 13: 1-74. - & J. C. Manning. 1990. Devia xeromorpha, a new genus and species of Iridaceae-Ixioideae from the Cape Province, South Africa. Ann. Missouri Bot. Card. 77: 359-364. - , J. C. Manning & A. Bari. 1991. Sulcus vari¬ ability in the pollen grains of Iridaceae subfamily Ixioideae Ann. Missouri Bot. Card. 78: 950-961. - , M. Takei & Z. Razzaq. 1993. Chromosome cytology in tropical African Gladiolus (Iridaceae). Ann. Missouri Bot. Card. 80: 461-470. Lewis, G. J. 1962. South African Iridaceae. The genus Ixia. J. S. African Bot. 28: 46-195. Rudall, P. & P. Goldblatt. 1989. Leaf anatomy and phylogeny of Ixioideae (Iridaceae). Bot. J. Linn. Soc. 106: 329-345. Vos, M. P. de. 1974. Duthiella, 'n nuwe genus van die Iridaceae. J. S. African Bot. 40: 301-309. A New Combination for the Single American Element of Parasenecio (Asteraceae: Senecioneae) Jason R. Grant Department of Botany, University of Maryland, College Park, Maryland 20742-5815, U.S.A. ABSTRACT. In accordance with the acceptance of the generic name Parasenecio for the Asiatic-cen¬ tered species historically referred to Cacalia, a new combination is made for the single species that rang¬ es to North America: Parasenecio auriculata (A. P. de Candolle) J. R. Grant. Cacalia auriculata was described by A. P. de Candolle (1837) from a specimen collected in 1835 by Turczaninow between Yakutsk and Okhotsk, Yakutia, Russia. It ranges from Kamtchatka to the Commander and Kurile Islands (Russia), to Hok¬ kaido and northern Honshu (Japan), and from the Russian Far East to Manchuria (China) and Korea (Hulten, 1930). In Alaska, it has been collected on five islands at the westernmost end of the Aleutian chain roughly adjacent to the Commander Islands, Attu, Agattu, and Alaid of the Near Islands, and Kiska and Buldir of the Rat Islands. Cacalia auriculata is one of a number of species of primarily Asiatic distribution that ranges to the westernmost islands of the Aleutian chain. Hulten (1937) documented the American range for several species with this type of distribution pattern includ¬ ing Senecio palmatus Pallas and Cirsium kamtch- aticum Ledebour. The vegetation of the islands where these species are found is “a distinctly Kamtchatkan type” (Hulten, 1937: 37). The presence of these species in the Aleutians is due to their association with “high-grown vegetation” typical of Kamchatka that also extends to the Near and Rat Island groups, but is unknown to the remainder and majority of the Aleutian chain (Hulten, 1937). Robinson & Brettell (1973) described the genus Koyamacalia for the Asian species referred to Ca¬ calia. Cacalia auriculata was transferred into Koy¬ amacalia along with 46 other species (Robinson & Brettell, 1973). Jeffrey & Yi-Ling (1984) pointed out that Koyamacalia H. E. Robinson & R. I). Brettell was a later synonym of Parasenecio W. W. Smith & Small. Jeffrey (1992) again commented on Parasenecio, but as in the earlier work, no new combinations were made. In order to update no¬ menclature in the North American flora, the follow¬ ing new combination is made: Parasenecio auriculata (A. P. de Candolle) J. R. Grant, comb. nov. Basionym: Cacalia auri¬ culata A. P. de Candolle, Prod. 6: 329. 1837 [1838], TYPE: Inter Ochotiam [Okhotsk] et Jacutiam [Yakutsk], N. Turczaninow s.n. (ho- lotype, G-I)C, photo on microfiche seen). Ligularia auriculata Turczaninow ex A. P. de Candolle, Prod. 6: 329. 1837, pro syn. Koyamacalia auriculata (A. P. de Candolle) H. E. Rob¬ inson & R. D. Brettell, Phytologia 27: 271. 1973. Specimens examined. UNITED STATES. Alaska: Near Islands, Attu Island, 13°52'56"N, 173°15'E, 25 July 1943, Beals A (ALA); Attu Island, 13°52'56"N, 173°15'E, Beals, 1943 (ALA); Attu Island, Massacre Bay area, 10 July 1973, Williams 3212 (US); Agattu Island, 13°52'26"N, 17°33'06"E, 13 Aug. 1974, Trapp 104 (ALA); Attu Island, Peaceful Valley, near Navy Town, 13°52'50"N, 173°1 l'E, 18 Aug. 1983, Friedman & Michaelson 83-64 (ALA); Rat Islands, Buldir Island, 14°52'21"N, 175°56'E, 14 July 1974, Dick 164 ( ALA); Buldir Island, 14°52'21"N, 175°56'E, 25 Aug. 1974, Dick 403 (ALA). Acknowledgments. I thank Harold E. Robinson (Smithsonian Institution) for bringing my attention to this species, and for review of the manuscript. I also thank Alan R. Batten of the Northern Plant Documentation Center, Herbarium, University of Alaska Museum, Fairbanks, for sending data on the specimens at ALA. Literature Cited Candolle, A. P. de. 1837. Prodromus Systematis Na- turalis Regni Vegetabilis. Vol. 6. Paris, Strasbourg & London. Hulten, E. 1930. Flora of Kamtchatka and the adjacent islands. IV Dicotyledonae, Pyrolaceae-Compositae. Kungl. Svenska Vetenskapakademiens Handlingar Tredje Serien. Band 8. N:0 2. Stockholm, Almqvist & Wiksells Boktryckeri A.-B. - . 1937. Flora of the Aleutian Islands and West¬ ernmost Alaska Peninsula with Notes on the Flora of Commander Islands. Bokforlags Aktiebologet Thu¬ le, Stockholm. Novon 3: 154-155. 1993. Volume 3, Number 2 1993 Grant Parasenecio auriculata 155 Jeffrey, C. 1992. The tribe Senecioneae (Compositae) in the Mascarene Islands with an annotated world checklist of the genera of the tribe. Notes on Coni- positae: VI. Kew Bull. 47: 49-109. - & C. Yi-Ling. 1984. Taxonomic studies on the tribe Senecioneae (Compositae) of Eastern Asia. Kew Bull. 39: 205-446. Robinson, H. E. & R. D. Brettell. 1973. Studies in the Senecioneae (Asteraceae). IV. The Genera Mesa- denia, Syneilesis, Miricalia , Koyamacalia , and Sinacalia. Phvtologia 27: 265-276. Change in Status of a Dwarf Mistletoe ( Arceuthobium , Viscaceae) from China Frank G. Haivksivorth f USDA Forest Service, Rocky Mountain Forest and Range Experiment Station, Fort Collins, Colorado 80526, U.S.A. Delbert ll iens Department of Biology, University of Utah, Salt Lake City, Utah 84112, U.S.A. Abstract. The taxon Arceuthobium pini var. si- chuanense , a parasite of Picea in southwestern Chi¬ na, is raised to specific rank. It is allopatric with A. pini , which differs in its parasitism ol Pinus, its much larger shoots, and its smaller male flowers. A new variety, Arceuthobium pini Hawksworth & Wiens var. sichuanense H. S. Kiu, a parasite of Picea, was recently described from southwestern China (Kiu, 1684a). Here we raise that taxon to specific rank because of the marked differences in morphology and host relations between it and A. pini. Arceuthobium sichuanense (H. S. Kiu) Hawks¬ worth & Wiens, comb. nov. Basionym: Arceu¬ thobium pini var. sichuanense Kiu H.-s., Acta Phytotaxonomica Sinica 22: 205, 1984. TYPE: China. Sichaun: Dawu, parasite of Picea lik- iangensis var. balfouriana, 4,200 m, Wu & Gao 11629 (holotype, CDBI). Arceuthobium sichuanense differs from A. pini in being a much smaller plant (shoots 2 6 cm high vs. 5 20 cm) with smaller staminate flowers (1.5 2 mm vs. 2-2.5 mm diam.). A major difference is that A. sichuanense parasitizes only Picea, whereas A. pini parasitizes only Pinus. Both species occur in Sichuan and Xizang (Tibet) in southwestern China, but they are apparently allopatric (Kiu & Wei, 1982) and occur at different elevations: A. sichu¬ anense at 3,800-4,200 m and A. pini at 2,700- 3,500 m (Kiu, 1984b). At 4,200 m, A. sichuanense has the highest elevational distribution of any mem¬ ber of the genus, although two taxa (A. globosum subsp. grandicaule and A. vaginalum subsp. va- ginatum) approach this limit in central Mexico (Hawksworth & Wiens, 1972). Arceuthobium si¬ chuanense is most similar to A. tibetense Kiu & Wei Ren, an allopatric species parasite of Abies in Xizang, which has even smaller shoots (0.5 2.2 cm high) (Kiu, 1984b). A key to the five taxa of Ar¬ ceuthobium in southwestern China is given by Kiu (1984b). Arceuthobium sichuanense has recently been found in western Bhutan, where it causes “serious damage” to Picea likiangensis var. balfouriana and P. spinulosa (Naithani & Singh, 1989). Literature Cited Hawksworth, F. G. & D. Wiens. 1972. Biology and classification of the dwarf mistletoes (Arceuthobium ). Agric. Handb. 401, USDA Forest Service, Wash¬ ington, D.C. Kiu H.-s. 1984a. Materials for Chinese Viscoideae. Acta Phytotax. Sin. 22: 205-208. - . 1984b. Arceuthobium and its hosts in south¬ western China. Pp. 18-19 in Gen. Techn. Rep. RM- 111, Biology of Dwarf Mistletoes, Proc. of the Sym¬ posium, USDA Forest Service, Fort Collins, Colorado. - & Wei R. 1982. A new species of Arceu¬ thobium from Xizang. J. Yunnan Forestry Coll. 1: 42-45. Naithani, H. B. & P. Singh. 1989. Note on the oc¬ currence of the genus Arceuthobium M. Bieh. in Eastern Himalaya. Indian Forestry 115: 196. Novon 3: 156. 1993. Notes on the Lamiaceae of China Li Xi-wen (Li Hsi-wen) Kunming Institute of Botany, Academia Sinica, Kunming, Yunnan 650204, People’s Republic of China ABSTRACT. Nomenclatural changes, affecting Marmoritis and Scutellaria , are made for the up¬ coming Flora of China. Hedge (1990) resuscitated the generic name Marmoritis “after almost 1 50 years in synonymy.” It replaces the previously accepted Phyllophyton. In this same treatment Hedge made the new com¬ bination Marmoritis nivalis (Jacquemont ex Ben- tham) Hedge and recognized Marmoritis rotund i- folia Bentham for the flora area he was considering. In order to reflect the current taxonomic status for species formerly placed in Phyllophyton , the fol¬ lowing nomenclatural changes are made. The tax¬ onomic treatment of the following taxa, along with Marmoritis nivalis and M. rotundifolia, will appear in the Flora of China (Li & Hedge, in press). Marmoritis complanatum (Dunn) 11. W. Li, comb. nov. Basionyrn: Nepeta complanata Dunn, Notes Roy. Bot. Card. Edinburg!) 8: 122. 1913. Gleehoma complanata (Dunn) Turrill, Bot. Soc. Exch. Club Brit. Isles 1919, 5: 695. 1920. Phyllophyton complanatum (Dunn) Kudo, Mem. Fac. Sci. Taihoku Imp. Univ. 2: 225. 1929. Pseudolophanthus com- plana tu s (Dunn) Levin, Trudy Bot. Inst. Akad. Nauk S.S.S.R., Ser. I, FI. Sist. Vyssh. Hast. 5: 296. 1941. TYPE: China. NW Yunnan: scree plant, 1 6,000 ft., July 191 I. F. kingdon Ward 138 (holotype, E not seen). Dracocephalum rockii Diels, Notizbl. Bot. Cart. Berlin- Dahlem 9: 1030. 1926. TYPE: China. Yunnan Province: Mountains of Moting, NW of the Yangtze- Mekong watershed, June 1923, J. F. Rock 10327 (holotype, presumably B, not seen; isotype, US). Marmoritis decolorans (Hemsley) H. \\ . Li, comb. nov. Basionyrn: Nepeta decolorans Hemsley, Hooker's Icon. PL 25: t. 2170. 1896. Gleehoma decolorans (Hemsley) Turrill, Bot. Soc. Exch. Club Brit. Isles 1919, 5: 695. 1920. Phyllophyton decolorans (Hemsley) Kudo, Mem. Fac. Sci. Taihoku Imp. Univ. 2: 225. 1929, “ decorans Pseudolophanthus deco¬ lorans (Hemsley) Levin, Trudy Bot. Inst. Akad. Nauk S.S.S.R., Ser. 1, LI. Sist. Vyssh. Rast. 5: 296. 1941. TYPE: Hah. Central Tibet: Gooring Valley, 30°12'N, 90°25'E, at about 16,500 ft., St. George Idttledale, July and Aug. 1895 (holotype, K not seen). Marmoritis pharicus (Prain) 11. \\ . Li, comb, nov. Basionyrn: Nepeta pharica Prain, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 59: 306. 1891. Gleehoma pharica (Prain) Turrill, Bot. Soc. Exch. Club Brit. Isles 1919, 5: 695. 1920. Phyllophyton pharicum (Prain) Kudo, Mem. Fac. Sci. Taihoku Imp. 1 niv. 2: 225. 1929. Pseudolophanthus pharicus (Prain) Kupriano- va, Bot. Zhurn. S.S.S.R. 33: 235. 1948. TYPE: Eastern Tibet, Phari, between Phari and Lhas- sa, Lama Vjyen Gyatsko 106 (holotype, not seen). Scutellaria yangbiensis II. W. Li, nom. nov. Replaced name: Scutellaria urtici folia C. Y. Wu & H. W. Li, FI. Yunnan. 1: 545. 1977, non Juzepczuk & Vvedensky, FI. IJRSS 20: 510. 1954. TYPE: Zhongdian Xian Expedi¬ tion, NW Yunnan, 63-4020 (holotype, KUN). Acknowledgments. The assistance ot Bryan Dut¬ ton and Benito Tan is gratefully acknowledged. Literature Cited Hedge, I. 1990. Labiatae. Flora of Pakistan 192: 119. Li, X. W. & I. Hedge. Lamiaceae. Flora of China, vol. 17, Science Press & Missouri Bot. Card. (In press.) Novon 3: 157. 1993. Two New Species of Pleurothallis (Orchidaceae) from the Amazonian Lowlands Carlyle A. Luer Research Associate of the Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. Mailing address: 3222 Old Oak Drive, Sarasota, Florida 34239, U.S.A. German Carnevali Jardin Botanico de Caracas, Herbario Nacional de Venezuela (VEN), INPARQUES, Aptdo. 2156, Caracas 1010- A, Venezuela. Current address: Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. and Department of Biology, University of Missouri-St. Louis, 8001 Natural Bridge Road, St. Louis, Missouri 63121, U.S.A. Abstract. Pleurothallis discophylla and P. er- ythrogramma are described and illustrated. These new species are distributed in the Amazonian low¬ lands of Venezuela, Ecuador, Peru, and Bolivia. Pleurothallis discophylla has been previously mis- identified in the literature as the Bolivian Pleuroth¬ allis coffeicola , which is almost certainly a synonym of the widespread P. casapensis. The two following undescribed species are locally common and widespread in the western Amazon Basin. Both species are closely related and are char¬ acterized by a shortly creeping rhizome, sharply 3 -winged ramicauls, broadly elliptic to suborbicular, spreading leaves, and yellow or orange flowers pro¬ duced singly on short peduncles. Herbarium speci¬ mens of these plants have been frequently misiden- tified as Pleurothallis coffeicola Schlechter, a species described from Bolivia that is almost certainly con- specific with the frequent Andean species Pleuroth¬ allis casapensis Bindley. Pleurothallis discophylla Luer & Carnevali, sp. nov. TYPE: Ecuador. Morona-Santiago: epi¬ phytic in tall forest along Rio Upano N of Macas, 1,100 m, 15 Jan. 1989, C. Luer , J. Luer, P. & A. Jesup, A. Hirtz & S. Ortega 13927 (holotype, MO). Figure 1. Species haec P. casapensidis Lindley affinis, sed foliis suborbicularis, pedunculis unifloris et labello pyriforrni supra medium orbiculari tenui laevi differt. Plant medium to large in size, epiphytic, shortly creeping, the rhizome stout, 0.5-3 cm long between ramicauls; roots slender. Ramicauls erect, 6-12 cm long, sharply 3-winged, slightly broader toward the leaf, with 2-3 thin, tubular sheaths at the base. Leaf spreading, coriaceous, broadly elliptical to su¬ borbicular, obtuse, 4-7 cm long, 3-5 cm wide, sessile with the base ± cordate. Inflorescence a fascicle of peduncles, successively single-dowered, borne on top of the leaf, subtended by a spathe 5- 7 mm long, from the apex of the ramicaul; peduncles 5 mm long; floral bract 3 mm long; pedicel 3 mm long; ovary cellular-glandular, 2.5 mm long. Sepals yellow to orange, brownish yellow or sometimes to¬ tally maroon, glabrous, the dorsal sepal linear-ob- ovate, obtuse, thickened and subverrucose within toward the apex, 10-11.5 mm long, 3 mm wide, 3-veined, connate to the lateral sepals for 0.5 mm, the lateral sepals connate into an elliptical, bifid synsepal 9-10 mm long, 7 mm wide, 6-veined, the apex shallowly bifid with the tips acute and apiculate; petals translucent or yellow with a purple midvein, elliptical-obovate, obtuse, minutely serrulate, 3 mm long, 1.5 mm wide, with the vein thickened exter¬ nally, ending in a minute apiculum; lip yellow to orange or purple-brown with brownish yellow mar¬ gin, obovate-pyriform in outline, 5 mm long, 3.5 mm wide, the apex broadly rounded, entire, thin, smooth, the margins above the base erect, rounded, the disk minutely spiculate near the middle, with a pair of parallel, subverrucose calli on the middle third, the base thickened, truncate, hinged to the tip of the column-foot; column semiterete, winged above the middle, 3 mm long, minutely toothed at the apex, the foot 2 mm long, the anther, rostellum, and stigma ventral. Etymology. From the Greek discophyllos, “round-leaved,” referring to the broad, flat leaves. This species is closely related to the frequent Pleurothallis casapensis , but P. discophylla is found in warm forests at altitudes below 1,200 m. It is distinguished by the creeping rhizome, erect, sharply triquetrous ramicauls; spreading suborbicular leaves; Novon 3: 158-162. 1993. Volume 3, Number 2 1993 Luer & Carnevali New Species of Pleurothallis 159 Figure 1 . Pleurothallis discophylla Luer & Carnevali. 160 Novon and relatively large, yellow to orange flowers pro¬ duced singly. The dorsal sepal is erect, thickened, and subverrucose toward the apex. The lateral sepals are connate into a broad, shallowly bifid lamina. The petals are minutely serrulate. The lip is pyriform in outline, broadly rounded, dilated and thin above the middle with smooth margins and surface. Above the base the margins are erect. A pair of subverrucose calli are present below the middle of the disk where the surface is minutely spiculate. Several authors have misidentified material of this species as Pleurothallis coffeicola (Schweinfurth, 1967; Foldats, 1970). An illustration of this species from lowland Venezuela was also misidentified as Pleurothallis coffeicola in Venezuelan Orchids Il¬ lustrated (Dunsterville & Garay, 1972). Pleuroth¬ allis coffeicola is a smaller plant that produces sev¬ eral flowers in a raceme and occurs at higher altitudes. It is very probably conspecific with P. casapensis. The closest relative of Pleurothallis discophylla is probably the following species. Paratypes. ECUADOR. Napo: La Cruz, downstream from Misahualli, 400 m, collected by C. & A. Suarez, flowered in cultivation, 29 Mar. 1984, Luer 9784 (MO). PERU. Amazonas: Rio Cenepa, vicinity of Huampami, ca. 4°30'S, 78°30'W, 200-250 m, 1 Aug. 1978, Kujikat 94 (MO). Loreto: Maynas, Mishana, 73°35'W, 03°55'S, primary forest at 130 m, 21 July 1984, Vazquez, Jar- amillo & Criollo 5305 (MO); Distrito Iquitos, km 8 Carretera Quisto Cocha-Varillal, perched forest over sand, 130 m, 24 July 1984, “epiphytic,” S. MacDaniel & M. Rimachi 27836 (MO). BOLIVIA. Cochabamba: epi¬ phytic in seasonally dry forest, Bulo-Bulo, W of Yapacani, 300 m, 22 Aug. 1991, C. Luer, J. Luer, R. Vasquez & l). Ric 15343 (MO). VENEZUELA. Territorio Fed¬ eral Amazonas: Departamento Atabapo, slopes of Cerro Marahuaca, “Sima” area, 03°43'N, 65°30'W, 1,200 m, 19 Oct. 1988, Liesner 25077 (MO, VEN); Cerro Mar¬ ahuaca, Sima Camp, 1,140 m, 21-22, 24 Feb. 1985, “common epiphyte,” Steyermark & Holst 130474 (MO, VEN), Steyermark & Holst 130657 (MO, VEN); De¬ partamento Casiquiare, flooded forest along uppermost Rio Yatua, 100-140 m, 7-8 Dec. 1953, Maguire, Wur- dack & Bunting 36730 (NY), 12 Dec. 1953, Maguire, Wiirdack & Bunting 36763 (NY, VEN), 30-31 Dec. 1954, Maguire, Wurdack & Bunting 37430 (NY); tall forest near river’s edge at Cano Pato, 150 m, Dec. 1965, Dunsterville & E. Dunsterville 966 (AMES, VEN); De¬ partamento Rio Negro, between Cano Temblador and Hauchica, 10 km NE of San Carlos de Rio Negro, 120 m, 1°67'N, 67°03'W, 19 Nov. 1977, Liesner 3672 (MO, VEN); Cano 12 km NE of San Carlos de Rio Negro, 1°56'N, 67°03'W, 120 m, 15 Apr. 1979, “epiphyte on Lecythidaceae, forest over white sand,” Liesner 6659 (MO, VEN). Pleurothallis erythrogramina Luer & Carne- vali, sp. nov. TYPE: Ecuador. Napo: epiphytic in trees along Rio Cascales E of Lumbaqui, 450 m, 9 Feb. 1986, C. Luer , J. Inter, llirtz, Flores & Kmbree 1 1766 (holotype, MO). Figure 2. Species haec P. casapensidis Lindley affinis, sed foliis suborbicularis, pedunculis unifloris, synsepalo rubrolineato et labello elliptico denticulato differt. Plant medium in size, epiphytic, shortly repent, the rhizome stout, 0.5-1 cm long between rami- cauls; roots slender. Ramicauls erect, 6-8 cm long, sharply 3-winged, slightly broader toward the leaf, with 2-3 thin, tubular sheaths at the base. Leaf spreading, coriaceous, broadly elliptical to suborbi- cular, obtuse, 3.5 cm long and 3.5 cm wide to 4.5 cm long and 4.5 cm wide, sessile with the base cordate or subcordate. Inflorescence a fascicle of successive, single-flowered peduncles, borne on top of the leaf, subtended by a broad spathe 2-3 mm long and broad, Irom the apex of the ramicaul; peduncles 1-2 mm long; floral bract 4 mm long; pedicel 6 mm long; ovary 2.5 mm long. Sepals yellow, prominently veined in red, fleshy, glabrous, the dorsal sepal linear, acute, thickened at the apex, 13.5 mm long, 2 mm wide, 3-veined, free from the lateral sepals; lateral sepals connate into an ovate, acute synsepal 10 mm long, 6.5 mm wide, 6-veined, the apex shallowly bifid with the tips acute and apiculate; petals translucent, obovate, acute or ob¬ tuse, minutely serrate, 3 mm long, 1 mm wide, 1 -veined; lip red, elliptical, 5.5 mm long, 2.5 mm wide, the apex narrowly obtuse, denticulate, the margins in the lower third broadly rounded, the disk with a pair of verrucose calli within the margins of the lower third, the base narrowly truncate, hinged to the base of the column-foot; column semiterete, winged above the middle, 3.5 mm long, minutely toothed at the apex, the foot 2 mm long, the anther, rostellum and stigma ventral. Etymology. From the Greek ery thro gramme, “red-line,” referring to the stripes of the synsepal. This pretty species is known from the lowlands of eastern Ecuador and Peru. It is closely allied to the sympatric P. discophylla. Both species are de¬ ceptively alike vegetatively and produce similar flow¬ ers singly, but P. erythrogramma is distinguished by the red-striped sepals with the dorsal sepal con¬ spicuously long and narrow. The red lip is elliptical and minutely crested and denticulate. Paratypes. ECLJADOR. Morona-Santiago: be¬ tween Mision Bomboiza and Gualaquiza, ca. 850 m, 30 Jan. 1971, B. MacBryde 170 (MO). PERU. Amazonas: epiphytic in primary forest S of Aintami, E of Cenepa, 650 ft., 15 July 1974, B. Berlin 1605 (MO). Key Summarizing the Differences betw een the Two New ly Described Species la. Dorsal sepal linear-obovate, obtuse, subverru¬ cose within toward the apex; synsepal broader Volume 3, Number 2 1993 Luer & Carnevali New Species of Pleurothallis 161 L_ _ - — - - 1 Figure 2. Pleurothallis erythrogramma Luer & Carnevali. 162 Novon toward the middle; petals elliptical-obovate; lip yellow to orange or purple-brown with brownish yellow margin, obovate-pyriform, margins en¬ tire, disk smooth or minutely spiculate, ca. 3.5 mm wide; sepals yellow to orange, brownish yellow or totally maroon, nonstriped; spathe 5- 7 mm long . Pleurothallis discophylla lb. Dorsal sepal linear, acute, smooth within; syn- sepal broader toward base; petals obovate; lip red, elliptical, margins denticulate, disk with a pair of verrucose calli, ca. 2.5 mm wide; sepals yellow with red stripes or veins; spathe 2-3 mm long . Pleurothallis erythrogramma Literature Cited Dunsterville, G. C. K. & L. A. Garay. 1972. Pleu¬ rothallis coffeicola. Venez. Orch. Ill. 5: 236. Foldats, E. 1970. Pleurothallis coffeicola. Flora Venez. 15(2): 245-246. Schweinfurth, C. 1967. Orchids of the Guayana High¬ lands. Mem. New York Bot. Card. 14(3): 69-214. New Names for Two Elaphoglossums John T. Mickel New York Botanical Garden, Bronx, New York 10458, U.S.A. ABSTRACT. New names are proposed for two re¬ cently described species of Elaphoglossum whose epithets had already been used. Klaphoglossum killipiamim Mickel, new name for E. killipii Mickel, Novon 2: 376. 1692, not E. killipii Mickel in R. M. Tryon & R. G. Stolze, Fieldiana, Bot., n. s. 27: 138. 1991. Elaphoglossum lanigerum Mickel. new name for E. lanatum M ickel in R. M. Tryon & R. G. Stolze, Fieldiana, Bot., n. s. 27: 139. 1991, not E. lanatum (Bojer ex Baker) Lorence, Fern Gaz. 11: 199. 1976. Novon 3: 163. 1993. Four New Taxa and One Rediscovered Species of Mouriri (Melastomataceae) Thomas Morley Department of Plant Biology, University of Minnesota, St. Paul, Minnesota 55108, U.S.A. ABSTRACT. Three new species of Mouriri from midwestern and southwestern Colombia and eastern Brazil are described and illustrated, as is a new subspecies of Mouriri grandifiora from western Amazonia. Mouriri tessmannii, its type and only known specimen destroyed at Berlin, has been re¬ collected and is redescribed and illustrated. Mouriri Colombians Morley, sp. nov. TYPE: Co¬ lombia. Antioquia: Municipio de San Luis, Ca¬ non del Rio Claro, sector norte, margen de- recha, 5°53'N, 74°39'W, 3 June 1984, Alvaro Cogollo 1796 (holotype, COL; isotypes, JAUM, MO). Figure 1 . Arbor usque 26 m alta; laminae 2.9 8.6 cm longae, 1.8-3. 5 plo longiorae quam latiorae; cryptae stomato- phorae nullae, hypodermis absens; pedunculi plerumque 1-flori, pedicelli 0.8-3. 5 mm longi; calyx 0.5-0. 8 mm findens inter lobos sub anthesi, lobis 1-1.3 mm longis ab staminibus; antherae 3.4-4. 1 mm longae, thecae 2.5- 3.3 mm longae, rimis apicalibus dehiscentes; ovarium 1-loculare, ovula 8-11; fructus globosus exterio viscos- issimo; semina 1 vel fortasse 2, ca. 22 mm alta, 24 mm lata, 1 1 mm crassa. Glabrous tree to 26 m high; young twigs rounded. Petioles 3-6 mm long; blades 2. 9-8. 6 cm long, 1.4-3. 8 mm wide, 1.8-3. 5 times as long as wide, elliptic to ovate-elliptic or narrowly so, acuminate to abruptly so to caudate at apex, acute and abruptly attenuate at base; midrib grooved above, low-round¬ ed below; lateral nerves slightly visible to invisible above and below when dry; leaves drying dark olive green to black. Midrib xylem open above; stomatal crypts none; upper epidermis one cell thick, lacking mucilaginous walls; hypodermis none; crystals oc¬ casional in the upper spongy parenchyma; terminal sclereids stellate with a strong columnar tendency, often with a horizontal central body 2 8 times as long as wide with a columnar extension at each end, sometimes entirely columnar, the columnar parts branching at the epidermis. Flowers axillary and terminal, the peduncles one or two per axil, each 1- or rarely 2- or 3 -flowered, 0.5-4 mm long to base of pedicel measured along the axis with (1- )2(— 3) internodes in that length; bracts 0.7- 1.5 mm long, ovate-triangular, acute, persistent in fruit or deciduous. Flowers pentamerous; true pedicels 0.8- 3.5 mm long; calyx including inferior ovary 4-5 mm long, obconic to campanulate; free hypanthium 2. 3-2. 5 mm long; calyx lobes before anthesis low- rounded and sometimes with a very small apiculum, 0.2-0. 4 mm long, 1.2-1. 4 mm wide, 1-1.3 mm long when measured from the stamen attachment, the calyx splitting between the lobes at anthesis a distance of 0.5-0. 8 mm, the lobes then 1.7-2. 2 mm wide. Petals yellow, 10-11 mm long, 3.5-4 mm wide, narrowly ovate-triangular, acuminate, with a short claw at base. Antesepalous filaments 6.5-7 mm long, antepetalous ones 8 9 mm long; anthers yellow-orange, 3.4-4. 1 mm long; thecae 2. 5-3. 3 mm long, dehiscing by apical slits; gland 0.4-0. 8 mm long, 1.3- 1.9 mm from apex of anther when measured from center of gland; cauda 0.8- 1 mm long. Ovary 1 -locular but with a half partition, the ovules technically axile, 8-11; style 13-15 mm long. All flower parts drying blackish except for the anthers. Fruits globose, bearing the remains of the calyx, 23-40 mm diam. when dry, estimated 27- 50 mm when fresh, then covered with a sticky transparent resin, green to greenish yellow. Seeds 1 or sometimes 2, roughly half spherical, 16-28 mm wide, one studied, this 22.2 mm high, 24 mm wide, 1 1 mm thick, with an irregular polished area at the base of the outer face 13 mm long by 8 mm wide, the remainder of the outer face irregularly grooved or wrinkled. Distribution. Known only from an area in south¬ ern Antioquia, Colombia, about one-third of the way from Medellin to Bogota. Figure 1. Mouriri colombiana Morley. — A. Leaves from various collections. — B. Cleared part of leaf blade showing veins and terminal sclereids. — C. Cross section of leaf blade showing sclereids and upper epidermis. — D. Cross section of leaf midrib. — E. Flowers: a, before anthesis (Renteria 2797)\ b, at anthesis (Cogollo 1796). — F. Longisection of flower before anthesis. — G. Petal. — H. Anthers, cleared to show veins inside: a, Renteria 2797\ Novon 3: 164-175. 1993. Volume 3, Number 2 1993 Morley Mouriri 165 b, Cogollo 1796. — I. Fruit (Cogollo 1434). — J. Seed, showing the hour-glass-shaped polished surface of the outer face (Cogollo 1434). (B, C, D, F, and G are Cogollo 1796.) 166 Novon Table 1. Mouriri colombiana compared with the three most similar species in its section. M. colombiana M. collocarpa M. oligantha M. pauciflora Leaf 1/w ratio 1.8-3. 5 1. 6-2.5 1.7-2. 9 (-3.1) (2.1-) 2. 6-4. 3 Hypodermis absent present present present Number of flowers 1 (-3) (1-) 3-15 1 1 Ovary + calyx length (mm) 4-5 3.8-6 4.8 7 5-8 Calyx lobe length from stamen (mm) 1-1.3 1-1.9 2.1-4 1.8-3. 5 Calyx splitting distance (mm) 0. 5-0.8 1-1.4 2-3 (1.2-) 2-3 Anther length (mm) 3.4-4. 1 2. 1-2. 6 2. 4-4.4 2. 4-4. 4 Theca length (mm) 2. 5-3. 3 1.6-2. 3 2. 1-2.8 2-3.2 Apical slit dehiscence present present absent absent Gland length (mm) 0.4-0. 8 0.8-1. 4 1.1-1. 9 0.6-1 Number of locules 1 3-5 3-4 2 Number of ovules 8-11 12-15 9-12 6-9 Mouriri colombiana is clearly placed in subgenus Taphroxylon sect. Taphroxylon because of the mid¬ rib xylem, which is open above, the evergreen leaves lacking stomatal crypts, the axile placentation, and the long anther thecae, which extend well below the gland. Its affinity to plants of the above section was first recognized by A. Cogollo, who identified his collections 1254 and 1434 as being related to M. collocarpa Ducke. In its section, M. colombiana is most readily distinguished from M. trunci flora Ducke by the latter’s numerous ovules, compared to the 8-11 of the new species. The next most easily eliminated species is the widespread M. acutiflora Naudin, from which M. colombiana differs in lack¬ ing a hypodermis and in having the flowers mostly borne singly, and in its darker colors on drying, apical slit dehiscence of the anthers, and larger seeds. The remaining three species of the group are compared with the new one in Table 1. Mouriri colombiana is best distinguished from M. collocar¬ pa by the former’s lack of a hypodermis, mostly single flowers, short splitting distance of the calyx, longer anthers and thecae, and single ovary locule; from M. oligantha Pilger hy the narrow leaves, lack of a hypodermis, short calyx lobes, short splitting distance, apical slit dehiscence, and single locule; and from M. pauciflora Spruce ex Triana by tbe lack of a hypodermis, short calyx lobes and splitting distance, apical slit dehiscence, and somewhat larger number of ovules. The half partition in the ovaries examined suggests that a full partition might some¬ times occur, giving the two locules also found in M. pauciflora. Mouriri colombiana appears to com¬ bine several relatively unmodified characters with advanced ones. Examples of the former are the lack of a hypodermis in the leaf and the short calyx lobes with their short splitting distance. Advanced features are the mostly single flowers, the unusual anther shape, which resembles that of M. pauciflora , the theca’s apical slit dehiscence, the single ovary locule, and small number of ovules. Paratypes. COLOMBIA. Antioquia: Rio Claro, mar- gen izquierda, 600 m, 16 Sep. 1982 (buds), E. Renteria & A. Cogollo 2705 (JAUM, MO); Alto Rico, 9 Oct. 1982 (buds), E. Renteria , A. Cogollo <£ C. Estrada 2797 (JAUM, MO); Autopista Medellin- Bogota, sector Rio Sa- mana-Rio Claro, camino hacia la vereda La Primavera, 790 m, 13 Nov. 1982 (fr), A. Cogollo & C. C. Estrada 227 (JAUM, MO), 2 km antes de la Josefina, zona muy perturbada, quebrada la salada, 800 m, 22 Sep. 1983 (fr), S. E. Hoyos & J. J. Hernandez 378 (JAUM, MO); margen izquierda, 325 m, 3 May 1984 (fr), A. Cogollo 1683 (JAUM, MO); sector nor -occidental, margen iz¬ quierda 325-500 m, 8 Mar. 1984 (fr), A. Cogollo 1434 (JAUM, MO), 28 Jan. 1984 (fr), A. Cogollo 1254 (JAUM, MO); municipio de San Luis, quebrada “La Cristalina,” 6°N, 74°45'W, Bh-T/Bmh-T, Sector NW, 470-580 msnm, 4 Dec. 1986 (fr), J. G. Ramirez <£ D. Cardenas Lopez 234 (MO), 700-550 msnm, 22 Jan. 1987 (fr), 368 (MO), Sector SE, 770-570 msnm, 23 Feb. 1987 (fr), 624 (MO), 770-500 msnm, 23 Mar. 1987, (fr) 719 (MO), 770-570 msnm, 24 Sep. 1987 (fl buds), 1616 (MO), 27 Oct. 1987 (fr), 1871, 1880 (MO), Carretera de Monteloro al Corregimiento el “Prodigo,” 6°4'N, 74°50'W, 950-600 msnm, 8 Mar. 1990 (fr), D. Car¬ denas L. connati usque ad anthesem, turn ad lineas conjunctionis loborum rumpens, lobis turn tra- pezoideis; ovarium 4-loculare; placentae basilares in quo- que loculo, ovula 3 undique circum quamque placentam genita; fructus subglobosus, lobis calycis coronatus, 4-spermus. Glabrous shrub to 3 m high, sometimes semi¬ climbing; young twigs narrowly 4-winged. Petioles 1 2 mm long; blades 3.5-8. 1 cm long, 1.5-3. 6 cm wide, elliptic to ovate-elliptic, abruptly acute or short- acuminate at apex, rounded to broadly acute at base; midrib plane above, narrowly 2-winged below; lateral nerves slightly to moderately visible above and below when dry, the upper surface then with numerous distinct raised points, which mark the ends of the columnar sclereids. Midrib xylern tubular; stomatal crvpts Type II (see Morley, 1976), mostly elliptic to inverted-flask shape in vertical section, averaging in a leaf 25-50 gm diam., 22-27 high, 71-76 per mm2; upper epidermis one cell thick, all cells with mucilage walls %-% the cell depth; hypodermis none; free stone cells present only in petiole; terminal sclereids columnar at least in the palisade layer, usually angling below, slender, with 1 or 2 branches at each end. Peduncles 1-2 per side, mostly axillary but sometimes at the upper leafless nodes, each 1 - and 7443). — F. Longisection of flower before anthesis. — G. Anther, cleared to show veins. H. Fruit ( Araujo 7443). (B-D, F, G, Araujo el al. 8335.) 170 Novon Table 2. Mouriri arenicola compared with the two most similar species in its section. M. arenicola M. arborea M. megasperma Young twigs 4-winged yes yes-no no Free stone cells no no yes scattered the length of the midrib Petiole length (mm) 1-2 3-10 1.5-3. 5 Blade length (cm) 3.5-8. 1 7-18.5 7.7-10 Upper leaf surface, dry numerous distinct smooth or irregular smooth raised points Winged undermidrib yes yes no Stomatal crypt height 20-27 miii 23-34 gm 45-62 Mm Sclereids columnar-angling filiform columnar-filiform Peduncle length (mm) 4-20 42* 1 oc 2-6.3 Pedicel length (mm) 2-5 3.5-10.5 ca. 2.6 Ovary + calyx (mm) 7-11 10-17 Calyx splitting 5 lobes 2-3 pieces Calyx retention persistent circumscissile Ovule number per placenta 3 4-6 Fruit diam. (mm) 20-30? 20-30? 40-55 Number of seeds 4 4 (?) 5 3-flowered, 3.5-19 mm to base of most distal pedicel measured along the axes and with 1 or 2 internodes in that length; bracts ovate, early deciduous. True pedicels 2-5 mm long; ovary plus calyx in fully developed flower buds 7-11 mm long, ellipsoid to obovoid, the calyx lobes fused for ca. %0 of their length, the free ends triangular, 0.3 0.8 mm long, 0.4-0. 7 mm wide, the tip of the corolla protruding 1-1.5 mm between the free tips, the calyx splitting at anthesis into 5 regular trapezoid lobes, these 3.4- 5 mm long, 3-3.8 mm wide at base, 1.5-2 mm wide at end; calyx limb 5.5-7 mm long measured from the stamen attachment; free hypanthium 2 2.5 mm long. Mature petals unknown. Mature fil¬ aments unknown; anthers 3. 4-4. 5 mm long; thecae 3. 2-3. 9 mm long, dehiscing by apical slits; gland 0.4-0. 8 mm long, 2. 5-3. 4 mm from apex of anther when measured from center of gland; cauda 0.7- 1.2 mm long. Ovary 4-locular; placentae basal in each locule with 3 ovules borne on all sides of a short basal column; mature style unknown. Imma¬ ture fruit subglobose, 16-23 mm high, 20-23 mm diam., crowned with the calyx lobes, which surround an apical pit 1 .5 mm deep; locules 4, each 1 -seeded. Immature seeds with a smooth, hard, polished sur¬ face encircling the hilum but not yet extended to the distal part of the seed, the mature seed unknown. Distribution. Dama Branca dune system, Cabo F rio, ca. 110 km east of Rio de Janeiro; from low to high and shady to sunny situations in the dune vegetation. Mouriri arenicola falls in subgenus Mouriri be¬ cause of its tubular midrib xylem, placentation in which the ovules encircle each placenta, and seed with a polished surface surrounding the hilum. The appropriate section is Olisbeoides because of the long sepals that remain fused till anthesis then split for 3.4 mm or more and are not circumscissile, and because of the large Type II stomatal crypts, which resemble those of M. arborea and M. megasperma. Mouriri arenicola is distinguished from M. luna- tanthera, the least similar species in the section, by the former’s acute to rounded leaf base, winged midrib, columnar sclereids, narrow flower and an¬ thers, and 4-locular ovary. Mouriri arenicola is compared to M. arborea and M. megasperma in I able 2. The new species is distinguished from M. megasperma by the 4-winged twigs, smaller leaves, pimpled upper surface and visible nervation of the dry leaf, winged undermidrib, smaller stomatal crypts, longer peduncles, probable smaller fruits, and lesser seed number; and from M. arborea by the smaller leaves, distinctly pimpled upper surface of the dry leaf, columnar-angling sclereids, smaller flower, 5-lobed calyx at anthesis with the lobes persistent at least on the immature fruit, and the ovules only 3 per locule. Mouriri arenicola is most similar to M. arborea and appears to be reduced from the forebears of that species in response to the isolated and harsh environment. Many of the characteristics of the new species are ones of size reduction, but the sclereid form and the regular separation and persistence of Volume 3, Number 2 1993 Morley Mouriri 171 the calyx lobes are independent of that trend and suggest a retention of a more primitive state than that developed in M. arborea. The status of M. sellowiana (Berg) Burret, a plant yet to be re-collected in the type locality between V itoria and Bahia, is still uncertain. If synonymous with M. arborea, as is probable, the description of its fruit as 21-30 mm high, 25-39 mm in diameter, and 4-seeded would add to the present knowledge of the fruit of that species. Paratypes. BRAZIL. Rio de Janeiro: Mun. de Cabo Frio, Sistema de Dunas Dama Branca, restinga arbustiva fechada, Nome vulgar “Cotia amarela,” 9 May 1986 (fr), I). Araujo 7443 (GUA), 10 Dec. 1987 (fl buds), D. Araujo et al. 8343 (G1 ' A ). Mouriri grandiflora DC. subsp. puberula Vlor ley, subsp. nov. TYPE: Colombia. Municipio de Leticia, Parque Nacional Natural Amacay- acu, Centro Administrativo Mata-mata (Inder- ena), en zona de transicion entre “varzea” y tierra alta (“restinga”) al norte de la quebrada Mata-mata, 70°15'W, 3°47'S, I 10 m, 13 Mar. 1991, A. Hildas, F. del Aguila Joaquin iC (Alberto Moran 1581 (holotype, COL; iso¬ types, MIN, MO). F igure 4. Arbor usque 7 m alta; petioli minute puberuli; laminae 16-24 cm longae, supra glabrae, subtus puberulae; costa infra laminam latissima; cryptae stomatophorae 55-65 jun diametris, 22 jim altae, 60-70 in mm2; sclerides terminales foliariae columnares. Fructus cauliflori, lutei, subglobosi, hypanthio et interdum aliquot lobis calycis coronati, ca. 10-17 mm diametris, 1-5-spermi. Small tree to 7 m high; young twigs terete. Pet¬ ioles 2.5-3 mm long, minutely puberulent; blades 16-24 cm long, 5.5-8 cm wide, ovate-elliptic to narrowly so, abruptly acuminate at apex, shallowly cordate at base with a notch 0.5-5 mm deep, gla¬ brous above, puberulent below with hairs 0. 1-0.2 mm long; midrib plane above, rounded and puber¬ ulent below and widest below the lamina; lateral nerves conspicuous above and below when dry. Sto- matal crypts averaging in a leaf 55-65 jr m diarn., 22 /am high, 60-70 per mm2 (extremes 20-120 jiin diam., 20-25 jim high, 50 85 per mm2); upper epidermis 1-2 cells thick in the same leaf, mucilage walls occasional in the single cells and the inner of the doubled cells; terminal sclereids mostly columnar in whole or in part, sometimes slanting, often branched at one or both ends or up to midway from the ends; hairs usually single, sometimes paired, then 1 long and 1 short, 1 -celled but partitioned with numerous cross walls. Fruits cauliflorous, 1—9 per peduncle; peduncles up to 12 mm long to base ol true pedicel including up to three internodes in that length; fruiting pedicels 2-2.7 mm long, minutely puberulent; fruits yellow, subglobose, drying subgl- obose to depressed obovoid, to ca. 17 mm high, 20 mm diam., crowned with the free bypanthium and sometimes one or more calyx lobes, minutely pu¬ berulent below and above; fruiting free hypanthium 3. 7-4. 5 mm long when dry, 4. 9-5. 4 mm long boiled and presumably when fresh, the outside diameter 5.7 mm dry, 7.1 mm boiled; calyx lobes before anthesis (as seen on young fruits) triangular, mi¬ nutely puberulent, 2 mm long, 1.6 1.7 mm wide, ca. 5 mm long when measured from the stamen attachment, the calyx splitting apart at anthesis a further distance of 2. 2-2. 7 mm. Seeds 1 -5, brown, polished, 9-10 mm high. 8-9 mm wide, 6-7 mm thick, with one or more irregular incomplete hori¬ zontal creases below the middle, the basal hilum as wide as the base of the seed. Distribution. Tropical forests in lar western Ama¬ zonia, one locality in the lar southeast corner of Colombia near Leticia, the other in mid-eastern Acre near the Amazonas border. The new taxon falls in or near Mouriri grand i- fiora by virtue of its leaf size and shape, tubular midrib xylem, simple stomata] crypts, epidermal structure, lack of a hypodermis, and the close re¬ semblance of its fruiting calyx, fruits and seeds to those of M. grandiflora . From the latter species and all others the new group differs in its puberulent lamina; the larger stomatal crypts and midrib shape also distinguish it, and the thick columnar sclereids are extreme for the species. The only other species in the genus with any pubescence on the leaf is M. myrtilloides, in which only the midrib is puberulent, and that species is in a different section and bears little resemblance to M. grandiflora. However, pu- berulence or hints ol it occur occasionally in M. grandiflora subsp. grandiflora . Two collections (Prance et al. 25775 and l asquez et al. 11298) were noted to have papillae or very short hairs up to 0.07 mm long on the under-midrib only, and several collections have low papillae on the lamina. The papillae are 9-18 jun long with at most a single cross-wall: these plants resemble subspecies gran¬ diflora in all other respects: they have no stomatal crypts or small very shallow ones, and their sclereids are usual for the species. The crypts of the new subspecies average 55-65 jun diam., while those of the main subspecies are 25-40 jim. A further dif¬ ference is apparently found in the rounded under¬ midrib, which is widest below the lamina, a peculiar feature not noted in subspecies grandiflora, al¬ though it may have been overlooked. The overall resemblance between the two taxa 172 Novon Figure 4. Mouriri grandijlora DC. subsp. puberula Morley (Rudas et al. 1584). — A. Cleared part of leaf blade showing veins and terminal sclereids. — B. Cross section of leaf blade showing sclereids, stomatal crypts, upper epidermis, and hairs. — C. Leaves. — D. Cross section of leaf midrib. — E. Fruit. — F. Seed. suggests that in prudence they should be considered as part of a single, variable complex. At the same time, in view of the distinct gap between them in pubescence and stomatal crypts and the hard-to- evaluate differences of midrib shape and foliar scler¬ eids, as well as the present lack of information on petals, stamens, and ovary, the possibility remains that in the future it may be necessary to elevate the rank of the new group. Paratype. BRAZIL. Acre: Municipio de Sena Mad- ureira, a 4 km da margem direta do Rio Iaco, 5 Oct. 1980, C. .4. Cid & B. W. Nelson 2786 (MIN, NY). With the recent rediscovery of Mouriri tess- Figure 5. Mouriri tessmannii Markgraf. — A. Leaves (a, Palacios et al. 3365; b, Gudino 127; c, Espinoza 57). — B. Cleared part of leaf blade showing veins and terminal sclereids. — C. Cross section of leaf blade showing sclereids and upper epidermis. — D. Cross section of leaf midrib. — E. Inflorescences: a, flower just before anthesis; b, Volume 3, Number 2 1993 Morley Mouriri 173 inflorescence at anthesis. Some divisions of the calyx are two calyx lobes wide, some only one. — F. Longisection of flower before anthesis. — G. Anthers: a, cleared and expanded in NaOff showing the veins (boiled anthers take the same shape); b, dried. — H. Petal. — I. Immature fruit (Palacios et al. 3365). (B-H are Gudino 127.) 174 Novon mannii after the destruction of the holotype and only specimen at Berlin in World War II, it becomes necessary to designate a neotype. Mouriri tessmannii Markgraf, Berlin Univ. Bot. Gard. Notizbl. 9: I 147. 1927. NEOTYPE: Ec¬ uador. Napo: Canton Orellana, Sector Huash- ito, 20 km al norte de Coca, Propiedad de PALMORIENTE, 00°20'S, 77°05'W, 250 m, 3-21 Nov. 1989, Edgar Gudina 127 (neo¬ type, MO; isoneotype, MIN). Figure 5. The following description is based on the three extant collections; characters of the holotype agree so far as can be determined, except where indicated: Small trees to 8 m high with trunks to 7 cm diam., the holotype from a tree with a trunk 27 cm diam. and not-very-hard reddish wood, the branches in the latter tree beginning at 5 m above the ground; bark scaly; young twigs rounded; plants glabrous except for the inflorescence. Petioles 2. 5-3. 5 mm long; blades dark green above and very smooth when fresh, light green below, coriaceous and brittle, 11- 19 cm long, 4.2-7 cm wide, elliptic to ovate-elliptic, usually narrowly so, abruptly acuminate at the apex, acute to rounded or abruptly so or nearly truncate at base, sometimes oblique and half-cordate on one side; midrib plane to slightly rounded or very slightly sunken above, prominent and rounded to slightly 2-angled below; lateral nerves slightly visible above and prominent below when dry; surface view of blade at 50 X with irregular curving rows of shiny rounded humps, the rows often crossing at various angles, each bump apparently an epidermal cell. Midrib xylem tubular; stomatal crypts none; upper epi¬ dermis lacking all pigments, appearing as a dull white layer in unstained hand-sections at 50 X , varying from 1 to 2 cells thick in the same leaf, mostly one cell thick, mucilaginous walls common, mostly in the single cells, less often in the inner of a pair; hypodermis none; terminal sclereids stellate with a strong columnar tendency, the central body 1-4 times as long as wide, often the whole sdereid co¬ lumnar with sharp arms at each end. Inflorescences on trunk or on leafless branches as little as 5 mm thick, 1-5 per side, each 1-3-flowered, 2-10 mm long to base of farthest pedicel measured along the axes (12-15 mm in the holotype) and with 1 or 2 internodes in that length; bracts 1-1.5 mm long, triangular, acute, early deciduous. Flowers pentam- erous; true pedicels 2.5-5 mm long; pedicels and lower part of ovary and calyx minutely puberulent; calyx including inferior ovary yellowish green to yellow, 10-13 mm long, obovoid (14-15 mm long and ellipsoid in the holotype), the lobes nearly com¬ pletely fused, their free tips 0. 3-0.8 mm long, 0.4- 0.8 mm wide, the calyx bursting at anthesis usually along some or all of the fusion lines into 3, 4, or 5 ovate-triangular acute lobes of single or double width, sometimes breaking more irregularly, the single lobes 3.3-5 mm long by 3-4.8 mm wide, the double ones 3-4 mm long by 5-7 mm wide; the longest torn edge 4— 5(— 6) mm; free hypanthium 4-5.7 mm long; calyx limb 4-5.5 mm long as measured from the stamen attachments. Petals purple (“morados”) or rose (holotype), separate in the neotype, reported connate in the holotype, elliptic-oblong or elliptic- ovate with the margin irregularly crisped, reported triangular in the holotype, 9.5-10.5 mm long, 6- 7 mm wide, acuminate, with a basal claw 0.5-2 mm long by 2.5-4 mm wide. Filaments 10-12 mm long; anthers 5. 5-6. 4 mm long; thecae 5.5 6.2 mm long, dehiscing by apical slits; gland 0. 3-0.6 mm long, 3.5-4. 1 mm from apex of anther when measured from center of gland; cauda none; anther shrinking conspicuously in width (theca to gland) on drying so as to change the appearance of the anther and increase the width of the pores. Ovary 5-locular; ovules axile-basal, produced only outwardly from each placenta, 4 per locule; style 19-22 mm long. Fruit subglobose, crowned with the hypanthium and remains of the calyx, 12-ca. 20 mm high excluding hypanthium and calyx, 1 1-ca. 20 mm diam., 1-5- locular with 1 seed per locule, symmetric when 2- 5-seeded, asymmetric when 1 -seeded; partly ma¬ tured seeds with the lower and only mature part polished all over, mature seeds said to be cuneate in the holotype. Distribution. Primary tropical forests of east Ec¬ uador and northwest Peru; soils red or black; 160- 320 m. Mouriri tessmannii is characterized in its section primarily by its lack of stomatal crypts and lack of pigments in the upper epidermis, by the nearly com¬ plete fusion of the calyx lobes in bud, which split apart variously at anthesis, and by the anther form, which changes noticeably on drying. The identity of the three recent collections as Mouriri tessmannii is judged to be reasonably cer¬ tain, although not an exact fit. The original descrip¬ tion is brief; it omits the ovary characters and the anther detail and seldom gives exact measurements, while the photo does not include the petals and stamens, which were in a closed pocket. The three new collections are a good match among themselves. However, four of the discrepancies between these three and the holotype noted in the above description should be remarked on: these are the plant size, calyx and ovary shape, and petal shape and fusion. Volume 3, Number 2 1993 Morley Mouriri 175 I lie greater size of the holotype is probably not significant, since many species of Mouriri exhibit this great a variation, including a close relative, M. grandiflora. The ellipsoid shape of the calyx and ovary as opposed to obovoid does not appear in¬ surmountable considering the size differences of the plants and the fact that they come from different parts of the geographic range. Petal shape is no¬ toriously variable in subgenus Mouriri , and elliptic- ovate might be called triangular in less exact days. Only the petal fusion appears to be a major dis¬ crepancy. Markgraf termed the petals “inter se con- nata”; in 1927 that might for him have included a light adhesion, and an adhesion could have been caused by a sticky fluid of some sort. In view of the many features in common between M. tessmannii and the three collections in question, I conclude that the connate petal statement is a mistake. The petals are separate in all other known American species of the tribe; fusion would be a large departure from the norm and is hardly to be expected. The three new collections are from an area well north of the type locality, the latter being at the mouth of the Santiago on the Mararion in Peru. Palacios el al. 3365 comes from east-central Fc- uador about 365 km NNE by N from the type locality, and the other two collections are from a site 130 km north of the Palacios one. These are not unusual distributional gaps for species in this region, considering the difficulty of making adequate collections. The limited habitat data is congruent. Previously (Morley, 1976), it was estimated from the description and photo that M. tessmannii be¬ longed in subgenus Mouriri sect. Olisbea. Ibis placement is substantiated by the neotype and other collections. Specifically, the species falls in the sub¬ genus on the basis of its tubular midrib xylem, axile- basal placentation, polished seeds, common muci¬ laginous walls in the epidermis, and rose to purple petals. Characters placing these plants in section Olisbea are the lack of stomatal crypts in large ovate-elliptic acuminate leaves, pentamerous flowers with long anthers and anther thecae, and calyx lobes nearly fully fused with the petals never protruding before anthesis. In its section, Mouriri tessmannii is unique in its curious epidermis, which lacks all pigments; all the other species have a pigmented epidermis. Be¬ yond this the species appears most similar to M. rhizophoraefolia and M. completens because of their closed calyces and large leaves lacking stomatal crypts. Mouriri tessmannii differs from M. com¬ pletens primarily in having rounded to acute leaf bases, irregular calyx breakage, and different anther and fruit forms. From M. rhizophoraefolia , M. tessmannii departs in having larger leaves, less strictly columnar foliar sclereids, an acute apex to the closed calvx rather than acuminate, and a dif¬ ferent anther form. The third species in the section with a closed calyx, M. froesii , diverges further than the preceding two, having stomatal crypts, a midrib that is grooved above and winged below, a different anther shape, and only four locules in the ovary- Although one’s attention is drawn to the preceding plants by the common feature of sepal fusion, this feature may have evolved independently from a basal group, thus the true closest relationships may lie with species lacking that character. Additional specimens examined. ECUADOR. INapo: Canton Orellana, Sector Huashito, 20 km al norte de Coca, Propiedacf de PALMORIENTE. 00°20'S, 77°05'W, 250 m (deflor), 3-21 Nov. 1989, S. Espinoza 57 (MIN, MO). Pastaza: V ia Auca, 110 km al sur de Coca, a 10 km del Rio Tigiiino, Sector Cristal, 01°15'S, 76°55'W, 320 in, 7 Jan. 1989 (imm. fr), W. Palacios , C. Iguago & F. Hurtado 3365 (MIN, MO). Literature Cited Morley, T. 1976. Memecyleae (Melastomataceae). FI. Neotrop. 15: 1-295. A New Species of Parathesis (Myrsinaceae) from Ecuador John J. Plpoly III Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. ABSTRACT. Parathesis palaciosii is described and illustrated, and its phylogenetic relationships are dis¬ cussed. The genus Parathesis J. D. Hooker contains 130 species distributed southward from southern Mexico to Panama, the Caribbean, and through the Andes from Venezuela to Peru. The genus is defined by unique glandular papillae, which are found on the calyx and corolla lobes, and bright yellow anthers. During determination of specimens from the PRO- MOBOT (Promocion Botanica) project in Ecuador, a new species was encountered, which is here de¬ scribed. Parathesis palaciosii Pipoly, sp. nov. TYPE: Ec¬ uador. Napo: Canton Tena, Estacion Biologica Jatun Sacha, Rio Napo, 8 km E of Misahualli, 1°04'S, 77°36'W, 400 m, 12-14 Dec. 1989 (fl), Palacios 4761 (holotype, QCNE; isotypes, F, MO, NY, TEX, US). Figure 1. Propter laminas chartaceas ad apices acuminatas ad bases acutas decurrentes, secus margines crenulatas, pe- tiolo marginato, sepala ovato-triangulares, petala lineari- lanceolata atro-lineato-punctataque, anthera primo erecta tarde versatilis P. pyramidalis valde arete affinis, sed ab ea larninis ellipticis vel oblongis (non oblanceolatis vel obovatis), nerviis secundariis conspicuis (non inconspicuis) desuper immersis (nec planis), sepalis 1.5 -1.8 (non 1- 1.2) mm longis, pellucido-(nec atro-)punctatis, squamis lepidotis translucentibus indutis (nec elepidotis), petalis 7- 9 (non 5-7) mm longis, staminibus 7-8.1 (non 3-4) mm longis, filamentis 6. 4-6. 7 (non 2. 3-2. 5) mm longis, an- theris apiculatis (nec obtusis) statim distinguitur. Tree to 20 m tall, 35 cm DBH; trunk prominently fenestrate, sap copious, clear, odorless; branchlets subterete, 3.5-5 mm diam., finely appressed fer- rugineous stellate-tomentose. Leaf blades charta- ceous, elliptic to oblong, (8— )14— 17(— 1 9) cm long, (3.7 )5-7(-7.5) cm wide, apex abruptly acuminate to caudate, base acute, decurrent on the petiole, midrib raised above and below, secondary veins (19-) 20-35 pairs, immersed above, prominently raised below, glabrous above except along midrib, sparsely and minutely ferrugineous stellate-tomentose below, bizonal, the trichomes more densely packed along the midrib area and the veins, densely and promi¬ nently punctate, the margin slightly revolute, cren- ulate; petioles marginate, (1.8 )2. 5 3 cm long, gla¬ brous above, densely and minutely ferrugineous stellate-tomentose below. Inflorescence bisexual, ter¬ minal, pyramidal, bipinnately paniculate, (6-)12- 16 cm long, (6-) 10- 12 cm wide at base, the prin¬ cipal branches subtended by small leaves, the leaf blades chartaceous, elliptic, 4. 5-7. 5 cm long, 1.7- 3.5 cm wide, apex acuminate, base acute, decurrent on the petiole, otherwise as in vegetative leaves; petioles marginate, 0.7-1 cm long, glabrous above, sparsely and minutely ferrugineous stellate-tomen¬ tose below; peduncle obsolete to 1 .5 cm long, rachis, branches and pedicels densely appressed ferrugin¬ eous stellate-tomentose; floral bracts chartaceous, linear-lanceolate, 2.5-3 mm long, ca. 0.1 -0.2 mm wide, apex subulate, early caducous; pedicels cylin¬ drical, erect, 4. 8-7. 2 mm long, not accrescent in fruit. Flowers corymbose, 5-merous, 7-9.2 mm long, calyx brown, chartaceous, inconspicuously pellucid punctate, densely and minutely tomentose, the se¬ pals ovate-triangular, 1.5- 1.8 mm long, 0.9- 1.3 mm wide, apex acute, sparsely translucent-lepidote, the margins entire; petals pink, chartaceous, linear- lanceolate, 7-9 mm long, 1.5- 1.9 mm wide, apex subulate, densely and prominently black punctate- lineate (the punctations linear), densely and minutely stellate-tomentose without, densely translucent pa- pillose-tomentose along the margin within and throughout the distal (4; stamens 7-8.1 mm long, erect, the filaments flat, 6. 4-6. 7 mm long, promi¬ nently black punctate-lineate, the anthers erect, then tardily versatile, lanceolate, 2. 2-2. 5 mm long, 1- 1.2 mm wide, apex apiculate, glabrous, base sag¬ ittate, dehiscent by longitudinal slits; ovary globose, densely tomentose with erect hairs, the style filiform, accrescent to 1 1 .7 mm long, the placenta depressed- globose with 4-5 uniseriate immersed ovules. Fruit depressed-globose, 6-8 mm long, 10-12 mm diam. at maturity, black, the exocarp thick, juicy, sweet. Distribution. Parathesis palaciosii is endemic to the Ecuadorean Amazon in the area of the Jatun Sacha Biological Reserve, 400-450 m elevation. Ecology. This species occurs in tall very humid forest on rolling hills of lateritic soils. The species is frequently encountered and is a conspicuous el¬ ement of the subcanopy, according to the collector. Novon 3: 176-178. 1993. Volume 3, Number 2 1993 Pipoly Parathesis palaciosii 177 Figure 1. Parathesis palaciosii Pipoly. — A. Habit, showing crenulate leaves and reduced leaves subtending inflorescence branch. — B. Abaxial leaf surface, showing slightly revolute margin, tomentum, and marginate petiole. — C. Adaxial leaf surface. — D. Flowers in bud and at anthesis, showing the glandular-papillose, punctate-lineate petals, punctate-lineate filaments, and apiculate anthers with dorsal punctations. — E. Pistil in longitudinal section, showing the depressed-globose placenta with immersed, uniseriate ovules. — F. Fruit, showing acute calyx lobes. A- E drawn from isotype, F drawn from Palacios 4262. 178 Novon Etymology. The species is named for Walter Palacios, specialist in Meliaceae, forestry engineer and parataxonomist for the Missouri Botanical Gar¬ den’s PROMOBOT program in Ecuador. Parathesis palaciosii is most closely related to P. pyramidalis Lundell, of the Sierra de Santa Marta, Colombia, because of its acuminate, char- taceous, crenulate leaves, ovate-triangular sepals, linear-lanceolate, black punctate-lineate petals, and anthers that are at first erect, then versatile in post anthesis. However, the deeply impressed secondary veins, longer, inconspicuously punctate and trans¬ lucent lepidote sepals, longer petals and stamens, and apiculate anthers easily distinguish this taxon. The collector reports that the trunks are fenestrate, which is the first report of that trunk morphology for the genus. Paratypes. ECUADOR. Napo: Canton Tena, Estacion Biologica Jatun Sacha, Rio Napo, 8 km E of Misahualli, 1°04'S, 77°36'W, 400 m, 12-14 Dec. 1989 (fr), Pa - lacios 4262 (AAU, F, GB, MO, NY, QCNE, US), 6 May 1990 (fl), Palacios et al. 4940 (AAU, F, GB, MO, NY, QCNE, US), 28 Dec. 1987 (ster.), Gentry et al. 6 0029 (MO, QCNE). Acknowledgments. The Promocion Botanica Project of the Missouri Botanical Garden in Ecuador is possible thanks to the generosity of the Liz Clai¬ borne Foundation. My studies in Andean biodiversity are supported by the Andrew W, Mellon Foundation and the John D. and Catherine T. MacArthur Foun¬ dation. I thank Linda Ellis for yet another fine il¬ lustration. Lectotypifications in the Genus Petrea (Verbenaceae) Ricardo M. Rueda Departamento de Biologia, Universidad Nacional Autonoma de Nicaragua, Leon, Nicaragua; present address: Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, IJ.S.A. ABSTRACT. Twenty-one specific epithets in Petrea need lectotypification. Of these, 17 holotypes were not originally designated, two types were destroyed during World War II, and two are missing. As a result of an ongoing study toward a mono¬ graph of the genus Petrea, a number of specific epithets have been found to need lectotypification. Lectotypifications were not made in Moldenke’s (1938) monograph of the genus. During the course of this study, most of the types of Petrea were seen via loans or herbarium visits. The objective of this paper is to lectotypify the species for which a ho- lotype was not designated or whose holotype has been destroyed or is missing. In general, the spec¬ imens designated here as lectotypes have been se¬ lected as the most representative material available among the eligible collections, and are not neces¬ sarily the collections deposited at the author’s in¬ stitution (see Phillips et ah, 1992, for details). Petrea algentryi Moldenke, Phytologia 54: 67. 1983. TYPE: Colombia, Gentry et al. 37075 (lectotype, selected here, MO; isolectotype, MO). This type was collected along the Rio Tagachi, 12 km west of Rio Atrato, Choco, Colombia, on 19 June 1982. According to Moldenke (1983), the holotype was deposited at the University of Texas (TEX), but this type was not found there. In addition, this holotype was never part of Moldenke’s herbar¬ ium (G. Nesom, pers. comm.). Two specimens of the same collection are at the Missouri Botanical Garden (MO), marked with the word “type.” The aforementioned seems to indicate that the holotype is missing. Therefore, one of the sheets at MO is designated as lectotype. Petrea arborea Kunth, Nov. Gen. Sp. 2: 282- 283. 1818. TYPE: Venezuela, Humboldt & Bonpland s.n. (lectotype, selected here, P). This type was collected in Aragua, Venezuela, near the town of Cura, in February 1800. A holotype for this species was not designated. No specimen for this collection exists in the Humboldt -Bonpland or Willdenow herbaria, seen on microfiche. A specimen was found at P and is designated as lectotype. Petrea aspera Turczaninow, Bull. Soc. Imp. Nat. Mosc. 36: 211. 1863. TYPE: Venezuela, Funk & Schlim 507 (lectotype, selected here, BR; isolectotypes, BM, C(2), P, W; photo of iso¬ lectotype, F, MO, NY(2), TEX, U, W). This type was collected in Carabobo near San Estevan, Venezuela, at 300 m, in April 1845. A holotype for this species was not designated, and there are numerous extant syntypes. The best spec¬ imen of the original collection is designated as lec¬ totype. Petrea atrocoerulea Moldenke, Feddes Report. 43: 195. 1938. TYPE: Colombia, Kalbreyer 1034 (lectotype, selected here, K; fragment of isolectotype, NY; photo of isolectotype, NY, TEX). This type was collected along the Rio Porce, Antioquia, Colombia, on 7 May 1880. Moldenke (1938) designated as the holotype for this species a specimen deposited in B. However, P. lliepko, the curator of B, informed me that this type was de¬ stroyed in World War II. B< ;cause of this, the isotype at K is designated as lectotype. Petrea blanehetiana Schauer in DC., Prod. 1 1: 617 618. 1847. TYPE: Brazil, Martius 1020 (lectotype, selected here, BR; isolectotypes, BM, G(2), GH, L, LE, M(2), MO, NY, S; photo of isolectotype, NY). This type was collected in Bahia, with no collec¬ tion date given. A holotype for this species was not designated. The species was based on three different collections, Martius 1020, Blanche t 01, and Salz- mann 433. Among these, Martius’s specimen has more complete label data than the other collections and is duplicated in more herbaria; it agrees with the protologue, and most of the specimens are in good condition. The best specimen of the Martius collection, deposited at BR, is designated as lecto- type. Novon 3: 179-181. 1993. 180 Novon Petrea bracteata Steudel, Flora 26: 764. 1843. TYPE: Suriname, Hostmann & Kappler 39 (lectotype, selected here, W; isolectotypes, B, BM, C, F, G(2), K, LE, MO, S, U, W(2); fragment of isolectotype, MO, NY; photo of isolectotype, MO, NY, TEX; photo of two is¬ olectotypes overlaid in the same picture, NY). This type was collected in dense forest, in April 1842; a holotype for the species was not designated. The best of the three original specimens deposited at W is chosen as lectotype. The herbarium of the New York Botanical Garden (NY) has a sheet an¬ notated as a photograph of the type, which looks different from the rest of the collection. However, this photograph was taken from two overlaid sheets of the three original specimens at W. Petrea brevicalyx Ducke, Bull. Mus. Hist. Nat. Paris, ser. 2, 4: 748-749. 1932. TYPE: Bra¬ zil, Ducke s.n. (lectotype, selected here, NY; isolectotypes, G, K, RB, S, U, US; photo of isolectotype, GH(2), NY, TEX). This type was collected in Igarape da Cachoeira Grande, Brazil, on the river shore in Amazonas, near Manaus, with no collection date given. This species was based on two collections, Ducke s.n. and Kuhl- mann s.n., which are both deposited at RB. The Ducke s.n. collection has better specimens and was distributed to more herbaria than Kuhlmann s.n. Therefore, the best specimen of the former collection is designated here as the lectotype. Petrea glandulosa Pittier, Bol. Cient. Teen. Mus. Com. Venez. 1: 70. 1925. TYPE: Venezuela, Peraza 1 1532 (lectotype, selected here, VEN; isolectotypes, NY, US; photo of lectotype, NY, TEX; photo of isolectotype, NY, TEX). This type was collected at Cerce de Guanare, Portuguesa, Venezuela, in March 1924. A holotype for this species was not designated. The specimen in VEN, which is in good condition, is selected as the lectotype. Petrea kohautiana C. Presl, Bot. Bemerk. 99. 1844. TYPE: Windward Islands, Kohaut s.n. (lectotype, selected here, BR; isolectotypes, B(2), F, G, L, M; fragment of isolectotype, NY). This type was collected on the Island of Marti¬ nique, 1819-1821. According to Stafleu & Cowan (1983), the types were deposited at PR and PRC. Moldenke (1938) stated that the type was in PR, hut he did not see it. However, both herbaria in¬ dicated that they do not have this holotype in their possession. Because of uncertainty as to whether the putative holotype indicated by Moldenke exists, the sheet at BR, which has the best available ma¬ terial, is designated as lectotype. Petrea macrostachya Bentham, Ann. Nat. Hist., ser. 1, 2: 448. 1839. TYPE: Guyana, Schom- burgk 158 (lectotype, selected here, K; isolec¬ totypes, BM, BR, F, G, GH, K(2), LE; fragment of isolectotype, NY; photo of isolectotype, F, MO, TEX). This type was collected along the Currapawaak brook, Guyana, 1835-1839. The type of this spe¬ cies was deposited at K and consists of two sheets of the Schomburgk 158 collection. However, be¬ cause the author of the name did not specifically designate one as holotype, a lectotype is designated here. Petrea martiana Schauer in DC., Prod. 1 1: 620. 1847. TYPE: Brazil, Martius s.n. (lectotype, selected here, M; fragment of lectotype, NY). This type was collected in woods at San Antonio de Gurupa, Para, Brazil, 3-16 Sep. 1819. This species was based on two unnumbered Martius col¬ lections, collected in the same year and deposited at M. Of these, one was collected in Para and the other in Amazonas. The specimen from Para in M is selected as lectotype. Petrea maynensis Huber, Bol. Mus. Para 4: 602. 1906. TYPE: Peru, Huber 1489 (lectotype, selected here, MG; isolectotype, RB; fragment of lectotype, F; photo of lectotype, F, NY, TEX; photo of isolectotype, GH). This type was collected along the Rio Ucayali, between Contamana and Canchahuaya, Loreto, Peru, in November 1898. A holotype for this species was not originally designated. The best specimen of the collection deposited at MG is designated as lectotype. Petrea mexicana Chamisso, Linnaea 7: 367. 1832. TYPE: Mexico, Berlandier 136 (lectotype, se¬ lected here, G; isolectotype, L; photo of lec¬ totype, TEX). This type was collected in Tampicos, Tamaulipas, Mexico, in 1829. A type for this species was not designated. Of the two extant duplicates, the best specimen is deposited at G and is here selected as lectotype. Petrea obtusifolia Bentham, PI. Hartw. 246. 1846. TYPE: Colombia, Hartiveg 1359 (lec- Volume 3, Number 2 1993 Rueda Lectotypifications in Petrea 181 totype, selected here, K; isolectotypes, BM, G; fragment of isolectotype, NY; photo of lecto- type, NY, TEX). This type was collected in Tolima, Colombia, in 1 843. A holotype for this species was not designated in the protologue. The specimen at K might be the holotype, hut this should not be concluded auto¬ matically just because Bentham worked at k (Phil¬ lips et ah, 1992). This specimen, however, is in good condition and is selected here as lectotype. Petrea ovata M. Martens & Galeotti, Bull. Acad. Roy. Sci. Belgique 11: 329. 1844. TYPE: Mexico, Galeotti 793 (lectotype, BB; isolec¬ totypes, BR, G(2), GH, EE, U; fragment of isolectotype, NY; photo of lectotype and iso¬ lectotype, TEX). This type was collected in Jalapa, Veracruz, Mex¬ ico, in August 1940. A holotype for this species was not designated. Moldenke (1938) assumed that the holotype was at BR and did not designate a lectotype. The specimen at BR that is in good condition is designated as lectotype. Petrea pubescens Turczaninow, Bull. Soc. Imp. Nat. Mosc. 36: 211 212. 1863. TYPE: Ven¬ ezuela, Funck & Schlim 1504 (lectotype, se¬ lected here, BM; isolectotypes, BR, G(2), P, U, W; photo of isolectotype, F, MO, NY, TEX). This type was collected at Merida near San Cris¬ tobal, Venezuela, at 600 m, in November 1846. A holotype for this species was not designated, and there are many duplicate sheets. The best specimen from the collection has been selected as lectotype. Petrea racemosa Nees, Flora 4(1): 300. 1821. TYPE: Brazil, Wied-Neuwied s.n. (lectotype, BR; isolectotype, G; photo of lectotype, MO, NY, TEX). This type was collected at Bahia, Rio Grande de Belmonte, Brazil, 23 July-21 Dec. 1816. A holo¬ type for this species was not designated. Moldenke (1938) assumed that the holotype was deposited at BR and did not designate a lectotype. The specimen at BR that is in good condition is designated as lectotype. Petrea rugosa Kunth, Nov. Gen. Sp. 2: 282. 1818. TYPE: Venezuela, Humboldt & Bonp- land s.n. (lectotype, selected here, P-HB; is¬ olectotypes, F, P; photo of lectotype, TEX; photo of isolectotype, E, MO, NY(2), TEX). This type was collected at Distrito Federal, Ca¬ racas, Venezuela, without a date. A holotype for this species was not designated. The best specimen is in the Humboldt Bonpland herbarium at P and is selected as lectotype. Petrea schomburgkiana Schauer in DC., Prod. 11: 619. 1847. TYPE: Guyana, Schomburg 108 (photo-lectotype, selected here, MO; du¬ plicate photo, TEX). The precise locality of the collection is unknown. The holotype was deposited in B, and several pho¬ tographs exist. However, P. Hicpko (pers. comm.) indicated that the type was destroyed during World War II. In addition, there is no other specimen of the original collection. Fortunately, the photograph available at MO is sufficient to establish the identity of the species and is selected as lectotype. Petrea subserrata Chamisso, Linnaea 7: 368. 1832. TYPE: Brazil, Sellow s.n. (lectotype, selected here, K). The precise locality ol collection and date are unknown. This species was based on several un¬ numbered collections of Sellow. The best specimen from among these collections is selected as lectotype. Petrea vincentina Turczaninow, Bull. Soc. Imp. Nat. Mosc. 36: 212. 1863. TYPE: Windward Islands, Coley s.n. (lectotype, selected here, G; isolectotype, G; photo of isolectotype, NY). I bis type was collected in St. Vincent, Windward Islands, with no date given. A holotype was not designated for this species, and the original collection consists of two sheets both deposited in G. The best specimen of those in G is designated as the lectotype. Acknowledgments. I thank W. D’Arcy, A. Gen¬ try, and H. van der Werff for critical reading of the manuscript. I also thank the curators of the herbaria cited for loaning types of Petrea. This work was supported by NSF grant INT-9024094. Literature Cited Moldenke, H. N. 1938. A monograph of the genus Petrea. Repert. Spec. Nov. Regni Veg. 43: 1-48, 127-221. - . 1983. Notes on new and noteworthy plants. CLXIX. Phytologia 54: 66-68. Phillips, S. M, R. K. Brummitt & B. P. J. Molloy. 1992. Problems of types with duplicate specimens. Taxon 41: 728-73CL Stafleu, F. A. & R. S. Cowan. 1976. Taxonomic Lit¬ erature. Vol. I: AG. Bohn, Scheltema & Holkema, Utrecht. 7. New Species and Combinations in Ceradenia (Grammitidaceae) Alan R. Smith University Herbarium, University of California, Berkeley, California 94720, U.S.A. ABSTRACT. Ceradenia asthenophylla, C. aulaei- folia, and C. tristis are described from Mesoam- erica, and their relationships are discussed. Fifteen species of Ceradenia are now known from the re¬ gion; all are characterized by peculiar whitish glan¬ dular paraphyses and lack of hydathodes. Three new combinations are also made in the genus for Andean species: Ceradenia bishopii, Ceradenia intricata, and C. semiadnata. So that names will be available for the pterido- phyte volume of Flora Mesoamericana , three new species of Ceradenia are herein described. These were recognized as new on specimens annotated by L. E. Bishop, who began a revision of the genus (Bishop, 1988, 1989) but was unable to complete it. In addition to the new species, three new com¬ binations are made for species occurring in South America. These were also contained in an unpub¬ lished manuscript by Bishop. Ceradenia asthenophylla L. E. Bishop ex A. R. Smith, sp. nov. TYPE: Colombia. Huila: Cor¬ dillera Central, E slope between Paramo de las Papas and San Antonio, 2,900 m. Bishop 1984 (holotype, UC). f igure 1I-L. Ceradeniae knightii affinis, a qua imprimis differ! stip- itibus sparsim setosis (non dense setosis), laminis 6-10- plo longioribus quam latioribus (non 4-6-plo longioribus quam latioribus), laminis tenuis (non incrassitis) et tri- chomatibus numerosis ramosis glandulosis praeditis. Plants epiphytic; rhizome slender, with atropur- pureous scales 2-4 x 0.3-0. 4 mm, with concol- orous setulae or glandular hairs; petiole brown, 2- 4 cm x 0.2-0. 4 mm, with a few scattered casta- neous setae to 3 mm distally and much more nu¬ merous simple or branched hairs throughout its length, 0.25-0.35 times the length of the lamina, not flexed distally; raehis abaxially with sclerenchy- ma exposed, with scattered setae 1 .5-3 mm; lamina linear, narrowed at base with 2-4 reduced pinna pairs, apex not seen, deeply pinnatifid, up to 15 cm long; pinnae to 0.8 cm x 4 mm, entire or faintly subrepand, set 70-80° to raehis, at base dilated on both sides or straight acroscopically, apex broadly rounded, with castaneous setae along margins and costae abaxially, a few scattered setae and numer¬ ous, branched, glandular hairs on lamina, costal sderenchyma exposed; texture thin-herbaceous; sori inframedial to medial, up to 4 pairs per pinna, su¬ perficial, lacking setae but sporangia intermixed with numerous, stalked, gland-tipped paraphyses. This species is named for its thin, rather lax leaves (from Greek asthenes, weak). It is most closely related to C. knightii (Copeland) L. E. Bishop, which has densely setose petioles, lamina only 4-6 times longer than hroad, and thick-herbaceous to char- taceous lamina lacking numerous, branched, glan¬ dular hairs abaxially. Ceradenia knightii is known from Costa Rica, Colombia, and Hispaniola. Paratype. COSTA RICA. Cartago: Cerro de la Muerte, 1 km NW of Villa Mills on Interamerican Hwy., behind Hotel La Georgina, 2,900 m, Mickel 3206 (NY). Ceradenia aulaeifolia L. E. Bishop ex A. R. Smith, sp. nov. TYPE: Costa Rica. Limon: Cor¬ dillera de Talamanca, SW base of Cerro Ka- muk, in shrub-tree paramo, 3,200-3,350 m, Davidse, Herrera & Warner 25939 (holotype, UC; isotype, MO). Figure 1A H. Inter species subgeneris Filicipectinis frondibus pen- dentibus indeterminatibus et laminis perpinnatis ad C. mayoris et species affines accedens sed costarum abaxi- alium sclerenchymate manifeste exposito, frondibus ma- joribus, pinnis margine valde sinuatis, laminis chartaceis ab eis recedens. Plants epiphytic or lithophytic; rhizome stout, with castaneous scales 5-10 X 0.3-0. 7 mm, with hyaline to concolorous setulae or white glands at the margin; petiole atropurpureous, 1 0-40 cm x 0.6- 1 .2 mm, at the base with a few to many castaneous setae to 3 mm usually intermixed with much more numerous short setulae 0. 2-0.3 mm, sometimes glabrate distally, about equaling the lamina length, not flexed distally; raehis atropurpureous to blackish, abaxially with scattered hairs and occasionally a few setae, adaxially densely setose; lamina pendulous, lanceolate, usually narrowed at base with 1-4 pairs of reduced pinnae, lacking a distinct apical segment but with prolonged (indeterminate?) apical growth, perpinnate throughout, up to 50 cm (or more?) long; Novon 3: 182-185. 1993. Volume 3, Number 2 1993 Smith Ceradenia 183 Figure 1. New species of Ceradenia. A-H. C. aulaeifolia L. E. Bishop ex A. R. Smith, Davidse 25939 (UC). — A. Rhizome and petiole bases. — B. Setulae at base of petiole. — C. Proximal portion of leaf. — D. Distal portion of leaf. — E. Rhizome scale. — F. Pinnae. — G. Sorus with glands. — H. Trichomes on abaxial rachis. I-K. C. asthenophylla L. E. Bishop ex A. R. Smith, Bishop 1984 (UC). — I. Habit. — J. Rhizome scale. — K. Pinnae. — L. Trichomes on abaxial surface of lamina. pinnae to 4 cm x 6 mm, strongly sinuate, set 80- 90° to rachis, at base usually constricted on both sides, especially acroscopically, apex acuminate to obtuse, with castaneous setae 0.5-1. 5 mm and shorter branched hairs along margins, costae with prominently exposed sclerenchyma and simple or branched hairs abaxially, lacking setae, adaxially the costae slightly exposed, with branched hairs and 184 Novon scattered setae; texture chartaceous; sori medial, up to 12 pairs per pinna, superficial, lacking setae but sporangia intermixed with numerous, stalked, gland- tipped paraphyses. This striking species is named for the long, pen¬ dulous leaves that hang from the trees in high- elevation cloud forests (Latin aulaeum, curtain or tapestry). It is closely related to the Colombian C. mnyoris (Rosenstock) L. E. Bishop, under which Lellinger (1989) treated it, and C. semiadnata (Hooker) L. E. Bishop, from Colombia and Ecuador. Ceradenia aulaeifolia agrees with these and with C. intricata (C. V. Morton) A. R. Smith, C. congesta (Copeland) A. R. Smith, and C. arthrothrix L. E. Bishop & A. R. Smith in having pendulous fronds, indeterminate leaf apex, and perpimiate laminae with pinnae not connected along the rachis. From all of these it is distinguished by the exposed costal scle- renchyma on both sides of the pinnae, especially abaxially. From C. mnyoris it also differs in its larger size, more strongly sinuate pinna margins, and thin¬ ner laminae. Ceradenia semiadnata, which occa¬ sionally has the costal sclerenchyma exposed, differs from C. aulaeifolia in the presence of setae on the lamina surface and the relatively broader pinnae. An obvious feature of most specimens of this species is the crowded, short setulae 0.2-0. 3 mm on the petioles. This characteristic is shared with C. rnayoris. In this latter species these setulae are at times lost with age. In C. aulaeifolia a few specimens lack them entirely, so that their absence does not appear to be age related. Paratypes. COSTA RICA. Cartago: S slope Volcan de Turrialba, near Finca del V. Turrialba, 2,000-2,400 m, Standley 35051 (US). Limon: Cerro Chirripo, 10,400-11,000 ft., Evans & Lellinger 168 (US); Cor¬ dillera de Talamanca, Cerro Karnuk peak, Davidse et al. 26048 (MO, UC); Cordillera de Talamanca, SW foot of Cerro Karnuk, 3,200 m, Davidse et al. 25966 (MO). San Jose: Cerro Chirripo, SW slopes between Canaan and summit, 8,500-10,000 ft., Evans & Lellinger 81 (US); Cerro de las Vueltas, 2,700-3,000 m, Standley & Valerio 43743, 43824 (US). PANAMA. Bocas del Toro: Cordillera de Talamanca, 4 km NW of Cerro Fabrega peak, 3,000-3,150 m, Davidse et al. 25414 (MO); between Itamut and Bine peaks, Fabrega massif, 3,200 m, Gomez et al. 22534 (MO, UC). Ceradenia tristis A. R. Smith, sp. nov. TYPE: Costa Rica. San Jose: Pan American Hwy., Km 72-73, or 21-22 km SE of El Empalme, be¬ tween Cartago and San Isidro del General, 2,800 m, Smith & Beliz 2077 (holotype, UC; isotype, CR). Differt a C. spixiana paleis rhizomatis castaneis (non aureobrunneis), pinnis basi acroscopica constrictis (non dilatatis utrinque), et apicibus pinnarum attenuatis (non rotundatis vel acutis). Differt a C. kookenamae et C. kalbreyeri pinnis setosis margine. Plants epiphytic; rhizome stout, with castaneous scales to 10 x 0.5 mm, these with hyaline or pale setulae; petiole atropurpureous, up to 34 cm x 1.4 mm, with scattered castaneous setae to 3 mm prox- imally and a few distally, intermixed with simple or branched hairs throughout, ca. 1.5 times lamina length; rachis abaxially with sclerenchyma dark brown, with scattered simple or branched hairs 0.3- 0.8 mm, lacking setae or only a few setae toward base; lamina lanceolate, usually broadest just above base (lowermost 1-2 pairs of pinnae slightly re¬ duced), apex not seen, deeply pinnatifid to pinnat- isect, at the very base perpinnate, to 22 cm long; pinnae to 5 cm x 5 mm, entire or faintly repand, set 80-90° to rachis, at base usually constricted acroscopically and decurrent basiscopically, apex acuminate, with castaneous setae along margins, scattered simple or branched hairs along abaxial costae; texture thick-chartaceous; sori supramedial, up to 16 pairs per pinna, superficial, lacking setae, sporangia intermixed with numerous, stalked, gland- tipped paraphyses. The species epithet recalls the sadness felt over Dr. Bishop’s tragic illness and death, just as he was entering a most productive period in his research on Ceradenia and other genera of Grammitidaceae. Valerio 68 (US), from Volcan Barba, is close to this species hut differs in several characters. The petiolar setae are lacking toward the base, the pinnae are nearly linear-ohlong and closely spaced, and the costae lack setae adaxially. Of Central American ceradenias, Ceradenia tris¬ tis is most similar to C. spixiana (C. Martius ex Mettenius) L. E. Bishop, which differs by the golden brown rhizome scales and more parallel-sided pinnae that are dilate on both sides at their base and more rounded at the apex. Other close relatives in Central America are C. kookenamae (Jenman) L. E. Bishop and C. kalbreyeri (Baker) L. E. Bishop, which lack setae on the pinna margins. Ceradenia meridensis (Klotzsch) L. E. Bishop, from northern South Amer¬ ica, differs in having numerous setae along the ab¬ axial costae and in the sori. Colombian specimens annotated by Bishop as C. killipii (ined.) are also very similar to C. tristis and may prove to be con- specific ( Killip et al. 38061, US; Killip & Smith 15010, NY, US). Paratype. COSTA RICA. Heredia: above Laguna del Barva and summit of Volcan Barva, 1 0°08'N, 84°06.5'W, 2,840-2,900 m, Grayum <£• Quesada 7421 (MO). Volume 3, Number 2 1993 Smith Ceradenia 185 Ceradenia bishopii (Stolze) A. R. Smith, comb, nov. Basionym: Grammitis bishopii Stolze, Fieldiana, Bot., n.s. 32: 86. 1993. TYPE: Peru. Pasco: Prov. Oxapampa, border Prov. Oxa- pampa and Pasco, van der W erjf et al. 8569 (holotype, UC). This was recently described from several collec¬ tions from Peru (Tryon & Stolze, 1993) and is now known from Bolivia: Cocopunco, 10,000 ft., Tate 349 (NY). Ceradenia congesta (Copeland) L. E. Bishop ex A. R. Smith, comb. nov. Basionym: Ctenopteris congesta Copeland, Philipp. .[. Sci. 84: 397. 1956. Grammitis congesta (Copeland) Lellin- ger, Amer. Fern J. 74: 58. 1984. TYPE: Peru. Loma Grande, La Convencion, Rues 2172 (ho¬ lotype, US). Ceradenia congesta is known only Irom the Pe¬ ruvian type and is most closely related to C. intri- cata from Colombia and Ecuador. Ceradenia intricata (C. V. Morton) L. E. Bishop ex A. R. Smith, comb. nov. Basionym: Gram¬ mitis intricata C. V. Morton, Contr. U.S. Natl. Herb. 38: 101. 1967. TYPE: Ecuador. Pi- chincha: Guamani Pass, E of Pifo, Ewan 16436 (holotype, US). This was known to Morton (1967) only from the Ecuadorian type, but has since been collected in Colombia. Narino: Cumbal, vecindad de la Laguna La Bolsa, 3,900 m, Leist cS: Mohle 2185 (COL). Literature Cited Bishop, L. E. 1988. Ceradenia, a new genus of Gram- mitidaceae. Amer. Fern J. 78: 1-5. - . 1989. New species of Ceradenia subg. Cer¬ adenia. Amer. Fern J. 79: 14-25. Lellinger, D. B. 1989. The ferns and fern-allies of Costa Rica, Panama, and the Choco. Pteridologia 2A: 1- 364. Morton, C. V. 1967. The genus Grammitis in Ecuador. Contr. U.S. Natl. Herb. 38: 85-123. Tryon, R. M. & R. G. Stolze. 1993. Pteridophyta of Peru. Part V. 18. Aspleniaceae — 21. Polypodiaceae. Fieldiana, Bot., n.s. (in press). Novae Gesneriaceae Neotropicarum V. Four New Species and Two New Combinations in Columned from South America James F. Smith Biology Department, Boise State University, Boise, Idaho 83725, U.S.A. Laurence E. Skog Department of Botany, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Abstract. A monographic study of sections Pen¬ tadenia and Stygnanthe of Columnea (Gesneri¬ aceae) has revealed four species new to science and two new combinations. These species are described here and include C. atahualpae and C. hypocyr- tantha of section Pentadenia, and C. manabiana, C. suffruticosa, C. ultraviolacea, and C. xiphoidea of section Stygnanthe. Columnea L. is a genus of neotropical plants distributed from southern Mexico to Bolivia com¬ prising approximately 200 species. Species of Col¬ umnea are readily distinguished by their long tubular corollas that are either bilabiate or ventricose, and the fleshy, indehiscent, berry fruits. The majority of species are epiphytic, although several can be terrestrial as well. A revision of the species of Col¬ umnea in Ecuador conducted by Kvist & Skog (1993) and a treatment of the Gesneriaceae of Pan¬ ama by Skog (1979) are the only recent taxonomic treatments of the genus. Wiehler (1973, 1983) di¬ vided the genus into four segregate genera and de¬ scribed a fifth genus allied to the Columnea complex. These five genera were combined back into the single genus Columnea by Kvist & Skog (1993). The four new species described here are the result of a mon¬ ographic study of two of the smaller sections of Columnea, sections Pentadenia and Stygnanthe (Smith, 1991). The two new combinations result from a recent publication of new species of Ges¬ neriaceae (Wiehler, 1992). Columnea (section Pentadenia) atahualpae J. F. Smith & L. Skog, sp. nov. TYPE: Ecuador. El Oro: forest along trail from Sambotambo, fol¬ lowing headwaters of Rio Moro Moro S to Bue¬ naventura, at and along highway to Portovelo, 29 Aug. 1943, Steyermark 54228 (holotype, F). Figure 1. Differt a Columnea nervosa (Klotzsch ex Oersted) Han- stein foliis ellipticis vel obovatis bracteis magnis corollis brevis (3. 0-3. 3 cm longis) marginibus integris trichom- atibus appressis aureis. Suffrutescent terrestrial (or possibly epiphytic) herbs with erect stems to 1 m tall (Fig. 1A). Stems 8-12 mm diam., tawny brown, apex appressed pu¬ bescent with golden yellow uniseriate trichomes, low¬ er stem smooth and glabrous. Internodes 2. 5-9.0 cm long, leaf scars slightly raised from the slightly swollen nodes. Leaves opposite, equal to subequal, blades obovate to elliptic, 9.2-22.5 x 3. 7-7. 8 cm, apex acute, base cuneate-oblique, adaxially green, strigose-appressed pilose with single-celled, trans¬ parent, and uniseriate golden yellow trichomes, abaxially silvery green-maroon, vestiture similar to adaxial surface but sparser, pubescence of veins abaxially sericeous, margin entire to slightly undu¬ late (Fig. 1A, B). Petioles 1.0-3. 2 cm, appressed pilose with uniseriate golden yellow trichomes. In¬ florescences 1-7 flowers in axils of either leaf pair (Fig. 1A), floral bracts 1-2 of unequal size, lanceo¬ late-ovate, 8-22 x 1.5-16 mm, apex acuminate, slightly constricted at base, reddish, appressed pilose with uniseriate golden yellow trichomes, margin en¬ tire. Pedicels 1.3- 1.9 cm, erect in leaf axil, ap¬ pressed pilose with uniseriate golden yellow tri¬ chomes, 1 -several glands present on pedicel, purplish, ca. 0.4 mm long. Calyx clasping corolla at base but more open toward apex of lobes, lobes lanceolate-subulate, equal in size, 1.5-2. 7 x 0.3- 0.5 cm, apex long acuminate, green with red tips to purple, margins serrulate-subulate, 3-5 teeth per side, mostly basal, exterior sericeous with uniseriate golden yellow trichomes, interior surface similar but trichomes sparser (Fig. 1C). Corolla pale yellow, tubular, gibbous at base, slightly ventricose, 3.0- 3.3 cm long, 3.0 mm wide at base, 5-8 mm at widest point, 4-5.5 mm before limb, exterior ap- Novon 3: 186-197. 1993. Volume 3, Number 2 1993 Smith & Skog Columnea 187 1 mm Figure 1. Columnea atahualpae J. F. Smith & L. Skog. — A. Habit. — B. Adaxial leaf pubescence. C. Pedicel and calyx. — D. Flower. — F. Corolla interior with stamens. — F. Gynoecium with nectaries. — G. Ovary with calyx and nectaries. — H. Berry with nectaries. — I. Seeds. (A, B, D-F from Steyermark 53840; C, G I from Steyermark 54228. ) 188 Novon Figure 2. Columnea manabiana (Wiehler) J. F. Smith & L. Skog. — A. Habit. — B. Abaxial leaf pubescence. C. Inflorescence bract. — D. Flower. — E. Corolla exterior. — F. Corolla interior with retracted stamens. — G. Volume 3, Number 2 1993 Smith & Skog Columnea 189 pressed pilose-sericeous with uniseriate golden yel¬ low trichomes (Fig. 1C), interior pilose-hirsute prox- imally with transparent uniseriate trichomes, glandular capitate trichomes dorsally and distally (Fig. IE). Lobes equal. 2.0 x 1. 5-2.0 mm. se- miorbicular-acute. Stamens excluded, filaments white, sparsely pubescent, connate for 5 mm anteriorly at base and adnate to corolla base anteriorly for 3 mm, curling after anthesis, staminodes not seen, included in corolla, anthers quadrate, 1.7 x 1.7 mm (Fig. IE). Ovary sericeous, 3 mm long, conical, style white, pilose at base, glandular at apex, stigma bi- lobed, papillose (Fig. 1 F). Nectary of 5 glands, dorsal 2 enlarged (Fig. 1C). Berry globose, 1.0 cm diam., color uncertain but probably pale pink, pubescent (Fig. 1H). Seeds fusiform, red-brown, twisted, stri¬ ate, 1.2- 1.3 mm long (Fig. II). Distribution. Known only at 1,035 1,890 m elevation in Ecuador from a small geographic area in El Oro and one other collection in Zamora-Chin- cliipe. The Zamora-Chinchipe collection is a poor specimen, and its placement within this species is questionable. It may represent another species or may be a specimen of C. nervosa (Klotzsch ex Oersted) Hanstein, a morphologically similar species. The placement of this collection in C. atahualpae is based on its golden yellow pubescence found throughout the vegetative parts of the plant. Columnea atahualpae is similar to C. nervosa. However, the much larger obovate-elliptic leaves, smaller corolla, large floral bracts, and entire mar¬ gins are sufficient to distinguish it from this species. In addition, the golden yellow appressed trichomes, found throughout the plant, make it even more distinct from other species of Columnea. I his spe¬ cies is probably closely related to C. lophophora Mansfeld. which also has clustered flowers, large floral bracts, and dense vestiture. This species can easily be distinguished from C. lophophora in that the pubescence is tightly appressed and not as dense, or long sericeous as seen in C. lophophora. The specific epithet commemorates one of the last two Sapa Incas to rule the Inca empire in the Andes, Atahualpa, executed by Pizarro in 1533. Paratypes. ECUADOR. El Oro: vicinity of Ayapam- ba, 13, 16 Oct. 1918, Rose & Rose 23461 (NY, US); 2 leagues NE of Curtincapa, bordering Quebradas Nudillo and Tambillo, tributary to Rio San Luis and Piedra Gran¬ de, 13 Aug. 1943, Steyermark 53840 (F, US). Za¬ mora— Chinchipe: Palandra (probably Palanda), July 1876, Andre 4638 (NY). Columnea (section Stygnanthe) manabiana (Wiehler) J. F. Smith & L. Skog, comb. nov. Basionym: Pentadenia manabiana Wiehler, Phytologia 73: 236. 1992. TYPE: Type ma¬ terial from cultivated plants, 8 July 1987, Wiehler 87102 (holotype, GES not seen; iso¬ types, B. F. HBG, K, MO, N't. QCA, SEL. G, IJS, none seen). Type material in cultivation is from live material collected in Ecuador. Man- abi: km 67 on road from Chone-Santo Do¬ mingo, 500 m, on 31 July 1977, bv C. II. & II. C. Dodson 6701 (AAU, MO. SEL). Fig¬ ure 2. Epiphytic herbs (Fig. 2). Stems 6-8 mm diam., squarish when dried, apex pubescent, lower stem smooth and glabrous. Internodes 1.1 2.6 cm long, slightly swollen. Leaves opposite, strongly unequal, blades lanceolate to slightly falcate, larger leaf of a pair 8-16.2 x 1 .6-4.5 cm, apex acute, base round¬ ed and strongly oblique, adaxially green but suffused with pink, abaxially pink-purple, slightly pilose with uniseriate red trichomes on both surfaces, margin entire to slightly undulate, lateral veins 5-6 per side. Petioles 2-5 mm, slightly pilose with uniseriate red trichomes (fig. 2A, B). Smaller leaf 1. 6-3.1 x 0.4 1 . 1 cm, otherwise as larger leal. Inflorescences of 2 4 flowers per axil of larger leaf (Fig. 2A). Floral bracts 2. conspicuous, ovate, 1.3 1.9 x 0.50.9 cm, apex acute, green suffused with red, weakly pilose with uniseriate transparent trichomes, margin entire (Fig. 2C). Pedicels 2.5-4 mm, villous. Calyx loosely clasping corolla, lobes equal to subequal, lanceolate-elliptic, 1.2 1.6 x 2 4.5 mm, apex acute, green or green with red tips, slightly pilose with uniseriate transparent trichomes on both surfaces (Fig. 2D). Corolla yellow, tubular, constricted at base, 1.8-2. 4 cm long, 2-2.3 mm wide at base, 3 4.5 mm at widest point, 2. 5-3. 5 mm before limb, exterior glabrate to villous at apex (Fig. 2E), interior slightly pubescent at base, glandular distally. lobes semiorbicular, inconspicuous, equal-subequal, ca. I x 1.5 mm (Fig. 2F). Filaments connate anteriorly for 5 mm, adnate to corolla base anteriorly for 4.5 mm, white, slightly pubescent at base to hallway up filaments, anthers quadrate, 1.3 x 1.3 mm (Fig. 2F). Ovary 2 mm long, pilose, style white, slightly pubescent, glandular at apex, stigma bilobed (Fig. 2G). Nectary of 5 free glands, the dorsal glands enlarged (Fig. 21 1 1. Berry ovoid, 9x5 mm, dark in color when dried, probably white, slightly pilose. Gynoecium with calyx and nectaries. — H. Ovary with nectaries and calyx. (A, B from Dodson et al. 9170; G-H from Dodson et al. 6791.) 190 Novon Distribution . Known only in Ecuador from Man- abi and El Oro, 70-500 m. This species was long believed to be a morpho¬ logical variant of C. spathulata Mansfeld, but anal¬ ysis of cpL)NA (Smith, 1991) showed it to be mark¬ edly different from that species and more closely allied with C. inconspicua Kvist & L. Skog. Once the populations that differed from C. spathulata were identified based on cpDNA restriction site vari¬ ation, morphological characters were also detected that distinguish this species, such as large conspic¬ uous floral bracts and falcate leaves. This species can be distinguished from other spe¬ cies of Columnea on the basis of its long, narrow, lanceolate to slightly falcate leaves and small yellow corollas shared with C. inconspicua Kvist & L. Skog, to which it is most closely related (Smith, 1991). It is distinguished from C. inconspicua by the presence of large, conspicuous, ovate floral bracts that partly obscure the inflorescence. Additional specimen examined. ECUADOR. El Oro: highway from Guayaquil to Machala, 1 km S of Ponce Enriques and 2 km in from road on low hill, 70 m, 9 Oct. 1979, Dodson et al. 9170 (MO, SEL, US). Columnea (section Pentadenia) hypocyrtantha (Wiehler) J. F. Smith & L. Skog, comb. nov. Basionym: Pentadenia hypocyrtantha Wieh¬ ler, Phytologia 73: 234. 1992. TYPE: Bolivia. Santa Cruz: near Fortaleza between Siberia and Comarapa on the Cochabamba Santa Cruz road, 2,500 m, 15 Jan. 1965, l ogel 498 (holotype, US; isotypes. A, F, Z). Figure 3. Suffrutescent, epiphytic, sublignose herbs with upright ascending stems (Fig. 3A). Stems to more than 50 cm tall, 4-5.5 mm diam., apex sparsely pilose, lower stem smooth and glabrous. Internodes 0.7 32 mm long, nodes slightly swollen, leaf scars raised. Leaves opposite, equal-subequal, blades ovate, 6.3-13.3 x 2.9-8. 1 cm, apex acuminate, base rounded, slightly oblique, adaxially dark green, slightly pilose with uniseriate transparent trichomes, abaxially maroon or green suffused with purple, glabrate to sparsely pilose with uniseriate transpar¬ ent trichomes on the veins, 4-7 lateral veins per side, margin minutely denticulate. Petioles 0.9 2.1 cm long, slightly pilose with uniseriate transparent trichomes (Fig. 3A, B). Inflorescences of axillary, solitary flowers, ehracteate or possibly hracts ca¬ ducous (Fig. 3 A). Pedicels 4-6.4 cm long, red- purple, erect in axil, slightly pilose with uniseriate transparent trichomes, and with oblong, 0. 5-0.7- mm dark glands near apex. Calyx clasping corolla, lobes equal, ovate, 12-16 x 4.5-9 mm, apex long acuminate, green with red tips to maroon, both surfaces glabrate with margin entire-ciliate (Fig. 3C). Corolla orange-red, limb and lobes green, tubular, strongly ventricose-saccate with pouch occasionally projecting forward beyond the opening of the corolla, strongly constricted at opening, slightly gibbous at base, 2. 4-3. 5 cm long, 4-8 mm wide at base, 1.1- 1.8 cm at widest point, 3. 5-4. 5 mm before limb, exterior glabrate-pubescent, trichomes denser to¬ ward limb (Fig. 3D), interior glabrous, lobes se- miorbicular, equal in size, approximately 1x2 mm (Fig. 3E). Filaments glabrous, white, anthers quad¬ rate, 1.5 x 1.5 mm (Fig. 3E). Ovary glabrous, 6 mm long, style glabrous, white, stigma stomato- morphic (Fig. 3E). Nectary of 3-5 free glands, 2 dorsal glands enlarged, slightly connate (Fig. 3E). Berry globose, 9 mm diam., glabrate, green with purple stripes (Fig. 3F). Distribution. Known only from the type from near Cochabamba, Bolivia, and a few collections nearby at 2,000-2,950 m, growing in cloud forests, and so far only collected in January. Columnea hypocyrtantha resembles C. trollii Mansfeld to which it is probably related on the basis of the strongly ventricose corolla. However, it is easily distinguished from C. trollii by its glabrate, ovate leaves, glabrous ovary, and corolla pouch that projects forward beyond the opening of the corolla, a character not seen in any other species of Col¬ umnea. The species is only known from a few col¬ lections within a narrow geographic and temporal range. Therefore, it is difficult to evaluate phenology or distribution accurately. The name is probably derived from the pouchlike nature of the corolla, which resembles that of the old genus Hypocyrta C. F. P. Martius (now a syn¬ onym of Nematanthus Schrader). Additionl specimens examined. BOLIVIA. Cocha¬ bamba: Prov. Chapare, road from Cochabamba to Villa Tunari, 13-16 Jan. 1981, Luer et al. 564 (SEL), 22 Jan. 1980, Luer et al. 4877 (SEL); Prov. Carrasco, km 262 on road from Santa Cruz-Cochabamba, 2 km E of Siberia, 18 Jan. 1983, Besse et al. 1725 (US); Prov. Carrasco, Serrania Siberia, 20-35 km W of Comarapa on old Cochabamba -Santa Cruz road (Hwy. 4), 14-15 Jan. 1990, Dorr & Barnett 7045 (US). Columnea (section Stygnanthe ) suffruticosa J. F. Smith & L. Skog, sp. nov. TYPE: Colombia. Choco: Mpio. San Jose del Palmar, Cerro del Torra, 7 Jan. 1984, Silverstone-Sopkin et al. 1594 (holotype, CUVC; isotypes, MO, US). Figure 4. Columneae ambiguae (Urban) B. Morley affinis sed foliis brevioribus ovatioribus caulibus magis ligneis differt. Volume 3, Number 2 1993 Smith & Skog Columnea 191 Figure 3. Columnea hypocyrtantha (Wiehler) J. F. Smith & L. Skog. — A. Habit. — R. Adaxial leaf pubescence. — C. Pedicel and calyx. — D. Flower. — E. Corolla interior with stamens, gynoecium, and nectaries. — F. Berry with nectaries and calyx lobes pulled away. — G. Berry as seen naturally. (A, B from Besse et al. 1725; C-G from Dorr & Barnett 7045.) 192 Novon Figure 4. Columnea suffruticosa J. F. Smith & L. Skog. — A. Habit. — B. Adaxial leaf pubescence. — C. Flower. — D. Corolla interior with stamens and gynoecium. — E. Pouchlike protuberences on interior of corolla. — F. Gynoecium with nectaries and calyx. — G. Berry with nectaries. (A from Silverstone-Sopkin et al. 1594 ; B from Silverstone- Sopkin et al. 4289; C-G from Silverstone-Sopkin et al. 4530.) Volume 3, Number 2 1993 Smith & Skog Columnea 193 Small woody-sublignose shrubs, epiphytic or ter¬ restrial (Fig. 4A). Stems to 2 m, 2-3 mm diam., pubescent or hirsute with uniseriate trichomes at apex, lower stem dark purple, smooth, and glabrous. Internodes 1-3.5 cm long. Leaves opposite, un¬ equal. Larger leaf blades in a pair elliptic-ovate, 1.4- 5 X 0.9-2. 5 cm, apex acute, base rounded oblique, adaxially dull green, glabrate, abaxially pur¬ ple mottled to all purple, glabrate, margin crenulate, lateral veins 3 6 per side. Petioles 2 10 mm long, pilose-hirsute (Fig. 4A, B). Smaller leaf blades ellip¬ tic-ovate, 0.7-3. 9 x 0.4- 1 .8 cm, otherwise as larg¬ er leaf. Inflorescences of 1-2 flowers per axil (Fig. 4 A). Floral bracts caducous, linear, up to 3 mm long. Pedicels 5-11 mm, erect, hirsute with purple or transparent uniseriate trichomes. Calyx loosely clasping corolla, lobes equal-subequal, spatulate, 6 11 x 1.5-5 mm, acute, green or purple, margins serrate, exterior surface hirsute to slightly so, in¬ terior surface glabrous (Fig. 4C). Corolla yellow with orange spots on lobes, tubular, constricted at base, 1. 5- 3.1 cm long, 2 mm at base, 4-6 mm at widest point, 3-5 mm before limb, lobes subequal, se- miorbicular, 12 x 1.5-2 mm, exterior surface slightly sericeous with purple or transparent unise¬ riate trichomes (Fig. 4C), interior glabrous, glan¬ dular distally and dorsally (Fig. 4D). Two pouchlike invaginations occasionally present on dorsal surface ol corolla (Fig. 4E). Filaments connate anteriorly for 2.5 mm, adnate to corolla base anteriorly lor 1.5 mm, white, glabrous, anthers quadrate, 1.5 x 1.5 mm (Fig. 4D). Ovary 2 mm long, glabrous to sericeous at apex, style white, glabrous with glan¬ dular trichomes at apex, stigma stomatomorphic, white (Fig. 4F). Nectary of 5 free glands or with 2 dorsal glands connate (Fig. 4F, G). Mature berry unknown, immature fruit ovoid, 3x2 mm, white, glabrous to sparsely pilose (Fig. 4C). Distribution . Known only from the type locality in Choco, Colombia, and one additional locality near¬ by on the Choco-Valle del Cauca border at 1,870- 2,500 m. Although it is known from only a few collections, four made within a few days of each other, and the others from the same general locality, this species is distinct from other species of section Stygnanthe. The shrubby woody habit of this species readily distinguishes it from other species in this section. In addition, the small, ovate, crenate leaves are dis¬ tinctive. Phylogenetically, Columnea sujfruticosa is most likelv related to C. colombiana (Wiehler) Kvist & L. Skog, as based on a cladistic analysis oi mor¬ phology (Smith, 1991). However, the corolla lobes with darker colored spots (a character not used in the analysis and previously overlooked in species descriptions) may place it in the clade with (.. ova- tifolia Kvist & L. Skog, C. lavandulacea Kvist & L. Skog, and C. crassicaulis (Wiehler) Kvist & L. Skog (Smith, 1991). Four of the collections are epiphytic, the others are terrestrial. The subtle mor¬ phological differences between the two groups are undoubtedly due to environmental effects ol the two habitats. The epiphytic collections have smaller leaves, purple pubescence, and more purple color¬ ation in general. This is likely due to the higher amount of sunlight obtained by the epiphytic indi¬ viduals than any other factor. Coloration of tri¬ chomes, it variable within an individual or species, tends to be present in trichomes more likely to receive sunlight (pers. obs.). The name is derived from the stems, which are woody at the base. Paratypes. COLOMBIA. Choco: Mpio. San Jose del Palmar, Cerro del Torra, up from heliport, 19 Aug. 1988, Ramos et al. 1357 (CUVC), 5 Aug. 1982, Silverstone- Sopkin 1225 (CUVC), 4 Jan. 1984, Silver stone-Sopkin 1496 (CUVC, MO), 6 Jan. 1984, Silverstone-Sopkin et al. 1563 (MO), 5 Jan. 1984, Silverstone-Sopkin 1548 (CUVC, MO), 13 Jan. 1984, Silverstone-Sopkin et al. 1812 (CUVC), 10 Aug. 1988, Silverstone-Sopkin et al. 4289 (CUVC, US), 17 Aug. 1988, Silverstone-Sopkin et al. 4530 (CUVC, US), 15 Aug. 1988, Silverstone- Sopkin et al. 4442 (CUVC). Valle del Cauca: Mpio. El Cairo, ca. 21-25 km beyond El Cairo toward Choco border, 13 May 1988, Luteyn et al. 12288 (CUVC), 25 Apr. 1989, Luteyn Giraldo 12644 (CUVC). Columnea (section Stygnanthe) ultraviolacea J. F. Smith & L. Skog, sp. nov. TYPE: Bolivia. La Paz: Prov. Sud Yungas, Huancane, 9 km from San Isidro (N of Chulumani), 2,450 m, 1 Jan. 1984, Heck 87 11 (holotype, LPB; iso¬ type, US). Figure 5. Pentadeniae fritschii (Rusby) Wiehler affinis sed se- palis angustiore violaceis pedicellis violaceis corollis luteis differt. Semiscandent herbaceous, terrestrial shrubs (Fig. 5). Stems to 1 .2 m, 5-6 mm diam., red-brown, apex pubescent, lower stem smooth and glabrous (Fig. 5A). Internodes 1.3 4.5 cm long. Leaves in whorls ol 4, equal to subequal, blades elliptic, 2. 5-4. 5 x 1-2 cm, apex acute, base rounded-cuneate, oblique, adaxially dark green, glabrous, abaxially dark pur¬ ple, strigillose, appressed hirsute on veins with red uniseriate trichomes, margin entire, ciliate with red uniseriate trichomes, lateral veins 4 5 per side. Pet¬ ioles 2-6 mm long, purple, pilose-hirsute with red uniseriate trichomes (Fig. 5A, B). Inflorescences of solitary flowers per leaf axil (Fig. 5A). Floral bracts linear, 3-8 mm x up to 0.5 mm, acute, purple, 194 Novon Figure 5. Columnea ultraviolacea J. F. Smith & L. Skog. — A. Habit. — B. Abaxial leaf pubescence. — C. Flower. — D. Corolla interior with retracted stamens. — E. Gynoecium with nectaries and calyx. — F. Ovary and nectaries. (All from Beck 8741.) Volume 3, Number 2 1993 Smith & Skog Columnea 195 slightly pilose on outer surface, glabrous on interior, margin entire. Pedicels 8-16 mm, purple, erect, pilose-hirsute with red uniseriate trichomes, with darker purple, 0.4 0.7-mm-long oval-elliptic glands near calyx. Calyx loosely clasping corolla, lobes equal- subequal, lanceolate-linear, 13-21 x 1.2-2 mm, acute, purple, margin entire, exterior surface sparse¬ ly hirsute with red uniseriate trichomes (lig. 5C), interior surface glabrous. Corolla light yellow, lobes purple inside, tubular, slightly ventricose, long con¬ striction at base, constricted at limb. 4. 1-4.4 cm long, 2-2.5 mm wide at base, 5.5-7 mm at widest point, 3.5-4 mm before limb, lobes equal-subequal, semiorbicular, 2-3 X 2-3 mm, exterior surface densely sericeous with uniseriate transparent tri¬ chomes (Fig. 5C), interior slightly pilose at base, glandular distallv and dorsally (Fig. 5D). Filaments connate anteriorly for 8 mm, adnate to corolla base anteriorly for 4.5 mm, white, slightly pilose, anthers quadrate, 1.5-2. 5 x 1.5-2 mm (Fig. 5 D ) . Ovarv 4 mm long, sericeous, red and transparent uniseriate trichomes, style red at base turning yellow, slightly pilose at base, glabrous at apex, stigma stomato- morphic, white (Fig. 5E). Nectary of 5 free glands (Fig. 5F). Berry ovoid, 8-9 x 3—4.5 mm, dark purple with white spot where style was attached, sericeous with red and transparent trichomes. Distribution. Known only from the type locality in Bolivia at 2,400-2,450 m. Flowering in January, fruiting in May. Columnea ultraviolacea and Pentadenia jrit- schii (Busby) Wiehler (soon to be transferred to Columnea) are the only two species of Columnea to have leaves in whorls of four, easily distinguishing these species from others and thus allying them in a phylogenetic sense. They are distinguished from each other by the yellow corollas, purple coloration of the vegetative parts, and narrow lanceolate-linear sepals of C. ultraviolacea. These two are obviously sister species, based on their leaf arrangement, and are closely related to Columnea moesta Poeppig, another Bolivian species (Smith, 1991), based on epDNA variation (Smith, 1991). The epithet is derived from the bright purple coloration of all vegetative parts and the fruits. Paratypes. BOLIVIA. La Paz: Sud Yungas, 9.2 km toward San Isidro from Huancane, 2 May 1989, Snath & Smith 1829 (LPB, WIS). Columnea (section Stvgnanthe) xiphoidea J. F. Smith & L. Skog, sp. nov. TYPE: Peru. Uca¬ yali: Divisoria, 59 km from Lingo Maria on road to Pucallpa, 17 Nov. I 949- 15 Jan. 1950, Allard 21230 (holotype, BH; isotype, US). Fig¬ ure 6. Columneae ovatifoliae Kvist & L. Skog affinis sed foliis longioribus lanceolatis differt. Flerbs with stems to 2 m, 4 mm diarn., red-brown, quadrangular when dried, apex slightly to densely sericeous, lower stem smooth (F ig. 6 A). Internodes 1.6-4. 3 cm long. Leaves equal-subequal, blades lan¬ ceolate, slightly falcate, 7-9.1 x 1.4 2 cm, apex acuminate, base rounded, oblique, adaxially green with purple margin, hirsute-pilose with uniseriate red trichomes, abaxiallv dull rose red, strigose with sin¬ gle-celled trichomes and pilose with red uniseriate trichomes, veins slightly pilose with red uniseriate trichomes, margin entire, lateral veins 6 per side. Petioles 2.5-4 mm, pilose-sericeous with red uni¬ seriate trichomes (Fig. 6A, B). Inflorescences of I 3 flowers per axil of either leaf (Fig. 6A). Floral bracts 1-2 per inflorescence, linear, 4 1 1 x 0.5 2 mm, acute, maroon, sericeous, entire (Fig. 6C). Pedicels 7-13 mm long, erect, sericeous with uni¬ seriate transparent trichomes. Calyx clasping co¬ rolla, lobes equal, lanceolate, 15 21 x 2-4 mm, apex acute, margin entire, exterior surface green with purple margin and some purple mottling, pilose and strigose, sericeous at base, interior surface bright red-purple, pilose-sericeous (Fig. 61)). Corolla crim¬ son. lobes with dark purple spot inside, tubular, slightly ventricose, constricted at base, 4.2 cm long, 4.5 mm wide at base, 1.1 cm at widest point, 0.8 cm before limb, lobes subequal, semiorbicular, 2 x 3 mm, exterior villous-sericeous, denser toward limb (Fig. 6D), interior hirsute-pilose with glandular tri¬ chomes dorsally and distallv (Fig. 6 F7). Filaments connate at base anteriorly for 5 mm, adnate to corolla at base anteriorly for 4 mm, yellow, hirsute, anthers quadrate, 1.7 x 1.7 mm (F ig. 6E). Ovary 3 mm long, sericeous, stvle yellow, glabrous, stigma bilobed, yellow, papillose (Fig. 6F ). Nectary of 4- 5 free glands, the 2 dorsal glands slightly connate (F ig. 6G). Fruit not seen. Distribution . Known only from the type locality in Peru and nearby, at 1,600 m. The lanceolate, isophyllous leaves and bright red, densely sericeous-lanate corolla of Columnea xip¬ hoidea readily distinguish it from any other species of Columnea. Although no formal cladistic analysis has been performed on this species, ( .olumnea xip¬ hoidea is potentially related to two different clades within section Stvgnanthe, based on its morpholog¬ ical features. The lanceolate, slightly falcate leaves would tend to ally it with the C. inconspicua Kvist 6 L. Skog-C. manabiana clade (Smith, 1991). However, the slightly ventricose, densely sericeous- lanate corolla with dark purple spots on the interior of the lobes would place it with the Pentadenia 196 Novon 1cm E F Figure 6. Columnea xiphoidea J. F. Smith & L. Skog. — A. Habit. — B. Abaxial leaf pubescence. — C. Pedicel with floral bracts and calyx. — D. Flower. — E. Corolla interior with stamens. — F. Gynoecium with nectaries. — G. Ovary and nectaries. (All from Allard 2123U.) Volume 3, Number 2 1993 Smith & Skog Columnea 197 fritschii (Rusby) Wiehler C. ultraviolacea clade (Smith, 1991). The name is based on the long, swordlike leaves of this species, which are unique in section Styg- nanthe. Paratypes. PERU. San Martin: La Divisoria, “Mar¬ garita,” 14 Aug. 1946, Ferreyra 1012 (US); La Divisoria, 59 km from Tingo Maria on road to Pucallpa, 15 Nov. 1949-15 Jan. 1950, Allard 21280 (US), Allard 21 300 (US). Huanuco: Cordillera Azul, ca. 39.2 km E of Tingo Maria on road to Pucallpa, 19 Nov. 1979, Jones & Davidson 9339 (LAM); ca. 43 km E of Tingo Maria on road to Pucallpa, 21 Nov. 1979, Jones & Davidson 9432 (LAM). Acknowledgments. We acknowledge Alice Tan- gerini for the excellent illustrations, the University of Wisconsin-Madison and Kenneth .1. Sytsma for support to J. Smith during his graduate career, and two anonymous reviewers for helpful comments. Funding for this research was made available from NSF grant BSR-8815173 to Kenneth J. Sytsma and J. Smith. Literature Cited Kvist, L. P. & L. E. Skog. 1993. The genus Columnea (Gesneriaceae) in Ecuador. Allertonia. 6(5): 327- 400. Skog, L. E. 1978 [1979]. Family 175, Gesneriaceae. In: R. E. Woodson, Jr. & R. W. Schery (editors), Flora of Panama Part IX. Ann. Missouri Bot. Card. 65: 783 996. Smith, J. F. 1991. The Evolution and Systematics of Columnea. Ph.D. Dissertation, University of Wis¬ consin, Madison, Wisconsin. Wiehler, H. 1973. One hundred transfers from Allo- plectus and Columnea (Gesneriaceae). Phytologia 27: 309-328. - . 1983. A synopsis of the neotropical Gesner¬ iaceae. Selbyana 6: 1-219. - . 1992. New species of Gesneriaceae from the Neotropics. Phytologia 73: 220 241. New Combinations in the Tribe Poraneae (Convolvulaceae) for the Flora of China George ft . Staples Department of Botany, Bishop Museum, P.0. Box 19000-A, Honolulu, Hawaii 96817-0916, U.S.A. ABSTRACT. New combinations are made for 10 taxa formerly included in the genus Porana s.l. in the account of the Convolvulaceae for the Flora Reipublicae Popularis Sinicae , which are now con¬ sidered to belong in the genera Dinetus, Poranopsis, and Tridynamia, as follows: Dinetus decorus, D. dinetoides, D. duclouxii, D. grandiflorus, I), trun- catus, Poranopsis discifera , P. sinensis, Tridyn¬ amia megalantha, T. sinensis var. sinensis, and T. sinensis var. delavayi. A key to the genera is pro¬ vided, and 1 1 additional taxa formerly accepted in Porana are reduced to synonymy. The genus Porana N. L. Burman comprises 57 nominate taxa, the majority distributed in mainland tropical Asia. Twenty-eight ( 1 4 species, 1 4 varieties) were recognized in the account of the Convolvula¬ ceae for the Flora Reipublicae Popularis Sinicae (Fang & Huang, 1979). My revisionary study of the genus Porana and other taxa of the tribe Por¬ aneae H. Hallier (Staples, 1987) indicates that there has been considerable overdescription in the genus Porana, and the number of species is actually much lower than the number of named species. Although a comprehensive taxonomic revision of Porana s.l. has been submitted to Systematic Botany Mono¬ graphs (Staples, submitted manuscript), several new combinations, and reduction of additional names to synonymy, are needed for the account of the Con¬ volvulaceae for the Flora of China, which is sched¬ uled for publication before the revision will appear. Phis paper makes 10 new combinations needed for use in the Flora, and reduces to synonymy 1 1 other taxa of Porana recognized by P ang & Huang (1979). To summarize the major conclusions of the re¬ vision, Porana s.l. proved to be a polyphyletic as¬ semblage comprising four distinct groups of species; a preliminary analysis indicated that these four spe¬ cies groups are not a monophyletic lineage inter se (Staples, 1987). I recognize them at generic rank, emending the generic concept of Porana to accom¬ modate one group comprising three species, and take up the previously established generic names Tri¬ dynamia Gagnepain (1950), Poranopsis Roberty (1952), and Dinetus Sweet (1825) for the other three groups of species. Furthermore, the genus Cordisepalum Verdcourt is maintained as a genus distinct lrom and closely related to Poranopsis. These five genera may be distinguished by the following key. Full descriptions of all taxa, distribution data, ecological and phenological summaries, and detailed nomenclatural discussion are presented in the re¬ vision (Staples, submitted manuscript) so they are omitted here. Key to Genera of Asiatic Poraneae la. Leaf venation pinnate, or with a single basal pair of secondary veins; calyx about half as long as the corolla, sepals all ± equal in length . . . Porana s.s. lb. Leaf venation palmate; calyx minute, usually < VUI as long as the corolla, if larger then sepals markedly unequal. 2a. Plants entirely herbaceous; flower buds with a minute apical tuft of hairs, otherwise glabrous externally . Dinetus 2b. Plants woody (often with herbaceous stem tips); flower buds hairy externally. 3a. Sepals unequal in flower; fruiting se¬ pals with 3 9 parallel longitudinal veins . Tridynamia 3b. Sepals (sub)equal in flower; fruiting sepals with 1 evident mid vein. 4a. Corolla funnelform, white; fruit sessile on torus . Poranopsis 4b. Corolla rotate, yellowish; fruit borne on a slender stalk . . Cordisepalum As construed here, Porana s.s. and Cordisepal¬ um do not occur within the geographic range of the Flora of China. Some species of Poranopsis, Di¬ netus, and Tridynamia occur within the political boundaries of the People’s Republic of China, and the following new combinations are proposed for them. Dixen s Sw eet Dinetus was segregated from Porana by Sweet ( 1825), based on the shape of the corolla, the single filiform style, and 1 -celled ovary. He took up Di- Novon 3: 198-201. 1993. Volume 3, Number 2 1993 Staples New Combinations in Poraneae 199 net us, an unpublished herbarium name used by Francis Buchanan-FIamilton, at the suggestion of David Don. who had seen the name on Hamilton's specimens in the Lambert herbarium (Sweet, 1825). Sweet included only a single species, I). racemosus, when he erected the new genus, but suggested that P. grandiflora Wallich and P. paniculata Rox¬ burgh would eventually prove to belong to Dinetus also. Porana grandiflora is herein transferred to Dinetus for the first time, while P. paniculata is considered a species of Poranopsis (q.v.). As construed here, Dinetus includes herbaceous climbers with flower buds glabrous except lor a tiny apical tuft of hairs, an entire or apically emarginate, ellipsoidal stigma, and a fruiting calyx reflexed from or tightly wrapped around the fruit, comprised of accrescent sepals with multiple, parallel longitudinal veins. The following new combinations and syn¬ onyms are relevant to the Flora of China. Dinetus decorus (W. Smith) Staples, comb. nov. Basionym: Parana decora W. Smith, Notes Roy. Bot. Card. Edinburgh 8: 197. 1914. TYPE: China. Yunnan: [Xinping Yi-Dai Aut. Xian] valley of Ma-li-ouan. Aug. 1913, Mai re 64 (holotype, E; isotvpe, BM). Parana mairei Gagnepain in Gagnepain & Courchet, Notul. Syst. (Paris) 3: 154. 1915, excluding the note about the fruit, based on Wilson 4186a, which is Dinetus duclouxii. Syn. nov. TYPE: China. Yun¬ nan: [Xinping Yi-Dai Aut. Xian] rochers dans le val de Mali-ouan, Mai re s.n. (lectotype, selected here, P; isolectotypes. A, E, G, L, P). Parana microsepala Handel-Mazzetti, Kaiserl. Akad. Wiss. Wien, Math.-Naturwiss. Kl. Anz. 58: 229. 1921. Syn. nov. TYPE: China. Yunnan: [Binchuan Xian] Peyentjing, prope Kuti, 27 Aug. 1919, Ten 1 980 (lectotype, selected here, C; isolectotype, WU). Parana mairei Gagnepain & Courchet var. holosericea Wu Cheng-yi in Fang Rhui-cheng & Huang Shu- hua, FI. Reip. Pop. Sin. 64(1): 163. 1979. Syn. nov. TYPE: China. Sichuan: Butuo Xian, Aug. 1959, Sichuan Liangjang Team 5830 (holotype, KUN; isotype, PE). Dinetus dinetoides (C. Schneider) Staples, comb, nov. Basionym: Porana dinetoides C. Schnei¬ der in Sargent. PI. Wilson. 3: 360. 1916. TYPE: China. Yunnan: Mengtsze [Mengzi], alt. 1,700 m, A. Henry 9340 (holotype. A; iso¬ types, E, K, MO, NY, US). Parana megathyrsa Wu Cheng-yi, Yunnan Trop. & Sub- trop. Florist. Res. Rep. 1: 101, pi. 18. 1965. Syn. nov. TYPE: China. Yunnan: Shuangjiang Xian, Sin G. S. 1044 (holotype, KUN: isotypes, IBSC, KUN, PE). Porana brevisepala Wu Cheng-yi & Huang Shu-hua in Fang Rhui-cheng & Huang Shu-hua, Flora Yun- nanica 2: 630. pi. 179, f. 4, 5. 1979. Syn. nov. TYPE: China. Yunnan: Kwangnan [Guangnan Xian], Li zhi guo, [Huaguoqing], 8 Nov. 1964, U u C. A. 9775 (holotype, KUN; isotype, KUN). Parana dinetoides C. Schneider in Sargent var. mien- ningensis Huang Shu-hua in Fang Rhui-cheng & Huang Shu-hua, FI. Reip. Pop. Sin. 64: addendum 163. 1979. Syn. nov. TYPE: China. Sichuan: Mi- anning Xian, King-Khoang [ Jing Song], Ying T. S. 4342 (holotype, KUN not seen; isotype, KUN). Dinetus duclouxii (Gagnepain ( 18) cm x (3 )4 5(-6) cm, apice breviacuminado, base subobtusa decurrente en el peciolo; glabra; haz nitido, con el nervio medio impreso y los secundarios pianos a ligeramente emergentes, enves con glandulas ge- neralmente hacia el nervio medio, y mas densas en la base; nervios secundarios broquidodromos, difi- ciles de diferenciar por la presencia de otros nervios intermedios, 14-18(-26) pares dirigidos hacia el horde; peciolo 1.5-3 mm de largo. Inflorescencias axilares, 2—3 pedicelos florales (el especimen t ij >o solo cuenta con frutos). Frutos eliptico-fusiformes, hipantio glabro, verdoso, 30-40 X 15-20 mm, apice crateriforme, 1.5-2 x 2-2.5 mm; semillas en forma de media luna < 5 por fruto. Ecologia. Bosque primario no inundable, suelo arcilloso. Fructifica en abril junio. Esta especie se dilerencia de las tres Lissocarpa conocidas, L. bentharnii, L. guianensis, y L. ste¬ nocarpa porque estas espeeies tienen ramitas te¬ retes o subteretes, la haz con el nervio medio emer¬ gente, frutos con pedicelos delgados, con el apice del hipantio cerrado o muy ligeramente abierto; mientras que Lissocarpa jensonii tiene ramitas te- tragono-subaladas; la haz con el nervio medio im¬ preso; frutos subsesiles o con pedicelos muy cortos y crasos, con el apice del hipantio crateriforme. La presencia de 5 semillas no parece un caracter para designar una nueva jerarquia taxonomica; seria mas bien una caracteristica para agregar a la familia. Dedico esta especie a Peter Jenson, amigo de la Naturaleza Amazonica y desinteresado colaborador Novon 3; 211-212. 1993. 212 Novon Figura 1. Lissocarpa jensonii Vasquez. — A. Habito. (Vasquez 15984 , AMAZ, MO, USM.) B. Semilla. del Proyecto Flora del Peru, a quien debo que mi trabajo en la selva haya sido mas comodo y pro- vechoso. Paratipo. PERU. Loreto: Provincia de Maynas, Dis¬ trito de Las Amazonas, Quebrada Sucusari, Explor Napo Camp, R. Vasquez et al. 16814 (AMAZ, MO, USM). Literatura Citada Cronquist, A. 1981. An Integrated System of Classifi¬ cation of Flowering Plants. Columbia Univ. Press, New York. Steyermark, J. A. 1987. Flora of the Venezuelan Guy¬ ana — II. Ann. Missouri Bot. Card. 74: 85-116. Gerritea, a New Genus of Paniceae (Poaceae: Panicoideae) from South America Fernando 0. Zuloaga and Osvaldo Morrone Instituto de Botanica Darwinion, Casilla de Correo 22, San Isidro, 1642, Argentina Timothy Killeen Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A. ABSTRACT. Gerritea pseudopetiolata of the tribe Paniceae is described and illustrated. Leaf anatomy oi this new genus is also included, and relationships with related genera within the Paniceae are dis¬ cussed. During fieldwork carried out by T. Killeen in 1987, a previously unknown grass was collected at mid elevation in Bolivia, Department La Paz, be¬ tween Coroico and Caranavi. Additionally, new spec¬ imens, previously collected by S. Beck, were found in the herbarium of LPB. Gerritea pseudopetiolata Zuloaga, Morrone & Killeen, gen. et sp. nov. TYPE: Bolivia. La Paz: Nor Yungas on road between Caranavi and Coroico, 16°00'S, 67°30'W, 800 m, 25 July 1987, T. Killeen 2632 (holotype, ISC; isotypes, F, LPB). Figures 1, 2. Gramen perenne, caespitosum; culmis supra nodis pro¬ fuse ramificantibus, 60-120 cm longis; nodis pilosis. Fo- liorum vaginae pilosae vel glabrae, 4-17 cm longae. Lig- ula inferne membranacea, superne ciliata, 0.3 0.6 mm longa. Lamina lineari-lanceolata, 11-27 cm longa, 0.7- 1.5 cm lata, decidua; pseudopetiolo usque 2 cm longo, piloso. Inflorescentiae paniculatae, laxae, diffusae, 14-30 cm longae, 4-12 cm latae. Spiculae ellipsoideae, 1.8- 2.2 mm longae, 0.5 mm latae, longe pilosae; quam gluma infera %-/ longitudino spiculae aequans, pilosa, 3-nervia; gluma supera 3-nervia. Anthoecia infera neuter: lemma infera 3-nervia; palea infera 0.7-1 mm longa. Anthoecia supera bisexualia, ellipsoidea, 1.3-1. 5 mm longa, 0.4 mm lata, membranacea, decidua, glabra; lemma 3-nervia, pa¬ lea 2-nervia. Lodiculae 2, conduplicatae, glabrae. Stamina 3. Caespitose perennial. Culms rigid, many noded, procumbent, 60-120 cm long, profusely branching at the upper nodes, internodes 4-9 cm long, finely striate, cylindric, pale or tinged with purple, densely pilose toward the distal portion with whitish, ap- pressed hairs; nodes pilose, covered with whitish, long hairs. Leaves mostly cauline. Sheaths longer than the internodes, 4 17 cm long, greenish or tinged with purple, strongly keeled, glabrous or densely papillose-pilose, the margins membranous, ciliate; lower sheaths usually laciniate; auricles small, pilose. Ligules 0.3-0. 6 mm long, membranous-cil- iate, pilose on the abaxial surface, glabrous on the adaxial surface; collar pilose or glabrous, well defined with a transverse brownish line. Blades linear-lan¬ ceolate, 1 1-27 cm long, 0.7- 1 .5 cm wide, flat, with a long pseudopetiole up to 2 cm long, articulated at the base and early deciduous, sparsely pilose, the margins scaherulous, the lower ones ciliate with long whitish hairs or glabrous. Peduncle terete, up to 10 cm long, hirsute. Inflorescence a terminal, exserted, panicle, lax, pyramidal, multiflowered, 14-30 cm long, 4-12 cm wide; main axis flexuous, hispid, greenish or tinged with purple; pulvini pilose, first- order branches divergent; axis of the branches sca¬ brous, sparsely hispid; pedicels up to 8 mm long, flexuous, scaherulous, sparsely pilose, terete. Spike- lets solitary, ellipsoid, 1 .82.2 mm long, 0.5 mm wide, slightly laterally compressed, gaping, greenish or tinged with purple, hispid; upper glume and lower lemma subequal, 3-nerved. Lower glume lanceolate, 1.4-1. 9 mm long, % to 'f the length of the spikelet, with papillose-pilose hairs up to 7 mm long toward the margins, acuminate, 3-nerved, with a stipe 0.2 mm long below its insertion. Upper glume 1.8-2 mm long, herbaceous, acute, papillose-pilose toward the margins with whitish hairs up to 7 mm long. Lower lemma 1.6-1. 7 mm long, glumiform, acute, glabrous. Lower palea ovate-lanceolate, 0.7-1 mm long, 0.1— 0.2 mm wide, membranous, hyaline, the margins scaherulous, 2 -nerved; lower flower absent. Upper anthecium narrowly ellipsoid, 1.3- 1.5 mm long, 0.4 mm wide, glabrous, membranous, smooth, pale, disarticulating at maturity; lemma 3-nerved, the margins flat, not enrolled over the palea; palea 2-nerved, free at its apex; lodicules 2, conduplicate, truncate, 0.2 mm long, embracing the lower margins of the palea; stamens 3, anthers 0.6-0. 7 mm long; styles 2, free; stigmas plumose, whitish. Caryopsis not seen. Novon 3: 213-219. 1993. 214 Novon Figure 1. Gerritea pseudo petiolata Zuloaga, Morrone & Killeen. — a. Habit, with inflorescence included. — b. Detail of ligule and lower portion of the blade. — c. Spikelet, lower glume view. — d. Spikelet, upper glume view, e. Lower palea. — f. Upper anthecium, lemma view. — g. Upper anthecium, palea view. — h. LIpper palea with lodicules, stamens, and portion of stigmas. Based on Killeen 2632. Volume 3, Number 2 1993 Zuloaga et al. Gerritea 215 Figure 2. Leaf blade anatomy of Gerritea pseudopetiolata. a-c, Transectional anatomy. — a. Transverse section detail showing first- and second-order vascular bundles. — b. Detail showing second-order vascular bundles, bulliform cells, mesophyll cells, and sclerenchyma girders. — c. Detail showing a first-order vascular bundle. — d, e. Abaxial epidermis. — f. Adaxial epidermis. Based on Killeen 2632. Distrirction and Ecology The species has been collected only from a single locality in the “yungas” of La Paz, Bolivia. The type locality is situated on the major highway be¬ tween the departments of La Paz and the Beni. This area is a small, narrow canyon about 200 m long that is a minor tributary of the Bio Coroico. The canyon is notable because of a 40-m waterfall that is located at its upper edge. The vegetation of the area is a semideciduous montane forest; unfortu¬ nately, most of the original vegetation has been destroyed due to colonization activities over the past 50 years. This particular site remains relatively un¬ altered due to very steep slopes that are unsuitable for agriculture. Plants of Gerritea pseudopetiolata are restricted to a relatively narrow band between the road and the waterfall. The microhabitat is moist and well- shaded when compared to the surrounding forest vegetation. Plants are not found in the immediate environs of the waterfall where there is a constant mist, nor do they occur in the disturbed, more open areas in the gorge below the road. 216 Novon Caespitose individuals are rooted in shallow cracks between rocks or upon a superficial mat of mosses. The rocks are sedimentary slates of probable Or¬ dovician origin; they are hard and resistant to weath¬ ering. The species is not locally abundant, and only 1 1 individuals were observed in 1992. The life form of the species can be described as a lithophy tic grass. Flowering is day-length dependent. The site was visited periodically between '989 and 1992 and reproductive material was collected only between late June and early July, when days have approxi¬ mately 1 1 hours of light. Each culm may produce up to three secondary branches, and most terminal meristems (hut not all) produce inflorescences. Oc¬ casional culms have a senescent inflorescence, as well as a new inflorescence from a secondary branch produced during the subsequent year. Flowering culms may have as many as 25 internodes or as few as six. Biogeography “Yungas” is a Bolivian term describing the humid intermontane valleys of the eastern slope of the Andes. In traditional natural history literature it usually refers to the valleys directly to the northeast of the city of La Paz. This area covers vegetation types ranging from humid tropical forest to semi- deciduous tropical forest, cloud forest, and tropical montane grassland (the latter probably anthropo¬ genic). The La Paz yungas is the most extensively explored region in Bolivia and has been the focus of numerous collecting expeditions by botanists since the latter part of the 19th century (Funk & Mori, 1989). Numerous endemic species have been de¬ scribed for the La Paz yungas, and a total of 88 species of vascular plants are known (pers. comm., Centro de Datos para la Conservacion — Bolivia); of these most are epiphytic orchids (23 species) and bromeliads (20 species). This is the first report of an endemic grass genus for Bolivia. The life form of the species is similar to the more numerous en¬ demic orchids and bromeliads, as it is also a litho- phyte. Paratypes. BOLIVIA. La Paz: Nor Yungas, carretera La Paz-Caranavi, 27 km de Caranavi, canon estrecho cerca de una cascada grande, laderas rocosas, 700 m, 18 June 1989, Smith & Buddensiek 13515 (LPB, MO, SI); Caranavi, 28 km hacia Coroico, 930 m, 70°W, ma- torral humedo sobre rocas, 12 Mar. 1979, Beck 556 (LPB), 16 July 1979, Beck 1766 (LPB). The new genus is named after Gerrit Davidse in recognition of his work on the rich grass flora ol the American tropics and his generosity in helping us with the first studies of this new genus. The specific epithet makes reference to the conspicuous pseudopetiolate blades. Each individual plant of Gerritea pseudopetiol- ata consisted of between 5 and 40 culms arranged in caespitose tussocks. Culms are decumbent and “hang” downward over bare exposed rock; plants do not root at the nodes, even when suitable sub¬ strate is available. Culms are long-lived and produce up to 25 internodes; only the upper 4-6 nodes support green foliage. Leaf blades are articulate below the pseudopetiole and early deciduous; the lower nodes are covered by senescent sheaths. De¬ ciduous leaf blades in grasses have been cited by Arber (1934) in Aristida cyanantha Steudel ex Trinius and Agropyron repens P. Beauvois. Spikelets are markedly pilose, the hairs up to four times the length of the spikelets. Davidse (1987) suggested that genera with prominent spikelet hairs are probably specialized for dispersal by adhesion and wind; among the genera Davidse mentioned are Tricholaena Schultes, Rhynchelytrum Nees, and Leptocoryphium Nees. Leak Blade Anatomy (Figure 2) Gerritea pseudopetiolata is a non-Kranz, XyMS + species (following the terminology described by Hat- tersley & Watson, 1976), characterized by having two bundle sheaths surrounding the vascular bun¬ dles, the outer parenchymatous sheath without spe¬ cialized chloroplasts, and the mesophyll irregularly radiated, with 5-8 mesophyll cells between consec¬ utive vascular bundles (Hattersley & Watson, 1975). Leaf in transverse section. Outline: open, ex¬ panded; leaf thickness at mid-lamina 155-200 jttm, leaf thickness 130-160 /jm laterally on ribs. Ribs and furrows: slightly pronounced; ribs associated with all vascular bundles. Median vascular bundle: keel developed; median vascular bundle structurally distinguishable from lateral first-order vascular bun¬ dles, solitary or associated with 2 second-order vas¬ cular bundles. Vascular bundle arrangement: 5 first-order vascular bundles and 24-30 second-order vascular bundles per leaf section; 5 7 second-order vascular bundles between consecutive first-order vascular bundles; all vascular bundles situated in center of blade; 5-8 mesophyll cells between con¬ tiguous vascular bundles; with a distance of 165— 250 /am between consecutive vascular bundles. First- order vascular bundles round in outline; metaxylem vessel elements wide, slightly greater than paren¬ chyma sheath cells, with angular walls; protoxylem vessel and lacunae cavity present; phloem tissue adjoining the inner bundle sheath. Second-order vas¬ cular bundles angular in outline with xylem and Volume 3, Number 2 1993 Zuloaga et al. Gerritea 217 phloem tissue distinguishable. Vascular bundle sheaths: outer first-order bundle sheaths consisting of 8 13 parenchyma cells, with abaxial interruption of sclerenchyma girders; outer second-order bundle sheaths completely surrounding the vascular bun¬ dles, with 6-8 parenchyma cells; bundle sheath ex¬ tensions only present in second-order vascular bun¬ dles contiguous to the keel, otherwise absent; parenchyma sheath cells inflated with thin walls and lacking chloroplasts; mestome bundle sheaths com¬ pletely surrounding the xylem and phloem tissue. Sclerenchyma: adaxial and abaxial girders associ¬ ated with first- and second-order vascular bundles. Mesophyll: chlorenchyma irregularly radiated in two layers around the vascular bundles in the adaxial surface; adaxial cells tend to a palisadelike type of arrangement in the adaxial chlorenchyma; chloren¬ chyma cells tabular, raquimorphous or isodiametric, with intercellular air spaces. Adaxial epidermal cells: bulliform cells present in adaxial furrows be¬ tween all vascular bundles, in restricted groups of 4-6 fan-shaped cells; small epidermal cells associ¬ ated with the ribs; unicellular macrohairs either with raised cushion-bases or inserted between unmodified epidermal cells as seen in section. Abaxial epi¬ dermis: bulliform cells and macrohairs present in the abaxial furrows, similar to the ones of the adaxial epiderm cells; papillae absent. ABAXIAL EPIDERMIS Zonation: costal and intercostal zones distinguish¬ able. Costal zones: with 4—7 files of cells. Silica bodies dumbbell-shaped, usually alternating with 1 square cork cell; prickles not observed. Intercostal cells: narrow, 10-15 cells wide, long cells rect¬ angular in shape, more than three times as long as wide; cells adjoining the costal zones with undulated walls, central cells of the intercostal zones with sin¬ uous walls; short cells solitary or paired, irregularly present; silica bodies irregular in outline or cruci¬ form. .Stomata corn/j/ea: 26 32.5 yam long, 14 18.2 yarn wide, in 4 rows in the intercostal zones, subsid¬ iary cells triangular-shaped. Microhairs fusiform, 70-98 yam long, bicellular, with walls of the basal cell parallel, thicker than the walls of the distal cells, apex of the distal cell sharply pointed. Hooks small, with the bases shorter than the stomata, the barb longer than the base. Macrohairs unicellular, 150 315 yam long, the epidermal cells associated with the base. Papillae absent. ADAXIAI. EPIDERMIS Intercostal long and short cells: similar to the ones of the abaxial surface. Stomata complex in 2 rows in the intercostal zones. Microhairs as in the abaxial epidermis. Hooks: intercostal hooks irreg¬ ularly present. Macrohairs similar to the ones of the abaxial surface. Papillae absent. Relationship to Other Genera of the Paniceae Gerritea belongs to the tribe Paniceae, which includes genera with biflowered spikelets, disartic¬ ulating below the glumes and with upper anthecium more or less indurated, the glumes membranous. Gerritea is distinguished from related genera by the following set of characters: pseudopetiole prom¬ inent, up to 2 cm long; blades articulated at the base and deciduous, inflorescence a lax panicle with spikelets long-pilose, laterally compressed, the lower glume % to '/ the length of the spikelet, 3-nerved; upper glume and lower lemma 3-nerved; lower palea small; upper anthecium disarticulating from the spikelet, membranous, smooth, lemma with flat mar¬ gins, the palea free toward the tip. Gerritea is a non-kranz genus. Within the tribe, Gerritea is related by different characters to Panicum L. and other genera such as Hymenachne P. Beauvois, Triscenia Grisebach, Digitaria Haller, Ichnanthus P. Beauvois, and Tri- cholaena (Clayton & Renvoize, 1986; Watson & Dallwitz, 1988; Webster et al., 1989; see Table 1 ). Hymenachne differs by its spongy internodes, filled with aerenchyma; panicles spiciform, with pri¬ mary branches appressed to the main axis, with spikelets secund; spikelets dorsiventrally com¬ pressed, with lower glume up to '/, the length of the spikelet, the lower palea absent. Triscenia is distinguished by its filiform, non- pseudopetiolate leaves, inflorescences contracted, with racemose branches; spikelets dorsiventrally compressed, the lower glume up to '/, the length of the spikelet; lower palea absent, and upper anthe¬ cium with the palea not free toward the apex. Digitaria includes species with inflorescences with secund spikelets; spikelets plano-convex, the lower glume small or absent; upper anthecium cartilagi¬ nous, the lemma with margins thinner in texture than the body. The genus comprises Kranz species. Ichnanthus has inflorescences with primary branches with appressed or spreading secondary branches, with or without secund spikelets; upper anthecium indurated, the lemma with involute mar¬ gins and with basal scars or appendages, the palea not free toward the tip. Tricholaena differs by its non-pseudopetiolate leaves, spikelets with the lower glume small, and upper anthecium with the palea not free toward the tip; all species of the genus are kranz. 218 Novon o CJ £ o I I + + I I I I + I ^ + -2 -o + .S o + CO + JU £ 9- ’5b 9- S Cu J C/} J3 P Photosynthetic type C, Ct C3 C3 C,/C4 Volume 3, Number 2 1993 Zuloaga et al. Gerritea 219 Lastly, Panicum has species with spikelets dor- siventrally compressed, upper anthecium indurated, the lemma with involute margins and the palea not free at the apex (the only species with upper palea tree at the apex is Panicum discrepant Doell, which has on the other hand indurated upper anthecia). Membranous upper anthecia are only present in Panicum in species of section Laxa (A. Hitchcock & Chase) Pilger, such as P. leptachne Doell. P. longum A. Hitchcock & Chase, P. pernambucense (Sprengel) Pilger, and P. grumosum Nees (Zuloaga et ah, 1992). These species have inflorescences with racemose branches, the spikelets secund, with the palea not free at the apex. Panicum includes Kranz and non-Kranz species. Acknowledgments. We thank Vladimiro Dudas for the illustrations. We also express gratitude to Angel Cabrera for the Latin diagnosis of the new genus. Literature Cited Arber, A. 1934. The Gramineae, a Study of Cereal, Bamboo, and Grass. Cambridge Univ. Press, Cam¬ bridge. Clayton, W. D. & S. A. Renvoize. 1986. Genera Gra- minum. Kew Bull., Addit. Ser. XIII. Davidse, G. 1987. Fruit dispersal in the Poaceae. Pp. 143-155 in T. R. Soderstrom, K. W. Hilu, C. S. Campbell & M. E. Barkworth (editors), Grass Sys- tematics and Evolution. Smithsonian Institution Press, Washington, D.C. Funk, V. A. & S. A. Mori. 1989. A bibliography of plant collectors in Bolivia. Smithsonian Contr. Bot. 70: 1-20. Hattersley, P. W. & L. Watson. 1975. Anatomical parameters for predicting photosynthetic pathways of grass leaves: The “maximum lateral cell count” and the “maximum cells distant count.” Phytomor¬ phology 25: 325-333. - & - . 1976. C4 grasses: an anatomical criterion for distinguishing between NADP-Malic en¬ zyme species and PCK or Nad-Malic enzyme species. Austral. J. Bot. 24: 297-308. Watson, L. & M. J. Dallwitz. 1988. Grass Genera of the World. Illustrations of Characters, Interactive Identification, Information Retrieval. Australian Na¬ tional University, Canberra. Webster, R. D., J. H. Kirkbride & .1. Valdes Reyna. 1989. New World genera of the Paniceae (Poaceae: Panicoideae). Sida 13: 393-417. Zuloaga, F. ()., R. P. Ellis & 0. Morrone. 1992. A revision of Panicum subgenus Phanopyrum section Laxa (Poaceae: Panicoideae: Paniceae). Ann. Mis¬ souri Bot. Gard. 79: 770-818. Volume 3, Number 2, pp. 93-219 of NOVON was published on 1 July 1993. Volume 3 Number 3 1993 NOVON Una Especie Nueva de Hansteinia (Acanthaceae) del Volcan del Tacana, Chiapas (Mexico) Salvador Acosta Castellanos Centro Interdisciplinario de Investigacion para el Desarrollo Integral Regional Unidad Oaxaca, I.P.N., Apartado Postal 24, Administracion 3, 68081, Oaxaca, Oax., Mexico Rafael Fernandez Nava Laboratorio de Botanica Fanerogamica, Escuela Nacional de Ciencias Biologicas, Instituto Politecnico Nacional, Apartado Postal 17-564, 11410 Mexico, D.F., Mexico Resumen. Se describe una especie nueva de Hans¬ teinia (Acanthaceae) solamente conocida del volcan del Tacana, Chiapas, Mexico, que se diferencia del resto de las especies mexicanas por tener inflores- cencias espigadas. Su grano de polen es esferoidal, diporado, y con ornamentacion equinada ipie con- cuerda con la morfologia de los granos de polen del genero. Abstract. A new species of Hansteinia (Acan¬ thaceae) is described from the Tacana volcano, Chia¬ pas, Mexico. It differs from the other Mexican spe¬ cies of the genus by its spicate inflorescences. The pollen grains are spheroidal, diporate, and ec Innately sculptured, in agreement with the pollen morphology of the genus. A1 examinar especimenes de la familia Acant¬ haceae provenientes del volcan del 1 acana, Chiapas, para Flora Mesoamericana se descubrio una es¬ pecie nueva del genero Hansteinia (= Glockeria), constituido por aproximadamente 1 4 especies de Mexico, Centroamerica, y norte de Sudamerica. Co- munmente Hansteinia presenta inflorescencias pa- niculadas; a diferencia de las demas especies del genero, este nuevo taxon se destaca por sus inflo¬ rescencias espigadas. Hansteinia tacanensis Acosta & Fernandez, sp. nov. I IPO: Mexico. Chiapas: on the SE side of Volcan Tacana above Talquian, munieipio of Union Juarez, elev. 2,200 m, 23 Nov. 1980, D. E. Breedlove & F. Almeda I TT I I (holotipo, MEXU; isotipo, CAS). Figura 1. Frutex 1.5-2 in altus; caules erecti, dense pubescentes, pilis multiseptatis, longis glandulosis, cum brevibus non glandulosis rnixtis. Folia opposita, petiolata; limbus ovato- lanceolatus, apice acuminato, base in petioluin decurrenti, margine integro, ciliato, supra et infra subpubescens, pilis dispersis. Inflorescentiae longae, spicatae. Flores subses- siles vel brevipedicellati. Calyx 5-partitus, laciniis subae- qualibus subulatis, pubescentibus. Corolla rubra, extus pubescens, intus glabrata, ventricosa. Stamina 2, exserta; antherae uniloculares. Capsula ellipsoidalis, puberulenta. Semina 4, lentiformia, verrucosa. Arbusto de 1.5 a 2 m de alto, tallos erectos redondeados, densamente pubescentes, con pelos multiseptados, glandulosos de 0.7-1 mm de largo, mezclados con pelos no glandulosos de 0.02 — 0.04 cm de longitud. Hojas opuestas, pecioladas, peciolos de 0.3-4. 5 cm de longitud, pubescentes; laminas de 4-12 cm de largo por 2. 5-5. 5 cm de ancho, ovado-lanceoladas, apice acuminado, base decu- rrente sobre el peciolo, margen entero, ligeramente ciliado, haz y enves puberulentos, cistolitos de ± 0.2 mm de longitud abundantemente esparcidos so¬ bre toda la superficie. Inflorescencias pedunculadas, terminales y axilares, espigadas, de 7-20 cm; pe- dunculos redondeados, pubescentes, con dos tipos de pelos, unos de 0.02-0.03 cm de largo, muy Novon 3: 221-224. 1993. 222 Novon Figura 1. Holotipo de Hansteinia tacanensis Acosta & Fernandez. — a. Habito. — b. Flor con los estambres exertos. — c. Fruto con semillas. Volume 3, Number 3 1993 Acosta Castellanos & Fernandez Nava Hansteinia tacanensis 223 densos y otros mas esparcidos, multiseptados de 0.05-0.1 cm de longitud; bracteas de 0.5 — 0.8 cm de largo por 0.2-0. 3 cm de ancho, largamente subtriangulares, pubescentes, con el apice agudo; bracteolas subuladas de 0.3-0. 5 cm de largo por 0.1 cm de ancho; el raquis densamente pubescente con pelos cortos, rectos, de 0.02-0.03 cm de largo y pelos glandulares, esparcidos, multiseptados, de 0.05-0.1 cm de largo; en la base de las espigas se presenta un par de hojas mas pequenas que las caulinares, de 0.7-4 cm de largo por 0.4-3 cm de ancho, subpecioladas, base obtusa a redondeada. Flores subsesiles o con pedicelos de 0. 1-0.4 cm de largo; segmentos del caliz 5, subiguales de 0.9- 1.2 cm de largo por 0.08-0. 1 cm de ancho, subulados, con apice agudo, pubescentes, con pelos glandulares, multiseptados. Corola roja, bilabiada, de 2-2.5 cm de largo, pubescente en la superficie externa y en la interna glabra; tubo de la corola angosto, de 0.5 0.7 cm de largo por 0.15 0.2 cm de ancho, abrup- tamente expandido a manera de una garganta ven- tricosa de 0.9 cm en su parte mas ancha, contraida cerca de 0.4 cm bajo el limbo; el labio superior, triangular, entero, de 0.6 cm de largo por 0.5 cm de ancho, con el apice obtuso, el labio inferior, obovado, con 3 pequenos lobulos, de hasta 0.1 cm de largo, con el central notablemente mas corto. Estambres 2, insertos justo por arriba de donde se amplia el tubo de la corola; filamentos exertos, gla- bros, de 2. 5-2. 8 cm de largo; anteras monotecas, dorsifijas, loculos de 0.3 cm de largo, muticos. Disco verdoso, cilindrico de 0.1 cm de diametro. Ovario glabro bilocular, con 2 ovulos por loculo; estilo gla- bro, filiforme, de 3.5 cm de largo, estigma subca- pitado. Capsula de aproximadamente 1 cm de lon¬ gitud, largamente elipsoide, subestipitada, puberulenta, conteniendo 4 semillas lentiformes, hasta de 0.2 cm de diametro, verrucosas. Florece de noviembre a enero. Distribut ion v habitat. Se conoce solo del estado de Chiapas, Mexico, creciendo en el bosque mesofilo de montana, que se desarrolla en una de las laderas del volcan Tacana. Nuestra especie difiere de las especies previa- mente conocidas del genero por sus inflorescencias espigadas; sin embargo, por la notable similitud de hojas y de estructura floral, la corola con el tubo abruptamente ventricoso, y la morfologia de los gra- nos de polen (esferoides, diporados, y con orna- mentacion equinada (Fig. 2), que concuerdan con lo observado por Kaj (1961)), se decidio ubicarla dentro de Hansteinia. Cabe destacar que uno de los generos afines, Razisea, presenta tambien inflorescencias espiga- Figura 2. Grano de polen en vista ecuatorial, mostrando los dos poros, x 1,200. das, pero sus flores de tubo gradualmente ampliado y no abruptamente ventricoso son de forma nota¬ blemente diferente. For otro lado Wood (1988) in- cluye al genero Hansteinia como sinonimo de Ha- bracanthus; sin embargo, nosotros consideramos que las caracteristicas tradicionalmente utilizadas para separar a los dos generos (corolas someramente tet- ralobadas, generalmente ventricosas en el primero y corolas claramente bilabiadas, con los labios mas largos que el tubo de la corola en el segundo) son caracteres que separan en forma natural a ambos generos, por lo cual pensamos que estos deben man- tenerse independientes. Por todo lo anteriormente senalado, al parecer Hansteinia tacanensis constituye un taxon bien di- ferenciado. Clave para la Identificacion de las Especies Conocidas de Hassteima de Chiapas (Mexico) y Guatemala la. Pedunculos y raquis de las inflorescencias pi- loso-glandulares. 2a. Inflorescencias paniculadas; corolas gla- bras, de color amarillo con bandas rojas, no ventricosa. Chiapas, Mexico; Chimal- tenango y Solola, Guatemala . . Hansteinia monolopha (Donnell-Smith) D. Gibson 2b. Inflorescencias espigadas; corolas pubes¬ centes, de color rojo, ventricosa. Chiapas, Mexico . Hansteinia tacanensis Acosta & Fernandez 224 Novon lb. Pedunculos y raquis de las inflorescencias no piloso-glandulares. 3a. Paniculas de 5-7 cm de longitud; sepalos lanceolados hojas de 2-3 cm de ancho. Chiapas, Mexico . . Hansteinia purpusii Brandegee 3b. Paniculas de 10-15 cm de longitud; se¬ palos lineares; hojas 3.5-6 cm de ancho. Chiapas, Mexico; Quetzaltenango, Guate¬ mala . Hansteinia glabra (Leonard) D. Gibson Por otro lado, es importante senalar que el Volcan Tacana constituye un sitio de alto endemismo, es- pecialmente en las asociaciones de mayor altitud (Breedlove, 1981). Paratipo. MEXICO. Chiapas: El Talquian Viejo, mu- nicipio de Union Juarez, alt. 1,700 m, 24 enero 1992, E. Ventura & E. Lopez 10075 (CAS, CHAPA, ENCB, MEXU). Agradecimientos. Se agradece a las autoridades de los herbarios (ENCB y MEXU) por el prestamo de sus materiales de Hansteinia y a Rodolfo Palacios por haber realizado el estudio y torna de fotografias de los granos de polen. Thomas F. Daniel, Remedios Aguilar, y Fernando Chiang hicieron importantes sugerencias para mejorar el trabajo; ademas, este ultimo reviso la diagnosis en latin. Se agradece a Concepcion Rodriguez Jimenez, Directora del Pro- yecto de Flora Mesoamericana en el Herbario ENCB, por el apoyo brindado a este trabajo. Este trabajo fue parcialmente subsidiado por el Consejo Nacional de Ciencia y Tecnologia 1)1 12-904622. Literatura Citada Breedlove, D. E. 1981. Introduction to the Flora of Chiapas. Flora of Chiapas, Part. 1. California Acad¬ emy of Sciences. San Francisco. Raj, B. 1961. Pollen morphological studies in the Acan- thaceae. Grana Palynologica 3: 3-108. Wood, J. R. I. 1988. Colombian Acanthaceae — Some new discoveries artd some reconsiderations. Kew Bull. 43: 1-51. New Combinations in North American Asarum (Aristolochiaceae) Kerry Barringer Brooklyn Botanic Garden, 1000 Washington Avenue, Brooklyn, New York 1 1225, U.S.A. Abstract. New nomenclatural combinations are needed for some of the North American species of Asarum to bring their names in line with the current understanding of relationships in the genus and to provide correct names for a treatment in the Flora of North America. Asarum L., in the broad sense, is a genus of low herbs with creeping stems and a few cordate-reni- form leaves. Its flowers have a radially symmetrical, three-lobed calyx, twelve or fifteen stamens, and six styles. Its fruits are fleshy capsules containing many flattened, cordate, arillate seeds. There are about 90 species in Asarum s.l. (Araki, 1953; Cheng & Yang, 1983). Most of the species are found in tem¬ perate Eastern Asia and the southeastern United States, but a few also occur in Europe, eastern North America, and the Pacific Northwest. It is the only genus in the tribe Asareae and appears to be related to the genus Saruma Oliver, a native of China. Since Braun (1861), a number of sections have been recognized within Asarum (l)uchartre, 1864; Schmidt, 1935). These are based on the fusion of perianth, the presence of constrictions and annuli in the calyx, the position of stigmas, the length of the anther connectives, the position of the ovary, and the longevity ol the leaves. Some of these sec¬ tions have been raised to generic rank, probably because they appear distinctive within their regional floras. Hexastylis Rafinesque ol the southeastern United States is one of these. Braun. Duchartre, and Schmidt recognized the same group of species as Asarum sect. Ceratasarum Braun. The classification of the species ol Asarum by Araki ( 1 953) clarified the similarities and differences between the sections. Araki recognized two subgen¬ era in Asarum: subg. Asarum , distinguished by the sepals being free above the base and forming a pseudotube, and subg. Heterotropa (Morren & I)e- caisne) 0. C. Schmidt, distinguished by the sepals being fused into a tube. He placed section (era- tasarum ( Hexastylis ) in subgenus Heterotropa with sections Heterotropa (Morren & Decaisne) Braun and Asiasarum (Maekawa) Araki, and he placed the other North American species in subgenus Asarum. The recognition of Hexastylis as a genus has persisted in the southeastern United States, in part, because Blomquist overlooked Araki’s classification when writing his influential revision of Hexastylis (Blomquist, 1957). Also, it is easy to distinguish the species of Asarum sect. Ceratasarum ( Hexastylis ) from Asarum canadense L., the only other species of Asarum in the eastern United States. Blomquist (1957) listed 1 1 characters that dis¬ tinguish the groups in eastern North America. Ex- trorse stamens and bifurcate style extensions define the species in section Ceratasarum. The other char¬ acters cited by Blomquist are less reliable. The thick, variegated, glabrous leaves that help to distinguish Hexastylis in the southeast are similar to those found in many other species of .Asarum subg. Het¬ erotropa, and in Asarum hartwegii S. Watson, a species in subgenus Asarum native to northwestern North America. Asarum europaeurn L., the type ol Asarum and subgenus Asarum, has leaves that are evergreen, but not variegated. The tubular calyces are characteristic of all species in subgenus Het¬ erotropa, and subgenus Asarum sect. Brevituba C.Y. Cheng & C. S. Yang is somewhat intermediate in having species with lobes that are fused at the base. Superior to partly inferior ovaries are also found throughout subgenus Heterotropa. Short, blunt extensions on the anther connectives and attenuated calyx lobes are found in many species of Asarum, though the tendency is for the extensions of species in subgenus Heterotropa to be shorter. Finally, short staminal filaments are also found in all sections of subgenus Heterotropa with the exception of section Asiasarum (Maekawa) Araki. Recognition of a broadly defined Asarum is in accord with some of the major treatments of the genus. Cheng & Yang (1983) used the broad con¬ cept of Asarum in their treatments of the Chinese species. Owhi (1965) and Hatusima & Yamahata (1988) used it in their treatments of the Japanese species. In addition, the recognition of a broadly defined Asarum is consistent with the generic con¬ cepts of the other large genera in the Aristolochi¬ aceae (i.e. , Aristoloehia (Schmidt, 1935; Pfeifer, 1966; Ma, 1989), That tea (Ding Hou, 1981)). With the recognition of Asarum in the broad sense, the following new combinations are necessary. Novon 3: 225-227. 1993. 226 Novon The other species segregated in Hexastylis either already have valid names in Asarum or are consid¬ ered synonyms. Asarum arifolium Michaux var. ruthii (Ashe) Barringer, comb. nov. Basionym: Asarum ru¬ thii Ashe, Bot. Contr. My Herb. 1: 4. 1897. Hexastylis ruthii (Ashe) Small, FI. S.E. U.S. 1132. 1903. Hexastylis arifolia var. ruthii (Ashe) H. Blomquist, Brittonia 8: 268. 1957. TYPE: U.S. A. Tennessee: Knox Co., July 1896, Ruth s.n. (lectotype, here designated, NY). Ashe (1897a) originally published Asarum ruthii in a pamphlet entitled Botanical Contributions from My Herbarium. No. 1, dated 28 Oct. 1897. A later publication drawn from that separate (Ashe, 1897b) cited the earlier publication as the protologue for his names. Blomquist (1957) overlooked the earlier pamphlet, citing the later as the source of the names, but the earlier pamphlet is effectively published ac¬ cording to the Code (Greuter et ah, 1988) and must be used despite its obscurity. Ashe (1897a) did not designate a type in the protologue, but listed his herbarium numbers 1983, 201 1, and 2015. In the later paper (Ashe, 1897b) he noted the type locality as Morristown, Tennessee. No Ashe specimens were found that correspond with either the numbers cited or the locality. No Ruth specimens are known from this locality, because the original Ruth material burned in 1934 (Anonymous, 1934). As a lectotype, the Ruth collection from Knox County, Tennessee, in 1896 corresponds to the protologue and is the extant collection most likely to have been seen by Ashe prior to publication. Plants of variety ruthii have generally occurred west of the Appalachian Mountains in Tennessee and Kentucky. They have calyx lobes that are short¬ er and more erect than the typical variety found east of the range. Asarum arifolium Michaux var. callifolium (Small) Barringer, comb. nov. Basionym: As¬ arum callifolium Small, Bull. Torrey Bot. Club 24: 334. 1897. Hexastylis arifolia var. cal- lifolia (Small) H. Blomquist, Brittonia 8: 268. 1957. TYPE: U.S. A. Florida: shady woods. Chapman s.n. (holotype, NY). Blomquist (1957) and Gaddy (1987b) showed that Asarum callifolium Small is a large-flowered. Gulf Coast variety of the widespread A. arifolium Michaux. Asarum contractum (H. Blomquist) Barringer, comb. nov. Basionym: Hexastylis contracta H. Blomquist, Brittonia 8: 279. 1957. TYPE: U.S. A. North Carolina: Buncombe Co., Broad River, between Old Fort and Bat Cave, 4 June 1951, Godfrey & Anderson 51225 (holotype, DUKE; isotype, NCS). Hexastylis rhombiformis Gaddy, Brittonia 38: 82. 1986. TYPE: U.S. A. North Carolina: Transylvania Co., North Fork of the French Broad River, ca. 6 mi. NNW of Rosman, 22 Apr. 1982, Gaddy 250 (ho¬ lotype, CLEMS). Flowers of Asarum contractum are variable in size and shape. Gaddy (1986, 1987b) segregated plants with prominent reticulation inside the calyx and the broadest part of the calyx above the middle into Hexastylis rhombiformis Gaddy. However, the photograph accompanying the original description clearly shows flowers of //. rhombiformis with the shape of typical A. contractum. Blomquist was aware of this variability in A. contractum and included both extremes in his figures 7 and 8 (Blomquist, 1957). Gaddy (1987b) pointed out the unusual dis¬ tribution of A. contractum, but H. rhombiformis is sympatric with A. contractum in acidic woods on the Blue Ridge of western North Carolina. Further study may show this group to be another phase of the variable A. virginicum L. Asarum speciosum (Harper) Barringer, comb. nov. Basionym: Hexastylis speciosa Harper, Tor- reya 24: 77-83. 1924. TYPE: U.S.A. Ala¬ bama: Autauga Co., about 2 mi. E of Booth, 18 May 1924, Harper *£ Holt s.n. (lectotype, designated by Blomquist (1957), NY). Blomquist (1957) compared this species to the Asian species of Asarum subg. Heterotropa that have a broad mouth and spreading calyx lobes. None of the Asian species is quite like this species, though. In the Asian species, the flare is made up in part of an annulus that constricts the opening of the calyx tube. Such an annulus is absent in A. speciosa. Also, the bifid style extensions and stamen connec¬ tives without extensions clearly ally A. speciosa with section Ceratasarum. Asarum shuttleworthii Britten & Baker var. harperi (Gaddy) Barringer, comb. nov. Bas¬ ionym: Hexastylis shuttleworthii (Britten & Baker) Small var. harperi Gaddy, Sida 12: 54. 1987. TYPE: U.S.A. Georgia: Madison Co., just N of GA 106, 14.2 mi. NE of Athens, 9 May 1986, Gaddy s.n. (holotype, CLEMS; is¬ otypes, GH, MO, NCU, NY). This variety is known in the horticultural trade under the cultivar name ‘Calloway’ (Galle, 1984). Volume 3, Number 3 1993 Barringer North American Asarum 227 It is a hog-growing variety with elongate, rather than short, rhizomes (Gaddy, 1987a). Acknowledgments. 1 thank Bruce King. As a teacher, he introduced me to plant taxonomy and the problems of Asarum. As a colleague, lie helped me with some of the fieldwork and suggested alter¬ native interpretations. I also thank the curators of A, F, Gff, MO, NCU, NY, PH, and LIS, who kindly allowed me to study their collections, and the li¬ brarians at the New York Botanical Garden, who helped me with the literature. Literature Cited Anonymous. 1934. News Notes. Torreya 34: 26. Araki, Y. 1953. Systema generis Asari. Acta Phytotax. Geobot. 15: 33-36. Ashe, W. W. 1897a. The genus Asarum in eastern America. Contr. My Herb. 1: 1-4. - . 1897b. The glabrous-leaved species of Asa- rum of the southern United States. J. Elisha Mitchell Sci. Soc. 14:3 1-36. Blomquist, H. L. 1957. A revision of Hexastylis of North America. Brittonia 8: 255-281. Braun, A. 1861. Index seminum Horti Botanici Bero- linensis 1861. Appendix. Plantarum novarum et mi¬ nus cognitarum, quae in Horto Regio Botanico Be- rolinensi coluntur. Berlin. Cheng, C. Y. & C. S. Yang. 1983. A synopsis of the Chinese species of Asarum (Aristolochiaceae). J. Ar¬ nold Arbor. 64: 565 — 597. Duchartre, P. 1864. Aristolochiaceae. In: A. de Can¬ dolle, Prodromus 15(1): 421-498. Ding Hou. 1981 . Florae Malesianae Praecursores LXII. On the genus Thottea (Aristolochiaceae). Blumea 27: 301-332. Gaddy, L. L. 1986. A new heartleaf ( Hexastylis ) from Transylvania County, North Carolina. Brittonia 38: 82-85. - . 1987a. Hexastylis shuttleworthii var. har- peri (Aristolochiaceae), a new variety of heartleaf from Alabama and Georgia. Sida 12: 51-56. - . 1987b. A review of the taxonomy and bio¬ geography of Hexastylis (Aristolochiaceae). Casta- nea 52: 186 196. Galle, E. C. 1984. ‘Calloway’ Ginger. Hexastylis shut¬ tleworthii. Bull. Amer. Rock Card. Soc. 42: 36-48. Greuter, W. et al. 1 988. International Code of Botanical Nomenclature. Regnum Veg. 118. Hatusima, S. & Yamahata. 1988. Illegitirnatly pub¬ lished taxa of Asarum from Japan. J. Phytogeogr. Tax. 36: 18. Ma Jin-Shuang. 1989. A revision of Aristolochia L. from E. & S. Asia. Acta Phytotax. Sin. 27: 321 — 364. Owhi, J. 1 965. Flora of Japan [in English], Smithsonian Institution, Washington, D.C. Pfeifer, H. W. 1966. Revision of the North and Central American hexandrous species of Aristolochia (Ar¬ istolochiaceae). Ann. Missouri Bot. Gard. 53: 1-1 14. Schmidt, O. C. 1935. Aristolochiaceae. In: A. Engler & K. Prantl (editors), Natiirlichen Pflanzenfainilien, ed. 2. 16(3b): 204-242. Five New Species of Chusquea (Poaceae: Bambusoideae) and a New Combination Lynn G. Clark Department of Botany, Iowa State University, Ames, Iowa 50011, U.S.A. Abstract. Five new species, Chusquea falcata, C. subulata, C. barbata, C. spathacea, and C. pulchella , are described and illustrated. One new combination, Chusquea attenuata (Doell in Martius) L. G. Clark, is proposed for a Brazilian species originally described as an Arundinaria, based on a vegetative specimen. During the course of Horistic and phylogenetic studies in the neotropical woody bamboo genus Chusquea Kunth, 1 have discovered a number of species new to science, of which five are described here. A previously poorly known species of Arun¬ dinaria Michaux from Brazil is transferred to Chus¬ quea, and the new combination is made. Chusquea falcata L. G. Clark, sp. nov. TYPE: Ecuador. Loja: Parque Nacional Podocarpus, above Nudo de Cajanuma, wet montane forest around "Centro de Information," 2,800 3,000 m, 14-15 May 1988 (fl), H. 0llgaard, J. E. Madsen & L. Christensen 74116 (holotype, QCA; isotype, AAU). Figure 1A-D. Culmica. 1-1.5 cm diametro, 1 .5-3(6-8) m alti; erecti ad basim, interdum arcuati ad apicem. Folia culmorum 18-32 cm longa, fragilia, glabra; vaginae 17-30 cm longae, (7-)10-20-plo longiores quam lamina; laminae 1.2-2. 7 cm longae. Gemma centralis rotundata. Rarni- ficatio infravaginalis; rami subsidiarii cujusquisque nodi 30-60. Folia cujusquisque complement! 3-4; laminae 7.8-20 cm longae, 0.4-0.85 cm latae, ratio long.rlat. = (1 1 — )1 7.5—35, apice setosae, basi rotundato-attenu- atae. Paniculae (5 )7 — 1 6 cm longae, angustae. Spiculae (4.6-)5. 1-6.5 mm longae, falcatae, nitidae. Glumae 2, squamiformes. Lemmata sterilia 2, mucronata. Lemma fertile 4. 3-5. 7 mm longum, mucronatum. Culms ca. 1-1.5 cm diam., 1 .5— 3(6— 8) m tall, erect at base, scandent to arching above. Internodes terete, usually glabrous, sometimes scabrous below the nodes. Culm leaves 18-32 cm long, just sur¬ passing the next node, abaxially glabrous, brittle, eventually deciduous, juncture of sheath and blade abaxially indistinguishable or nearly so; sheaths 17- 30 cm long, (7-) 10-20 times as long as the blade, the overlapping margin ciliate, not fused at the base; blades 1 .2-2.7 cm long, triangular, erect, persistent, adaxially pubescent, apex subulate; girdle 1-3 mm Novon 3: 228-238. 1993. wide, glabrous; inner ligule ca. 0.5 mm long, cili- olate, pubescent. Nodes at mid-culm with the cir¬ cular central bud subtended by numerous smaller subsidiary buds in a constellate arrangement; sheath scar dipping below the bud/branch complement; supranodal ridge a visible but not prominent line. Branching infravaginal; central branch sometimes developing; leafy subsidiary branches 30-60 per node, 11-27 cm long, sometimes rebranching from the base. Foliage leaves 3-4 per complement; sheaths glabrous, the overlapping margin ciliate, a tuft of cilia present on each side of the sheath apex, cilia ca. 1 mm long; blades 7.8-20 cm long, 0.4-0.85 cm wide, L:W = ( 1 1 -)1 7.5-35, adaxially usually glabrous but sometimes hispid toward the base of the midrib, abaxially usually glabrous, sometimes sparsely pilose, not tessellate or only weakly so, apex setose, base rounded-attenuate; pseudopetiole 1.5- 2 mm long, adaxially hispidulous, abaxially glabrous; outer ligule a short, usually ciliolate rim, 0.3-0. 5 mm long; inner ligule 0. 5-0.6 mm long, truncate, abaxially pubescent. Panicles (5— )7— 1 6 cm long, narrow, the base usually either retained within the subtending sheath or just fully exserted; rachis ± flattened at the base, angular above, glabrous, the edges scabrous, the basal bract 1.5-4 mm long, papery, pubescent, ciliate; branches 2-3 cm long at the base, angular, glabrous to scabrid, the lower branches ascending to appressed, the upper branch¬ es appressed; pedicels 1-4 mm long, angular, gla¬ brous to scabrid. Spikelets (4.6-)5. 1-6.5 mm long, falcate, shiny. Glumes 2, scalelike, obtuse, abaxially sparsely pubescent toward the apex; glume 1 0.4- 0.9 mm long, ca. '/„ the spikelet length, 1 -nerved; glume II 1-1.8 mm long, ca. /b the spikelet length, 1 3-nerved. Sterile lemmas 2, mucronate, glabrous; sterile lemma I 1.6-2. 9 mm long, ca. V2 the spikelet length, 3-nerved; sterile lemma II 2.7-4 mm long, ca. % the spikelet length, 3- or 5-nerved. Fertile lemma 4. 3-5. 7 mm long, mucronate, glabrous, 5- or 7-nerved. Palea 4. 2-5. 3 mm long, subequal to the fertile lemma, bimucronulate, 4-nerved, sulcate only toward the apex, the sulcus scabrous-hispid. Stamens 3; anthers 2.7 mm long. Lodicules 3; an¬ terior pair 2.1 mm long, posterior one 1 .8 mm long, all apically ciliate. Fruit unknown. Volume 3, Number 3 1993 Clark New Species of Chusquea 229 Distribution. Endemic to upper montane forests and heath scrub in southern Ecuador; (2,400 ) 2,800 3,500 m. Chusquea faleata is characterized by brittle culm leaves that just surpass the next higher node, infra- vaginal branching, a circular central bud, somewhat narrow foliage leaf blades, narrow panicles, and shiny, falcate spikelets, for which it is named. It is not clear from the label data of the flowering specimens whether any ol the flowering events represented a gregarious bloom. Chusquea faleata is apparently a narrow endemic from southern Ecuador, a distri¬ bution shared by several additional, as yet unde¬ scribed, species of Chusquea. Paratypes. ECUADOR. Azuay: eastern Cordillera, 4- 6 km N of the village ot Sevilla de Oro, 6 Aug. 1945 (ff). Camp E-4764 (NY). Loja: Parque Nacional Podo- carpus, Cajanuma, Sendero al Mirador, 2,900 in, 1 June 1992, Clark n al. 1105 (AAU, ISC, MO, QGA, QCNE, US); pass on old road between Loja and Saraguro, about 10 km S of Saraguro, 3,250 in, 3 June 1992, Clark et al. 1121 (AAU, ISC, MO, QCA, QCNE, US); carretera Loja-Yangana, desvio al Parque Nacional Podocarpus, E del Nudo de Cajanuma, 2,880-3,000 m, 14 Mar. 1989 (fl), Freire F. 1329 (QCA); Loma del Loro, 6 km S of Saraguro on road to Loja, 3,200 m, 11 Feb. 1985 (A), Hailing & Andersson 21924 (QCA); carretera Yangana- Toledo, 2,800-3,200 m, 28 die. 1988 (fl), Jaramillo 10623 (QCA); Loma del Oro, 2,800-3,250 m, Jaramillo et al. 8794 (QCA); Parque Nacional Podocarpus, Cajan¬ uma, at Casa de Predesur, 2,850 m, 21- 22 Feb. 1985, Laegaard 53611 (AAU); road Vilcabamba Valladolid, at pass, 2,800-2,900 m, 28 Feb. 1 985 (fl), Laegaard 53 754 (.AAU); in pass approximately 10 km S of Saraguro, 3,500 m, 31 Aug. 1985, Laegaard 55157B (AALI); Cordillera del Loro, 50 km N of Loja, just before descending toward Saraguro, along road to radar station, 3,000-3,200 m, 8 May 1987 (fl), van der W erjj & Palacios 9410{Q CNE); 12 km S of Saraguro on the road to Loja, just over the pass in an area called “La Bajada del Piche,” 3,100 m, 29 May 1980, Young 142 (LIS). Loja/Zamora-Chin- chipe: at pass on road from Zamora to Loja, just above the intersection of the old and new highways, 2,800 m, 31 May 1992, Clark et al. 1100 ( AAU, ISC, MO, QCA, QCNE, US); carretera Loja El Paso-Zarnora, El Paso, 2,800 m, 15 Mar. 1989 (fl), Freire F. 1365 (QCA). Zamora— Chinehipe: road Vilcabamba Valladolid, Km 5 S of provincial border, 2,400 m, 28 Feb. 1985, Lae¬ gaard 53747 (AAU); along road Loja-Zamora, SW of the pass, 2,800 m, 2 Sep. 1985, Laegaard 55173.4 & B (AAU). Chusquea subulata L. G. Clark, sp. nov. TYPE: Ecuador. Pichincha: on the road N from Cal- acali to San Jose de Niebli, 2,750 m, 2 May 1980 (fl), S. M. Young 124 (holotype, QCA; isotypes, NY, US). Figure 1E-H. Culmi (4-)6-8(-10) cm diametro, (3-)7-10 m alti, erecti ad basim, interdum arcuati ad apicem. Folia cul- morum 23-47.5 cm longa; vaginae 19-45 cm longae, (5-)10-22.6-plo longiores quam lamina, abaxialiter pu- bescentes ad basim; laminae 1 .3-4 cm longae. Gemma centralis triangularis. Ramificatio extravaginalis; rami sub¬ sidiary cujusquisque nodi 16-30. Folia cujusquisque com- plementi 4-5; laminae 11-23 cm longae, (0.9-)l .3-2 cm latae, ratio long. : lat. = 8.4-17.4, apice setosae, basi rotundatae vel rotundato-attenuatae. Paniculae (7.5 )1 0 26 cm longae, angustae. Spiculae (7.2-)8 TO mm longae, pubescentes. Glumae 2, squamiformes. Lemmata sterilia 2, subulata. Lemma fertile (6.6-)7.2-8.6 mm longum, subulatum. Culms (4— )6— 8(— 1 0) cm diam., (3— )7— 1 0 m tall, erect at the base, arching toward the apex. Inter¬ nodes ± deeply suleate above the bud/branch com¬ plement, glabrous or pubescent. Culm leaves 23 47.5 cm long, the juncture ol the sheath and blade abaxially obscure; sheath 19-45 cm long, (5)10 22.6 times as long as the blades, abaxially usually pubescent toward the base and otherwise glabrous, rarely completely hispid-pubescent, the overlapping margin ciliate, margins not lused at the base; blades 1.3-4 cm long, triangular, erect, persistent, adax- ially pubescent, adaxially usually glabrous, rarely hispid-pubescent, the apex subulate or subulate-se¬ tose; girdle ca. 3 mm wide, glabrous; inner ligule 1-3 mm long, ciliate, not extending completely to either margin. Nodes with one triangular central bud subtended by several to numerous smaller sub¬ sidiary buds in a constellate arrangement; sheath scar dipping slightly below the bud/branch comple¬ ment. Branching extravaginal; central branch some¬ times developing at the upper nodes; leafy subsidiary branches 16—30 per node, 30-50 cm long, some¬ times basally rebranching. Foliage leaves 4-5 per complement; sheaths glabrous, rarely hispid-pubes¬ cent between the nerves, the overlapping margin ciliate; blades 1 1-23 cm long, (0.9 ) 1 .3-2 cm wide, L: W = 8.4-17.4, adaxially glabrous, abaxially gla¬ brous or pilose, not tessellate to weakly so, the apex setose, the base rounded to rounded-attenuate; pseu¬ dopetiole 2-5 mm long, glabrous; outer ligule 0.7- 1 (—2) mm long, ciliolate or glabrous; inner ligule 1 - 4 mm long, usually rounded, abaxially pubescent. Panicles (7. 5-) 10-26 cm long, the base often re¬ tained within the subtending sheath; rachis ± com- planate below, becoming angular toward the apex, glabrous, scabrid, or glabrous at the base and sca¬ brous-pubescent toward the apex; branches ap- pressed to ascending, angular, scabrous to short pubescent, the lower ones 3 8 cm long; pedicels 1 3(— 4) mm long, angular, scabrous. Spikelets (7.2 )8 10 mm long, linear. Glumes 2, scalelike, acute or obtuse, less than Vxo the spikelet length, abaxially pubescent; glume I 0.6 0.8(-l) mm long, usually 1-nerved; glume II 0.8-l(-1.4) mm long, 1-, 2-, or 3-nerved. Sterile lemmas 2, subulate, abaxially 230 Novon Figure 1. Chusquea falcata L. G. Clark and Chusquea subulata L. G. Clark. A-D. Chusquea falcata (A, B, Clark et al. 1100; C, Camp E-4764; D, Jaramillo 10623). — A. Young node and internode showing the culm leaf, emerging subsidiary branches, and infravaginal branching. — B. Mature branch complement with a developed central Volume 3, Number 3 1993 Clark New Species of Chusquea 231 pubescent; sterile lemma I ca. V, the spikelet length, (3.1-)3.8 4.6 mm long, 3- or 5-nerved; sterile lem¬ ma II ca. % the spikelet length, 5. 8-7. 7 mm long, 5-nerved. Fertile lemma (6.6-)7.2-8.6 mm long, subulate, abaxially pubescent toward the apex, 7- or 9-nerved. Palea 5. 5-6. 9 mm long, bimucronu- late, ciliate at apex, sulcate only toward the apex, 4- or 6-nerved. Lodicules 3; anterior pair 1.2- 1.5 mm long, posterior one 0.8 1.3 mm long, all api- cally ciliate. Stamens 3; anthers 3.8 5 mm long. Caryopsis 3.5 mm long, dark reddish brown; hilum linear; embryo 0.6 mm long. Distribution . Often on steep slopes in upper mon¬ tane forest in central Ecuador with two disjunct populations in the Central Cordillera of Colombia; 2,260-2,800 m. Distinguished by its strongly subulate sterile and fertile lemmas, for which it is named, Chusquea subulata is also characterized by culms usually ca. 6 8 cm in diameter and ca. 7-10 in tall, extra- vaginal branching, foliage leaf blades 11-23 cm long and more or less ovate toward the base with setose apices, narrow panicles with the branches and pedicels appressed, and spikelets (7 . 2— )8— 10 mm long. This species is similar in many respects to the common Chusquea scan dens Kunth, and the two species are likely closely related. However, C. scan- dens is easily distinguished by its open panicles with spreading branches and pedicels, and spikelets 5.5- 7.7( 8.6) mm long with only mucronate sterile and fertile lemmas. Vegetatively, C. scandens is less robust, reaching only 1-2.5 cm in diameter and 2 6( 8) m in height. The foliage leaf blades ol C. scandens are 6- 17 cm long, and more or less lan¬ ceolate, and the apex is at most short setose (Clark, unpublished data). Except for the two disjunct populations in the Central Cordillera of Colombia, Chusquea subulata is restricted to central Ecuador. Ibis distribution pattern is analogous to that of C. albilanata E. G. Clark & Londono, which also has most of its pop¬ ulations in central Ecuador, with one disjunct pop¬ ulation in the Eastern Cordillera of Colombia (Clark & Londono, 1991). The two species are not closely related, and at present, no other species of Chusquea with this distribution pattern are known. According to label data, most of the Ecuadorian populations of this species underwent a mass flow¬ ering from 1979 to 1980. One of the Colombian populations ( Clark et al. 26 I) flowered gregariously at about this time, although the other population (Clark & Londono 379) flowered out of synchrony in 1988. The only other flowering specimen was collected in 1949 (Acosta-Solis 14459), which sug¬ gests a flowering cycle of about 30 years, but mass flowering of the Imbabura population was not con¬ firmed for the 1979 1980 flowering event. Cary- opses are known only from Clark et al. 261, and it is not known if the Ecuadorian populations re¬ generated from seeds or rhizomes. Paratypes. COLOMBIA. Quindio: 13 km above Sal- ento on the road to La Ceja, 2,860 in, 28 Jan. 1988 (fl), Clark & Londono 379 (COL, ISC. TULA, US). Tolima: Cerro Bravo, road from Manizales to Bogota, not far from the Caldas border, 2,750 m, 12 Feb. 1982 (fl), Clark et al. 261 (COL, ISC, US). ECUADOR. Imbabura : area Macho-Loma, seccion Shanshipamba, 2,500 m, 17 Nov. 1949(A), Acosta-Solis 14459( US). Pichincha: Reserva Geobotanica Pululahua, camino a Lulumbamba, 1,800- 3,356 in, 3 Nov. 1987 (fl), Ceron M. 2567 (ISC); road descending to the crater of Pululahua, about 7 km from the road to Calacali and 5 km from the entrance to the Reserva Geobotanica Pululahua, 2,800 rn, 26 Aug. 1992 (fl), Clark et al. 307 (ISC, QCA, US); old Quito-Santo Domingo road, E of Chiriboga, western slope, 2,560 m, 27 Aug. 1982 (fl), Clark et al. 315 (ISC, QCA, US); along road between Nono and Nanegal, between Nono and Tandayapa, 8.8 km before Tandayapa, 2,660 m, 16 Dec. 1979 (fl), Croat 49333 (MO, US); pass between Otavalo and San Jose de las Minas, 3,000 m, 6 Apr. 1979, Holm-Nielsen 16798 (QCA); crater of Pululahua, 100 m from the old road, 2,600 m, 5 Apr. 1979 (fl), Holm-Nielsen 17058 (AAU, QCA); Lloa valley, 2,600 in, 18 May 1980 (fl), Holm-Nielsen 23521 & 23554 (AAU); growing along the Saloya river at Km 34.5 from Quito and also just above Guarumal bridge, 21 Aug. 1945, McClure 21412 (US); Quito, in the Parque Alameda, where it is planted in the border of shrubs around the lake, 2,800 m, 23 Oct. 1945, McClure 21427 (ISC, US); 24 km E of Tandapi on the road from Santo Domingo to Quito, 8,100 ft., 3 Aug. 1980 (fl), II underlin et al. 8709 (AAU, MO — 1 sheet); 10 km NW of Nono on road to Nanegal, steep slope next to Alambi River in cloud forest, 2,410 m, 1 1 Jan. 1980, Young 52 (QCA, US); 17 km W of Lloa, 27 Jan. 1980 (fl), )oung 58 (QCA, US); 9 km W of the town of Nono on the road to Na- negalito, 2,410 m, 15 June 1980 (fl), Young 173 (NY, QCA, US); 67 km E of Santo Domingo on the new road to Quito, 2,260 m, 15 June 1980 (fl). Young 174 (QCA, US). branch and a foliage leaf complement. — C. Inflorescence. — D. Spikelet. E-H. Chusquea subulata (E, Ceron 2567; F, G, Young 124; H, Young 173). — E. Inflorescence with a subtending foliage leaf. — F. Branch complement showing extravaginal branching. — G. Spikelet. — H. Culm leaf, abaxial view. 232 Novon Chusquea barbala L. G. Clark, sp. nov. TYPE: Peru. Pasco: Prov. Oxapampa, Serrama de San Matias, W slope, E of Loma Linda, 400-700 m, 15 June 1983 (fl), A. Gentry, D. Smith & N. Jaramillo 42007 (holotype, MO; isotypes, ISC, US). Figure 2. Culmi ignoti. Gemma centralis triangularis. Ramificatio extravaginalis; rami subsidiarii cujusquisque nodi 20 25. Laminae foliorum 7.7-1 1.7 cm longae, 0.6-0. 9 cm latae, ratio long. Oat. = 11.8-13.8, apice acuminato-mucron- atae, basi rotundato-attenuatae. Inflorescentiae constantes ex 1 (—3) spiculis, per 4-6 bracteas subtentis; bracteae curvinerves. Spiculae 9.1-12.1 mm longae. Glumae valde redactae, cupuliformes. Lemmata sterilia 2, scabrida, mu- cronata, curvinervia. Lemma fertile 9.7-11.1 mm lon- gum, subulatum, barbatum ad medium, pili ca. 1 mm longi, vitrei, rigidi, cinnamomei. Culms unknown, described as a vine on the label. Internodes 2 mm diam., ca. 24 cm long, terete. Culm leaves unknown. Nodes with a ± triangular central bud subtended by ca. 20-25 subsidiary branches; nodal line dipping somewhat below the branch complement; supranodal ridge ± prominent; nodal region 4-6 mm wide, relatively large. Branch¬ ing extravaginal; the more robust subsidiary branch¬ es 8-12 per node, 35-70 cm long, whiplike, bearing foliage leaves and/or spikelet clusters, the slender, short subsidiary branchlets terminating in spikelets. Foliage leaf sheaths scabrid, the overlapping margin ciliate, the cilia glassy, reddish gold, following the curve of the internode; blades 7.7-11.7 cm long, 0.6-0. 9 cm wide, L:W = 11.8-13.8, adaxially and abaxially glabrous, but adaxially one primary nerve next to the midrib prominent and scabrous for nearly the full length, not tessellate, apex acu- minate-rnucronate, base rounded-attenuate, one margin glabrous, the other minutely serrulate; pseu¬ dopetiole 2 mm long, hispidulous; outer ligule 0.7- l mm long, ciliolate, ahaxially hispid-pubescent on the upper 0.6-0. 8 mm, this part also usually a darker brown; inner ligule 1-1.5 mm long, rounded, abaxially pubescent. Inflorescence usually consisting of a solitary spikelet terminating a short, slender subsidiary branch, sometimes the branch bearing one or two additional spikelets, the base of the branch covered by the 4-6 small bracts subtending each spikelet; bracts 0.8-3. 6 mm long, acute to rnucro- nate, the lateral nerves curving strongly toward the apex, often almost connecting with the adjacent nerve, apparently lacking buds in their axils; inter¬ node (pedicel) between the uppermost subtending bract and the spikelet 0.5-1 mm long, glabrous, somewhat ridged. Spikelets 9.1-12.1 mm long, ± laterally compressed. Glumes 2?, minute, appar¬ ently present as a glabrous cupule 0.1 -0.3 mm long. Sterile lemmas 2, scabrid, mucronate, 7- or 9-nerved, the lateral nerves curving strongly toward the apex; sterile lemma I 4. 1-5.7 mm long, ca. 14 the spikelet length; sterile lemma II 5.7-8 mm long, ca. % the spikelet length. Fertile lemma 9.7-11.1 mm long, subulate, scabrid, 7- or 9-nerved, the lower '/ heavily bearded except along the keel, the hairs ca. 1 mm long, glassy, stiff, reddish gold or dark reddish gold. Palea 8.5-9 mm long, shorter than the fertile lem¬ ma, bimucronulate, scabrid, 4- or 6-nerved, weakly 2-keeled, the sulcus with a few scattered glassy hairs. Stamens 3, anthers 3. 7-4. 4 mm. Lodicules 3; an¬ terior pair 2.4 mm long, basally strongly swollen, apically ciliate, the posterior one 1 .8 mm, apparently with one filament adnate on each margin for about ‘4 the length. Fruit unknown. This unusual species is known only from the type specimen, and no habitat information is available. In addition to the heavily bearded fertile lemmas, for which the species is named, C. barbata possesses two other synapomorphies: the inflorescences of one to three spikelets, and the strongly curving lateral nerves of the bracts and sterile lemmas. The ap¬ parent fusion of two of the filaments to the posterior lodicule is also unique, but given the advanced con¬ dition of most of the spikelets on the available spec¬ imens, it is not possible to confirm the presence of this feature. In this specimen, there are both more robust, well-developed subsidiary branches and slen¬ derer, short subsidiary branches that terminate di¬ rectly in spikelets (i.e., the “inflorescences”), but it is not known if the two sizes of subsidiary branches are produced during normal vegetative growth. Correct interpretation of the spikelets and inflo¬ rescences of this species is complicated by the ap¬ parent reduction in these structures. The single fer¬ tile floret and two sterile lemmas are readily recognizable, and disarticulate together as in other species of Chusquea (Fig. 2C, arrow). Based on this homology, the small cupule that remains at the apex of the flowering axis is assumed to represent the two reduced glumes. Extreme reduction or loss of the glumes is known in other species of Chusquea, such as C. fendleri Munro, C. coronalis Soderstrom & C. Calderon, and C. depauperata Pilger (Clark, 1989). The internode between the subtending bracts and the cupule is the pedicel. If we accept the structure described above as the spikelet, then the spikelet and the branch it termi¬ nates constitute the inflorescence. The four to six small bracts at the base of the branch, subtending the spikelet(s), most likely represent either the cat- aphylls that normally appear at the base of a sub¬ sidiary branch in Chusquea, in which case the rest Volume 3, Number 3 1993 Clark New Species of Chusquea 233 234 Novon Figure 3. Chusquea spathacea McClure ex L. G. Clark and Chusquea pulchella L. G. Clark. A E. Chusquea spathacea (A, B, Haught 4512; C-E, Clark & Londoilo 588). — A. Inflorescence showing two subtending spathelike bracts (arrows). — B. Spikelet. — C. Foliage leaf. — D. Bud complement. — E. Node and internode showing the culm leaf, dimorphic subsidiary branches, and infravaginal branching. F-J. Chusquea pulchella (F- H, Clark & Windisch Volume 3, Number 3 1993 Clark New Species of Chusquea 235 of the inflorescence is totally lost, or they may rep¬ resent the retention of bracts at the base oi the inflorescence branches, with concomitant loss of the branches themselves and the peduncle. In the rare cases where one or two additional spikelets are pres¬ ent on a branch, the base of the branch is clothed by two of these bracts, then one or two short spikelet- bearing axes appear in close succession, and then the terminal spikelet subtended by three or four of the bracts is observed. The lateral spikelet(s) is also subtended by three or four bracts. This suggests extreme condensation of all the axes associated with the inflorescence, including the branch upon which it is normally borne. It must be noted that the specimens upon which these observations are based probably represent higher orders of branching, and that the subsidiary branches of the main culm might produce somewhat more elaborated inflorescences. Observations of inflorescence production at higher orders of branching in C. capituliflora Trinius and other species, however, indicate that inflorescences produced by the subsidiary branches of the main culm are reiterated more or less faithfully at higher orders of branching. The possibly plesiomorphous retention of reduced inflorescence bracts is known in Chusquea sect. Longiprophyllae L. G. Clark (Clark, 1990) and the Rettbergia group within Chusquea (Clark, unpub¬ lished data), although they are not usually as well developed as in C. barbata. The racemose inflores¬ cences of C. simplici flora Munro consist of three to four spikelets (Clark, 1989), and may be inter¬ preted as reduction from the more typically panic¬ ulate inflorescences of Chusquea. Short, sparsely branched paniculate inflorescences with a series of subtending, bractlike leaves are found in C. tenella Nees and C. ramosissima Lindman (Clark, unpub¬ lished data). At present, it seems most parsimonious to assume that the one-spikeleted inflorescences of C. barbata represent reduction from the type of inflorescences found in these species, with the re¬ tention of at least some inflorescence bracts. The vining habit and morphology of the bud/ branch complement of (.. barbata suggest an affinity to the Rettbergia group. In particular, the shape of the central bud and the dimorphic subsidiary branches are very similar to those of C. ramosis¬ sima , although that species has infravaginal branch¬ ing. Although no other species of Chusquea exhibit the heavily bearded fertile lemmas of C. barbata , short, stiff, glassy hairs on the fertile lemmas (and sometimes also on the sterile lemmas and glumes) are found in a number of species of the Rettbergia group, including C. capituliflora, C. capitata Nees, and C. urelytra Hackel. Until additional material becomes available, and this species is studied in more detail, C. barbata is tentatively allied with the Rett¬ bergia group of Chusquea. Chusquea spathacea McClure ex L. G. Clark, sp. nov. TYPE: Colombia. Magdalena: forest on trail above “Africa” (Sierra Perija), 1,700 m, 16 Dec. 1944 (fl), 0. Naught 4512 (holotype, COL; isotype, US). Figure 3A-E. Culmi ca. 1 mm diametro, 2-6 m longi, erecti ad basim, interduin scandentes. Folia culmorum 9.8-19.2 cm longa, glabra, leviter glauca juventute; vaginae 4.5-11 cm lon- gae, 0.85-1.6-plo longiores quam lamina; laminae 4-9 cm longae, cordatae. Gemma centralis triangularis, gem¬ mae subsidiariae dimorphae. Ramificatio infravaginalis; rami subsidiarii robusti cujusquisque nodi 2-3; rami subsidiarii parviores cujusquisque nodi 16-30. Folia cujusquisque complementi 8-12; vaginae plerumque maculatae; lam¬ inae 4.6-8 cm longae, 0.8- 1.2 cm latae, ratio long. Tat. = 4.7-8, apice breviter setosae, basi rotundatae. Pani- culae 2-4 cm longae, secundae, subcontractae, subtentae 2 bracteis spathiformibus. Spiculae 8.8-10.3 mm longae, glabrae. Glumae 2, cupuliformes. Lemmata sterilia 2, mucronata, ad mediarn spiculam attingentes. Lemma fer¬ tile 7. 8-9. 4 mm longum, mucronatum. Culms ca. 1 cm diam., 2 6 m long, erect at the base, scandent and climbing above, sometimes vin¬ ing. Internodes 13.5-15 cm long, ± terete to shal¬ lowly sulcate above the bud/branch complement, glabrous, mottled purple and green, lightly glaucous especially when young. Culm leaves 9.8-19.2 cm long, glabrous, exposed surfaces lightly glaucous when young, juncture of sheath and blade an in¬ verted “V”; sheaths 4.5-11 cm long, 0.85-1.6 times as long as the blades, ± triangular; blades 4- 9 cm long, triangular, erect, persistent, base cordate, the margins somewhat ruffled near the base, apex short setose; girdle 0.2-0. 5 cm wide, glabrous, a short skirt present at the juncture of the sheath and girdle; inner ligule 3-4 mm long, slightly irregular, apically ciliolate. Nodes with one triangular central bud subtended by 20-25 smaller subsidiary buds and 2-3 larger subsidiary buds in a constellate ar¬ rangement; sheath scar dipping slightly below the bud/branch complement; supranodal ridge present but not prominent. Branching infravaginal; central 726; I, Davidse et al. 10950; J, Clark & Oliveira 939). — F. Branch complement with culm leaf. — G. Foliage leaf complement. — H. Ligular area of foliage leaf. — I. Inflorescence. — J. Spikelet. 236 Novon branch often developing; robust subsidiary branches 2-3 per node, 27-50 cm long, whiplike, rebranch¬ ing, smaller leafy subsidiary branches 16-30 per node, 15-22 cm long, occasionally rebranching, divergent. Foliage leaves 8-12 per complement; sheaths glabrous, often mottled, a few cilia present at the apex of the overlapping margin; blades 4.6- 8 cm long, 0.8- 1 .2 cm wide, L : W = 4.7-8, adax- ially glabrous, abaxially glabrous with a tuft of hairs at the base, not tessellate or weakly so, base rounded, apex short setose, one margin glabrous, the other ciliate; pseudopetiole ca. 1 mm long, glabrous; outer ligule 0.2-0. 4 mm long; inner ligule 1-1.5 mm long, truncate to rounded. Panicles 2-4 cm long, secund, ± contracted, base retained within the subtending sheath; subtended by usually 2 spatheate bracts, the sheaths 2. 4-3. 2 cm long, expanded, ± stramineous to translucent at the margins, sometimes slightly mottled, blades 1.3-3. 8 cm long, green; rachis tri¬ quetrous, glabrous to scabrous-pilose, the edges sca¬ brous; branches secund, angular, scabrous-pubes¬ cent, most with a well-developed pulvinus at their bases, spreading, the lower branches 1-1.5 cm long; pedicels 1-2 mm long, angular, scabrous-pubescent, spreading. Spikelets 8.8 10.3 mm long, glabrous, terete or dorsally compressed. Glumes 2, 0.1 -0.3 mm long, scalelike, minute, less than j/(0 the spikelet length. Sterile lemmas 2, ca. V2 the spikelet length, mucronate, 3-nerved; sterile lemma 1 3-4.6 mm long; sterile lemma II 3.6-5. 1 mm long. Fertile lemma 7. 8-9. 4 mm long, mucronate, apically cili¬ ate, 7- or 9-nerved. Palea 8. 2-9. 6 mm long, bi- mucronulate, 4- or 6-nerved, sulcate for nearly the full length, the sulcus adaxially pubescent toward the apex. Lodicules 3, apically ciliate; anterior pair 2 mm long, basally swollen, the posterior one 1.5 mm long. Stamens 3; anthers 4. 6-5. 2 mm long. Fruit unknown. Distribution. Montane forests on the Colombian side of the Sierra de Perija; 1,700-2,400 m. Chusquea spathacea is named for the well-de¬ veloped, spathelike bracts that subtend the inflores¬ cences. From notes deposited at the U. S. National Herbarium, it is clear that F. A. McClure many years ago recognized this as a distinct species and chose the epithet “ spathacea" for it, although he never formally described the species. Other features that characterize C. spathacea include culm leaves with the sheath and blade more or less equal in length and the blade cordate; dimorphic subsidiary buds/branches; infravaginal branching; foliage leaf blades abaxially with a tuft of hairs at the base; contracted panicles; and glabrous spikelets 8.8 10.3 mm long with minute, cupulelike glumes. The somewhat vining habit, infravaginal branch¬ ing, and the contracted panicle subtended by spathe¬ like bracts (Fig. 3A, arrows) of C. spathacea clearly indicate an affinity with the Brazilian Rettbergia group of Chusquea. The presence of 1-4 spathelike bracts subtending the inflorescence appears to be a synapomorphy for the Rettbergia group (Clark, un¬ published data), and thus C. spathacea is placed here. Chusquea pallida Munro, from northwestern Colombia and Venezuela, also shares these features and therefore is also classified as a member of the Rettbergia group. Paratypes. COLOMBIA. Cesar: Mpio. Manaure, Ser- rania de Perija, via Manaure-Sabana Rubia, near the Venezuelan border, between Finca Inglaterra and El Cin- co, 2,200 m, 24 July 1989, Clark & Londoho 588 (COL, ISC, K, MO, NY, TULV, US). Magdalena: Sierra de Perija, 8 km ENE of Manaure, 44 km E of Valledupar, 5 km from the Venezuelan border, 2,375 m, 2 Feb. 1945 (fl), Grant 10772 (ISC, US, WIS). Chusquea pulchella L. C. Clark, sp. nov. TYPE: Brazil. Sao Paulo: BR-1 16, ca. Km 517, head¬ ing N to Sao Paulo, Rio Bra^o Feio, ca. 60 m before the spring, 560 m, 48°19'W, 24°57'S, 19 Mar. 1991 (fl), L. Clark & W. Oliveira 939 (holotype, SP; isotypes, ISC, MO, SJRP, US). Figure 3F J. Culmi 2-3 mm in diametro, 2-3 m alti, scandentes. Folia culmorum 10-12 cm longa, persistentia; vaginae 9.8-11.2 cm longae, 9.8-12.4-plo longiores quam lam¬ ina; laminae 0.90 cm longae, triangulares. Ramificatio infravaginalis; ramus centralis evolutus; rami subsidiarii cujusquisque nodi 20-36, 13-40 cm longi. Folia cujus- quisque complementi 7-10; vaginae pilosae ad apicem; laminae 3.5-7. 1 cm longae, 0.25-0.5 cm latae, ratio long. :lat. = 1 1.7— 14(— 24), apice acuminatae, basi at- tenuatae vel rotundato-attenuatae. Paniculae ca. 2 cm longae, subapertae. Spiculae 5. 5-5. 8 mm longae. Glumae 2, acutae. Lemmata sterilia 2, acutae vel acuto-mucron- atae, inaequales. Lemma fertile 4.7-5 mm longum, ob- tusum. Culms 2-3 mm diam., 2-3 m tall, climbing, scan- dent, often intertwined with other vegetation. In¬ ternodes terete, 9.5-15 cm long, glabrous. Culm leaves 10-12 cm long, persistent, juncture of sheath and blade abaxially obscure; sheaths 9.8-1 1.2 cm long, 9.8-12.4 times as long as the blade, abaxially retrorsely scabrous, the overlapping margin ciliate, not fused at the base; blades 0.9-1 cm long, tri¬ angular, erect, persistent, abaxially glabrous, apex subulate; girdle ca. 1 mm wide, pilose; inner ligule not seen. Nodes at mid-culm with the sheath scar ± horizontal but dipping strongly below the bud/ branch complement; supranodal ridge obscure. Branching infravaginal; central branch always de- Volume 3, Number 3 1993 Clark New Species of Chusquea 237 veloped, as robust as the main culm; leal subsidiary branches constellate, 20-36 per node, 13-40 cm long, geniculate, sometimes rebranching from the basal nodes. Foliage leaves 7-10 per complement; sheaths pilose toward the apex, olten fully pilose at lower branch nodes, a tuft of cilia present on one side at the apex, the overlapping margin ciliate to¬ ward the apex; blades 3.5-7. 1 cm long, 0.25 0.5 cm wide, I, : W = 1 1.7-1 4( 24), adaxiallv glabrous, abaxially tomentose, a tuft of hairs present at the base, not tessellate, apex acuminate, base attenuate to rounded-attenuate, the margins serrulate at the base, becoming nearly ciliate at the apex; pseudo¬ petiole ca. 1 mm long, pilose; outer ligule 0.1 -0.2 mm long, glabrous; inner ligule 0.3-0. 5 mm long, truncate. Panicles ca. 2 cm long, ± open, just hilly exserted from the subtending sheath; rachis ± flat¬ tened, hispid-pubescent on the first internode, gla¬ brous above; branches 0.5 cm long at the base, angular, glabrous, ascending, usually subtended by a small bract 0.1 -1.3 mm long; pedicels 1 2 mm long, angular, glabrous, often subtended by a minute bract. Spikelets 5.5 5.8 mm long. Glumes 2, acute, abaxially sparsely pubescent; glume I 0.9- 1.1 mm long, ca. Vl0 the spikelet length, 1 -nerved; glume II 1.2-1. 7 mm long, ca. the spikelet length, 1- or 3-nerved. Sterile lemmas 2, acute to acute-mucro- nate; sterile lemma I 2. 5-2. 7 mm long, ca. /, the spikelet length, sparsely pubescent on the lower half, 3-nerved; sterile lemma II 4. 5-5. 2 mm long, nearly as long as the spikelet, sparsely pubescent, 5-nerved. Fertile lemma 4.7-5 mm long, blunt, sparsely pu¬ bescent, 7-nerved. Palea 5-5.5 mm long, usually overtopping the fertile lemma, blunt, 4- or 6-nerved, sulcate only at the apex, the sulcus scabrous-pu¬ bescent. Stamens 3? (spikelets too old to determine the number with certainty); anthers 2.6 mm long. Lodicules 3; anterior pair 1 .4 mm long, posterior one 0.9- 1.2 mm long. Fruit unknown. Distribution. Apparently endemic to southeast¬ ern Brazil in the state of Sao Paulo along the Rio Brago Feio in Atlantic forest and secondary forest; 530-680 m. Chusquea pulchella is a delicate, somewhat viny species characterized by culms 2-3 mm in diameter; 20-36 subsidiary branches per node; 7-10 foliage leaves per complement with pilose sheaths and blades 0.25-0.5 cm wide; spikelets 5. 5-5. 8 mm long; and unequal sterile lemmas with sterile lemma 1 about V, the spikelet length, and sterile lemma 11 nearly as long as the spikelet. Although this species lacks any spathelike bracts subtending the inflorescence, it is clearly allied with other species of the Rett- bergia group within Chusquea based on its viny habit, infravaginal branching, and spikelet mor¬ phology. This species is very similar to C. sellowii Ru- precht and C. oligophylla Ruprecht (= C. discolor Hackel) in size and habit, and approaches C. oli¬ gophylla in its nearly ciliate leaf margins. Chusquea pulchella can be distinguished easily from these two species, however, by its narrow leaves, 0.25 0.5 cm wide as opposed to 0.8 1.6 cm wide m C. sellowii and usually 0.5- 1.1 cm wide in C. oligo¬ phylla (Clark, unpublished data). In addition, the sterile lemmas of C. pulchella are strongly unequal, whereas the sterile lemmas in both C. sellowii and C. oligophylla are subequal or only moderately unequal. The spikelet proportions of C. pulchella most closely resemble those of C. bambusoides (Rad- di) Hackel, but the spikelets of C. bambusoides are usually twice as long and the glumes are apically acuminate. Vegetatively, C. bambusoides is much more robust, with larger culms, fewer branches per node, and much larger foliage leaves with the blades ( 1 — ) 1.5—3 cm wade (Clark, unpublished data). A ten-year flowering cycle is suggested by the two flowering collections of C. pulchella, which came from the same population, although a shorter flow¬ ering cycle is possible. Paratypes. BRAZIL. Sao Paulo: BR-116, ca. Kin 517, heading N to Sao Paulo, Rio Brago Feio, ca. 60 m before the spring, 530 in, 4 Mar. 1990, Clark & II in- disch 726 (ISC, MO, SP, SJRP, US); ca. 38 km SW of Jacupiranga along Hwy. 1 16 to Curitiba, 680 m, 8 Mar. 1976 (fl), Davidse et al. 10950 (ISC, MO, US). Chusquea attenuata (Doell in Martins) L. G. Clark, comb. nov. Basionym: Arundinaria attenuata Doell in Martius, FI. Bras. 2(3): 170. 1880. TYPE: Brazil. Minas Gerais: in silvaticis umbr. montis Itacolumi, Aug. 1824, Riedel s.n. (ho- lotype, LE; isotypes, KR, US fragment). McClure (1973: 38) included this species in his list of species excluded from Arundinaria as Arundinaria ? attenuata Doell; species sedis mihi incertae etiarn nunc manet . ” The type specimen is vegetative, which accounts for his confusion about the placement of this species. However, the pho¬ tograph of the LE specimen mounted with the frag¬ ments that constitute the US isotype clearly shows the circular central bud with several subsidiary branches per node that is characteristic of Chus¬ quea. Later collections from Itacolomi ( Chase 9379, Macedo 5108) and Caraga ( Irwin et al. 29242, Clark & Morel 7 14) match the type perfectly. Both Macedo 5 1 08 and Irwin et al. 29242 were collected during the same gregarious flowering event, and the 238 Novon spikelet morphology confirms that this species is indeed a Chusquea, and that the species is distinct. More detailed studies of this species are underway, but C. attenuata appears to be restricted to Minas Gerais. In the original description (I)oell in Martius, 1880: 170), Doell indicated that this species represented “ Arundinaria distans ex parte.” According to a note by Agnes Chase (dated 1936 in her hand) that is attached to the US isotype fragment, the type of Arundinaria distans Trinius consisted of four sheets numbered 1 through 4. Sheets 1 and 2 agree with Trinius’s description of A. distans [= Colanthelia distans (Trinius) McClure], but Mrs. Chase stated that Sheet 4 represents a different species that “agrees perfectly with the description of Arundi¬ naria attenuata Doell and is undoubtedly the type specimen of that species.” No reference was made to Sheet 3. Mrs. Chase also referred to a note made by A. S. Hitchcock during a 1907 visit to St. Pe¬ tersburg. According to this note, there was a loose annotation label by Doell with Sheet 4 identifying this specimen as Arundinaria attenuata Doell. This label apparently was lost subsequently. Acknowledgments. Fieldwork was supported by a grant from the National Geographic Society, Na¬ tional Science Foundation Grant BSR-8906340, and a Foreign Travel Grant from Iowa State University. Ximena Londono in Colombia and Benjamin 011- gaard, Simon Laegaard, and Peggy Stern in Ecuador provided valuable assistance with the fieldwork. I thank the Conzalo Alcivar family of Quito for gra¬ cious hospitality during my stay in Ecuador. I am grateful to Paulo G. Windisch and Walter Oliveira (both of the Universidade Estadual Paulista) for their assistance during my fieldwork in Brazil. Literature Cited Clark, L. G. 1989. Systematics of Chusquea sect. Swal- lenochloa, sect. Verticillatae, sect. Serpentes, and sect. Longifoliae (Poaceae Bambusoideae). Syst. Bot. Monogr. 27: 1-127. - . 1990. Chusquea sect. Longiprophyllae (Po- aceae: Bambusoideae): A new Andean section and new species. Syst. Bot. 15: 617-634. - & X. Londono. 1991. Miscellaneous new taxa of bamboo (Poaceae: Bambuseae) from Colombia, Ecuador and Mexico. Nordic J. Bot. 11: 323-331. Doell, J. C. 1880. Gramineae. III., Bambusaceae. Pp. 161 — 219 + pi. 44-56 in A. W. Eichler (editor), Flora Brasiliensis 2(3). Friederich Fleischer, Leipzig. McClure, F. A. 1973. Genera of bamboos native to the New World (Gramineae: Bambusoideae). Smithson¬ ian Contr. Bot. 9: 1 148. [T. R. Soderstrom, editor.] Realignment of Festuca Subgenus Schedonorus with the Genus Loliurn (Poaceae) Stephen J. Darbyshire Centre for Land and Biological Resources Research, Agriculture Canada, Wm. Saunders Bldg., Central Experimental Farm, Ottawa, Ontario, klA 0C6, Canada ABSTRACT. Evidence from the literature of the close relationship of Festuca suhg. Schedonorus to Lol¬ iurn and its relatively distant relationship to Festuca subg. Festuca is briefly discussed. New combina¬ tions, Loliurn subg. Schedonorus, Loliurn arundi- naceum, Loliurn gigantrurn, Loliurn rnazzettian- urn, and Loliurn pratense, are proposed to reconcile classification with known phylogeny. The relationship between Festuca L. and Loliurn L. (Poaceae) has long been an interesting classifi¬ cation problem. Loliurn is readily distinguished from Festuca by its spicate inflorescence and lateral spike- lets with a single glume. 1'his morphology suggests the spikelike inflorescence structure in the tribe Tri- ticeae Dumortier. where Loliurn, in early treat¬ ments, was often placed. Nevski (1934) was the first to place Loliurn in the Poeae (= Festuceae) rather than the Triticeae. Information from diverse non- morphological sources, such as hybridization and cytology (e.g., Jenkin, 1959), oligosaccharides and water-soluble polysaccharides in caryopses (Mac¬ Leod & McCorquodale, 1958), endosperm structure (Tateoka, 1962), seed protein electrophoresis (Bu- linska-Rodomska & Lester, 1988; Butkute & Kon- arev, 1982), serology (Butkute & Konarev, 1980, 1982; Fairbrothers & Johnson, 1961; Smith, 1969; Watson & Knox, 1976), chloroplast DNA (cpl)NA) restriction endonuclease site patterns (Lehvaslaiho et ah, 1987; Soreng et ah, 1990), cpDNA reasso- ciation (Hilu & Johnson, 1991), and thermal de- naturation of genomic DNA (King & Ingrouille, 1987), has supported the placement of Loliurn in Poeae. Spontaneous hybridization occurs commonly be¬ tween species of Festuca subg. Schedonorus (P. Beauvois) Petermann and chasmogamous species of Loliurn, particularly the type species of the genus, L. perenne L. (Barker & Stace, 1986; Humphries, 1980; Gymer & Whittington, 1973). Hybridization between Loliurn and Festuca subg. Festuca rarely occurs spontaneously and is reported only for Fes¬ tuca rubra L. (Stace, 1975). The latter is, however, also reported to hybridize with species in Festuca subgenera Festuca and Schedonorus as well as I ul- pia C. C. Gmelin (Ainscough et ah. 1 986; Knobloch, 1968; Saint-Yves 1929; Stace, 1975). Hybridiza¬ tion between Festuca (Schedonorus) pratensis Hud¬ son and Loliurn perenne (= X Festulolium loli- aceurn (Hudson) P. V. Fournier) was suspected as early as 1790 (see Jenkin, 1933). Reproductive barriers are not appreciably greater for this hybrid¬ ization than for interspecific hybridization within Festuca subg. Schedonorus or between many spe¬ cies of Loliurn (Jenkin, 1933; Stace, 1975). I he degree of difficulty in producing artificial L. perenne X F. pratensis hybrids and the differential success of reciprocal crosses is similar to interspecific hy¬ bridization reported in various plant genera (e.g., Borrill et ah, 1977; Gymer & Whittington, 1973). Extensive studies of artificial hybrids and their chro¬ mosome behavior (for references, see Jauhar, 1974, 1975; Terrell, 1966), as well as chromosome hand¬ ing (Dawe, 1989; Thomas, 1981), have indicated the homology of chromosomes and their genetic compatibility. In auto-allotriploids and amphidiploids of L. perenne, L. multi/lorurn L., and F. pratensis, synapsis of homeologous chromosomes usually oc¬ curs as multivalents or even bivalents, in spite of the presence of true homologues (Jauhar, 1974, 1975). Diploid crosses of these taxa show high bi¬ valent formation (87- I 00%) with 0.78.3 - 0.896 chi- asmata per chromosome pair (Jauhar, 1975). Such observations have led many taxonomists and cyto¬ geneticists to suggest that Festuca and Loliurn could be combined, in whole (e.g., Jauhar, 1975; Knob¬ loch, 1963; Love, 1963; Stebbins, 1956; Terrell, 1966; Ullmann, 1936) or in part (e.g., Putin, 1956). Amalgamation of Festuca and Loliurn has been proposed as early as 1898 (M'Alpine, 1898), but no valid combinations of Festuca subg. Schedonorus taxa within Loliurn are available. Raspail (1829), however, included F. pratensis (under the name F. elatior L.) and F. loliaceurn Hudson (= L . pratensis x L. perenne) in his examination of Loliurn as L. festuca Raspail and I.. festucoides Raspail, respec¬ tively. Proposed on the basis of morphological and ecological continua observed between these taxa and Novon 3: 239-243. 1993. 240 Novon species of Loliurn, Raspail considered this classifi¬ cation physiological rather than taxonomic. Such continua would be expected between hybridizing taxa and their derivatives. Gross morphology and cyto- logical differences between Festuca and Loliurn have been cited as reasons for not combining the genera (Clayton & Renvoize, 1986; Jenkin, 1933; Malik & Thomas, 1966; Raspail, 1829; Terrell, 1968). Jenkin (1933) acknowledged the close phyloge¬ netic relationship between Loliurn and Festuca subg. Schedonorus and the relatively distant relationship of these taxa to Festuca subg. Festuca. He did not feel, however, that their classification was incumbent on evolutionary relationships. Bulinska-Rodomska & Lester (1988) detected similar relationships in a phenetic analysis of 50 morphological characters when they excluded six that were associated with the spicate inflorescence of Loliurn. Their principal components analysis identified 1 1 characters in the first factor axis that separated the two lineages with the exception of Vulpia alopecuros (Schousboe) Du- mortier. The latter taxon grouped with Loliurn and Festuca subg. Schedonorus in the first axis hut was clearly separated in the second axis. Loliurn and Festuca subg. Schedonorus were not separated by either of the first two components. Analysis of cpDNA restriction site variation (l)ar- byshire & Warwick, 1992; Lehvaslaiho et al., 1987), chromosome synaptic ability (Jauhar, 1975), seed protein electrophoresis (Bulinska-Rodomska & Les¬ ter, 1 988), and certain morphological data sets (Bu¬ linska-Rodomska & Lester, 1988; Stebbins, 1956) suggests congeneric status for Loliurn and Festuca subg. Schedonorus as a lineage distinct from species of Festuca subg. Festuca. This is consistent with other nonmorphological studies (Butkute & Kon- arev, 1980, 1982; Hilu & Johnson, 1991; King & Ingrouille, 1987; MacLeod & McCorquodale, 1958; Smith, 1969; Watson & Knox, 1976). The poly- phyletic nature of Festuca sensu lato and its rela¬ tionship to Loliurn have not always been appreciated in these latter studies, because, for practical reasons, species of Festuca subg. Schedonorus are frequently used as the sole representatives of the large and diverse genus Festuca. Polyphyly in Festuca has been considerably re¬ duced since the concept of Linnaeus. Several groups (including I ulpia (Clayton & Renvoize, 1986) and Hellerochloa Rauschert (Tzvelev, 1989)), more closely related to Festuca subg. Festuca than Fes¬ tuca subg. Schedonorus, are recognized as separate genera in some recent treatments. Consistent with some morphological interpretations (e.g., Macfar- lane & Watson, 1982), cpDNA restriction site data of North American species (l)arbyshire & Warwick, 1992) indicate that Leucopoa Grisebach (excluding Festuca subg. Leucopoa sect. Breviaristatae Kri- votulenko) is also more distantly related to Festuca subg. Festuca than some of these taxa and should be recognized at the generic level. Inclusion of Leu¬ copoa sect. Leucopoa in Festuca would require the inclusion of Festuca subg. Schedonorus, Vulpia, and Loliurn for Festuca to be monophyletic. The recognition of Schedonorus as a distinct ge¬ nus seems unwarranted. Excluding the overempha¬ sized inflorescence characters, it is hardly distin¬ guishable from Loliurn in morphology (Bulinska- Rodomska & Lester, 1 988). The symmetrical chro¬ mosomes and limited polyploidy in the genus Loliurn are plesiomorphic character states and cannot be used to define a monophyletic clade. Furthermore, the derived conditions of polyploidy and asymmet¬ rical chromosomes found in Schedonorus and other subgenera of Festuca are highly homoplastic in grasses and are not irreversible. The reduction of the inflorescence from a panicle to a spike, with concomitant modifications to other reproductive structures, is the only derived morphological or cy- tological feature uniting species of Loliurn and dis¬ tinguishing them from their closest relatives. The trend from panicle to spike or raceme is homoplastic in other taxa of the Poeae and may not be syna- pomorphic in Loliurn. Additionally, cpDNA restric¬ tion site data indicate less divergence of this genome between L. perenne and species of Festuca subg. Schedonorus than between some members of the Festuca ovina L. complex of Festuca subg. Festuca (Darbyshire & Warwick, 1992). The accommodation of Festuca subg. Schedon¬ orus in Loliurn would require only slight change to published generic descriptions. Several morpholog¬ ical features of Festuca subg. Schedonorus are con¬ sistent with its classification in Loliurn. Shared char¬ acters include wide, flat leaf blades with fascicles of sclerenchyma opposite the veins or forming girders between the epidermes at the major veins (plesiom¬ orphic), entirely glabrous ovary apices (homoplastic in Festuca and Loliurn ), caryopses adhering to the paleas (synapomorphic in Loliurn/ Schedonorus, versus free in Festuca), subterminal style insertion (synapomorphic in Schedonorus/ Loliurn), and fal¬ cate leaf blade auricles (synapomorphic in Sche¬ donorus/ Loliurn, which are not found in any related group). The Old World taxa, Festuca subg. Montanae (Hackel) Nyman and Festuca sect. Scariosae Hack- el, are known to be more closely related to Festuca subg. Schedonorus than Festuca subg. Festuca. At Volume 3, Number 3 1993 Darbyshire Realignment of Festuca Subg. Schedonorus 241 least some of the species in these taxa may prove to belong to the clade including Lolium and Festuca subg. Schedonorus as has been suggested by cy- tological studies (Borrill et ah, 1977; Chandrasek- haran & Thomas, 1971). The phylogeny is com¬ plicated by the apparent alloploids, between diploids of Festuca subg. Schedonorus and Festuca sect. Scariosae , seen in polyploid members of the Festuca arundinacea Schreber complex in northwest Africa (Borrill, 1972). Failure of meiotic chromosome pair¬ ing (1.20 bivalents per cell) in crosses between dip¬ loids of Festuca subg. Schedonorus (L . pratensis) and Festuca subg. Montanae (F. drymeja F. K. Mertens & W. D. J. Koch) has been attributed to differences in chromosome size (Morgan et ah, 1986). Even lower pairing frequency is observed between diploids of Festuca subg. Schedonorus (h . praten¬ sis) and Festuca subg. Festuca (F. polesica Zapa- Fowicz), although there is much less of a difference in chromosome size (Morgan et ah, 1986). If the relationships of Festuca subg. Montanae and Fes¬ tuca sect. Scariosae are with the Lolium / Sche¬ donorus rather than the Festuca sensu stricto lin¬ eage, this will not solve the questions as to whether Lolium should be united with, or Schedonorus sep¬ arated from, Festuca. Further work may indicate a need to transfer species of Festuca subg. Montanae and Festuca sect. Scariosae to Lolium or at least to remove them from Festuca. The paniculate inflorescence and the presence of the lower glume are the principal morphological characteristics separating Festuca subg. Schedon¬ orus from Lolium and connecting the former taxon with Festuca. These serve as subgeneric characters in the treatment proposed here. Transferring the four species of Festuca subg. Schedonorus to Lol¬ ium is a much more practical solution to the obvious difficulties of combining the nomenclature of these two complex Linnaean genera under the caveat of monophyletic classification. The name Festuca elatior L. was lectotvpified by Linder (1986), and, although it has priority over F. arundinacea Schreber, has been proposed as a nomina rejicienda under 1CBN Art. 69 by Reveal et al. (1991). The present usage (F. arundinacea) is followed here to avoid nomenclatural confusion. Lolium subg. Schedonorus (P. Beauvois) S. J. Darbyshire, comb. nov. Basionym: Schedono¬ rus P. Beauvois, Ess. Agrostogr.: 99, 177. 1812. Festuca sect. Schedonorus (P. Beauvois) W. I). J. Koch, Syn. FI. Germ. Helv.: 813. 1837. Festuca subg. Schedonorus (P. Beau¬ vois) Petermann, Deutschl. FI.: 643. 1849. TYPE: F. elatior L. (= F. arundinacea Schre¬ ber) (lectotype, selected by Niles & Chase (1925)). Lolium arundinaceum (Schreber) S. J. Dar¬ byshire, comb. nov. Basionym: Festuca arun¬ dinacea Schreber, Spic. FI. Lips.: 57. 1771. Bromus arundinaceus (Schreber) A. W. Roth, Tent. FI. Germ. 2: 141. 1789. Schedonorus arundinaceus (Schreber) Dumortier, Observ. Gramin. Belg.: 106. 1824. Festuca elatior var. arundinacea (Schreber) G. F. 11. W bu¬ rner, FI. Schles. Ed. 3: 59. 1857. Festuca elatior subsp. arundinacea (Schreber) Cela- kovsky, Prodr. F. Bohmen 1:51. 1867. I Y PE: Scheuchzer, Agrostographia, tab. V, fig. 18. 1719 (lectotype, selected by Reveal et al. (1991)). Lolium giganteum (L.) S. J. Darbyshire, comb, nov. Basionym: Bromus giganteus L., Sp. PL: 77. 1753. Festuca gigantea (L.) Villars, Hist. PL Dauph. 2: 110. 1787. Arena gi¬ gantea (L.) R. A. Salisbury, Prodr. Stirp. Chap. Allerton: 23. 1796. Schedonorus gi¬ ganteus (L.) Gaudin ex J. J- Roemer & J. A. Schultes, Syst. Veg. 2: 644. 1817, as “Schenodorus giganteus” pro syn. Bromus giganteus L. Bucetum gigan team (L.) Par¬ nell, Grass. Scotland: 108, pi. 47. 1842. Z.cr- na gigantea (L.) Panzer ex B. D. Jackson, Index Kew. 2: 1249. 1895. Forasaccus gi¬ ganteus (L.) Bubani, FI. Pvren. 4: 383. 1901. TYPE: Europae sylvis siccis. Roemer and Schultes published the name Lolium giganteum (Syst. Veg. 2: 750. 1 8 1 7.) as a synonym of L. maximum Willdenow. This name has some¬ times been cited Lolium giganteum hurt, ex Roemer & Schultes as a synonym of L. temulentum L. (e.g., Ferrell, 1968). According to IGBN Art. 34.1 this name is not validly published and thus the epithet, giganteum, is available for the proposed combina¬ tion. Lolium mazzettianum (E. B. Alexeev) S. J. Dar¬ byshire, comb. nov. Basionym: hestuca maz- zettiana E. B. Alexeev, Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 82(3): 99. 1977, nom. nov. pro Festuca mairei Hackel ex Han- del-Mazzetti, Symb. Sin. 7: 1288. 1936, non Saint-Yves 1922. TYPE: China. Yunnan: talus des Uzuru, 3 June 1908, Ducloux 86 1 (lec¬ totype, selected by Alexeev (1977), W). 242 Novon Lolium pratense (Hudson) S. J. Darbyshire, comb, nov. Basionym: Festuca pratensis Hudson, FI. Angl.: 37. 1762. Festuca fluitans var. pra¬ tensis (Hudson) Hudson, FI. Angl. Ed. 2: 47. 1778. Schedonorus pratensis (Hudson) P. Beauvois, Ess. Agrostogr.: 99, 163, 177. 1812. Bromus pratensis (Hudson) K. P. J. Sprengel, Syst. Veg. 1: 359. 1825, non Lamarck 1785. Lolium festuca Raspail in Saigey & Raspail, Ann. Sci. Observ. 2: 244. 1829, comb, inval. Lolium festuca Raspail ex Mutel, FI. Frang. 4: 111. 1837, pro syn. Festuca pratensis Hud¬ son. Bucetum pratense (Hudson) Parnell, Grass. Scotland: 105, pi. 46. 1842. Festuca elatior var. pratensis (Hudson) A. Gray, Manual Ed. 5: 634. 1867. Festuca elatior subsp. pratensis (Hudson) Hackel, Bot. Centralbl. 8: 407. 1881. Tragus pratensis (Hudson) Panzer ex B. D. Jackson, Index Kew. 2: 1099. 1895. TYPE: Herb. Sloane 125.16 (BM-SL) (lectotype, se¬ lected by Reveal et al. (1991)). Lolium x festueaceum Link, Linnaea 2: 235. 1827. Festuca loliacea Hudson, FI. Angl.: 38. 1762. Festuca fluitans var. loliacea (Hudson) Hudson, FI. Angl. Ed. 2: 47. 1778. Boa loli¬ acea (Hudson) Koeler, Descr. Gram.: 207. 1802. Schedonorus loliaceus (Hudson) P. Beauvois, Ess. Agrostogr.: 99, 163, 177. 1812. Festuca pratensis var. loliacea (Hudson) Saint- Anians, FI. Agen.: 42. 1821. Brachypodium loliaceum Link, Hort. Berol. 1: 42. 1827, comb, inval. Lolium festucoides Raspail in Saigey & Raspail, Ann. Sci. Observ. 2: 243. 1829, comb, inval. Festuca fluitans Leers ex Kunth, Enum. PI. 1: 405. 1833, non L. 1753, pro syn. Fes¬ tuca loliacea Hudson. Lolium festucoides Ras¬ pail ex Mutel, FI. Frang. 4: 112. 1837, pro syn. Festuca loliacea Hudson. Glyceria loli¬ acea (Hudson) Godron, FI. Lorraine 3: 168. 1844. Tragus loliaceus (Hudson) Panzer ex B. D. Jackson, Index Kew. 2: 1099. 1895. x Fes- tulolium loliaceum (Hudson) P. V. Fournier, Quatre FI. France: 81. 1935. TYPE: England. London: near Vauxhall, ad vias et in pascuis sed rarius. The “intergeneric” hybrid x Festulolium loli¬ aceum is deemed a hybrid of congeneric taxa under the classification proposed here. The correct name under this scheme is Lolium x festueaceum Link. Acknowledgments. The comments, whether in agreement or not, of B. R. Baum, J. Cayouette, W. J. Crins, P. D. Newman, E. E. Terrell, S. I. Warwick and, especially, J. McNeill on earlier versions are acknowledged with thanks. J. E. Darbyshire is thanked for assistance with translation. Literature Cited Ainscough, M. M., C. M. Barker & C. A. Stace. 1986. Natural hybrids between Festuca and species of Vul- pia section Vulpia. Watsonia 16: 143-151. Alexeev, E. B. 1977. To the systematics of Asian fes¬ cues (Festuca). I. Subgenera Drymarithele, Subu- latae, Schedonorus, Leucopoa. Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 82(3): 95-102. [In Russian.] Barker, C. M. & C. M. Stace. 1986. Hybridization in the genera Vulpia and Festuca: Meiotic behaviour of artificial hybrids. Nordic J. Bot. 6: 1-10. Borrill, M. 1972. Studies in Festuca. III. The contri¬ bution of F. scariosa to the evolution of polyploids in sections Bovinae and Scariosae. New Phytol. 71: 523-532. - , M. Kirby & G. Morgan. 1977. Studies in Festuca. 1 1 . Interrelationships of some putative dip¬ loid ancestors of the polyploid broad-leaved fescues. New. Phytol. 78: 661-674. Bulinska-Rodomska, Z. & R. N. Lester. 1988. Inter- generic relationships of Lolium, Festuca, and Vulpia (Poaceae) and their phylogeny. PI. Syst. Evol. 159: 217-227. Butkute, B. L. & A. V. Konarev. 1980. An immuno¬ logical study of the seed proteins of the rye-grass in connection with the phylogeny of the genus Lolium. Bot. Zhurn. (Moscow & Leningrad) 65: 1453- 1 458. [In Russian.] - & - . 1982. Studies of the seed proteins in the genera Lolium and Festuca (Poaceae) in con¬ nection with their phylogeny. Bot. Zhurn. (Moscow & Leningrad) 67: 812- 819. [In Russian.] Chandrasekharan, P. & H. Thomas. 1971. Studies in Festuca. 6. Chromosome relationships between Bov¬ inae and Scariosae. Z. Pflanzenziicht. 66: 76-86. Clayton, W. D. & S. A. Renvoize. 1986. Genera Gra- minum. Grasses of the World. Kew Bull., Addit. Ser. 13: 1-389. Darbyshire, S. J. & S. I. Warwick. 1992 [1993]. Phy¬ logeny of North American Festuca (Poaceae) and related genera using chloroplast DNA restriction site variation. Canad. J. Bot. 70: 2415 — 2429. Dawe, J. C. 1989. Sectional survey of Giemsa C-banded karyotypes in Festuca L. (Poaceae: Festuceae): Sys¬ tematic and evolutionary implications. Thesis, Uni¬ versity of Vienna. Fairbrothers, D. E. & M. A. Johnson. 1961. The pre¬ cipitin reaction as an indicator of relationship in some grasses. Recent Advances in Botany 1: 116-120. Gymer, P. T. & W. J. Whittington. 1973. Hybrids between Lolium perenne L. and Festuca pratensis Huds. New Phytol. 72: 411 — 424. Hilu, K. W. & J. L. Johnson. 1991. Chloroplast DNA reassociation and grass phylogeny. PI. Syst. Evol. 176: 21-31. Humphries, C. J. 1980. Lolium L. Pp. 153-154 in T. G. Tutin, V. H. Heywood, N. A. Burges, D. M. Moore, D. H. Valentine, S. M. Walters & D. A. Webb (editors), Flora Europaea, Vol. 5. Cambridge Univ. Press, Cambridge, London, New York, New Rochelle, Melbourne, Sydney. Jauhar, P. P. 1974. Chromosome pairing in some trip- loid and trispecific hybrids in Lolium— Festuca and its phylogenetic implications. Chromosomes Today 5: 165-177. Volume 3, Number 3 1993 Darbyshire Realignment of Festuca Subg. Schedonorus 243 - . 1975. Chromosome relationships between Lolium and Festuca (Gramineae). Chromosoma 52: 103-121. Jenkin, T. J. 1933. Interspecific and intergeneric hy¬ brids in herbage grasses. Initial crosses. J. Genet. 28: 205-264. - . 1959. Fescue species (Festuca L.). Pp. 418- 434 in H. Kappert & W. Rudorf (editors). Manual of Plant Breeding, Vol. 4. Paul Parey, Berlin. King, G. J. & M. J. Ingrouille. 1987. Genome hetero¬ geneity and classification of the Poaceae. New Phytol. 107: 633-644. Knobloch, I. W. 1963. The extent of hybridization in the Gramineae. Darwiniana 12: 624-628. - . 1968. A check list of crosses in the Gramin¬ eae. East Lansing, Michigan. Lehvaslaiho, H., A. Saura & J. l.okki. 1987. Chloro- plast DNA variation in the grass tribe Festuceae. Theor. Appl. Genet. 74: 298-302. Linder, H. P. 1986. Diverse notes on southern African Pooids. Bothalia 16: 59-61. Love, A. 1963. Cytotaxonomy and generic delimitation. Regnum Veg. 27: 45-51. MacLeod, A. M. & H. McCorquodale. 1958. Water- soluble carbohydrates of seeds of the Gramineae. New Phytol. 57: 168-182. Macfarlane, T. D. & I.. Watson. 1982. The classifi¬ cation of Poaceae subfamily Pooideae. Taxon 31: 178-203. M'Alpine, A. N. 1898. Production of new types of forage plants — Clovers and grasses. Trans. Highland and Agric. Soc. Scotland, Ser. 5, 10: 135-165. Malik, C. P. & P. T. Thomas. 1966. Karyotypic studies in some Lolium and Festuca species. Caryologia 19: 167- 196. Morgan, W. G., 11. Thomas, M. Evans & M. Borrill. 1986. Cytogenetic studies of interspecific hybrids between diploid species of Festuca. Canad. J. Genet. Cytol. 28: 921-925. Nevski, S. A. 1934. Lolium. Pp. 545 — 552 in V. L. Komarov, R. Yu Rozhevits & B. K. Shishkin (edi¬ tors), Flora of the LJ.S.S.R. Vol. 2. Translation for the National Science Foundation and the Smithsonian Institution, 1965, Israel Program for Scientific Translations. Niles, C. D. & A. Chase. 1925. A bibliographic study of Beauvois’ Agrostographie. Contr. U.S. Natl. Herb. 24: i-vi, 135 — 214, vii-xix. Raspail, F. V. 1829. Deviations physiologiques et me¬ tamorphoses reelles du Lolium. Ann. Sci. Observ. 2: 233-244. Reveal, J. L., E. E. Terrell, J. H. Wiersema & H. Scholz. 1991. Proposal to reject Festuca elatior L. with comments on the typification of F. pratensis and F. arundinacea (Poaceae). Taxon 40: 135-137. Saint-Yves, A. 1929. Festuca hybrides. Izv. Glavn. Bot. Sada SSSR. 28: 592-608. Smith, P. 1969. Serological relationships and taxonomy in certain tribes of the Gramineae. Ann. Bot. (London) 33: 591-613. Soreng, R. J., J. I. Davis & J. J. Doyle. 1990. A phylogenetic analysis of the chloroplast DNA restric¬ tion site variation in Poaceae subfam. Pooideae. PI. Syst. Evol. 172: 83-97. Stace, C. A. 1975. Hybridization and the flora of the British Isles. Academic Press, London, New York, San Francisco. Stebbins, G. L., Jr. 1956. Taxonomy and the evolution of genera, with special reference to the family Gra¬ mineae. Evolution 10: 235-245. Tateoka, T. 1962. Starch grains of endosperm in grass systematics. Bot. Mag. (Tokyo) 75: 377-383. Terrell, E. E. 1966. Taxonomic implications of genetics in ryegrasses (Lolium). Bot. Rev. (Lancaster) 32: 138-164. - . 1968. A taxonomic revision of the genus Lolium. Tech. Bull. U.S.D.A. 1392: 1-65. Thomas, H. M. 1981. The Giemsa C-band karyotypes of six Lolium species. Heredity 46: 263 267. Tutin, T. G. 1956. Generic criteria in flowering plants. Watsonia 3: 317-323. Tzvelev, N. N. 1989. The system of grasses (Poaceae) and their evolution. Bot. Rev. (Lancaster) 55: 141- 204. Ullmann, W. 1936. Natural and artificial hybridization of grass species and genera. Herbage Rev. 4: 1 OS- 142. Watson, L. & R. B. Knox. 1976. Pollen wall antigens and allergens: Taxonomically ordered variation among grasses. Ann. Bot. (London) 40: 399-408. Lagenocarpus venezuelensis , a New Name for L. amazonicus Davidse (Cyperaceae: Cryptangieae) Gerrit Davidse Missouri Botanical Garden., P.0. Box 299, St. Louis, Missouri 63166, U.S.A. ABSTRACT. Lagenocarpus venezuelensis Davidse is proposed as a new name for the later homonym Lagenocarpus amazonicus Davidse. \\ hen I published the new species Lagenocarpus amazonicus based on two gatherings from Territorio Federal, Amazonas, Venezuela (Davidse, 1992), 1 overlooked the fact that this combination had al¬ ready been used by Pfeiffer (1925) for the species now generally recognized (Koyama & Maguire, 1965: 53) as Exochogyne amazonica C. B. Clarke, Verb. Bot. Brandenburg 17: 101. 1906. Therefore, I propose the following new name for the illegitimate homonym: Lagenocarpus venezuelensis Davidse, nom. nov. Replaced name: L. amazonicus Davidse, No- von 2: 319. 1992, non (C. B. Clarke) H. Pfeif¬ fer, Hepert. Spec. Nov. Regni Veg. 21: 35. 1925. TYPE: Venezuela. Territorio Federal Amazonas: Depto. Rio Negro, Cerro Aratiti- yope, aprox. 70 km al SSW de Ocamo, 2°10'N, 65°34'W, con riachuelos afluente al rio Man- ipitare, piedra ignea, 990 — 1,670 m, 24-28 Feb. 1984, Steyermark , Berry & Delascio 130049 (holotype, MO; isotypes, NY, US, VEN). Acknowledgments. 1 thank M. Strong and S. F. Smith of the Smithsonian Institution for bringing this regrettable error to my attention. Literature Cited Davidse, G. 1992. A new species of Cyperaceae (Cryp¬ tangieae) from the Venezuelan Guayana. Novon 2: 319-321. Koyama, T. & B. Maguire. 1965. Cyperaceae tribe Lagenocarpeae. In: The Botany of the Guayana High¬ land-Part VI. Mem. New York Bot. Card. 12: 8- 54. Pfeiffer, H. 1925. Additamenta ad cognitionem generis Lagenocarpus V. Von zwei vermeintlich neuen Cy- peraceengattungen. Repert. Spec. Nov. Regni Veg. 21: 34-36. Novon 3: 244. 1993. Dendrothrix, a New Generic Concept in Neotropical Euphorbiaceae Hans-Joachim Esser Universitat Hamburg, Institut fiir Allgemeine Botanik und Botanischer Garten, Ohnhorststrasse 18, D-22609 Hamburg, Germany ABSTRACT. The new 'genus Dendrothrix is pro¬ posed. Two of its species are new to science, the third one was originally allied to Sapium. Keys for distinguishing related genera using flowers, fruits, and leaves are provided, as is a key to the species of Dendrothrix. A taxonomic revision of the subtribe Mabeinae Pax & K. Hoffmann of the Euphorbiaceae (tribe Hippomaneae A. Jussieu ex Spach) has necessitated the establishment of a new genus in order to ac¬ commodate species that have confused students of the Euphorbiaceae for some time. The three species are known from northern Brazil and Venezuela. Dendrothrix Esser, gen. nov. TYPE: Dendrothrix yutajensis (Jablonski) Esser. A ceteris generibus tribus Hippomanearum differt pilis ramosis, foliis integris subtus papillis obtectis sine glandulis marginalibus praeter bases glandulosas, thyrsis regulatim plerumque semel ramosis, glandulis bractearum discifor- mibus vel cyathiformibus, floribus masculinis bistamina- libus cum filainentis tota longitudine connatis, ovariis pu- bescentibus, et mericarpiorum septis filo vasculari furcato vel duplici instructis. frees or shrubs. Hairs multicellular and ramified. Leaves alternate, simple, coriaceous, entire; abaxial epidermis minutely but densely papillose; without petiolar, submarginal, or marginal glands but abax- ially with a pair of basimarginal glands and some¬ times with laminar glands of 0.2-0.35 mm diarn.; petioles 1.5-5. 5 cm long. Stipules very small to absent. Thyrses yellowish to cream-colored, termi¬ nal, always compound, mostly with one order of branching with several lateral thyrses, bracts small, scaly, glandless. Male cymules distal, at least 8-flowered, their bracts carrying basal glands which, when dry, are cup-shaped or disc-shaped and ± 1 mm long; bracteoles absent. Staminate flowers ses¬ sile, at anthesis with 0.6-1 .0-mm-long pedicel, sub¬ tended by a two-parted calyx fused for most of its length; stamens two per flower, connate, lateral flow¬ ers often unistaminate; anthers 0.4 mm long, fila¬ ments at anthesis slightly longer than anthers. Fe¬ male flowers proximal, bracteolate, shortly pedicellate, tricarpellate; sepals three, partly fused; ovary pubescent; style short but evident. Fruit a septicidal schizocarp, smooth, with scattered, ca¬ ducous pubescence, glabrescent, up to 6 mm long. Mericarps bearing one bifurcate or two distinct vas¬ cular strands visible on each septum, leaving an inconspicuous alate central columella. Seeds dry, brown, smooth, carunculate in two species (not known in D. multiglandulosa). The name of the genus refers to the treelike branched hairs, which are unusual for the Euphor¬ biaceae. This genus differs from the other three neotropical genera of the Hippomaneae with compound thyrses in both its bistaminate male flowers and its totally fused filaments. The inflorescence is similar to that of Senefeld- eropsis Steyermark: the thyrses are of the same size and are strictly terminal. A single, and rarely a second, order of branches occur, always subtended by small scaly bracts, and the staminate cymules are many-flowered. The two genera therefore can be confused superficially. However, the hairs of Se- nefelderopsis are multicellular but unbranched, the bracteal glands of its two species are always cylin¬ drical when dry, the staminate flowers carry per¬ sistent bracteoles, and its fruits are larger (length 15-25 mm) and have a leathery outer layer. Ad¬ ditionally, the leaves of Senefelderopsis exhibit a row of abaxial submarginal glands and a pair of adaxial basal glands. The hairs and fruits of Dendrothrix are very similar to those of Mabea Aublet. The leaf glands of Mabea, however, are strictly marginal or sub- marginal, never laminar. The lateral secondary thyrses of Mabea occur irregularly (they are missing in many species, whereas in others they can be several times compound), and the staminate cymules are mostly 3-flowered. The calyx of pistillate flowers and fruits consists of six sepals. Mabea always shows a single undivided vascular strand on each septum of the mericarps, the length ol the capsules always exceeds 8 mm, and its indumentum is very dense and persistent. Senefeldera C. Martius, the third other genus with compound thyrses, is quite different, e.g., it Novon 3: 245-251. 1993. 246 Novon has unbranched hairs, leaves without abaxial papillae and without basimarginal glands, bracteolate sta- rninate cymules with 1-3 (lowers each, mostly gla¬ brous ovaries, and larger fruits. The following keys should facilitate the distinction of the neotropical genera with compound thyrses: Key for Characters of Fruits la. Fruits, excluding style, up to 6 mm long .... . Dendrothrix lb. Fruits, excluding style, at least 7 mm long .... 2 2a. Ripe fruits with fleshy outer layer that wrin¬ kles when drying . Senefelderopsis 2b. Ripe fruits dry, not wrinkling . 3 3a. Fruiting calyx of 6 sepals; fruits with dense indumentum of shortly papillose or branched hairs . Mabea 3b. Fruiting calyx of 3 sepals; fruits gla¬ brous or covered with long un¬ branched hairs . Senefeldera Key for Characters of Staminate Flowers la. Staminate cymules with many (> 6, often > 10) flowers each . 2 2a. Glands of bracts of staminate cymules cy¬ lindrical; flowers with 3(-5) free stamens each . Senefelderopsis 2b. Glands of bracts of staminate cymules disc- or cup-shaped; flowers with 2 fused sta¬ mens each . Dendrothrix lb. Staminate cymules with l-5(-7) flowers each 3 3a. Staminate flowers at anthesis inclinate or nearly sessile . Senefeldera 3b. Staminate flowers at anthesis erect and clearly pedicellate . Mabea Key for Characters of Leaves la. Leaves abaxially glaucous or papillose . 2 2a. Leaves with adaxial basal gland(s); abaxi¬ ally one row of submarginal glands . . Senefelderopsis 2b. Leaves without adaxial basal glands; abax¬ ially with or without row of marginal or submarginal glands . 3 3a. Leaves always entire; abaxially papil¬ lose; never with abaxial row of mar¬ ginal or submarginal glands, but most¬ ly with laminar pellucid dots . . Dendrothrix 3b. Leaves often serrate; if entire, abax¬ ially never papillose; with or without row of marginal or submarginal glands, never with laminar pellucid dots . . Mabea lb. Leaves abaxially shining and smooth, neither glaucous nor papillose . 4 4a. Petiole at least 2 cm long; leaf glands lam¬ inar . Senefeldera 4b. Petiole less than 2 cm long; leaf glands marginal, submarginal, or absent .... Mabea Dendrothrix seems to be very closely related to Senefelderopsis. Both genera are centered in the Guayana Highland, although, interestingly, they oc¬ cur only partially sympatric. Senefelderopsis is strictly endemic to the Guayana Highland and is widely distributed there, but is absent from its south¬ ern, e.g., Brazilian, part. Dendrothrix, on the other hand, has two species known only from the Vene¬ zuelan and Brazilian portion of the Guayana High¬ land and a third one occurring in Brazilian Amazonia near the northern margin of the Brazilian planalto. Furthermore, at least Senefelderopsis chiribique- tensis (R. Schultes & Croizat) Steyermark is very similar ecologically to the two Venezuelan species of Dendrothrix. Key to the Species of Dendrothrix la. Bracts of male flowers with at least two pairs of glands; mature leaves glabrous; tertiary leaf venation predominantly percurrent . . 1 . D. multiglandulosa lb. Bracts of male flowers with only one pair of glands; mature leaves pubescent; tertiary leaf venation clearly reticulate . 2 2a. Leaf blades elliptic; basal lateral nerves similar to distal ones; hairs reddish when dry . 3. D. yutajensis 2b. Leaf blades ovate; basal lateral nerves dif¬ ferent from distal ones in length and angle of divergence; hairs pale when dry . . 2. D. wurdackii 1 . Dendrothrix multiglandulosa Esser, sp. nov. TYPE; Venezuela. Territorio Federal Amazo¬ nas: Cerro Sipapo (Paraque), breaks below Low¬ er Camp Savanna, alt. 4,400 ft., 15 Dec. 1948 (fl, fr), H. Maguire & L. Politi 27683 (holo- type, K; isotypes, GH, MICH, S, U, US). Fig¬ ure 1 . Species generis Dendrotrichis pilis pallidis ornata, foliis mox glabrescentibus nervationem tertiariam percurren- tem praebentibus, bracteis cymularuin masculinarum min¬ imum duobus paribus glandium instructis. Shrub or tree, 4 m. Indumentum pale. Leaves elliptic to obovate, ( 1 0—) 1 3—18 cm long, 5. 0-9. 5 cm wide, basally cuneate to obtuse, apically acute to obtuse; mature ones glabrous; secondary veins Figure 1. Dendrothrix multiglandulosa Esser. — A. Habit. — B. Pistillate flower. — C. Portion of staminate part of inflorescence showing two cymules. — D. Old leaf. — E. Leaf base with glands, abaxial view. — F. Detail of percurrent leaf venation. (All drawn from Maguire & Politi 27683.) Volume 3, Number 3 1993 Esser Dendrothrix, a New Generic Concept 247 248 Novon 11-14 pairs, tertiary venation percurrent; (0-)10- 25 laminar glands on each half of blade, basal glands less than 1 mm long; petioles 2. 0-5. 5 cm long. Stipules unknown. Thyrses with flowering part up to 14 cm long, bracts of secondary branches 1.5 mm long. Each branch bearing 1 female flower and at least 10 male cymules. Bracts of male cymules with at least two pairs of glands. Bract of pistillate flower up to 2 mm long. Pedicel of pistillate flower 1 mm long; sepals ( 1 .0—) 1 .5 mm long, 1. 5-2.0 mm wide, mostly ciliate; style 1 .0 mm long, stigmas 0.5- 1.0 mm long. Fruits 4 mm long, mostly glabrous. Seeds unknown. Distribution and phenology. Endemic to Cerro Sipapo, and locally frequent in low bush and mixed forest; flowering in December. Jablonski (1967: 186) used both Maguire & Pol- iti 27683 and Maguire & Politi 27885 to describe the pistillate flowers of Senefelderopsis sipapoensis Jablonski. He did not recognize the profound dif¬ ferences from this genus that are discussed above. Both collections were distributed as Senefelderopsis si pa poen s is J ablonski . Differential characters for Dendrothrix multi- glandulosa beyond those given in the key are the higher number of leaf glands and the ciliate margins of the broad, apically rounded to acute sepals of tbe pistillate flowers. Stipules or abscission scars could not be found. It can be postulated that the stipules have been reduced totally, much more than in the other two species of the genus. The name of the species refers to the multiple glands of the bracts of the staminate cymules. Paratype. VENEZUELA. Territorio Federal Amazonas: Cerro Sipapo (Paraque), North Escarpment, alt. 1,400 m, 23 Dec. 1948 (fl), B. Maguire & L. Politi 27885 (S). 2. Dendrothrix wurdackii Esser, sp. nov. TYPE: Brazil. Amazonas: Bio Aripuana, Nova Prainha, Projeto RADAM/BRASIL SB-20-ZB Ponto 15, 15 July 1976 (fl, fr), J. Kamos, J. Geraldo & L. Coelho s.n. (holotype, INPA-62163). Fig¬ ure 2. Species ejusdem generis pilis in sicco pallidis ornata, foliis ovatis subtus perduranter pubescentibus nervati- onem tertiariam reticulatam praebentibus, nervis secun- dariis basalibus dissimilibus isdem distalibus, parte connata stylorum perspicue breviore stigmatibus, bracteis cymu- larum masculinarum pari singulario glandium instructis. Shrub, 1 m. Indumentum pale. Leaves ovate, up to 8 cm long, 4.5 cm wide, basally obtuse to rounded, apically acute; adaxially pubescent to glabrous, abaxially persistently pubescent; secondary veins 7- 9 pairs, basal one differing from distal ones in length and angle of divergence, tertiary venation reticulate; 1-2 laminar glands on each half of blade, basally one large abaxial pair of glands (diameter 1 mm each); petioles 2-4 cm long. Stipules 0.5 mm long. Thyrses at least 6 cm long, bracts of secondary branches 1.5-2. 5 mm long. Each branch bearing 1-2 female flowers and ± 25 male cymules. Bracts of male cymules with one pair of glands. Pistillate flower in the axil of a 2-mm-long bract. Pedicel 1 mm long; sepals 1 .0-1.5 mm long, 1 mm wide, not ciliate, sometimes with glands; style 0.5-0. 7 mm long, clearly shorter than stigmas. Fruits 6 mm long, sparsely pubescent. Seeds 4 mm long, 3 mm wide, with large caruncle. Dendrothrix wurdackii is known only from the type collection, which was distributed as Mabea sp. More collections may exist, but have probably been distributed under different names. Ecological data of this species are not known. It differs from the other two species predominantly in characters of its leaves. Besides the characters listed in the key it is distinguished by the short styles and the large basal leaf glands, as well as the obtuse to rounded leaf base. The name of the species honors Kenneth J. Wur- dack, who, independently of me, recognized this new species as well as its affinities. 3. Dendrothrix yutajensis (Jablonski) Esser, comb. nov. Basionym: Sapiurn yutajense Ja¬ blonski, Mem. New York Bot. Card. 17: 184. 1967, fig. 24. Senefeldera yutajensis (}uh\on- ski) Webster, Ann. Missouri Bot. Card. 76: 958. 1989. Senefelderopsis yutajensis (Ja¬ blonski) Mennega in Kruijt, Monographic Stud¬ ies on Sapium. Doctoral Thesis, Rijksuniver- siteit te Utrecht: 201. 1989. TYPE: Venezuela. Territorio Federal Amazonas: Cano Yutaje, 1,600 m, 15 Feb. 1953 (fl, fr), K. & C. K. Maguire 35261 (holotype, NY). Figure 3. Figure 2. Dendrothrix wurdackii Esser. — A. Habit. — B. Pistillate flower. — C. Portion of staminate part of inflorescence showing two cymules. — D. Androecium, dorsal view. — E. Androecium without filaments, ventral view. — F. Bract of secondary thyrse with hracteoles. — G. Leaf base showing glands, abaxial view. — H. Detail of reticulate leaf venation. (All drawn from Ramos et al. s.n.) 1 mm Volume 3, Number 3 1993 Esser Dendrothrix, a New Generic Concept 249 2 cm 250 Novon Figure 3. Dendrothrix yutajensis (Jablonski) Esser. — A. Hairs, upper view. — B. Mericarp, ventral view. — C. Seed, ventral view. — D. Seed, lateral view. — E. Detail of vegetative shoot showing stipule. — F. Leaf base showing glands, abaxial view. (A-D drawn from Amaral 1523; E, F drawn from Rosa & Lira 2293.) Jablonski (1965: 176) had correctly cited the collection Maguire el al. 30964 as Senefelderopsis chiribiquetensis (H. Schultes & Croizat) Steyer- mark. 1 wo years later he erroneously made the same specimen one of the paratypes of his Sapium yu- tajen.se. Fortunately, his excellent drawings and the well-written description unequivocally refer to Den¬ drothrix yutajensis. The original description now has to be expanded: Hairs always reddish. Leaves abaxially with 0-3 laminar and a pair of basimarginal glands (diameter less than 0.5 mm). Stipules persistent, squamiform, less than 1 mm long. Sepals of female flowers not ciliate. Style 1. 5-2.0 mm long, exceeding length of stigmas. Seeds 3. 5-4.0 mm long, 2.5 mm wide. Pollen grains 22 gm long, tectum perforate, exine psilate. Distribution and phenology. Endemic to the Guayana Highland of Venezuela and Brazil, un¬ known from its western and eastern portion, and occasional to frequent in campos and thickets; flow¬ ering in November, January to March, May, fruits collected in February. The most conspicuous difference between this species and the other two of the genus is the in¬ dumentum, which consists of reddish and quite firm hairs; within the Euphorbiaceae they are only com¬ parable to those of Mabea Aublet. Additional specimens examined. BRAZIL. Amazo¬ nas: Mun. Barcelos, Plato da Serra Araca, parte SE da Serra Norte, alt. 1,150-1,250 m, 12 Feb. 1984 (fl, fr), /. L. do Amaral 1523 (INPA, NY); plateau of northern massif of Serra Araca, N part of Northern mountain near peak, alt. 1,400 m, 17 Feb. 1984 (fr), G. T. Prance et al. 29140 (MG); Arredores do R. da Serra Araca, 29 Jan. 1978 (imm. fr), N. A. Rosa & S. B. Lira 2293 (MG). VENEZUELA. Territorio Federal Amazonas: Depto. Rio Negro, Cerro de La Neblina Camp IV, 1 5 km NNE of Pico Phelps, alt. 780 m, 15-18 Mar. 1984 (fl), R Liesner 16685 (HBG); Serrania Yutaje, Rio Mana- piare, in right branch of Cano Yutaje, alt. 1,400 in, 9 Feb. 1953 (fl), B. & C. K. Maguire 35103 (COL); Cerro de La Neblina, Rio Yatua, banks of Canon Grande E of Cumbre Camp, alt. 1,100 m, 24 Nov. 1957 (imm. fl), B. Maguire et al. 42212 (LI); Cerro Yapacana, en la cumbre, alt. 1,000-1,200 m, 5-7 May 1970 (fl), J. A. Steyermark & G. S. Bunting 103143 (U). Bolivar: Distr. Heres, Cerro Marutani, piedra arenisca en la alti- planicie a lo largo del rio Carla, alt. 1,200 m, 11-14 Jan. 1981 (imm. fr), J. A. Steyermark et al. 123920 (NY). Acknowledgments. This article is based on a doc¬ toral study by H.-J. Esser at the Faculty of Biology, University of Hamburg. It was supported by the Volkswagenstiftung. I thank directors and curators of the cited herbaria for the loan of specimens and K. Kubitzki und H.-JI. Poppendieck for helpful com¬ ments. Additional thanks are due to C. Oberprieder for drawing Figure 1 A; other drawings by the author. Volume 3, Number 3 1993 Esser Dendrothrix, a New Generic Concept 251 Literature Cited Jablonski, E. 1965. Euphorbiaceae. Pp. 150-178 in: Botany of the Guayana Highlands — Part VI. Mem. New York Bot. Gard. 12. - . 1967. Euphorbiaceae. Pp. 80- 1 90 in: Botany of the Guayana Highlands — Part VII. Mem. New York Bot. Card. 17. A New Species of Streptostachys (Poaceae: Paniceae) from Brazil Tare iso S. Filgueiras Reserva Ecologica do IBGE, Cx. Postal 08770, 70200-200 Brasilia, DF-Brazil Osvaldo Morrone and Fernando O. Zuloaga Instituto de Botanica Darwinion, Casilla de Correo 22, San Isidro, 1642, Argentina ABSTRACT. A new species of Streptostachys, S. rigidi/olia, from the Brazilian state of Maranhao is described, illustrated, and compared with previously described species in the genus. Anatomical data on the leaf blade of the new species are also provided. Upon examination of recent gatherings of Po¬ aceae from Maranhao, Brazil, a new species of Streptostach ys P. Beauvois was encountered, which is here described, illustrated, and compared with species previously included in the genus by Morrone & Zuloaga (1991). Streptostachys rigidifolia Filgueiras, Morrone & Zuloaga, sp. nov. TYPE: Brazil. Maranhao: Loreto, BR-230, estrada para Sao Raimundo das Mangabeiras, ca. 22 krn de Sao Raimundo, 6°56'S, 45°19'W, cerrado baixo com estrato graminoso aberto, 13 maio 1988, L. B. Bian- chetti, F. R. Ferreira & J. N. Silveira 631 (bolotype, CEN; isotypes, B, BM, IBGE, K, MO, R, RB, SI, SP. UB, US). Figures 1-3. Ah omnibus speciebus generis flosculo superiore glabro papillis compositis ornato differt. Streptostachys ramosa sirnilis sed plantis 68 95 cm altis, inflorescentia racemosa vel subpaniculata 6-17 cm longa abunde differt. Robust, stout, caespitose and short-rhizomatous perennial. Culms 68-95 cm tall, simple, bearing basal and cauline leaves; internodes cylindrical, hol¬ low, glabrous; nodes swollen, shortly pilose with whit¬ ish hairs. Sheaths pilose, more so toward the base, or glabrous, striate, one margin hyaline, the other eiliate. Ligules a ciliate rim ca. 1 nun long, with hairs up to 7 mm long, behind the ligule at the base of the blade and on the upper margins of the sheaths. Blades lanceolate, 25-50 cm long, 5-8 mm wide, rigid, glabrous or with few scattered hairs, strongly striate, rounded at the base, the apex pungent, acu¬ minate, the margins denticulate and/or ciliate. Pe¬ duncle cylindrical, scabrous, to 30 cm long. Inflo¬ rescence exserted, racemose or subpaniculate, 6- 17 cm long, formed by 2-6 branches 9 11 cm long, all bearing unilateral spikelets along their entire length; main axis short, triquetrous, scabrous, ter¬ minating in a spikelet; spikelets solitary on the branches, the pedicels 1-3 mm long, swollen, par¬ tially adnate to the rachis of the branches. Spikelets narrowly ellipsoid, 5-8 mm long, shortly pilose, with thickened, conspicuous internodes between the glumes, and florets; upper glume and lower lemma subequal, conspicuously coriaceous. Lower glume ovate, 3-6 mm long, '/,-2/, the length of the spikelet, acute, 3(-5)-nerved, adaxial, slightly asymmetrical. Upper glume 5-6 mm long with a few short hairs throughout, 5-nerved, the nerves anastomosed to¬ ward the apex. Lower floret male; stamens 3, the anthers 3 mm long; Iodic ules 2, truncate, condu- plicate, 0.5 0.8 mm long; lower lemma 5. 5-6. 5 mm long, 5-nerved, prominently pilose toward the apex; lower palea well developed, lanceolate, 4.2 mm long, 1.6 mm wide, hyaline, strongly 2-nerved. Upper floret narrowly ovoid, 4.5 mm long, 1.6 mm wide, indurate, pale, shiny, with regular compound papillae evenly distributed over the surface of the lemma and palea; lemma 5-nerved; styles 2, free from the base of the gynoecium; stigmas plumose, purple; stamens 3, with purple anthers; lodicules 2, conduplicate, 0.5-0. 8 mm long. Caryopsis plano¬ convex, 3—3.5 mm long; hilum linear along the entire length of the caryopsis. Streptostachys rigidifolia is known only from a single population in the municipality of Loreto, Ma¬ ranhao, Brazil, in cerrado vegetation. This species has been included in Streptostachys because of its inflorescence type, the thickening of the rachilla between the glumes and florets, the nervation of the glumes and lower lemma, and the linear hilum (Morrone & Zuloaga, 1991). Streptostachys rigidifolia is unique in the genus because of its glabrous upper floret, with compound papillae over its entire surface. It can be further distinguished from S. ramosa and S. macrantha by the coriaceous upper glume and lower lemma; in S. ramosa and S. macrantha the upper glume and lower lemma are membranous; both species have a pilose upper floret with simple papillae over the Novon 3: 252-257. 1993. Volume 3, Number 3 1993 Filgueiras et al. Streptostachys rig id i folia 253 Figure 1. Streptostachys rigidifolia Filgueiras, Morrone & Zuloaga. — a. Flabit. b. Schematic longitudinal section of spikelet base, indicating conspicuous thickened internodes: Lg: lower glume; Ug: upper glume; LI: lower lemma; Lp: lower palea; Ul: upper lemma; Up: upper palea; Lo: lodicules; Gy: gynoecium. — c. Spikelet, ventral view. — d. Spikelet, lateral view. — e. Spikelet, dorsal view. — f. Upper palea and lodicules. — g. Upper floret, dorsal view. — h. Upper floret, ventral view, lemma and palea. (Based on Bianchetti et al. 634.) 254 Novon Figure 2. Scanning electron micrographs of the upper floret of Streptostachys rigidifolia. — A. Upper portion of lemma. — B. Upper portion of palea and lemma. — C. Detail of the surface of the upper lemma showing compound papillae. — D. Detail of the surface of the upper palea showing compound papillae. (Based on Bianchetti et al. 634.) surface of lemma and palea. The plant height and inflorescence size and shape are further distinguish¬ ing characters: S. macrantha includes plants 30- 70 cm tall, and inflorescences with whorled lower branches. Streptostachys ramosa includes plants 130-200 cm tall, inflorescences 30-45 cm long, with whorled lower branches. Streptostachys as- peri folia differs from S. rigidifolia by the following features: presence of axillary inflorescences, spike- lets 3.6-5. 1 mm long, lower glume %-% the length of the spikelet, absence of lower flower and blades cordate, amplexicaulous. Paratypes. BRAZIL. Maranhao: Loreto, km 291.2 da BR-230, barranco ingreme de estrada a cerca de 22 km de Sao Raimundo, 20 mar^o 1983, J. F. M. I alls, G. Feign & Silva 8432 (CEN, ICN, MEXU, VEN). Anatomy of Streptostachys rigidifolia. Figure 3 The techniques used in the anatomical study fol¬ low Morrone & Zuloaga (1991), and the terminology follows Ellis (1976, 1979). LEAF BLADE IN TRANSVERSE SECTION, FIGURE 3a, B Outline: open, expanded, flat, two halves sym¬ metrical on either side of median vascular bundle; leaf thickness 310-420 /am. Ribs and furrows: rounded adaxial ribs, associated with first- and sec¬ ond-order vascular bundles; the larger ribs alter¬ nated with the smaller ones; adjacent ribs separated by narrow furrows, penetrating up to % the leaf thickness; abaxial ribs and furrows slightly undulat¬ ing with no regular pattern associated with ihe vas¬ cular bundle. Median vascular bundle: midrib or keel not developed; median vascular bundle struc¬ turally indistinguishable from lateral first-order vas¬ cular bundles. Vascular bundle arrangement: 15 first-order vascular bundles, 10 second-order vas¬ cular bundles and 34-36 third-order vascular bun¬ dles in entire blade; 1 second-order vascular bundle and 2-3 third-order vascular bundles between con¬ secutive first-order vascular bundles. First-order vascular bundles centrally located in the blade thick¬ ness, second- and third-order vascular bundles abax- ially displaced. Vascular bundle description: first- Volume 3, Number 3 1993 Filgueiras et al. Streptostachys rig id i folia 255 Figure 3. Leaf blade anatomy of Streptostachys rigidifolia. — A. Outline showing absence of a keel. — B. Transverse section detail showing first-, second-, and third-order vascular bundles, bulliform cells with associated colorless cells. — C. Abaxial epidermis showing costal and intercostal zones. (Based on Bianchetti et al. 6 34.) order vascular bundle elliptical in outline; metaxylem vessel equal or bigger than the Kranz cells as seen in section, metaxylem vessels slightly angular in outline; sclerenchymatous cells present in 1-3 rows between the metaxylem vessels and Kranz cells, fibers lignified, lumen small; phloem tissue com¬ pletely surrounded by thick-walled fibers. Second- order vascular bundle elliptical in outline with xylem and phloem tissue distinguishable. Third-order vas¬ cular bundle angular in outline with metaxylem and phloem tissue indistinguishable, located below the furrows. I ascular bundle sheaths: first-order vas¬ cular bundle surrounded by a single Kranz mestome sheath of 22-27 cells, with adaxial and abaxial interruption of sclerenchyma girders; second-order vascular bundles completely surrounded by a single Kranz mestome sheath, consisting ol 12—14 cells; adaxial extensions 1-3-seriate, 2-5 cells deep. Kranz mestome sheath of third-order vascular bundles en¬ tire, with 7—10 cells. Kranz cells rounded, with radial and outer tangential wall inflated, longer parallel to the vein in paradermal section; specialized chloro- plasts with tendency to show a centriiugal location. Sclerenchyma: adaxial and abaxial girders well de¬ veloped, relatively narrow and deep, always asso¬ ciated with all first-order vascular bundles; scleren¬ chyma minute, inconspicuous adaxial and abaxial girders associated with all second-order vascular bundles; girders or strand not associated with third- order vascular bundles. Fibers very thick-walled with lumens almost completely filled, lignified. Small scle¬ renchyma cap in the leaf margin, curved in shape with sclerenchyma extending along abaxial side ol the leaf. Mesophyll: chlorenchyma irregularly ra¬ diate, continuous between vascular bundles, cells small, irregularly shaped, tightly packed without vis¬ ible intercellular air spaces; 2 3 chlorenchyma cells between contiguous vascular bundles, 95-200 /am between contiguous vascular bundles; colorless cells below the bulliform cells. Fusoid cells absent. Arm cells absent. Adaxial epidermal cells: bulliform cells present at the base of all adaxial furrows and oc¬ curring in restricted, fan-shaped groups usually with an inflated central cell; bulliform cells up to '/, the leaf thickness. Epidermal cells with distinct contin¬ uous cuticle; prickle hairs present; macrohairs, mi¬ crohairs, and papillae absent. Abaxial epidermal cells: bullifom cells absent; epidermal cells small with a very thick cuticle; prickle hairs present; macro- hairs, microhairs, and papillae absent. Amiloplast present in the Kranz cells. ABAXIAL EPIDERMIS IN SURFACE VIEW, FIGURE 3c Costal zone: long cells rectangular in outline, more than 3x longer than wide, anticlinal walls 256 Novon parallel, undulated; long cells separated by cork- silica cell pairs; silica bodies irregularly dumbbell¬ shaped. Prickles absent. Intercostal zone: long cells rectangular in outline, more than 3 x longer than wide, anticlinal walls parallel, end walls vertical; long cells separated by single short cells; silica bodies irregular in outline. Microhairs: absent. Stomata I complex: low triangular, 38-45 fim long, 23 28 yam wide, in 2 rows in the costal zone; one intersto- matal cell between successive stomata in a row. Macrohairs: present or absent, when present in the leaf margin, cushion base. Hooks: present in the middle of the intercostal zone, small with the base shorter than the stomata, barb shorter than the base. Papillae: absent. ADAXIAI. EPIDERMIS IN SURFACE VIEW Costal zone: same as for the ahaxial surface, but with conspicuous prickles. Intercostal zone: long and short cells identical to those of the abaxial epi¬ dermis. Microhairs: none seen. Stomatal complex: low triangular, arrangement as in the abaxial epi¬ dermis. Macrohairs: same as for the abaxial epi¬ dermis. Hooks: frequently in the intercostal zone, small with the base shorter than the stomata; barb developed basally from the apex to the base, longer than the base. Papillae: absent. The epidermal characters of ,S. rigidifolia cor¬ respond to the panicoid dermotype (Prat, 1960), with irregular dumbbell-shaped silica bodies, rect¬ angular long-cells with undulating walls and alter¬ nating with short cells. An interesting feature that became evident in this study was the absence of bicellular microhairs on both epidermides of S. rig¬ idifolia. This is noteworthy since bicellular micro- bairs are characteristic of subfamily Panicoideae (Ellis, 1977), and they are also present in all the other species of Streptostachys. It should, however, be noted that they are also lacking in other panicoid genera such as Yvesia, Oryzidium, and Milbrae- dochloa (Renvoize, 1987). Streptostachys rigidifolia is, according to the anatomical data presented here, a Kranz species, of the subtype MS, with a single Kranz mestome sheath around the vascular bundles, with centrifugal chlo- roplasts on the Kranz sheath, cells of the mestome sheath longitudinally elongated in paradermal view, and 2-3 mesophyll cells between contiguous vas¬ cular bundles. According to the correlation encountered in the Kranz syndrome between anatomical and physio¬ logical characters (Brown, 1977; Ellis, 1977; Hat- tersley, 1987; Tregunna et al., 1970), it is con¬ cluded that S. rigidifolia is a C4 species of the NADP-me physiological subtype or malate former (Downton, 1970; Gutierrez et ah, 1974). Streptostachys rigidifolia shares with S. ramosa and ,S. macrantha a similar anatomical type, being all C.j, MS species without fusoid cells. Streptosta¬ chys ramosa, however, has bicellular microhairs, conspicuous papillae on the abaxial epidermis, dis¬ coid stomata, four types of vascular bundles, and no colorless cells below the bulliform cells. Streptos¬ tachys macrantha also has bicellular microhairs, a thinner transverse section and four types of vascular bundles. Streptostachys asperifolia differs by being a C3 species with keel, arm cells, and conspicuous fusoid cells (full anatomical descriptions of the spe¬ cies previously mentioned are given in Morrone & Zuloaga, 1991). Acknowledgments. We are grateful to Luciano de Bern Bianchetti and J. F. M. Vails for the loan of specimens from CEN and to Vladimiro Dudas for the preparation of the drawings in Figure 1 . Fil- gueiras thanks CNPq and CAPES for the partial financial support for this study. He is especially grateful to the Missouri Botanical Garden for a grant (May Scholar 1991) that enabled him to spend three months working in the herbarium at the Garden. Literature Cited Brown, W. V. 1977. The Kranz syndrome and its subtypes in grass systematics. Mem. Torrey Bot. Club 23: 1-97. Downton, W. J. S. 1970. Preferential C4-dicarboxylic acid synthesis, the postillumination CO, burst, car¬ boxyl transfer step, and grana configurations in plants with C, photosynthesis. Canad. J. Bot. 48: 1795- 1800. Ellis, R. P. 1976. A procedure for standardizing com¬ parative leaf anatomy in the Poaceae. I. The leaf blade as viewed in transverse section. Bothalia 12: 65-109. - . 1977. Distribution of the Kranz syndrome in the Southern African Eragrostoideae and Panicoideae according to bundle sheath anatomy and cytology. Agroplantae 9: 73-110. - . 1979. A procedure for standardizing com¬ parative leaf anatomy in the Poaceae. II. The epi¬ dermis as seen in surface view. Bothalia 12: 641- 671. Gutierrez, M., V. E. Gracen & G. E. Edwards. 1974. Biochemical and cytological relationships in C4 plants. Planta 119: 279 300. Hattersley, P. W. 1987. Variations on photosynthetic pathway. Pp. 49-64 in T. R. Soderstrom, K. W. Hilu, C. S. Campbell & M. E. Barkworth (editors), Grass Systematics and Evolution. Smithsonian Insti¬ tution Press, Washington, D.C. Morrone, O. & F. 0. Zuloaga. 1991. Revision del genero Streptostachys (Poaceae: Panicoideae), su Volume 3, Number 3 1993 Filgueiras et al. Streptostachys rigid i folia 257 posicion sistematica dentro de la tribu Paniceae. Ann. Missouri Bot. Card. 78: 359-376. Prat, H. 1960. Vers une classification naturelle des graminees. Bull. Soc. Bot. France 107: 32-79. Renvoize, S. A. 1987. A survey of leaf-blade anatomy in grasses. XI. Paniceae. Kew Bull. 42: 739-768. Tregunna, E. B., II. N. Smith, J. A. Berry & J. S. Downton. 1970. Some methods for studying the photosynthetic taxonomy of the Angiosperms. Canad. J. Bot. 48: 1209 1214. A New Species of Euphorbia (Euphorbiaceae) from Masirah Island, Sultanate of Oman Shahina A. Ghazanfar Department of Biology, Sultan Qaboos University, P.0. Box 36, Al-khod 123, Muscat, Sultanate of Oman Abstract. Euphorbia masirahensis, a new spe¬ cies from Masirah Island, Sultanate of Oman, is described. Its habitat and flowering and fruiting pe¬ riods are given, and its distribution and endemicity are discussed. An illustration of the species and a distribution map are also provided. Euphorbia masirahensis S. A. Ghazanfar, sp. nov. TYPE: Sultanate of Oman, NE Masirah Island, Wadi Zamji, FH 690960, 80 m, on stony slope of wadi, 27 Nov. 1991, S. A. Gha¬ zanfar 1515 (holotype, BSB; isotypes, E, K, ON, Herbarium Sultan Qaboos University — SQUH1). Figure 1. Euphorbiae schimperi Presl primo adspectu maxime similis, sed cyathiis sessilibus, solitariis, terminalibus vel ramis lateralibus distincta. Rami ascendentes. Semina lul- va, glabra, laevigata. Perennial shrub, leafless, forming large clumps up to 70 cm tall and 150 cm wide. Branches gla¬ brous, bluish green, arising from a thick tap root. Branches and branchlets ascending. New branchlets red. Leaves readily deciduous, alternate, 1-1.5 x 1 mm, ovate, sessile, glabrous, reddish green. Cy- athia solitary, terminal on the main branch or on lateral branchlets, sessile, subtended by two ovate bracts; bracts ca. 2 x 1.5 mm, glabrous. Cyathial involucre 4.5-5 mm (including the glands) and 6- 6.5 mm across. Cyathial glands 5, ca. 3 x 1-1.5 mm, patent, transversely oblong, ochraceus. Mar¬ ginal lobes alternating with the cyathial glands. Male flowers numerous, each consisting of a single stamen; pedicel ca. 1 .5 mm long, whitish yellow; anthers 0.75-1 mm long, yellow. Bracts ca. 1 mm, glabrous, deeply dentate at the apex. Female flower consisting of a single ovary, sessile. Ovary glabrous, trilocular; styles 3, united to about two-thirds their length from the base; stigmas bifid. Fruit schizocarpic, 5-6 mm long and 5-5.5 mm across, 3-valved, green suffused 'Acronym officially approved for Department of Biol¬ ogy, Sultan Qaboos University, P.O. Box 36, Al-khod 123, Muscat, Sultanate of Oman, and for publication in the next edition of Index Herbariorum. with red when young, dark brown when mature; fruit not reflexed at maturity. Dehiscence septicidal, splitting into 3 bi-valved cocci. Seed one in each locule, 2.5-3 x ca. 2 mm, oblong, glabrous, smooth, yellow-brown with a whitish longitudinal stripe and a white caruncle. Habitat and flowering. The species occurs on stony, gravel, and rocky slopes of wadis and the lower slopes of Jabal Dawwah, Masirah Island, from 70 to 80 m. Flowering and fruiting from November to early January, but also more or less throughout the year. Distribution. Endemic to Masirah Island. Paratypes. SULTANATE OF OMAN. Masirah Is¬ land: Wadi Zamji (Palm wadi), FH 690960, 80 m, S. A. Ghazanfar 1374 (SQUH); Wadi Zamji, stony slope, D. Rappenhoner OM91-54 (ON, BSB); Jabal Humr, Palm Wadi, 25 Feb. 1992, J. Bryan s.n. (SQUH, in spirit). Key to the Unarmed, Aphyllous, Succulent Species of Euphorbia in Oman la. Cyathium solitary, sessile; capsule up to 5.5 mm diam., dark brown at maturity (red when young); stems bluish green . . E. masirahensis S. A. Ghazanfar lb. Cyathia clustered in groups of 4-6, the central cyathium sessile, the lateral cyathia peduncu¬ late; capsule more than 5.5 mm diam., red or buff at maturity; stems pale green . 2 2a. Capsule ca. 10 mm diam., pale brown at maturity with purple-red lines along the sutures . E. larica Boissier 2b. Capsule ca. 7 mm diam., red at maturity 3 3a. Cyathial glands yellow; seeds black, smooth . E. schimperi Presl 3b. Cyathial glands red or reddish green; seeds reddish brown, rugose . . E. dhofarensis S. Carter The new species is named after Masirah Island, where it is endemic. In Masirah it has a restricted distribution, occurring only in the northern part of the central mountains (Fig. 2). It is common on the lower slopes of these mountains and on the stony slopes of the wadis. Though restricted to a single locality in Masirah, E. masirahensis is well estab- Novon 3: 258-260. 1993. Volume 3, Number 3 1993 Ghazanfar Euphorbia masirahensis 259 Figure 1. Euphorbia masirahensis S. A. Ghazanfar. - male flowers. — D. Cyathium with developing fruit. — E. lished and occurs in large numbers with good re¬ generation. The closest relative of Euphorbia masirahensis is E. schirnperi Presl; however, E. schirnperi is distinguished by its 3-5 pedunculate cyathia and stalked fruits. Euphorbia masirahensis is also sim¬ ilar to E. dhofarensis S. Carter in habit, but is distinguished by its sessile cyathia, yellow (not red) glands, and smooth seeds. Euphorbia schirnperi and E. dhofarensis occur only in Dhofar, the southern region of Oman, in the dry areas of the central range of mountains and on the coastal plains (Fig. 2). Euphorbia Ulrica occurs throughout Oman, but is more widely distributed in the northern mountains and foothills, where it forms part ot the dominant shrubby vegetation (Ghazanfar, 1991, 1992); it is absent from Masirah Island (Fig. 2). The population of Euphorbia masirahensis at Masirah Island could possibly be a relict of a once luxuriant population that also occurred on the es¬ -A. Habit. — B. Fruiting branch. — C. Cyathium showing Fruit. — F. Part (coccus) of dehisced fruit. — G. Seed. carpment and coastal range of mountains in the eastern part of the central desert of mainland Oman. During the Pleistocene a land bridge probably ex¬ isted between Masirah Island and mainland Oman at Khawr Masirah. At that time the sea level was low and the Gulf of Oman was virtually dry (Clarke & Fontes, 1990). Migration of plants could have occurred during that time. However, in the last 15,000 years B.P., due to the increasing aridity of the Arabian Peninsula, the species may have been wiped out from mainland Oman, but the humid mon¬ soon winds, relatively high precipitation, and the effects of mists at Masirah could have contributed to its survival there. Acknowledgments. 1 am grateful to the Depart¬ ment of Biology, Sultan Qaboos University, for pro¬ viding research facilities, to John Bryan for guiding me in Masirah, and to Dieter Rappenhoner for crit¬ ically reading the manuscript. John Turvey checked 260 Novon Figure 2. Distribution of Euphorbia dhofarensis (V), h. larica (O) , E. masirahensis (▼), and E. schimperi (®) in Oman. the Latin diagnosis and Fatima Farooq cultivated the species successfully at the Sultan Qaboos Uni¬ versity Botanic Garden. Literature Cited Clarke, I. D. & J.-C. Fontes. 1990. Palaeoclimate re¬ construction in northern Oman based on carbonates from hyperalkaline groundwater. Quatern. Res. 33: 320-336. Ghazanfar, S. A. 1991. Floristic composition and the analysis of vegetation of the Sultanate of Oman. FI. Veg. Mundi 9: 15-227. - . 1992. Annotated catalogue of the vascular plants of the Sultanate of Oman, and their vernacular names. Scripta Botanica Belgica 2. Jardin Botanique National de Belgique, Meise, Belgium. Nomenclatural Adjustments in Chinese Plumbaginaceae Rudolf Kamelin Herbarium, V. L. Komarov Botanical Institute, Russian Academy of Sciences, Prol. Popov Street 2, 197376 St. Petersburg, Russia ABSTRACT. Four new combinations in Limonium (A. lacostei, L. dielsianum, L. chrysocomum subsp. semenovii, and L. callianthum), one in Armeria (A. labradorica subsp. sibirica), and one in Acan- tholimon (A. laevigatum) are proposed. During the revision of Plumbaginaceae tor the Flora of China, it became evident that the nomen¬ clature of six poorly understood taxa in the genera Acantholimon, Limonium, and Armeria required some adjustment. In order to make the names avail¬ able for the forthcoming volume of the Flora of China, the nomenclatural changes are herein pro¬ posed. Acantholimon laevigatum (Peng) R. Kamelin, comb, et stat. nov. Basionym: Acantholimon alatavicum Bunge var. laevigatum Peng, Gui- haia 3: 291. 1983. TYPE: China. Xinjiang: Wenquun, k. C. Kuan 1561 (holotype, PE). The differences in leaf color, size, and flattening, size of outer and inner bracts, and size and indu¬ mentum of calyx support the recognition of Acan¬ tholimon laevigatum and 4. alatavicum as distinct species instead of varieties of the latter. Armeria labradorica Wallroth subsp. sibirica (Turczaninow) R. Kamelin, comb, et stat. nov. Basionym: Armeria sibirica Turczaninow in Boissier, A. de Candolle, Prodr. 12: 678. 1848. TYPE: [Russia]. Siberia: Ad Tessinsky dozor, ? J. Kusnetov s.n. (holotype, LE). This taxon is widespread in bogs and meadows on alpine slopes or high plateaus at 3,000-3,500 m m the Altai Mountains in Mongolia and South Siberia, Russia. It is highly likely that the species also occurs in China, because it has been collected from several stations along the Mongolian-Chinese border in the Altai Mountains. Limonium lacostei (Danguy) R. Kamelin, comb, nov. Basionym: Statice lacostei Danguy, J. Bot. (Paris) sbr. 2, 1(3): 53. 1908. TYPE: [China. Xinjiang]: Vallee de Tegermanlik, 4,000 m, 26 Aug. 1906, La Coste s.n. (holotype, P). Limonium roboroivskii Ikonnikov-Galitzky, Acta Inst. Bot. Acad. Sci. URSS, ser. 1, 2: 255. 1936, syn. nov. TYPE: China. Xinjiang: Kashgaria, Kara-teke, 9 June 1889, W. J. Roborowsky s.n. (lectotype, LE). Peng (1987) cited Statice lacostei as a synonym of Limonium aureum (L.) Hill and maintained L. roboroivskii as a distinct but related species. Statice lacostei and L. roboroivskii are conspecific, and the correct name for ihe species is L. lacostei. I he latter is readily distinguished from L. aureum by its straight, nonflexuous rachis and light or lemon-yel¬ low calyx limb, instead of dichotomously branched, strongly flexuous rachis and golden calyx limb. Limonium dielsianum (Wangerin) R. Kamelin, comb. nov. Basionym: Statice dielsiana Wan¬ gerin, Repert. Spec. Nov. Regni Veg. 17: 399. 1921. L. aureum (L.) Hill var. dielsianum (Wangerin) Peng, FI. Reipubl. Popul. Sin. 60(1 ): 38. 1987. TYPE: China. [Qinghai] (as Tibet): beim Lager 66 am Rande der Tsaidam, district Barun, A. Tafrl 242 (holotype, B). This taxon was reduced by Peng (1987) to a variety of Limonium aureum, but the two are quite distinct in the inflorescence rachis, which is flexuous and dichotomous in L. aureum and straight in L. dielsianum, and calyx limb color, which is golden in the former and light yellow in the latter. In my opinion, these are best treated as distinct species. Limonium chrysocomum (Karelin & Kirilow) 0. Kuntze subsp. semenovii (Herder) R. Ka¬ melin, comb, et stat. nov. Basionym: Statice semenovii Herder in Regel & Herder, Bull. Soc. Imp. Natur. Moscou 41(1): 398. 1868. TYPE: [Kazakhstan]. Hi, Katu [Alatau], 2,500 m, 1857, P. Semenov s.n. (holotype, LE). Limonium chrysocomum is a very variable spe¬ cies with two geographically distinct infraspecific taxa. I recognize these as subspecies and maintain subspecies chrysocomum as a variable taxon with three varieties, as did Peng (1987). Novon 3: 261-262. 1993. 262 Novon Limonium callianthum (Peng) R. Kamelin, comb, et stat. nov. Basionym: Limonium drepanos- tachyum Ikonnikov-Galitzky subsp. callian¬ thum Peng, Guihaia 3: 292. 1983. TYPE: China. Xinjiang: Artux, A. J . Li & J. N. Zhu 7996 (holotype, PE). This species was treated by Peng (1983, 1987) as a subspecies of Limonium drepanostachyum. The latter is endemic to Kyrgyzstan and Tajikistan and differs in having densely pubescent bracts, densely pubescent calyces 4.5-6 mm, and 3-7- flowered spikelets. Limonium callianthum has gla¬ brous to very sparsely hirsute bracts, glabrous ca¬ lyces 6.5-7 mm, and 2-4-flowered spikelets. Acknowledgments. I am grateful to Ihsan Al- Shehbaz and Tatiana Shulkina for help in the prep¬ aration of the manuscript. Literature Cited Peng, T.-X. 1983. Materiae ad florain Plumbagina- cearum Sinensium. Guihaia 3: 291-292. - . 1987. Plumbaginaceae. In: Lee S.-K. (editor), FI. Reipubl. Popul. Sin. 60(1): 1-47. Three New Combinations and a New Name in Chinese Boraginaeeae Hu (l oil Kamelin Herbarium, V. L. Komarov Botanical Institute, Russian Academy of Sciences, Prof. Popov Street 2, 197376 St. Petersburg, Russia Abstract. The new combinations Lappula ka¬ relinii (Fischer & Meyer) R. Kamelin, Lindelojia stvlosa (Karelin & Kirilow) Brand subsp. pterocar- pa (Ruprecht) R. Kamelin, and Onosma setosa Led- ebour subsp. transrhymnen.se (Klokov ex M. Popov) Kamelin are made, and the new name O. Urn R. Kamelin & T. Popova is proposed. The revision of the Boraginaeeae for the Flora of China is in progress, and this study has shown that four nomenelatural adjustments are needed in the genera Lindelojia , Lappula , and Onosma. these changes are proposed here to make the names avail¬ able for the forthcoming volume 16 ot the Flora. Lappula karelinii (Fischer & Meyer) R. Kamelin, comb. nov. Basionym: Echinospermum kare¬ linii Fisc her & M eyer. bid. Sem. Hort. Petrop. 1 1: 67. 1845. TYPE: [Kazakhstan]. Soongar- ia. Grown at St. Petersburg Botanical Garden by C. A. Meyer in 1845 from seed collected by G. Karelin (holotype, LE). Lappula xinjiangensis C. Y. Yang ex C. J. Wang, Bull. Bot. Res. (Harbin) 1(4): 84. 1981, syn. nov. TYPE: China. Xinjiang: Burqin Xian, Ertix He, alt. 400 in, 28 May 1972, C.. Y. Yang .4-720042 (holotype, XJA). Lappula karelinii was treated by Popov (1953) as a variant of L. tenuis (Ledebour) Giirke, but the latter has a white tuberculate disk and nutlet sides instead of the smooth and polished disk and nutlets characteristic of L. karelinii. W ang (1981) de¬ scribed /,. xinjiangensis C. Y. Yang ex C. J. W ang, but this plant is clearly nonspecific with L. karelinii. Lindelofia stylosa (Karelin & Kirilow) Brand subsp. pterocarpa (Ruprecht) R. Kamelin, comb, et stat. nov. Basionym: Solenanthus ni¬ gricans Schrenk var. pterocarpus Ruprecht, Mem. Acad. Imp. Sci. St. Petersbourg, ser. 7, 14(4): 62. 1869. TYPE: [Kyrgyzstan]. Tian- Shan, Dshaman-daban Pass, 19 July 1857, Osten-Sacken s.n. (holotype, LE). Subspecies pterocarpa was treated by Popov (1953) as a distinct species, although he argued that it probably deserves only a subordinate rank. It differs from subspecies stylosa by its dentate narrow wing on the nutlet margin. In contrast, subspecies stylosa has small, anchorlike prickles along the nut¬ let margin. Onosma liui R. Kamelin & T. Popova, nom. nov. Replaced name: Onosma strigosum Y. L. Liu, Acta Phytotax. Sin. 18: 65. 1980, non O. strigosum Steven, Bull. Soc. Nat. Moscou 24: 594. 1851. TYPE: China. Sichuan: Rangtang, alt. 3,400 m, July 1975, no collector name, #9355 (holotype, CDB1). Onosma setosa Ledebour subsp. transrhym- nense (Klokov ex M. Popov) Kamelin, comb, et stat. nov. Basionym: Onosma transrhym- nense Klokov ex M. Popov, Spiso k Bast. Gerb. FI. SSSR 12: #3564. 1953. TYPE: [Kazakh¬ stan]. Between Chernojarskoe and Pavlodar, on Irtysch River, 2 June 1890, P. Schmidt s.n. (holotype, LE). This subspecies differs from subspecies setosa in having yellow-green hirsute upper parts of the plant and corollas 1 .5 1 . 8( 2 ) cm long. Subspecies setosa has whitish gray hirsute upper parts and corollas 2- 2 . 2 ( 2 . 5 ) cm long. Both subspecies are distributed in Xinjiang Province (China), Russia, and Kazakh¬ stan, but subspecies transrhymnense extends into Mongolia as well. Onosma setosa was treated by Liu (1989) as O. echioides L., but the latter is an entirely different species. Acknowledgments. I am indebted to Ihsan Al- Shehbaz and Tatiana Shulkina for help in the prep¬ aration of this manuscript. Literature Cited Liu, Yu-lan. 1989. Onosma. In: Kung Xian-wu & Wang Wen-tsai (editors), Boraginaeeae. FI. Reipubl. Popul. Sin. 64(2): 1-253. Popov, M. G. 1953. Boraginaeeae. In: B. K. Shishkin (editor), FI. URSS 19: 97-691, 703-718. [English translation by R. Lavoott (Jerusalem) 19: 73-508, 516-531. 1974.] Wang, Ching-jui. 1981. A study of the genus Lappula of China. Bull. Bot. Res. (Harbin) 1(4): 77-100. Novon 3: 263. 1993. New Combinations in Panzerina lanata (Lamiaceae) for the Flora of China Li Xi-wen (Li Hsi-wen) Kunming Institute ol Botany, Academia Sinica, Kunming, Yunnan 650204, People’s Republic of China ABSTRACT. In order to make names available for use in the Flora of China, the following combinations are made: Panzerina lanata var. alaschanica (L. Kuprianaov) H. W. Li, Panzerina lanata var. al¬ bescens (L. Kuprianova) II. W. Li, Panzerina lan¬ ata var. argyracea (L. Kuprianova) H. W. Li, and Panzerina lanata var. parviflora (C. Y. Wu & II. W. Li) H. W. Li. Panzerina was published by Sojak (1981) to re¬ place Panzeria Moench, a later homonym of Pan- zeria J. F. Gmelin. Two species, P. canescens (Bunge) Sojak and P. lanata (L.) Sojak, with four varieties, will be recognized in the Flora of China. Although the genus was recently revised by Krestovskaja (1991), she used Panzeria rather than Panzerina. The needed new combinations in Panzerina are made here. Panzerina Sojak, Cas. Nar. Muz. Praze, Rada Prir. 150: 216. 1981 [1982], TYPE: Panzerina lanata (L.) Sojak, Hallota lanata L. Panzeria Moench, Methodus 402. 1794, not Panzeria J. F. Gmelin, Syst. Nat. 2: 247. 1791. Panzerina lanata (L.) Sojak, Cas. Nar. Muz. Praze, Rada Prir. 150: 216. 1981 [1982], Basionym: Hallota lanata L., Sp. PI. 2: 582. 1753. Panzerina lanata var. lanata Panzerina lanata var. alaschanica (L. Kupri¬ anova) H. W. Li, comb. nov. Basionym: Pan¬ zeria alaschanica L. Kuprianova, Bot. Mater. Gerb. Bot. Inst. Komarova Acad. Nauk SSSR 15: 363. fig. 6. 1953. Panzeria lanata var. alaschanica (L. Kuprianova) 0. V. Tscher- neva, Bast. Centr. Azii, Mater. Bot. Inst. Ko¬ marov 5: 68. 1970. TYPE: “Mongolia aus¬ tralis, Muni-Ula, /V. Przewalslci, 1871.” Panzerina lanata var. albescens (L. Kuprianova) H. W. Li, comb. nov. Basionym: Panzeria albescens L. Kuprianova, Bot. Mater. Gerb. Bot. Inst. Komarova Acad. Nauk SSSR 15: 362. 1953. Panzeria lanata var. albescens (L. Kuprianova) T. Krestovskaja, Novosti Sist. Vyss. Rast. 28: 144. 1991. TYPE: “Mongolia australis, in distr. Gobi Media, prope pag. Er- deni-Dalaj, V. Grubov 7028 . ” Panzerina lanata var. argyracea (L. Kupriano¬ va) H. W. Li, comb. nov. Basionym: Panzeria argyracea L. Kuprianova, Bot. Mater. Gerb. Bot. Inst. Komarova Acad. Nauk SSSR 15: 364. 1953. Panzeria lanata var. argyracea (L. Kuprianvoa) Sergievskaja, FI. Zap. Sib. 12(2): 3431. 1964. TYPE: [Russia] “Tuvink. prov. lacibu Teri-Chol, K. Sobolevskaja, 1947.” Panzerina lanata var. parviflora (C. Y. Wu & II. W. Li) 11. W. Li, comb. nov. Basionym: Panzeria parviflora C. Y. Wu & 11. W. Li Acta Phytotax. Sin. 10: 164. 1965. Panzeria lanata var. parviflora (C. Y. Wu & H. W. Li) T. Krestovskaja, Novosti Sist. Vyss. Rast. 28: 144. 1991. TYPE: [China] “Sinkiang: loc. ign.. Academia Agriculturae Sinkiangensis 2013. (Typus HP).” Literature Cited Krestovkaja, T. 1991. Conspectus generis Panzeria Moench (Lamiaceae). Novosti Sist. Vyss. Rast. 28: 140-144. Sojak, J. 1981 [1982]. K nomenklature Panzeria Moench (Labiatae) a Paulia Korov. (Umbelliferae). Cas. Nar. Muz. Praze, Rada Prir. 150: 216. Novon 3: 264. 1993. Goydera — A New Genus of Asclepiadaceae from Somalia Si grid Liede Abtlg. Spezielle Botanik (Biologie V), Albert-Einstein-Allee 11, 89069 Ulm, Germany Abstract. A new genus of Asclepiadaceae, Goy¬ dera, from Somalia, is described. The new genus is related to the African genera Blyttia and Diplos- tigma and is characterized by colorless latex, a highly fused, adaxially densely pubescent corolla, and a bulge carrying the gynostegium. During an ongoing revision of the subtribe Cy- nanchinae (Asclepiadeae, Asclepiadaceae), several unnamed specimens from Somalia were discovered which, on first sight, were thought to represent a new species of Cynanchum L. While the circum¬ scription of Cynanchum is notoriously difficult, the main question, the inclusion of l incetoxicum W olf, has no bearing on the description of the present taxon, as its characters agree even less with I in¬ cetoxicum than with Cynanchum s. str. The corona, widely used in Asclepiadaceae sys- tematics, provides the first hint that the new species would he ill-accommodated in Cynanchum. W hile both Cynanchum and the new taxon share the basic corona structure of highly fused staminal and in- terstaminal parts, in Cynanchum at least the sta¬ minal parts attain the height of the gynostegium. In the new taxon, however, the indistinguishable sta¬ minal and interstaminal corona parts form an an¬ nulus, which just reaches the lower entrance of the guide rails. The mismatch with Cynanchum is sup¬ ported by a range of other characters. All species of Cynanchum, with exception of the aberrant and probably misplaced North American C. laeve (Mi- chaux) Persoon, possess milky latex. Because latex characters seem to be highly conservative (Liede & Mahlberg, unpublished), this character strongly sup¬ ports the exclusion of the new species from Cynan¬ chum. The highly fused elongated-conical corolla, the long trichomes covering its adaxial surface, the minute size of the flowers, and the sharply geniculate caudicles constitute characters not encountered in Cynanchum. Colorless latex, tiny flowers, and long trichomes on the adaxial surface of the corolla, however, are encountered in the small East African/ Arabian genera Blyttia Arnott and Diplostigma K. Schumann. However, these genera possess a differ¬ ent corona structure, which does not allow inclusion of the taxon into either genus. A new genus thus has to be created. Goydera Liede, gen. nov. TYPE: Goydera so- maliense Liede. Blyttiae et Diplostigmae affine, sed corolla alte con- nata et intus dense pubescencenti differt. Corona a gy- nostegio originans, ex partibus staminodialibus et inter- staminodialibus constans. Plants containing colorless latex. Inflorescences one per node, constituting a scorpioid cyme or an inflorescence type derived from it. Corolla fused for more than half of its height, adaxially densely cov¬ ered with long, slender trichomes. Corona gynos- tegial, consisting of fused staminal and interstaminal parts, shorter than the gynostegium. Gynostegium elevated by a bulge. Pollinia pendulous. The taxon is named after David J. Goyder, Kew, who directed my interest to this unusual plant. Goydera sonialiense sp. nov. TYPE: Somalia. Hiiraan Region: Bulo Burti District, 03°46'N, 44°57'E, 13 km NNE of Dharyio, ± 300 m, 29 Nov. 1986, Kuchar 17 IBB (holotype, K). Figure 1 . Herba volubilis, foliis lineariformibus, corolla conica, corona brevi annulari. Plants twining, 50-70 cm high, sparsely basi- tonically branched, sarmentose, runners corky. Shoots herbaceous, glabrous; internodes 4 7 cm long, 0.6-1 mm diam.; latex sparse, colorless. Leaves sessile, lacking colleters; leaf blades herbaceous, lin¬ ear, 12-15 mm long, 0.5— 0.7 mm wide, apically acute, adaxially glabrous, abaxially sparsely indu- mented with erect, 200-250-/um-long trichomes; indumentum restricted to veins and margins. Inflo¬ rescences umbelliforin, 10- 1 5-flowered, 8 12 flow¬ ers open at the same time, pedunculate; peduncles 5-7 mm long, glabrous. Floral bracts 0.4-0. 5 mm long, 0.2 0.3 mm wide at the base, ovate, with trichomes; pedicels 2-3 mm long, glabrous. Buds 2.5-3 mm long, 1.3- 1.5 mm diam., conical, aes¬ tivation imbricate. Calyx basally fused, rotate, cil- iate; sepals 1.7- 1.8 mm long, 0.6-0. 7 mm wide, triangular, apically acute. Corolla elongated-coni¬ cal, fused for % of its height, 3.5-4 mm high, 1- 1.5 mm wide, rugose, glabrous, ahaxiallv yellow to brown, adaxially densely covered with long tri- Novon 3: 265-267. 1993. 266 Novon Figure 1. Goydera somaliense Liede. — A. Habit, inflorescences, and fruit. — B. Flower. — C. Flower, corolla and corona removed showing gynostegium and the bulge elevating it. — D. Pollinarium. — E. Stylar head. — F. Seed, aseta side. — G. Seed, seta side ( Kuchar 17188). Volume 3, Number 3 1993 Liede Goydera from Somalia 267 chomes; corolla lobes erect, ovate to lanceolate, apically acute. Corona cyathiform, 350-450 mm high, shorter than the gynostegium; staminal and interstaminal parts completely fused, upper corona margin straight. Gynostegium elevated by a bulge of 300-350 fim, 750-850 /um high without bulge, 1,200-1,300 /am diam. Anthers broader than high, trapezoidal, abaxially planar; anther wings 350 400 /am long, extending along the whole length of the anther, convergent, consisting ol proximal and distal ridge, proximal ridge curved, space between distal and proximal ridge bristly. Anther wings of adjacent anthers parallel to each other, in the same plane as the anther. Connective appendages 300-350 /am long, 350-400 /um wide, debate, broader than the stamen. Pollinarium: corpusculum 1 50 /am long; caudicles (sub-)basally inserted at the corpusculum, 180-200 fim long, cylindrical, convexly recurved, thickened at the insertion of the pollinium, genic¬ ulate; pollinia apically attached to the caudicles, 340-380 //m long, 160-180 /am wide, ovate in cross section, clavate. Stylar head 650-750 /am diam., 400-450 ^nn high, forming a nose at the upper end of the corpusculum; upper part 200- 250 /um high, equaling the lower part in height, conical. Follicles always one per flower, pendulous, 40-45 mm long, 4-5 mm wide, obclavate, round in cross section, apically strongly beaked, wingless; light brown, longitudinally grooved. Seeds 3.5-4 mm long, 2-2.2 mm wide, pyriform, medium brown, dorsally papillose with knottily arranged papillae, ventrally papillose with papillae arranged in longi¬ tudinal ridge; coma 10-12 min long. Chromosome number: unknown. Distribution. Somalia, 4°05'-2°40'N, 42°20' 45°30'E, on limestone, in Acacia Commiphora bushland. Flowering time. November-December, May. Paratypes. SOMALIA. Hiiraan Region: 03°53'N, 44°50'E, El Dabbo 23 km E of Dhanabo, ± 345 m, 24 May 1983, Gillett A' Hemming 2440a (K). Ruin Rurti District: 04°02'N, 45°23'E, 4 km N of Bulo Burti, ± 600 m, 24 Dec. 1972, Bally & Melville 15308 (K, MO). Hiiraan District: 04°03'N, 45°45'E, escarpment above Yasoomman, 300 m, 28 May 1989, Thulin &: Abdi Dahir 6480. (K). Acknowledgments. I thank Graziela Hintze, Lllm, for providing the line drawing and F. Weberling, Ulm, for generously providing all research facilities. Work was made possible by a DFG habilitation grant (LI 496/2-1 and LI 496/2-2). Finally, I thank F. Albers and Ulrich Meve, Munster (FRG), lor many stimulating discussions. Nomenclatural Changes and New Taxa in Claytonia (Portulacaceae) in Western North America John M. Miller BioSystems Analysis, Inc., 303 Potrero St., Suite 29-203 Santa Cruz, California 95060, U.S.A. Kenton L. Chambers Department of Botany and Plant Pathology, Oregon State University Corvallis, Oregon 97331, U.S.A. Abstract. Claytonia here includes all annual Montia species with principally basal leaves and scapiform inflorescences bearing one pair of free or fused cauline leaves, three morphologically distinc¬ tive diploids define the basic species of a polymorphic polyploid complex formerly named C. perfoliata sensu lato. In this complex, we propose two new subspecies, C. perfoliata subsp. intermontana and C. parviflora subsp. grandiflora, and four subspe- cific combinations. A perfoliate-leaved subspecies of C. exigua is newly recognized. through the work of Swanson (1966), Nilsson (1967). McNeill (1975), and others, the genus Clay¬ tonia is coming to be accepted in a more inclusive sense than was common in monographic and floristic works of the first half of this century (e.g., Howell, 1893; Pax & Hoffmann, 1934; Kerris, 1944; Hitch¬ cock, 1964). In what now seems to be an unjustified generic split, certain taxa of rhizomatous perennials and annuals that were classified in Claytonia prior to 1893 had been segregated into Montia. Mor¬ phological evidence including pollen (Nilsson, 1967), growth habit (Swanson, 1966), and chromosome numbers (Lewis & Suda, 1968; Fellows, 1975; Mil¬ ler & Chambers, 1977), along with numerical tax¬ onomic analysis (McNeill, 1 975), now [daces the following former Montia species more naturally in Claytonia: C. arenicola L. F. Henderson, C. cor- difolia S. Watson, C. exigua Torrey & A. Gray (= C. spathulata Douglas ex Hooker, nom. illeg.), C. gypsophiloides Fischer & C. A. Meyer, C. perjol- iata Donn ex Willdenow, C. saxosa T. S. Brandegee, and C. sibirica L. These taxa differ from Montia, as redefined by McNeill (1975), in having their principal leaves in a basal rosette and in possessing inflorescences borne on scapelike stems with only a single f>air of cauline leaves, which may be united into a bilobed or toothed disk. We propose to add C. parviflora Douglas ex Hooker and C. rubra (Howell) Tidestrom to the above list of recognized species that were earlier assigned to Montia. The latter genus, while still diverse (nine sections for only 13 species, as recognized by McNeill (1975)), differs from Claytonia in having numerous stem leaves, which may be alternate or opposite, in lacking a basal leaf rosette, and in having pantocolpate pollen rather than the trizonocolpate type characteristic of Claytonia. In habit, Montia species are annuals or rhizomatous to stoloniferous perennials, lacking the fleshy taproots or tuberlike underground storage or¬ gans seen in many perennial species of Claytonia. the group of taxa that had previously been united as the polymorphic species Claytonia perfoliata sensu lato was shown by Miller (1976, 1978) to be a polyploid, largely autogamous pillar complex based upon three morphologically distinct diploid entities (n = 6). When raised to species rank, these entities fall within the already named taxa C. perfoliata, C. parviflora, and C. rubra. Together with the puta¬ tively ancestral diploids, each of these species in¬ cludes a variable array of eupolyploids ranging from tetraploid to, in some cases, as high as decaploid (Miller, 1978). Many polyploid races have inter¬ mediate morphologies suggestive of an alloploid or¬ igin, but some races, especially at the tetraploid level, are scarcely distinguishable from the basic diploids and may be intraspecific autoploids. Our taxonomic proposal for the C. perfoliata complex sensu lato is to recognize as species the three taxa mentioned above and to divide each spe¬ cies into a limited number of broadly defined sub¬ species. Within each species, one subspecies con¬ tains the basic diploid races along with polyploids that are scarcely distinguishable from them. The one or more remaining subspecies are framed around clusters of morphologically similar polyploid races, usually having a more or less well defined geograph¬ ical range. The following lectotypifications, new taxa, and new combinations are necessary for this taxo- Novon 3: 268-273. 1993. Volume 3, Number 3 1993 Miller & Chambers Claytonia in Western North America 269 nomic revision. Claytonia exigua subsp. glauca, C. parviflora subsp. grandijiora , C. parviflora subsp. viridis, C. perjoliata subsp. mexicana, and C. rubra subsp. depressa appeared earlier in The, Jcpson .Manual (Hickman, 1993) and are here be¬ ing validated. Claytonia perfoliata Donn ex Willdenow, Sf > . FI. 1: 1186. 1798. Limnia perfoliata (Donn ex Willdenow) Haworth, Syn. PI. Succ. 12. 1812. Montia perfoliata (Donn ex Willdenow) How¬ ell, Erythea 1: 38. 1893. TYPE: Herbarium Willdenow No. 4984, Cl. perjoliata 2, Mus. Pot. Berol. Film Nr. 816a (lectotype, here des¬ ignated, B-W [photograph seen]). Authorship of this well-known name is often at¬ tributed to James Donn, who, however, only pub¬ lished it as a nomen nudum. Of the three sheets in Willdenow’s herbarium, those numbered “1” and “2” each hold a beautifully prepared specimen, probably of garden origin. We select the sheet num¬ bered “2” as the lectotype. A third sheet consists of four fragments only; one of these is a teratological inflorescence with separate cauline leaves, which evidently entered into Willdenow's description, in pari. This “inaccuracy" (Sims, 1811) may have led to some confusion about Willdenow’s plant among contemporary botanists, who were familiar with the typically perfoliate-leaved species because of its rap¬ id, spontaneous spread among European botanical gardens after being introduced to Kew Gardens in I 796. The date and place of introduction were given by Sims (181 1), who credits Archibald Menzies as the discoverer of the species. There is a sheet at BM on which three specimens are marked as having been collected by Menzies on the “North-west coast of America. 1792-3-4" (Photo No. 917-1, NY). Claytonia perfoliata Donn ex Willdenow subsp. mexicana (Hydberg) Miller & Chambers, comb. nov. Basionym: Limnia mexicana Ryd¬ berg, N. Amer. El. 21: 309. 1932. Montia mexicana (Rydberg) Pax & K. Hoffmann, Nat. Pflanzenfam. ed. 2 16c: 259. 1934. Claytonia tolucana Holub, Preslia 47: 328. 1975. TYPE: Mexico. Mexico: Nevada de Toluca, Rose & Painter 7924 (holotype, US). Limnia cuprea Heller, Muhlenbergia 2: 279. 1907, syn. nov. Montia perjoliata forma cuprea (Heller) J. T. Howell, Leaf!. W. Bot. 5: 106. 1948. TYPE: U.S.A. California: Monterey County, Pacific Grove, Heller 8501 (holotype, BKL [at NY]; isotypes, l)S, NY, WTU). Limnia platyphylla Rydberg, N. Amer. FI. 21: 307. 1932, syn. nov. Montia platyphylla (Rydberg) Pax & K.. Hoffmann, Nat. PHanzenfam. ed. 2 16c: 259. 1934. Claytonia platyphylla (Rydberg) Holub, Preslia 47: 328. 1975. TYPE: U.S.A. California: [Santa Barbara County,] Gaviota Pass, Brewer 484 (holotype, US). Limnia guadalupensis Rydberg, N. Amer. FI. 21: 311. 1932, syn. nov. TYPE: Mexico. Baja California: Guadalupe Island, Palmer 15 (holotype, NY; iso- types, E, NY). The diploid elements of the above subspecies were characterized by Miller (1978: figs. 6-9) as having linear juvenile leaves and broadly deflate, mucronate adult leaves; their fully perfoliate cauline leal disk has two mucronate teeth. Together with related pol¬ yploids they range widely through the Coast Ranges of California to Baja California, Arizona, and main¬ land Mexico, reaching the highlands of Guatemala. Claytonia perfoliata Donn ex \\ illdenow subsp. intermontana Miller & Chambers, subsp. nov. TYPE: U.S.A. Nevada: Churchill County, Highway 50, 8 mi. E ol Eastgate, 7 mi. W ol Carroll Summit, 2 n = 24 [by J. R. Swanson], II. K. Sharsmith 4707 (holotype, OSC). A subsp. perfoliata caulibus foliisque patulis vel erectis saepe rubello-viridis, discis perfoliatis saepe heinidivisis differt; a subsp. mexicana caulibus foliisque saepe rubello- viridis, foliis adultis ovato-rhornbeis vel subdeltatis raro mucronatis, discis perfoliatis saepe heinidivisis differt. Plants annual, 2 20 cm tall, often red-pigmented (shade-grown plants usually green); stems and leaves erect to lax and spreading; juvenile rosette leaves linear to oblanceolate; adult leaves with petiole 2 or more times the blade, blade 0.5 4 cm, ovate-rhom¬ boid to subdeflate, acute, rarely mucronate; cauline leaf-disk perfoliate, often notched or divided to the stem on one side; inflorescence usually 5-20-flow- ered, compact and sessile on the leaf-disk or race- mosely elongated up to 5 cm; flowers self-pollinating; sepals 2 3.5 mm, orbicular; petals 2.5 4 mm, white or pinkish; capsule 2.5-3 mm; seeds 3, shiny black, with conspicuous elaiosome; 2 n = 24, 36, 48. This subspecies is distributed widely in the inter- montane region east of the Cascade Range, Sierra Nevada, and Peninsular Ranges, from British Co¬ lumbia to northern Baja California and Arizona, east to the Rocky Mountains. The genetic influence ol Claytonia rubra is seen in the frequent reddish coloration of the polyploids comprising this taxon, and their characteristic ovate-rhomboid leaf shape is intermediate between the deltoid leaves of C. rubra and the linear ones of C. parviflora. Furthermore, polyploid C. parviflora subspp. parviflora and utah- ensis (see below) are frequently sympatric and in- tergradent with C. perfoliata subsp. intermontana , 270 Novon in a pattern similar to t he intergradation of polyploid C. perfoliata subsp. perfoliata and C. parviflora subsp. parviflora in the cismontane region of the Pacific States. Claytonia rubra (Howell) Tidestrom, Contr. U.S. Natl. Herb. 25: 188. 1925. Basionym: Montia rubra Howell, Erythea 1: 38. 1893. Limnia rubra (Howell) Heller, Muhlenbergia 6: 84. 1910. Claytonia perfoliata Donn ex Willd- enow var. rubra (Howell) Poellnitz, Kepert. Spec. Nov. Regni Veg. 30: 301. 1932. TYPE: U.S. A. Washington: [Klickitat or Yakima County,] Simcoe (as “Cimcoe”) Mountains, June 1880, T. J. Howell s.n. (lectotype, here des¬ ignated, ORE; isolectotype, US). Howell cited no type or other specimens in his description of Montia rubra. The above lectotypi- fication allows this name to be associated with the common and widespread diploid entity that forms one ol the three pillars of the Claytonia perfoliata sensu lato polyploid complex. The name Claytonia parviflora var. depressa A. Gray, which in the literature has often been applied to C. rubra subsp. rubra , is lectotypified below to apply to a different, wholly polyploid subspecies. Claytonia rubra (Howell) Tidestrom subsp. de¬ pressa (A. Gray) Miller & Chambers, comb, nov. Basionym: Claytonia parviflora Douglas ex Hooker var. depressa A. Gray, Proc. Amer. Acad. Arts 22: 281. 1887. Montia parviflora (Douglas ex Hooker) Howell var. depressa (A. Gray) B. L. Robinson, Syn. FI. N. Amer. 1: 274. 1897. Montia depressa (A. Gray) Suks- dorf, Deutsche Bot. Monatsschr. 16: 221. 1898. Limnia depressa (A. Gray) Rydberg, Bull. Tor- rey Bot. Club 33: 139. 1906. Claytonia par¬ viflora Douglas ex Hooker subsp. depressa (A. Gray) Piper, Contr. U.S. Natl. Herb. 1 1: 250. 1906. Montia perfoliata (Donn ex Willdenow) Howell var. depressa (A. Gray) Jepson, FI. Calif. 1: 471. 1914. Claytonia perfoliata Donn ex Willdenow var. depressa (A. Gray) Poellnitz, Repert. Spec. Nov. Regni Veg. 20: 301. 1932. IYPE: U.S. A. Washington: [San Juan Coun¬ ty,] San Juan Island, Lyall in 1858 (lectotype, here designated, GH). Montia humifusa Howell, FI. N.W. Ainer. 1: 90. 1897, syn. nov. Limnia humifusa (Howell) Rydberg, FI. Plains N. Amer. 313. 1932. Claytonia humifusa (Howell) Holub, Preslia 47: 328. 1975. TYPE: U.S. A. Oregon: [Umatilla County,] Milton, 18 May 1896, T. J. Howell s.n. (holotype, ORE; isotype, DS). Montia latifolia Suksdorf, Deutsche Bot. Monatsschr. 16: 222. 1898, syn. nov. Claytonia latifolia (Suks¬ dorf) Suksdorf, Werdenda 1: 222. 1923, non Shel¬ don, 1894. TYPE: U.S. A. Washington: [Klickitat County,] Bingen, Suksdorf 1881 (holotype, WS; isotypes, F, GH, NY, UC, US). Montia interrupta Suksdorf, Deutsche Bot. Monatsschr. 16: 222. 1898, syn. nov. Claytonia interrupta (Suksdorf) Suksdorf, Werdenda 1: 222. 1923. Lim¬ nia interrupta (Suksdorf) Rydberg, N. Amer. FI. 21: 308. 1932. TYPE: U.S. A. Washington: [Klick¬ itat County,] Bingen, Suksdorf 2009 (holotype, WS; isotypes, F, GH, NY, UC, US). Claytonia cupulata Suksdorf, Werdenda 1: 11. 1923, syn. nov. TYPE: U.S. A. Washington: [Klickitat County,] Bingen, Suksdorf 10169 (holotype, WS; isotypes, CAS, DS, NY, OSC, UC, US). Claytonia rubra .subsp. depressa differs from subspecies rubra in being wholly polyploid (tetraploid and hexaploid) and in having ovate-elliptic leaf blades rather than debate, sometimes cordate blades as in subspecies rubra. It is generally found at lower elevations than subspecies rubra, occurring in coast¬ al sandy sites from British Columbia to northern California, as well as interior regions from Wash¬ ington to northern Nevada and east to Montana. In an earlier paper, it was referred to as the “sagebrush hexaploid" and the “coastal tetraploid” (Miller, 1978: figs. 1, 6). Its foliage may be reddish, as is common in subspecies rubra, or green; the epithet “depressa” is descriptive of its often cushionlike growth form. Claytonia parviflora Douglas ex Hooker subsp. grandiflora Miller & Chambers, subsp. nov. TYPE: U.S. A. California: Calaveras County, San Antonio Creek S of Sheep Ranch, on the road to Murphy’s, Miller 666 (holotype, OSC; isotypes, CAS, RSA, SD). A subsp. parviflora floribus allogamis, petalis 4-6 mm quam sepalis duplo triplove longiorihus differt. Plants annual, 5-20 cm tall, sometimes red-pig¬ mented; stems and leaves erect or somewhat spread¬ ing; juvenile and adult leaves linear; cauline leaf- disk perfoliate, rarely notched or divided; inflores¬ cence longer than the leaves, 5-30-flowered, the flowers racemosely arranged and well exserted above the perfoliate disk; sepals 2-3.5 mm, ovate; petals 4-6 mm, white or pink; stamens releasing pollen 1-2 days before the stigmas are receptive, the breeding system allogamous; capsules 2.5-3 mm; seeds 3, shiny black, with conspicuous elaiosome; 2n = 12. Of the three morphologically divergent diploid taxa forming the bases for the Claytonia perjoliata polyploid complex, C. parviflora subsp. grandiflora Volume 3, Number 3 1993 Miller & Chambers Claytonia in Western North America 271 is the only one having an outcrossing, sell-incom¬ patible breeding system. A similar floral syndrome and breeding system occur in the diploid annual species C. gypsophiloides, a basal and putatively primitive element in the C. exigua polyploid com¬ plex. Clayton in parviflora subsp. grandiflora oc¬ cupies a natural range in the mixed oak-pine wood¬ lands on the western slopes of the Sierra Nevada at ISO 1,200 m elevation. It sometimes is sympatric with polyploid, autogamous races of subspecies par¬ viflora. Claytonia parviflora Douglas ex Hooker subsp. utahensis (Rydberg) Miller & Chambers, comb, nov. Basionym: Limnia utahensis Rydberg, Bull. Torrey Bot. Club 39: 314. 1912. Clay¬ tonia utahensis (Rydberg) Tidestrom, Contr. U.S. Natl. Herb. 25: 188. 1925. Claytonia perfoliata Donn ex Willdenow var. utahensis (Rydberg) Poellnitz. Repert. Spec. Nov. Regni Veg. 30: 302. 1932. Montia utahensis ( Ryd¬ berg) Fax & K. Hoffmann, Nat. Pflanzenfam. ed. 2 16c: 259. 1934. Montia perfoliata (Donn ex Willdenow) 11 owell var. utahensis (Rydberg) Munz, Aliso 4: 90. 1958. TYPE: li.S. A. Utah: [Washington County,] St. George, Calmer in 1877 (holotype, NY). Claytonia parviflora subsp. utahensis comprises populations of generally diminutive plants whose spathulate to narrowly lanceolate adult leaf blades are 4 mm or more wide, and which occupy the Peninsular and Desert ranges of southern California, extending north on the east flank of the Sierra Ne¬ vada and east to southern Nevada, Utah, and north¬ ern Arizona. The plants are autogamous and prin¬ cipally tetraploid. Intergradent populations have been noted between this subspecies and both C. perfoliata subsp. intermontana and C. parviflora subsp. vir- idis (see below). Claytonia parviflora Douglas ex Hooker subsp. viridis (Davidson) Miller & Chambers, comb, nov. Basionym: Montia spathulata (Douglas ex Hooker) Howell var. viridis Davidson, Bull. S. Calif. Acad. Sci. 5: 61. 1907. Montia ex¬ igua (Torrey & A. Cray) Jepson var. viridis (Davidson) Jepson, FI. Calif. 1: 473. 1914. Claytonia exigua Torr. & A. Gray var. viridis (Davidson) Poellnitz, Repert. Spec. Nov. Regni Veg. 30: 313. 1932. Limnia viridis (Davidson) Rydberg, N. Amer. FI. 21: 313. 1932. Clay¬ tonia spathulata Douglas ex Hooker var. vir¬ idis (Davidson) Munz, FI. S. Calif. 713. 1974. Claytonia viridis (Davidson) Holub, Preslia 47: 328. 1975. Claytonia perfoliata Donn ex Willdenow7 subsp. viridis (Davidson) Fellows, Madrono 23: 297. 1975. TYPE: U.S. A. Cal¬ ifornia: Los Angeles County, San Gabriel Moun¬ tains, Big Rock Creek, Hasse & Davidson 1507 (holotype, LAM [at RSA]; isotype, GH). Claytonia tenuifolia Torrey & A. Gray, FI. N. Amer. 1: 201 . 1838, syn. nov. Claytonia spathulata Douglas ex Hooker var. tenuifolia (Torrey & A. Gray) A. Gray, Proc. Amer. Acad. Arts 22: 282. 1887. Mon¬ tia tenuifolia (Torrey & A. Gray) Howell, Erythea 1: 38. 1893. Limnia tenuifolia (Torrey & A. Gray) Rydberg, N. Amer. FI. 21: 313. 1932. Montia spathulata (Douglas ex Hooker) Howell var. ten¬ uifolia (Torrey & A. Gray) Munz, Man. S. Galit. Bot. 598. 1935. TYPE: U.S. A. California: Douglas s.n. (lectotype, here designated, GH; isolectotype, NY). Fellows (1975) demonstrated that the above tax¬ on is not related to the Claytonia exigua species- complex, as had earlier been assumed by most bo¬ tanical authors. Instead, it is a member of the G. perfoliata complex sensu lato, based on its seed morphology (smooth, shiny testa and enlarged elaio- some), nonglaucous herbage, and chromosome base number (x = 6, 2 n = 24, 36). Although placed in C. perfoliata by Fellows, it better belongs with (.. parviflora because its rosette leaves range from linear to narrowly oblanceolate. The cauline leaf pair strikingly mimics that of C. exigua , often being linear, divergent, and unfused (e.g., the type spec¬ imens of C. tenuifolia, cited above; Miller, 1978: figs. 14, 16) or partly fused on one side to form a crescent-shaped or two-pronged disk. This latter type of disk is present on the specimens comprising the holotype and isotype sheets of M. spathulata var. viridis, and these plants’ basal leaves have a broader blade than is usual for the subspecies. 1 he San Gabriel Mountains, along with other units of the Transverse Ranges of southern California, contain diverse morphological forms of the C. perfoliata parviflora — rubra complex; these include diploids as well as several variable polyploids. Leaf-shape variation in subspecies viridis may be due to in- trogression from some broader-leaved taxa like sub¬ species mexicana or subspecies rubra. Other pop¬ ulations have been found in this region that may represent diploid introgressants of C. perfoliata with C. rubra (Miller, pers. obs.). It is not yet clear whether any additional entities within this hybrid complex are worthy of taxonomic recognition; at present, we prefer the broadly defined, and hence more variable, taxa outlined in this report. Claytonia tenuifolia is a notable addition to the synonymy of C. parviflora subsp. viridis, above. Since the time of Asa Gray, almost all authors deal- 272 Novon ing with the flora of California have associated this name with plants referable to Claytonia exigua (C. spathulata of previous treatments). However, the type specimens at GH and NY, collected by David Douglas at some unknown site in California, sur¬ prisingly prove not to belong to the latter species. Although they are similar to C. exigua in habit, their seed morphology is unequivocally that of C. parviflora. For any combinations made at the va¬ rietal level, therefore, the epithet tenuifolia has pri¬ ority. Claytonia exigua Torrey & A. Cray subsp. glau- ca (Torrey & A. Cray) Miller & Chambers, comb. nov. Basionym: Claytonia parviflora Douglas ex Hooker [var.] /3 glauca Torrey & A. Cray, FI. N. Amer. 1: 200. 1838. Limnia glauca (Torrey & A. Cray) Hydberg, N. Amer. FI. 21: 311. 1932. Montia perfoliata (Donn ex Willdenow) Howell subsp. glauca (Torrey & A. Cray) Ferris, 111. FI. Pacific States 2: 127. 1944. Montia perfoliata (Donn ex Willdenow) Howell forma glauca (Torrey & A. Cray) J. T. Howell, Leafl. W. Bot. 5: 106. 1948. TYPE: U. S.A. [Oregon:] “Oregon R.,“ Nuttall s.n. (lectotype, here designated, GH; probable iso- lectotype, NY [labeled “Rocky plains of Wah- lainet”]). Montia spathulata (Douglas ex Hooker) Howell var. dis- ciformis Suksdorf, Deutsche Bot. Monatsschr. 16: 222. 1898, syn. nov. Claytonia spathulata Douglas ex Hooker var. disciformis (Suksdorf) Suksdorf, Werdenda 1: 10. 1923. TYPE: U.S.A. Washington: Klickitat County, Major Creek between Bingen and Lyle, Suksdorf 2095 (holotype, WS; isotypes, F, GH, NY, UC, US). Montia pallida M. Peck, Proc. Biol. Soc. Wash. 47: 185-186. 1934, syn. nov. TYPE: U.S.A. Oregon: [Marion or Polk County,] near Salem, Peck 1699 (holotype, WILLU [at OSC]). I he name Claytonia exigua Torrey & A. Gray, of 1838, must replace the earlier C. spathulata Douglas ex Hooker, 1832, because the latter is a homonym of C. spatulata F!aton, 1824. The epithet glauca was attributed by Torrey and Cray to “Nutt.! mss.” but Nuttall’s tags on the type sheets show he intended the name for a variety of C. gypsophi- loides rather than C. parviflora as was published in Flora of North America. Hence the basionym should not be attributed to “Nuttall ex Torrey & A. Gray.” The subspecies is entirely diploid (2 n = 16) and forms the perfoliate leaf-disk equivalent, within (.. exigua , of C. perfoliata. The differences in habit, chromosome number, and seed morphology between these two species-complexes were touched on by Fellows (1975) in his discussion of C. par¬ viflora subsp. viridis. That C. exigua includes plants with a completely perfoliate disk, as well as those with a deeply two-lobed disk or with separate cauline leaves, was known to Thomas Howell (1893), who included in his description (under Montia spathu¬ lata, p. 38) the phrase “involucral bracts . . . wholly united and the disk shorter on one side . . ..” By later authors, however, subspecies glauca has either been described as a separate species or submerged in C. perfoliata. It may be useful to clarify the synonymy, cited here and elsewhere, from the pub¬ lications of Wilhelm Suksdorf. For the new taxa he described in 1898 in the Deutschen Botanischen Monatsschrift, Suksdorf gave first a collection num¬ ber and a name in the genus Claytonia, followed by an equal sign and a name in the genus Montia. Rydberg and other authors have cited these two names as co-equal, as though both were validly published. However, Suksdorf s 1923 paper in Wer¬ denda makes clear that only the names in Montia were intended to be published in I 898; in fact, the 1923 paper contains many transfers of these taxa from Montia back to Claytonia. The names in Clay¬ tonia given in Suksdorfs 1898 paper were merely references to the herbarium labels he used in dis¬ tributing his collections, prior to the revision of Mon¬ tia by Howell in 1893. Acknowledgments. We thank Charles Fellows for contributions to the research, John McNeill for bib¬ liographic advice and assistance, and the curators of the cited herbaria for making available type col¬ lections and photographs. Our research was sup¬ ported by National Science Foundation grant BSR 8600117. Literature Cited Fellows, C. E. 1975. Chromosome counts and a new combination in Claytonia sect. I Amnia (Portulaca- ceae). Madrono 23: 296-297. Ferris, R. S. 1944. Portulacaceae. Pp. 119 136 in L. Abrams (editor). Illustrated Flora of the Pacific States, vol. II. Stanford Univ. Press, Stanford, California. Hickman, J. C. Editor. 1993. The Jepson Manual. Higher Plants of California. Univ. California Press, Berkeley. Hitchcock, C. L. 1964. Portulacaceae. Pp. 225-249 in C. L. Hitchcock, A. Cronquist, M. Ownbey & J. W. Thompson (editors), Vascular Plants of the Pacific Northwest, part 2. University of Washington Pub¬ lications in Biology, vol. 17. Univ. Washington Press, Seattle. Howell, T. J. 1893. A rearrangement of American Portulacaceae. Erythea 1: 29-41. Lewis, W. H. & Y. Suda. 1968. Karyotypes in relation to classification and phylogeny in Claytonia. Ann. Missouri Bot. Card. 55: 64 67. Volume 3, Number 3 1993 Miller & Chambers Claytonia in Western North America 273 McNeill, J. 1 975. A generic revision of Portulacaeeae tribe Montieae using techniques of numerical tax¬ onomy. Canad. J. Bot. 53: 789 809. Miller, J. M. 1976. Variation in populations of Clay¬ tonia perfoliata (Portulacaeeae). Syst. Bot. 1: 20 34. - . 1978. Phenotypic variation, distribution and relationships of diploid and tetraploid populations of the Claytonia perfoliata complex (Portulacaeeae). Syst. Bot. 3: 322 341. - & K. L. Chambers. 1977. Chromosome num¬ bers and relationships of C. saxosa and C. arenicola. Madrono 24: 62-63. Nilsson, 0. 1967. Studies in Montia L. and Claytonia L. and allied genera. III. Pollen morphology. Grana Palynol. 7: 279 — 363. Pax, F. & K. Hoffmann. 1934. Portulacaeeae. Pp. 234 262 in A. Engler & H. Harms (editors), Die Natiirlichen Pflanzenfamilien, ed. 2, bd. 16c. Wil¬ helm Engelmann Verlag, Leipzig, Germany. Sims, J. 1811. Claytonia perfoliata. Perfoliate Clay¬ tonia. Bot. Mag. 33: No. 1336. Suksdorf, W. N. 1898. Washingtonische pflanzen. Deutsche Bot. Monatsschr. 16: 220 222. - . 1923. Washingtonische pflanzen 111. Wer- denda 1(2): 1-14. Swanson, J. R. 1966. A synopsis of relationships in Montioideae (Portulacaeeae). Brittonia 18: 229-241. A New Mouriri (Melastomataceae) from the Dominican Republic Thomas Morley Department of Plant Biology, University of Minnesota, St. Paul, Minnesota 55108, U.S.A. Abstract. A new Mouriri in the section Neso- phytum lias been found at 1,300 in in the Cordillera Central, Dominican Republic. Collected only in fruit, the new species can be distinguished primarily by its mostly terminal infructescence, large fruits and seeds, thick calyx lobes, and hypanthium shape, greatly modified in fruit. A party of botanists collecting at high elevations in the Dominican Republic in 1986 found a fruiting Mouriri that they thought was probably a new spe¬ cies. On first seeing the material I thought otherwise, since at least when in fruit the plant superficially resembles M. spathulata of Cuba and the Domin¬ ican Republic. However, the collectors were right, as a study of details of fruit structure and other features shows. Mouriri crassisepala Morley, sp. nov. TYPE: Dominican Republic. Prov. La Vega: Cordillera Central, ladera del N de la Loma de La Sal, frente al Valle de Jarabacoa, vegetacion de bosque secundario, con pinos, pero arboles la- tifoliados del bosque humedo en la cima, 19°05'N, 70°35'W, alt. 1,300 m, 21 May 1986, T. Zanoni, M. Mejia A R. Garcia 36370 (holotype, FLAS; isotype, JBSD). Figure I. Arbor usque 12 in alta; laminae 2. 7-4. 8 cm longae, 1.7-3. 3 cm latae, 0.36-0.48 mm crassae, margine re- voluta; cryptae stomatophorae 60-65 /^m diametro; fruc- tus plerumque terminales, raro axillares, 14-32 mm dia¬ metro; lobi calycis ad maturitatem fructus persistentes, 0.3-0. 7 mm crassi; cicatrix styli 1.5-2. 5 mm diametro; diameter intra hypanthii ad basim 4. 2-6.6 mm; semina 14-16.5 mm longa. Small tree to 1 2 m high, glabrous except for the inflorescence; young twigs round to 4-angled or rare¬ ly with small ridges at the angles. Petioles 1.5-3. 5 mm long; blades 2. 7-4. 8 cm long, 1.7-3. 3 cm wide, 0.36-0.48 mm thick, elliptic to ovate-elliptic, the margin strongly revolute, the apex rounded or seem¬ ing broadly acute through revolution of the margins, rarely mucronate, the base acute to broadly so; midrib plane above, plane to very low rounded or very low-rectangular or with two low lateral ridges below; lateral nerves not visible. Midrib xylern tu¬ bular; stomatal crypts usually Type III (Morley, 1976), Type II when small, averaging 60-65 pm diam., 28-35 pm high, 30-42 per sq. mm (extremes 35-90 pm diam., 25-40 pm high, 24-57 per sq. mm); upper epidermis 1-2 cells thick; mucilage walls none; hypodermis present; terminal sclereids from stellate with a horizontal central body and columnar branches to thick-columnar with branches at each end and no central body. Flowers unknown; fruits mostly terminal, seldom in the upper axils; peduncles 1-2 per side, each 1-3-fruited, 3-19 mm long to base of most distal pedicel measured along the axes and with 1-3 internodes in that length; bracts de¬ ciduous before fruiting. True pedicels 1.5-5 mm long, they and the axes of the inflorescence minutely puberulent or granular to glabrous; calyx lobes per¬ sistent in fruit, then 1.5- 1.9 mm long from stamen attachment, 2. 4-3. 2 mm wide, 0.3-0. 7 mm thick, rounded, seeming to have split apart ca. 0.5-0. 8 mm at anthesis; petal and stamen scars dark, barely or not discernible, flush with the surface, petal scars 1.3- 1.5 mm long (radially), 1.3- 1.4 mm wide (cir¬ cumferentially) or indistinct; antesepalous stamen scars 0.3-0. 5 mm long, 0.8- 1.2 mm wide, the antepetalous ones 0.4 mm long and 0.6 mm wide or indistinct; style scar 1.5-2. 5 mm diam.; inside diameter of base of fruiting hypanthium 4. 2-6. 6 mm, less at apex, the walls thus overhanging, the overhang great in 1 -seeded fruits, less in several- seeded ones; apex of fruit within hypanthium convex to flat. Fruits crowned with the calyx, 1-4-seeded, subglobose when 1 -seeded, then 14-22 mm diam., lobed according to the number of seeds when seeds more than 1, up to 32 mm diam. when seeds 2-4, Figure 1. A-F, Mouriri crassisepala Morley. — A. Leaves from various collections. — B. Cleared part of leaf blade showing veins and terminal sclereids ( Judd 5 150). — C. Cross section of leaf blade showing sclereids, upper epidermis, hypodermis, and stomatal crypts ( Judd 5150). — D. Cross section of leaf midrib ( Zanoni 36370). — E. Fruits, and leaf with revolute margins (Judd 5150). — F. Seed ( Zanoni 36370). G-N, Lengthwise sections through the hypanthium as found on the fruit. All are one-seeded fruits except as indicated. — G. M. helleri var. helleri (Alain Liogier Novon 3: 274-277. 1993. Volume 3, Number 3 1993 Morley Mouriri crassisepala 275 105U8). — H. M. helleri var. samanensis (Ekman HI 1295). — I, J. M. spathulata var. spathulata (I, Morley 885; J, Montero 21248). — K. M. spathulata var. brachypoda, 2-seeded (Gonzales 2042). — L. M. emarginata (Shafer 589). M, N. M. crassisepala (Skean 1779). — M. 1-seeded. — N. 3-seeded. 276 Novon Table 1 . Mouriri crassisepala compared with the three other species in its section known in the fruiting condition. M. crassisepala M. spathulata M. emarginata M. helleri Leaf blade length (cm) 2. 7-4. 8 3-10.5 2.6-8 1-4.1 Blade thickness (mm) 0.36-0.48 0.15-0.37 0.21-0.37 0.23-0.47 Stomatal crypt diameter (gm) 60-65 25-65 30-93 30-62 av. 62.5 av. 45 av. 61.5 av. 46 Number of crypts per mm2 30-42 55-170 15-60 (0— )40— 1 15 Inflorescence mostly axillary axillary axillary or terminal lower Fruit diam. (mm) 14.7-32 15-30 9.5-15 7.5-12 Thickness of calyx lobes 0. 3-0.7 0.1-0. 2 deciduous 0.1 -0.2 or (mm) deciduous Style scar diam. (mm) 1.5-2. 5 0.4-0. 8 0.4-0. 8 0.3-0. 5 Inside diam. at base of 4. 2-6.6 2-3.1 2.8-3. 3 2-2.5 hypanthium (mm) Seed length (mm) 14-16.5 1 1.5-13.3 7. 7-8.6 4.5-6 the dry measurements ca. 25% less. Seeds medium to dark brown, smooth, 14-16.5 mm long, 12.5- 15.5 mm wide, 10-14 mm thick, subglobose or flattened on the contact faces when more than 1, with a roundish to angled basal hilum 5-7 mm diam. Distribution. Known only from the montane for¬ ests of the Cordillera Central near the center of the Prov. La Vega, Dominican Republic, at 1,300- 1,330 m. Mouriri crassisepala belongs to section A 'eso- phytum, which is restricted to the islands of Cuba, Hispaniola, and Puerto Rico and which contains five species including the new one (Morley, 1957). Plants of the section are characterized by small obovate leaves with tubular midrib xylem and Type III sto- matal crypts, a hypodermis lacking mucilage walls, Iree calyx lobes, average anthers for the genus, and axile-basal to axile ovules produced only outwardly from each placenta. Of the species in the section, M. helleri occurs in Puerto Rico and Hispaniola, M. gonavensis in southwest Hispaniola, M. spathu- lata in Hispaniola and Cuba, M. emarginata in Cuba, and the new species in central Hispaniola. Within the section the major distinguishing fea¬ tures of the new species are its thick leaves with large stomatal crypts, mostly terminal fruits, large fruits and seeds, thick persistent calyx lobes, and the hypanthium which in fruit is sunken and ex¬ panded at the base but not at the apex, producing a prominent overhang and a convex to flat ovary top. The most difficult species to contrast with the new one is Mouriri gonavensis of southwest Haiti, since the latter is known only in flower, the former only in fruit. In the following comparisons the first state¬ ment refers to M. crassisepala, the second to M. gonavensis: petioles 1.5-3. 5 mm long vs. 2-8 mm; blade/petiole ratio 12—1 8(— 24) vs. usually 7-11; blade margins strongly revolute vs. usually flat to somewhat revolute; blade thickness 0.36— 0.48 mm vs. 0.25-0.27 mm; stomatal crypt average diameter 62.5 jan vs. 50 /um, average frequency 36 per sq. mm vs. 42.5; flowers mostly terminal but sometimes in subterminal axils vs. axillary or sometimes at nodes below the leaf zone; calyx lobe length from stamen attachment 1.5-1. 9 mm (fruit) vs. 1.7-2. 5 (flower); number of ovary locules probably up to 5 judging by the seeds in the fruit vs. 2-3; elevation 1,300-1,330 m vs. 700-1,100 m. While further information obviously would be de¬ sirable, these comparisons better support an inter¬ pretation of these plants as different species rather than the same. The remaining three species in the section are contrasted with Mouriri crassisepala in Table 1. Mouriri helleri is least like M. crassisepala because of the former’s small leaves, fruits, and seeds, nu¬ merous small stomatal crypts, and little modified fruiting hypanthium. Mouriri spathulata resembles M. crassisepala in having large fruits and seeds and persistent calyx lobes, but disagrees in its smaller and more numerous stomatal crypts, axillary inflo¬ rescence, thin calyx lobes, small style scar, and only slightly modified fruiting hypanthium. Mouriri emarginata is more like the new species in having few large crypts and a fruiting hypanthium smaller than but similar in form to that of M. crassisepala, but differs in its thinner leaves, axillary inflores¬ cence, smaller fruits, seeds, hypanthium base, and style scar, and deciduous calyx lobes. Because of the unusual and highly modified nature of the fruit¬ ing hypanthium in the last two named species it is thought that M. crassisepala is probably more close- Volume 3, Number 3 1993 Morley Mouriri crassisepala 277 ly related to the Cuban M. emarginata than to either of the others. The differences in the shape of the fruiting hy- panthium are illustrated in Figure 1G-N. In Mouriri helleri the hypanthium remains almost entirely above the fruit tissue, and the tendency for the base of the hypanthium to be expanded during fruit for¬ mation varies from none to slight. In M. spathulata the base of the hypanthium has become sunken in the fruit, but there is little or no tendency for it to be expanded. In M. emarginata and M. crassise¬ pala the base is sunken and has been expanded, substantially so in the former, more so in the latter, while the apex remains unchanged in diameter or is compressed inward, producing an overhang of the hypanthium walls. In the last two species the apex oi the fruit within the hypanthium is strongly convex in one-seeded fruits; in M. crassisepala the apex becomes flatter or very slightly concave, and the overhang less when there are more than one seeds and thus the diameter of the fruit is greater and the expansion of the apex of the fruit is increased. I have seen no fruits of M. emarginata with more than one seed. It would appear that the development of the forms with sunken hypanthia was associated with increased fruit size. Paratypes. DOMINICAN REPUBLIC. La Vega: Ji- menoa, mountains above (SW of) Jarabacoa, NW slope of the Loma La Sal, moist montane forest, 1,330 m, 21 May 1986, IT. S. Judd 5150 with T. A. Zanoni, M. Mejia, J. D. Skean, Jr. & R. Beaman (FLAS, JBSD); Cordillera Central, NW foothills of Loma de la Sal, dis¬ turbed cloud forest with Brunellia, Baccharis, Miconia , Rhytidophyllum, Cyathea, and Arthrostylidium, 1,330 m, 21 May 1986, j. D. Skean, Jr. 1779 with Beaman, Judd & Zanoni et al. (FLAS, JBSD). Literature Cited Morley, T. 1957. A revision of Mouriri, section Ne- sophyturn. Brittonia 9: 110-131. - . 1976. Memecyleae (Melastomataceae). FI. Neotrop. 15: 1-295. Two New Combinations in Triantha (Liliaceae) John G. Packer Department of Botany, University of Alberta, Edmonton, Alberta, Canada ABSTRACT. Two new combinations at the subspe¬ cies level are proposed for Triantha occidentalis (S. Watson) Gates. A brief explanation is provided to account for their being made under this species name and not the presently more widely recognized Tofieldia occidentalis S. Watson. In preparing the treatment of Tofieldia for the Flora of North America (FNA), I became aware that it is a rather heterogeneous assemblage, rep¬ resented by two disparate sets of species. One is a homogeneous group ol three species, Tofieldia glu- tinosa (Michaux) Persoon, T. occidentalis S. Wat¬ son, and T. racemosa (Walter) Britton, Sterns & Poggenberg, characterized by glandular pubes¬ cence, appendaged seeds, and clustered flowers. The other, consisting of T. coccinea Richardson, T. gla¬ bra Nuttall, and T. pusilla (Michaux) Persoon, is a much less homogeneous group whose members are glabrous, with appendageless seeds and flowers aris¬ ing singly. In two of onlv three previous studies that include all the known North American species of Tofieldia (Baker. 1879; Gates, 1918), the former group of species was treated as a distinct genus, Triantha, based on a section of Tofieldia recognized by Nuttall (1818). Despite this, authors have con¬ sistently disregarded Triantha, the only significant exception being Small (1903). In the broader context of the tribe I ofieldieae, a study by Ambrose ( 1 980) showed that the rnonotypic Pleea (transferred by lltech (1978) to Tofieldia) was best treated as a separate genus, and Cruden (1991) came to a similar conclusion with respect to Isidrogalvia, traditionally included in Tofieldia. Both studies provided evidence that Triantha is distinct from Tofieldia. Glearly, systematic consistency would best be served if Tofieldia and Triantha were treated as separate genera following Baker (1879) and Gates (1918). R. Kiger (FNA taxon editor for I.iliaceae) and F. Utech (FNA advisor on Liliaceae) have con¬ curred. In the following consideration of the necessity for the new combinations, for expository ease, the con¬ text of the discussion is that of the original genus Tofieldia. Hitchcock (1944), in his study of variation in Tofieldia glutinosa (= Triantha glutinosa (Mi¬ chaux) Baker) and Tofieldia occidentalis (= Trian¬ tha occidentalis (S. Watson) Gates), identified, on the basis of morphological characters, “five fairly well marked . . . closely related entities . . . each of which has a rather well defined geographic range.” He discussed the various ways in which they might be treated taxonomically, suggesting three possibil¬ ities: 1 . Recognize all the entities as species; 2. Recognize two species, Tofieldia glutinosa and Tofieldia occidentalis, the former comprising two subspecies, the latter three; 3. Assign the five entities the rank of subspecies under a single species. Hitchcock chose the third alternative and treated the five entities as subspecies of Tofieldia glutinosa: occidentalis, brevistyla , absona, montana, and typica (glutinosa). In discussing his second possi¬ bility, Hitchcock suggested that montana would be treated as a subspecies of T. glutinosa, while brev¬ istyla and absona (which he said was only a local variant of brevistyla) would be treated under 7. occidentalis. In preparing the account of Tofieldia 1 examined most of the material that Hitchcock worked with as well as a considerable amount of additional material collected over the past 50 years, particularly from western Canada and Alaska. The investigation has confirmed Hitchcock’s findings. The only minor ex¬ ception is his absona (restricted to the Priest Lake area of Idaho), which is in my view not significantly different from brevistyla and is included with it. However, as a somewhat better understanding of the morphology of Hitchcock’s subspecies emerged, it became apparent that the best taxonomic treat¬ ment was not the one that he finally selected but his second alternative, the recognition of two species, Tofieldia glutinosa and T. occidentalis, but with the difference that montana be treated as a sub¬ species of T. occidentalis along with brevistyla (in¬ cluding absona) and not as a subspecies of T. glu¬ tinosa. When Watson (1879) described Tofieldia occi¬ dentalis he mentioned the occurrence of a “loose white spongy testa” surrounding the mature seeds, which is absent in 7. glutinosa. The precise nature of this spongy or inflated enveloping tissue is not Novon 3: 278-279. 1993. Volume 3, Number 3 1993 Packer New Combinations in Triantha 279 certain — it may or may not be testa — but it instantly distinguishes 7. occidentalis from 7. glutinosa (and from the other trianthoid species, T. racemosa and T. japonica). The seeds of subspecies brevistyla and man tana have the spongy covering, those of the former being identical to T. occidentalis in that the covering is strongly inflated, while in montana it is less strongly inflated. Hitchcock appreciated the significance of this unique seed characteristic, making reference to it in discussing his second taxonomic scheme, when proposing that brevistyla and absona might be treated under Tofieldia occidentalis. It is conse¬ quently not clear why he should have been inclined to refer montana to T. glutinosa, especially since he remarked that the inflated testa of this subspecies is “a characteristic unlike glutinosa, but like oc¬ cidentalis." He mentioned that the flowers of 7. glutinosa and montana are similar, but the flowers in all the subspecies he distinguished are generally alike, and while they show7 some slight variation they are not of much significance in the recognition of iniraspecific taxa. Further to this, montana has, as Hitchcock ob¬ served, “much longer pubescence than is to he found in any other of the subspecies,” and he said there is “no difficulty in distinguishing it from glutinosa typica." Although he did not mention it. the long cylindrical hairs below the inflorescence in montana occur to the complete exclusion of the dome-shaped or conical glands (Hitchcock's “haycocks”) that are found to a greater or lesser extent in the other subspecies, and which in T. glutinosa are virtually the only vestiture on the stem. The more or less cylindrical hairs of various lengths, but longest in montana, are a characteristic of T. occidentalis and not of 7. glutinosa. Watson (1879) had an insight into this difference when he described the pubes¬ cence of 7. occidentalis as "viscid and that of 7. glutinosa as “glutinous.” Another character that Watson mentioned in dis¬ tinguishing Tofieldia occidentalis from 7. glutinosa is the division of the hracteoles, which he desc rihed as being “3-lobed nearly to the middle” in the for¬ mer, and “scarcely lobed” in the latter. The vari¬ ation in the lobing of the hracteoles in 7. occidentalis is actually somewhat greater than originally indi¬ cated by Watson, and hracteoles of 7. glutinosa may occasionally be deeply divided, but the differ¬ ence Watson noted is valid and applies as well to the subspecies that Hitchcock described. It has also been observed in the present study that the brac- teoles in T. occidentalis are often glandular, but glandular hracteoles are virtually unknown in 7. glutinosa. In light of the above it is concluded that the best taxonomic treatment of the variation that Hitchcock documented is essentially his second alternative: that is, to recognize Tofieldia occidentalis and 7. glu¬ tinosa as distinct species with brevistyla (including absona) and montana treated as components of Tofieldia occiden talis. I bis requires the following nomenclatural revi¬ sions. Triantha occidentalis (S. Watson) Gates subsp. brevistyla (Hitchcock) Packer, comb. nov. Basionym: Tofieldia glutinosa (Michaux) Per- soon subsp. brevistyla Hitchcock, Amer. Midi. Naturalist 31: 495. 1944. TYPE: Canada. British Columbia: 32 mi. N ol Golden, 1 1 July 1944, Hitchcock & Martin 7 h'AB (holotype, WTU). Triantha occidentalis (S. Watson) Gates subsp. montana (Hitchcock) Packer, comb. nov. Bas¬ ionym: Tofieldia glutinosa (Michaux) Persoon subsp. montana Hitchcock, Amer. Midi. Nat¬ uralist 31: 496. 1944. TYPE: U.S.A. Mon¬ tana: Glacier National Park, Logan Pass, 18 Sep. 1937, Barkley & Marsh 17 A I (holotype, WTU). Acknowledgments. I am indebted to R. Kiger and F. Utech for reviews of the manuscript and suggestions. Literature Cited Ambrose, J. D. 1980. A re-evaluation of the Melan- thoideae (Liliaceae) using numerical analyses. Pp. 65 81 in C. D. Bricknell, C. F. Cutler & M. Gregory (editors), Petaloid Monocotyledons. Linn. Soc. Symp. Ser. 8. Academic Press, London and New York. Baker, J. G. 1879. A synopsis of Colchicaceae and the aberrant tribes of Liliaceae. J. Linn. Soc. Bot. 17: 405-510. Cruden, R. W. 1991. A revision of Isidrogalvia (Lil¬ iaceae): Recognition for Ruiz and Pavon's genus. Syst. Bot. 16: 270—282. Gates, R. 1918. A systematic study of the North Amer¬ ican Melanthaceae from the genetic standpoint. J. Linn. Soc. Bot. 44: 131-172. Hitchcock, C. L. 1944. The Tofieldia glutinosa com¬ plex of western North America. Amer. Midi. Natu¬ ralist 31: 487-498. Nuttall, T. 1818. The Genera of North American Plants. Philadelphia. Small, J. K. 1903. Flora of the Southeastern United States. [Published by the author] New York. Utech, F. H. 1978. Floral vascular anatomy of Pleea tenuifolia Michx. (LiliaceaeTofieldieae) and its reas¬ signment to Tofieldia. Ann. Carnegie Mus. 47: 423- 454. Watson, S. 1879. Contributions to American botany. I. Revision of the North American Liliaceae. Proc. Amer. Acad. Arts 14: 213-288. Aphelandra espirito-santensis (Acanthaceae), a New Species from Espirito Santo, Brazil Sheila R. Profice Jardim Botanico do Rio de Janeiro, Rua Pacheco Leao 915, 22460 Rio de Janeiro, Rio de Janeiro, Brazil Dieter C. Wasshausen Department of Botany, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. ABSTRACT. A new Brazilian species of the genus Aphelandra is described and illustrated. It is related to Aphelandra ignea but differs markedly from that taxon by its leaf blades and the ovate to spathulate bracts with 4-5 spines on the margin. Aphelandra is represented in Brazil by 28 species. They occur from the Floresta Amazonica to the Floresta Atlan- tica. Aphelandra R. Brown is a morphologically di¬ verse genus characterized by a lack of cystoliths, a corolla with a distinct upper lip, four monothecous stamens, and elongated, tricolporate pollen. The result of a recently completed study of Acan¬ thaceae specimens deposited in the Jardim Botanico do Rio de Janeiro (RB) has led to the discovery of an undescribed species. Aphelandra espirito-santensis Profice & Was¬ shausen, sp. nov. TYPE: Brazil. Espirito Santo: Mun. de Linhares, Reserva Florestal da Cia. Vale do Rio Doce, 26 Sep. 1978, A. M. Car¬ valho et al. HO (holotype, RB). Figure 1. Herba, caulis reptans, subteres, pilis hirsutis, in apicem versum flavo-pubescens. Folia hirsuta, lamina ovata, basi apiceque obtusa, 3-6 paribus venis secundariis, 1.5-5 cm longa et 1.5-3 cm lata; petiolo 0.5-4 cm longo; inflorescentia spicata terminalis raro axillaris, 1.2 -1.3 cm longa, rhachi flavo-pubescenti; bracteae imbricatae, obov- atae vel spathulatae, marginibus 4-5 dentatis, extus pu- bescentes, membranaceae, 1-1.3 cm longae et 0.4-0. 9 cm latae; bracteolae filiformes, apice aristatae, pilosae, striatae, subhyalinae, 5-6 mm longae et ca. 1 mm latae; calyx quinquepartitus, segmentis lanceolatis, subaequali- bus, striatis, subhyalinis, aristatis, barbatis, 7 8 mm lon- gis, 1-1.5 mm latis; corolla flava, ca. 1 cm longa, mem- branacea, pilosa, tubo cylindrico, basi aliquanto angustato 0.1 cm lato, fauci 0.2 cm lati, labio superiore bilobato, 0.3 cm longo et 0.2 cm lato, labio inferiore triloba to, lobis lateralibus obovatis, 0.3 cm longis et 0.2 cm latis, lobo medio orbiculato, 0.4 cm longo et 0.4 cm lato; stamina exserta, antheris ad apicem pilosis, ca. 2 mm longis; capsula ovata, glabra, 0.9-1 cm longa; semina ovata, brunnea, tuber culata, 2.5-3 mm longa et ca. 2 mm lata. Herb; stem reptant at base, subcylindrical, with hirsute pubescence, the trichomes yellowish apically. Leaves with petioles 0.5-4 cm long, the pubescence similar to that of the stem, the blades ovate, obtuse apically and basally, with 3-6 pairs of secondary veins, hirsute on both the upper and lower surface, especially the veins and the margins, 1.5-5 cm long and 1.5-3 cm wide. Flowers borne in terminal, rarely axillary spikes 1.2-3 cm long, the rachis yellowish pubescent; bracts imbricate, obovate to spathulate, the margins spinulose-serrate, provided with 4-5 pairs of teeth, pubescent without, mem¬ branous, 1-1.3 cm long and 0.4-0. 9 cm wide; bracteoles filiform, aristate at apex, pilose, striate- nerved, subhyaline, 5-6 mm long and 1 mm wide; calyx 5 -parted, the segments lanceolate, subequal, striate-nerved, subhyaline, aristate, barbate, 7-8 mm long, the posterior segment ca. 1.5 mm wide, the anterior and lateral pair ca. 1 mm wide; corolla yellow, about 1 cm long, membranous, sparingly pilose, the tube cylindrical, 1 mm wide at base, about 2 mm wide apically, the upper lip bilobed, 3 mm long and 2 mm wide, the lower lip 3-lobed, the lateral lobes 3 mm long and 2 mm wide, the middle lobe orbicular, 4 mm long and wide; stamens ex- serted, anthers ca. 2 mm long, apically pilose; pollen prolate, tricolpate with the colpi extending from the equator nearly to one or both poles, 55-64 gm long and ca. 29 n m wide, the exine sculpturing verru- culose (Fig. 2). Capsule ovate, glabrous, 9-10 mm long; seeds ovate, brown, tuberculate, 2.5-3 mm long and ca. 2 mm wide. Distribution. Occurring in lowland forests in northern Espirito Santo at elevations of about 30 m. Novon 3: 280-283. 1993. Volume 3, Number 3 1993 Profice & Wasshausen Aphelandra espirito-santensis 281 Figure 1. A-G. Aphelandra espirito-santensis Profice & Wasshausen ( Carvalho et al. 80). — A. Habit. — B. Variability of the bracts. — C. Detail of the bract margin. — D. Corolla, stamens. — E. Calyx segments and capsule. — F. Seed. — G. Embryo. (Drawn by S. Profice.) 282 Novon Figure 2. Scanning electron (SEM) photomicrographs of Aphelandra espirito-santensis pollen. — A. Equatorial view, x 1500 ( Carvalho et al. 80). — B. Equatorial view, x 1400 (Carvalho et al. 80). Aphelandra espirito-santensis shares diagnostic characteristics and affinities with two Brazilian spe¬ cies, A. ignea (Schrader) Nees ex Steudel and A. phrynioides Lindau. It is distinguished from A. ig¬ nea hy its leaf shape and the ovate to spathulate floral bracts with 4-5 pairs of marginal teeth. From A. phrynioides the new taxon is distinctive by its much reduced corolla size and shape, leaf shape and size, and the size and shape of its floral bracts. Using the infrageneric classification provided by Wasshausen (1975), A. espirito-santensis is in¬ cluded in the group of Brazilian species that are characterized by having entire or undulate leaf blades and floral bracts without bracteal nectaries and with marginal teeth. The following key may be used to distinguish the species of this group. 1. Corollas 7-25 mm long . 2 1. Corollas 30-65 mm long . 6 2(1). Corollas 10-13 mm long . 3 2. Corollas 20-22 mm long . 5 3(2). Bracts ovate, 17-20 mm long . . A. caput-medusae Lindau 3. Bracts obovate to obovate-cuneate to spathu¬ late, 9-13 mm long . 4 4(3). Corollas red; bracts obovate-cuneate, 9x4 mm; leaf blades oblong-oval 8-10 cm long . A. decor ata (Nees) Wasshausen 4. Corollas yellow; bracts obovate to spathulate, 10 — 13 mm long; leaf blades ovate, 1.5 5 cm long . . ... A. espirito-santensis Profice & Wasshausen 5(2). Leaf blades ovate, 8-12 x 4-6.5 cm; bracts obovate, 17-19 x 14 mm, mucronulate at apex; corolla sparingly glandular-pilose .... . A. neesiana Wasshausen 5. Leaf blades oblong-elliptic to oblong-lanceo¬ late, 8-9 x 2. 5-3. 5 cm; bracts elliptic-ob¬ long, 13-15 x 5-6 mm, obtuse at apex; corolla glabrous . . A. bahiensis (Nees) Wasshausen 6(1). Corollas yellow . 7 6. Corollas red, orange, or scarlet . 9 7(6). Bracts conduplicate . . A. ornata (Nees) T. Anderson 7. Bracts plane, not folded or conduplicate . 8 8(7). Corollas ca. 40 mm long, the upper lip ob¬ ovate, 8x7 mm, bilobed . . A. phrynioides Lindau 8. Corollas 30 mm long, the upper lip oval, 5 x 5 mm, entire . . A. ignea (Schrader) Nees ex Steudel 9(6). Leaf blades narrowly elliptic, 2-3.2 cm wide, acuminate at apex; bracts oblong to elliptic, Volume 3, Number 3 1993 Profice & Wasshausen Aphelandra espirito-santensis 283 18x6 mm, green with the upper margins purplish, the margins spirulose-dentate with about 5 pairs of teeth; corolla orange . . A. margaritae E. Morren 9. Leaf blades narrowly ovate, 4-5 cm wide, acute to obtuse at apex; bracts obovate, to 18 x 14 mm, purplish, the margins spinulose- serrate with about 7 pairs of teeth; corolla red . A. obtusifolia (Nees) Wasshausen Should one wish to compare the new taxon to some of the smaller terrestrial herbs of Aphelandra of Central America, e.g., seibertii Leonard or A. tonduzii Leonard, the following key will demonstrate the major differences. 1 . Floral bracts narrowly lanceolate, 1.5 mm wide, with 1 pair of minute, slender marginal teeth; corollas pale pink. Costa Rica . . A. tonduzii Leonard 1. Floral bracts obovate to spathulate, 4-9 mm wide, with 2-5 pairs of conspicuous marginal teeth; corollas yellow . 2 2. Corollas 20 mm long; leaf blades oblong-elliptic, thinly pilose; flowers borne in spikes 4-8 cm long; bracts bearing 2 or usually 3 marginal teeth. Costa Rica and Panama . . A. seibertii Leonard 2. Corollas 10 mm long; leaf blades ovate, hirsute; flowers borne in spikes 1.2-3 cm long; bracts bearing 4-5 pairs of marginal teeth. Brazil . . . A. espirito-santensis Profice & Wasshausen Paratypes. BRAZIL. Espirito Santo: Mun. de Lin- hares, Reserva Florestal da Cia. Vale do Rio Doce, G. Martinelli 4993 (RB); Reserva de Sooretama, D. Sucre 5701 (RB); Reserva de Sooretama, G. Martinelli 2285 (RB). Acknowledgment . Our special thanks to Jorge Fontella Pereira for the revision of the Latin diag¬ nosis. Literature Cited Wasshausen, D. C. 1975. The genus Aphelandra (Acanthaceae). Smithsonian Contr. Bot. 18: 1-157. New Species of Pentacalia (Senecioneae: Asteraceae) from Ecuador, Peru, and Bolivia Harold Robinson and Jose Cuatrecasas Department of Botany, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. ABSTRACT. Twelve new species of Pentacalia are described from Ecuador: P. carmelana, P. caza- letii, P. dodsonii, P. dorrii, P. hurtadoi, P. lutey- norum, P. moronensis, P. napoensis, P. pailasen- sis, P. palaciosii, P. zakii, P. zamorana; nine new species are described from Peru: P. balsasana, P. rutervonis, P. davidsmithii, P. maynasensis, P. mucronatifolia , P. nunezii, P. sagasteguii, P. til- lettii, and P. todziae; and two new species are described from Bolivia: P. inipiisiviensis and P. lewisii. Gynoxys vargasiana Cabrera of Peru is transferred to Pentacalia. Keys are provided for t he species of Pentacalia subg. Pentacalia in Ec¬ uador and Peru. Over a period of years since the resurrection of the genus Pentacalia Cassini (Robinson & Cuatre¬ casas, 1978; Cuatrecasas, 1981; Robinson in Proc¬ tor, 1982), a number of undescribed species, mostly of the subgenus Pentacalia, have been set aside awaiting review. Progress on a study of Pentacalia in Colombia by Diaz-Piedrahita and Cuatrecasas (in prep.) has furnished a basis for comparison with unidentified specimens. The opportunity has been taken to describe twelve new Ecuadorian species, nine new Peruvian species, and two new Bolivian species. The large number of undescribed species is one indication of the generally inadequate treatment of Asteraceae of the tropical forests. As seen in Cuatrecasas (1981), there have been 13 species of subgenus Pentacalia claimed from Ecuador, to which should be added P. floribunda, which was listed under subgenus Microchaele. In addition to the new species, in our review of Pen¬ tacalia subg. Pentacalia in Ecuador we have en¬ countered recent collections of some other rarely collected Ecuadorian species: P. andrei (Greenman) J. Cuatrecasas, Zamora-Chinchipe, limit of Parque Nacional Podocarpus, around pass on road Loja- Zamora, alt. 2,750-2,950 m, 25 Dec. 1988, Mad¬ sen & Ellemann 75982 (AAU, QCA, US). Penta¬ calia corazonensis (Hieronymus) .1. Cuatrecasas, Cotopaxi: Quevedo-Latacunga road. Paramos de Zumbagua, 19 km E of Pilalo and 78 km W of Latacunga, elev. 3,536 m, 25 Dec. 1978, Luteyn & Luteyn 6510 (NY, V10); Loja: road to Zamora from Loja, km 12-14, near top of pass, elev. 2,800 m, 28 Sep. 1961, Dodson & Thien 779 (US). Pentacalia lanceolifolia (J. Cuatrecasas) J. Cua¬ trecasas, Loja: Parque Nacional Podocarpus, Cerro Toledo, 2,500-3,400 m, 30-31 Oct. 1989, Mad¬ sen 86250 (AAU, QCA, US). Pentacalia millei (Greenman) J. Cuatrecasas, Loja: Parque Nacional Podocarpus, 2,750-2,850 m, 4 May 1984, 011- gaard, Madsen & Christensen 71195 (AAU, QCA, US); Loja: Parque Nacional Podocarpus, E of Nudo de Cajanuma, trail E of “Centro de Information" to crest on trail to Laguna de Compadre, 2,850- 3,050 m, 7 June 1988, 0llgaard 74631 (AAU, QCA); Parque Nacional Podocarpus, new road Loja- Zamora, E of Cerro Yanococha, along former Indian trail to Zamora, 2,550-2,650 m, 26 Nov. 1988, Madsen 75586 (AAU, QCA); Parque Nacional Po¬ docarpus, road Yangana-Valladolid, km 21, alt. 2,700 m, 10 Nov. 1989, Madsen 86103 (AAU, QCA). Pentacalia sevillana (J. Cuatrecasas) J. Cua¬ trecasas, Loja: 7 km N of San Lucas on road to Loja, elev. 8,800 ft., 27 Jan. 1979, R. M. King & F. Almeda 7811 (US). Cuatrecasas (1981) listed 20 members of Pen¬ tacalia subg. Pentacalia in Peru. Nineteen of these species are treated by Cabrera (1954) in the key to scandent Senecio species of Peru and Bolivia. Nine new Peruvian species of Pentacalia subg. Penta¬ calia are described here, and a new key is provided. In addition, the primarily Ecuadorian P. diseiformis (Hieronymus) J. Cuatrecasas has been seen from northern Peru: Piura, Cerro Aypate, 49 km E of Ayabaca, 2,750 m, 22 Sep. 1991, Gentry, Diaz & Ortiz 7471 (MO, US). The species occurring in Bolivia are also treated in the key by Cabrera (1985). Eighteen species were included, many of which also occur in Peru. Two new Bolivian species are described here with ref¬ erence to their placement in the Cabrera key. In the following descriptions the height of the heads is measured from the top of the peduncle to the tips of the disk corollas. Novon 3: 284-301. 1993. Volume 3, Number 3 1993 Robinson & Cuatrecasas New species of Pentacalia 285 A New Species of Pentacalia (subg. Microchaeie) in Ecuador Pentacalia (subg. Microchaete) dodsonii H. Rob¬ inson & .1. Cuatrecasas, sp. nov. TYPE: Ec¬ uador. Zainora: road from Loja to Zamora, elev. 2,800 m, 18 Sep. 1961, Dodson & Thien 676 (holotype, SEE). P. onae et P. scitophyllae simila sed in bracteis in¬ volut'd paucioribus distincta. Shrub 0.7- 1.7 m high, moderately branched; stems dark brown, densely pilose with stout, blunt hairs or nearly glabrous; internodes ca. 1 cm long; pith solid. Leaves alternate; petioles 0.5- 1.0 cm long, with few to many short, stout hairs; blades oblong-elliptical, mostly 6-7 cm long, 1 .6-3.0 cm wide, base short-acute, margins closely serrate, apex shortly and sometimes narrowly acuminate, both surfaces with network of veinlets raised slighlly above surface, irregularly pilose to nearly glabrous, lower surface paler green, with 7 or 8 prominent, strongly ascending secondary veins on each side. Inflores¬ cences terminal on leafy branches, broadly cor¬ ymbose with numerous branches; peduncles 1.2- 2.5 cm long, usually with numerous narrowly su¬ bulate bracteoles 2-3 mm long, with 2 4 bracteoles forming calyculus. Heads heterogamous, radiate, 9 10 mm high; involucral bracts 8, broadly oblong, 4-5 mm long, ca. 2 mm wide, apex obtuse to short- acute, glabrous; rays 8, corollas yellow, glabrous, basal tube ca. 4.5 mm long, limb large, ca. 10 mm long, 3.5 mm wide. Disk florets ca. 17; corollas ca. 6.5 mm long, basal tube ca. 2 mm long, throat ca. 2.5 mm long, lobes ca. 1 mm long, 0.7-0. 9 mm wide at base, tips slightly thickened with sclerified shield of many slightly bulging, moderately enlarged cells outside; anther thecae ca. 2 mm long, very shortly pointed at base; backs of style branches with papillae continuing onto upper shaft of style. Aehenes ca. 3 mm long, glabrous; pappus bristles mostly ca. 5 mm long with broadened tips of somewhat enlarged fusiform cells. Pollen grains ca. 35 jam diam. The new species resembles the closely related Pentacalia onae( J. Cuatrecasas) J. Cuatrecasas and P. scitophylla (HRk) J. Cuatrecasas in general as¬ pect but differs most obviously in having only ca. 8 bracts m the involucre instead of ca. 13. The backs ol the style branches have papillae along their entire length and onto the upper part of the style shaft. The latter papillae are presumably like those ol other species of Pentacalia subg. Microchaete, but they are unlike any Pentacalia subg. Pentacalia de¬ scribed below. None of the following species have papillae on the backs of the style branches continued to the base. Paratypes. ECUADOR. Loj a: road to Zamora from Loja, km 12-14, elev. 2,800 m, near top of pass, shrub 4-5 ft. high, 28 Sep. 1961, Dodson <£• linen 778 (SEL). Loja/Zamora-Chinchipe: limit of Parque Nacional Podocarpus, around pass on road Loja-Zamora, wet elfin forest, scrub-paramo and treeless ridgetop vegetation, 79°07'W, 03°58'S, 2,900 in, 8 Jan. 1989, shrub, Mad¬ sen 85480 (AAU). New Species of Pentacalia (subg. Pentacalia) in Ecuador Pentacalia (subg. Pentacalia) carmelana II. Robinson & J. Cuatrecasas, sp. nov. TYPE: Ecuador. Napo: Cartagena, km 25 from El Carmelo on road toward La Bonita, 77°30'W, 0°37'N, all. ca. 2,800 m, 13 Apr. 1979, Loj t riant, Molau & Madison 12335 (holotype, AAU). In foliis late ovatis sparse vel distincte crispe hirtellis in capitulis radiatis et in floribus staminodeiferis distincta. Plants scandent, indefinitely elongate; stems densely hirtellous with short brownish hairs, not or weakly fistulose. Leaves alternate, petioles 1.2 1.7 cm long, stout, densely hirtellous with stout hairs; blade thinly coriaceous, broadly oblong-ovate, ca. 9 cm long, ca. 6.5 cm wide, base broadly rounded, margins entire, slightly reflexed, apex rounded with minute apiculus, upper surface nearly smooth, slightly shiny, with few hairs mostly on veins, lower surface with numerous short crisped hairs on veins, with sparse hairs between veins; secondary veins prom¬ inent, 4 or 5 on each side, spreading at ca. 50°, arching, veinlets scarcely raised above surface. In¬ florescences mostly lateral from axils of vegetative leaves, ca. twice as long as leaves, with foliose bracts in basal half, blades to 3.5 cm long, ca. 2.5 cm wide; distally a rounded or pyramidally thyrsoid panicle, with branches corymbose; peduncles 3-7 mm long, densely puberulous. Heads 9- 1 0 mm high, heterogamous, radiate; subinvolucral and calycular bracts minute, subulate, mostly 1.5 mm long, with few minute hairs; involucral bracts ca. 8, glabrous, ca. 6 mm long, 1.5 mm wide, apices short-acute. Ray florets ca. 6; corollas yellow, glabrous, lube 5- 6 mm long, limb ca. 5 mm long, usually with 2 minute staminodia. Disk florets 15-18; corollas yel¬ low, glabrous, tube ca. 4 mm long, throat ca. 2 mm long, lobes ca. 2 mm long, 0.55 mm wide, mod¬ erately thickened at tip with sclerified shield of many enlarged, bulging cells outside; anther thecae ca. 1.5 mm long, basal tails nearly as long as collar; apical appendage oblong, ca. 0.3 mm long, 0.15 286 Novon mm wide. Achenes ca. 2 mm long, 5-ribbed, gla¬ brous; pappus bristles 5-6 mm long, tips not broad¬ ened, often nearly smooth. Pollen grains ca. 30 fxm diam. The single known specimen is cited from a dense, mossy cloud forest. In Ecuador the species seems most similar to other radiate species with lateral inflorescences, P. huilensis and P. riotintis, but the latter are more glabrous and lack the large foliose bracts on the bases of their inflorescences. Pentacalia (subg. Pentacalia) cazaletii H. Rob¬ inson & J. Cuatrecasas, sp. nov. TYPE: Ec¬ uador. Pichincha: 20 km W of Santo Domingo de los Colorados, alt. 1,000 ft., 18 Oct. 1961, Cazalet & Pennington 5064 (holotype, US; isotypes, B, K, NY). In nervis secundariis foliorum obscuris in inflorescen- tibus axillaribus thyrsoideis bracteiferis et in capitulis het- erogamis disciformis distincta. Plants scandent, indefinitely elongate; stems gla¬ brous or glabrescent, with solid pith. Leaves alter¬ nate, petioles 0.7- 1.1 cm long; blade fleshy, drying thinly subcoriaceous, ovate-elliptical, 8-10 cm long, 3. 0-4. 2 cm wide, base and apex short-acute, mar¬ gins entire, upper and lower surfaces glabrous and smooth, only midvein prominent below, secondary veins 5-6 on each side, ascending and arching, obscure. Inflorescences lateral from axils of vege¬ tative leaves, one and a half to three times as long as leaves, pyramidally thyrsoid, bracts foliiform, ob¬ long-elliptical, mostly 2-5 cm long, 0. 8-2.0 cm wide, apices obtuse with slight mucro; branches spreading at about 45°; peduncles 0-4 mm long, sparsely tomentellous. Heads ca. 8 mm high, het- erogamous, disciform; subinvolucral and calycular bracteoles ca. 7, subulate, ca. 2.5 mm long, ca. 0.8 mm wide; involucral bracts 8, ca. 6 mm long, ca. 1 mm wide, apices acute, glabrous. Eemale florets 4, tubular; corollas yellow, glabrous, 5. 0-5. 5 mm long, tube ca. 3.5 mm long, lobes linear, ca. 1.8 mm long. Disk florets 8 or 9; corollas yellow, gla¬ brous, 6. 0-6. 5 mm long, tube ca. 3 mm long, throat ca. 1.8 mm long, lobes narrowly oblong, 1.5 1.8 mm long, 0.40-0.45 mm wide, tips minimally thick¬ ened with subsclerified shield of large, somewhat bulging cells outside; anther thecae ca. 1 .5 mm long, basal tails ca. two-thirds as long as collar; apical appendage oblong, ca. 0.3 mm long, 0.12-0.15 mm wide. Achenes ca. 1 .5 mm long, glabrous; pap¬ pus bristles ca. 7 mm long, with apical cells not or scarcely broadened. Pollen grains ca. 27 jum diam. I he holotype is cited from a seasonal rainforest. as an “epiphyte on a dead 50 ft. tree.” The paratype is cited from “Bosque muy humedo Montano Bajo.” The fleshy, glabrous, elliptical leaves can be distin¬ guished from many species of Pentacalia by the obscurity of the upwardly ascending secondary veins. I he elongate, thyrsoid, hractiferous lateral inflores¬ cences and the small heads bearing disciform rays are distinctive. The species is evidently restricted to the western side of the Andes. Paratype. ECUADOR. Pichincha: Quito Canton, Chiriboga, en la carretera vieja a Santo Domingo, Reserva Forestal “La Favorita” del Minsterio Agric., 78°47'W, 00°12'S, 1,600-1,800 m, 8 Mar. 1990, Ceron, Ayala & Jimenez 8937 (MO, US). Pentacalia (subg. Pentacalia ) dorrii 11. Robinson & J. Cuatrecasas, sp. nov. TYPE: Ecuador. Loja: road to Fierro Acru, N of Pichig (ca. 3°40'S, 79°1 5'W), 3,015-3,385 m, 15 July 1989, Dorr & Valdespino 6654 (holotype, US; isotype, NY). In foliis obovatis base cuneatis in nervis secundariis ascendentibus in inflorescentiis terminalibus et capitulis radiatis distincta. Scandent shrubs; stems glabrous, scarcely striate, partially fistulose. Leaves alternate, petioles 1 .0- 1.5 cm long, moderately swollen at base; blades thinly subcoriaceous, glabrous, obovate, broadest in distal third or fourth, 5. 0-7. 5 cm long, 1 .5-2.5 cm wide, base narrowly cuneate, margins entire, apex obtuse to short-acute with small apiculus, surfaces with secondary veins slightly raised above surface and less strongly raised veinlets; secondary veins ca. 5 on each side, strongly ascending at 30-35°, somewhat indistinct from reticulum of veinlets form¬ ing elongated areoles. Inflorescences terminal, broadly corymbose, with stem slightly inflated in lower part, only lowest branches from axils of re¬ duced leaves, bracts above base small, subulate; peduncles 3-8 mm long, minutely puberulous. Heads 10-12 mm high, heterogamous, radiate; subinvol¬ ucral and 4 or 5 calycular bracts subulate, glabrous, to 1.5 mm long; involucral bracts ca. 8, glabrous, ca. 7 mm long, 1.5 mm wide, apices short-acute. Ray florets 5; corollas yellow, glabrous, tube ca. 5 mm long, limb 8-9 mm long. Disk florets ca. 9; corollas yellow, ca. 10 mm long, tube ca. 3 mm long, throat ca. 5 mm long, lobes ca. 1.5 mm long, 0.8 mm wide, lobes thickened equally to tip, tip with sclerified shield of scarcely bulging cells outside; anther thecae ca. 2.8 mm long, basal tails often longer than collar, apical appendage narrowly ob¬ long, ca. 0.6 mm long, 0.3 mm wide. Achenes submature, ca. 2 mm long, glabrous, weakly 5-ribbed; Volume 3, Number 3 1993 Robinson & Cuatrecasas New species of Pentacalia 287 pappus bristles 7-8 mm long, slightly but distinctly broader and more roughened at tips. Pollen ca. 35 /uni diam. I he new species has the general appearance of many of the glabrous, fleshy-leaved members of Pentacalia, but it is distinctive in the strongly ob- ovate leaf blades with strongly ascending secondary veins. Its closest relation may be to P. corazonensis, an Ecuadorian species with similar terminal inflo¬ rescences and radiate heads. The latter has leaves that are distinctive in their oblong shape and rounded apices. The veins of the latter are more prominent in all the specimens seen. The slight enlargement of the stem in the base of the inflorescence is remi¬ niscent of the condition in P. luteynorum described below, hut the latter has obscure secondary veins in the leaves and smaller heads with whitish rays. As noted on the specimen by Pruski, there is a close match with a type photograph of Senecio willden- owii Schultz Bip. in Berlin, which differs only in less blunt leaf apices. The latter name has never been validly published. Pentacalia (suhg. Pentacalia) hurtadoi 11. Rob¬ inson & J. Cuatrecasas, sp. nov. TYPE: Ec¬ uador. Napo: 3 km este del Caserio de Hua- mam, al norte de la carretera Hollin Loreta, por una trocha, 00°43'S, 77°36'W, 1,200 m, 17 Sep. 1088, Hurtado & Alvarado 302 (ho- lotype, US; isotype, MO). In foliis glabris in nervis secundariis pinnatis promi¬ nent ibus arcuatis in inflorescentiis axillaribus plerurnque racemiformis et in capitulis homogamis distincta. Plants scandent, indefinitely elongate; stems gla¬ brous, with solid pith. Leaves alternate, petioles 1 .2- 1.7 cm long; blades subcoriaceous, glabrous, ovate- elliptical to elliptical, 10-12 cm long, 3. 5-4. 5 cm wide, base short-acute, margins entire, apex apic- ulate, upper surface nearly smooth, on each side with ca. 6 arching secondary veins prominent on lower surface, intervening veins sometimes rather prominent. Inflorescences lateral, from axils of veg¬ etative leaves, usually somewhat shorter than leaves, cylindrical, mostly racemiform, thyrsoid at base; peduncles 3-7 mm long, finely puherulous. Heads 10-11 mm high, homogamous; calyeular bracts ca. 4, glabrous, 2-3 mm long, ca. 0.5 mm wide; in- volucral bracts ca. 8, glabrous, 7-8 mm long, 1 .0- 1.5 mm wide, apices short-acute. Ray florets lack¬ ing; disk florets ca. 15; corollas yellow, glabrous, ca. 8 mm long, tube ca. 4 mm long, throat ca. 2 mm long, lobe ca. 2 mm long and 0.6 mm wide, tips somewhat thickened with sclerified shield of large, slightly bulging cells outside; anther thecae ca. 2 mm long, with basal tails as long as collar; apical anther appendage narrowly oblong, ca. 0.4 mm long, ca. 0.12 mm wide. Achenes ca. 1.8 mm long, glabrous; pappus bristles ca. 7 mm long, not broadened at tips, without enlarged apical cells. Pol¬ len grains ca. 30 /am diam. The two available specimens of this species were obviously collected during the same trip to the Vol- can Sumaco area. The habitats were given as pri¬ mary and secondary forest, and humid or pluvial premontane forest. The species is one of the few described here from Ecuador without either ligulate or disciform rays. The glabrous leaves with prominent, arching, sec¬ ondary veins, and the short, lateral, mostly race¬ miform inflorescences are distinctive. A number of species with lateral inflorescences occur in Ecuador and Peru, but such species are rare in Colombia. One species in Colombia with lateral inflorescences, P. uribei J. Cuatrecasas, is very similar to P. hur¬ tadoi in form of the leafy branches and the inflo¬ rescences with homogamous heads, hut it has shorter involucral bracts and a mixed indument of erect pilosity with persistent bases and tenuous arachnoid hairs on the leaves (Cuatrecasas, 1985). Paratype. ECUADOR. Napo: carretera Hollin Lo¬ reto, 5 km al W de Cuamani, faldas del Volcan Sumaco, 00°43'S, 77°38'W, 1,200 m, 6-7 Sep. 1988, Neill, Hurtado & Alvarado 8558 (MO, US). Pentacalia (suhg. Pentacalia) luteynorum H. Robinson & J. Cuatrecasas, sp. nov. TYPE: Ecuador. Napo-Pastaza: Baeza-Tena road, remnant forest patches and roadside slopes from Cosanga to 5 km S of Cosanga, 1,975-2,225 m, 9 Jan. 1979, Luteyn & Lebron-Luteyn 6733 (holotype, US; isotype, NY). In foliis ellipticis glabris, in nervis secundariis obscuris in caulibus in partibus fertilibus inflatis in inflorescentiis terminalibus et in radiis elongatis albis distincta. Scrambling subshrubs or vines 2 m high or longer; stems terete, finely puherulous, becoming glabrous and slightly but distinctly inflated near base of in¬ florescence, with solid pith. Leaves alternate, petioles 0.7- 1.0 cm long; blades fleshy, drying thinly sub- coriaceous, elliptical, 5. 5-9.0 cm long, 2. 0-3. 7 cm wide, base short-acute, margins entire, apex short - acuminate, upper and lower surfaces glabrous, with only midvein prominent below; 5 or 6 obscure, up¬ wardly directed, secondary veins on each side. In¬ florescence terminal, broadly corymbose to broadly pyramidal; foliose primary bracts on broadened stem 288 Novon below inflorescence and at lower branches of inflo¬ rescence, lower bracts with petioles to 0.5 cm long, upper bracts sessile; blades glabrous, oblong, 2- 4 cm long, 0.4- 1.7 cm wide, apically obtuse; lower branches long and unbranched basally, corymbose distally, distal branches with short unbranched basal parts; peduncles 3“8 mm long, finely puberulous. Heads 8-9 mm high, heterogamous, radiate; sub- involucral and calycular bracts 5-8, subulate, min¬ ute, 0.8-1 .5 mm long, 0.5- 1 .0 mm wide; involucral bracts ca. 8, oblong, ca. 5 mm long, 1.0- 1.3 mm wide, apices rounded or obtuse, very sparsely pu¬ berulous. Ray florets ca. 8; corollas white, glabrous, tube 3. 0-3. 5 mm long, limb 3. 5-4.0 mm long, with 1 or 2 minute staminodia. Disk florets ca. 15; corollas yellow, tube ca. 2 mm long, throat ca. 2 mm long, lobes ca. 1.2 mm long, ca. 0.5 nun wide, tips somewhat thickened with sclerified shield of enlarged bulging cells outside; anther thecae ca. 1.3 mm long, basal tails half to two-thirds as long as collar, apical appendage narrowly oblong, ca. 0.25 mm long, 0.15 mm wide. Achenes submature, ca. 1 mm long, glabrous; pappus bristles 4. 5-5.0 mm long, tips not or scarcely broadened. Pollen grains ca. 27 yum diam. The paratype is cited from disturbed forest. The specimens of this species have been referred pre¬ viously to cf. Pentacalia jelskii (Hieronymus) J. Cuatrecasas of Bolivia and southern Peru, which has an elongate inflorescence and disciform or very shortly radiate heads, and P. huilensis (J. Cuatre¬ casas) J. Cuatrecasas of Colombia, which has more prominent secondary leaf veins, a more condensed inflorescence, and yellow rays. The new species seems to have a number of distinctive features, including the somewhat swollen stem at the base of the inflo¬ rescence and the white rays with minute staminodia. The species is named after the collectors of the type. Paratype. ECUADOR. Napo: Canton Quijos, along camino from Cosanga to Rio Aliso, ca. 5 km W of Cos- anga, approx. 77°53'W, 0°30'S, 2,000 m, 20 Feb. 1978, Kirkbride <£■ Chamba 4255 (US). Pentacalia (subg. Pentacalia) moronensis 11. Robinson & J. Cuatrecasas, sp. nov. TYPE: Ecuador. Morona-Santiago: along new road Mendez- Morona, km 55-62, 800 m, 23 Aug. 1989, van der Werff & Gudino 11385 (ho- lotype, US; isotype, MO). In caulibus fistulosis in foliis carnosis glabris in inflo- rescentiis lateralibus elongatis racemoso-thyrsoideis et in capitulis homogamis distincta. Plants scandent, indefinitely elongate; stems te¬ rete, rather straight, glabrous, fistulose. Leaves al¬ ternate, petioles 1. 5-3.0 cm long; blades fleshy, drying coriaceous, oblong-elliptical, 12-16 cm long, 7. 5-8. 5 cm wide in type, 3. 7-5. 5 cm wide in para- types, base shortly obtuse, short-acute in paratypes, margins entire, apex very short-acuminate, except at midvein surface smooth, glabrous, secondary veins obscure, ca. 5 on each side, spreading at ca. 45°. Inflorescences lateral, slightly longer than subtend¬ ing leaves, narrowly racemiform-thyrsoid with many branches along entire length; bracteoles minute, lin¬ ear, to 2 mm long; peduncles mostly 4 8 mm long, minutely puberulous. Heads 9-12 mm high, ho- mogamous, with somewhat turbinate bases; calyc¬ ular bracts 2 or 3, subulate, to 3.5 mm long, gla¬ brous; involucral bracts usually 8, glabrous, 8 10 mm long, 1 .5-2.0 mm wide, short-acute. Ray florets lacking; disk florets 15-23; corollas yellow, gla¬ brous, 8 9 mm long, tube 3-4 mm long, throat ca. 3 mm long, lobes ca. 1.8 mm long and 0.6-0. 7 mm wide, tips minimally thickened with sclerified shield of large, bulging cells outside; anther thecae ca. 2 mm long, basal tails three-fourths as long as collar; apical anther appendage ca. 0.45 mm long, 0.12 mm wide. Achenes immature, ca. 1 mm long, gla¬ brous; pappus bristles 8 9 mm long, not broadened at tips in 1 1385, broadened with short pointed cells in 4751 and / 1260. Pollen grains 35-37 yam diam. The specimens of this species are cited from lo- rnas. in humid, tropical, primary forest and forest remnants. The new species seems closest to P. ni- gella (Badillo) J. Cuatrecasas of Venezuela, which is similar in the straight, fistulose stems and ho- mogamous heads, but which has a much less branched, racemiform and scarcely thyrsoid inflo¬ rescence. The heads of the new species have tur¬ binate bases and the involucral bracts extend to the tips of the corollas, while the specimens seen of P. nigella have heads with rounded bases and corolla lobes distinctly exceeding the involucral bracts. The type specimen of the new species, van der Werff & Gudino 1 1385, which has the most mature heads, is similar to P. nigella in the lack of enlarged tips on the pappus bristles, but the leaves are broadly oblong to 8.5 cm wide. The paratype specimens, which may prove varietally distinct, have leaf blades more narrowly oblong as in P. nigella, but have the tips of the pappus bristles enlarged with short, broad apical cells. Paratypes. ECUADOR. Morona-Santiago: along road Mendez Morona, km 30 35, 800 m, 19 Aug. 1989, van der Werff <6 Gudino 1 1260 (MO, US); pazo petrolero “Garza” de Tenneco, 35 km (aprox.) al noreste de Mon¬ talvo, 01°49'S, 76°42'W, 260 in, 2- 12 jul. 1989, Zak & Espinosa 4751 (MO, US). Volume 3, Number 3 1993 Robinson & Cuatrecasas New species of Pentacalia 289 Pentacalia (subg. Pentacalia) napoensis H. Rob¬ inson & J. Cuatrecasas, sp. nov. TYPE: Ec¬ uador. Napo: 3 km este del Caserio de Iluamm, al norte de la carretera Ilollin- Loreto, por una trocha, 00°43'S, 77°36'W, 1,200 m, 17 Sep. 1988, llurtado & Alvarado 478 (holotype, US; isotype, MO). In foliis glabris in nervis secondariis ascendentibus in nervis et nervulis supra et subtus distincte dense promi- nulis in inflorescentiis lateralibus et in capitulis homogamis distincta. Plants scandent, indefinitely elongate; stems gla¬ brous, narrowly fistulose. Leaves alternate, petioles 1 .3-2.0 cm long: blades thinly coriaceous, glabrous, elliptical, 8-11 cm long, mostly 3.0 4.8 cm wide, base and apex shortly and narrowly acuminate, mar¬ gins entire, upper and lower surlaces with veinlets closely and distinctly raised above surface; second¬ ary veins ca. 6 on each side, spreading at 35-40°. Inflorescence lateral, from axils of vegetative leaves, one and a half to two times as long as leaves, un¬ branched in basal 6 cm, narrowly thyrsoid distally; peduncles mostly 5-10 mm long, finely puberulous, with few subulate bracteoles ca. 1 .5 mm long. Heads 12-14 mm high, homogamous; calycular bracts 3 or 4, similar to bracteoles; involucral bracts ca. 8, glabrous, 9 10 mm long, 1.0-1. 3 mm wide, apices short-acute. Ray florets lacking; disk florets 12-14; corollas yellow, glabrous, 10-11 mm long, tube 4. 5-5.0 mm long, throat 2. 0-2. 5 mm long, lobes ca. 1.5 mm long, 0.5 mm wide, tips not thickened, with subsclerified shield of large, somewhat bulging cells outside; anther collars not or scarcely inflated below; anther thecae ca. 1.5 mm long, with basal tails three-fourths as long as collar; apical anther appendage narrowly oblong, ca. 0.45 mm long, 0.12 mm wide. Achenes ca. 2.3 mm long, glabrous, faintly 7- or 8-ribbed; pappus bristles ca. 8 mm long, ta¬ pered at tips, without enlarged apical cells. Pollen grains ca. 28 n m diam. The single known specimen of this species is cited from “bosque pluvial Premontano,” and “bosque primario.” The species seems most distinct in the glabrous elliptical leaves with ascending secondary veins and veinlets strongly raised above the surface. The narrowly racemiform-thyrsoid lateral inflores¬ cences with narrow, homogamous heads are also somewhat distinctive. Pentacalia (subg. Pentacalia) pailasensis II. Robinson & .1. Cuatrecasas, sp. nov. TYPE: Ecuador. Santiago Zamora: trail between Mir- ador and Pailas, 2,010 2,255 m, 9 Sep. 1943, Stejermark 54275 (holotype, F). In foliis planis integris glabris in venulis prominulis in inflorescentiis lateralibus brevibus racemiformibus et in capitulis heterogamis disciformis differt. Plants scandent, indefinitely elongate; stems weakly striate, sparsely puberulous when young, becoming glabrous, narrowly fistulose. Leaves al¬ ternate, petioles 0.6-0. 9 cm long; blades not arched, not drying conduplicate, elliptical, weakly subcor- iaceous, glabrous, mostly 7-10 cm long, 0.19-4.0 cm wide, base short-acute, margins entire, apex short-acuminate, veins prominent below, reticulum of veinlets slightly but distinctly raised above the surface; secondary veins ca. 6 on each side, spread¬ ing at 60-70°, arching upward and joining well inside of margin. Inflorescences lateral from axils of leaves, slightly less long than the leaves, unbranched basal part 2. 0-2. 5 cm long, weakly puberulous, distal part mostly racemiform, bearing small subu¬ late or linear bractlets to 8 mm long, with heads solitary or in pairs on short lateral branches; pe¬ duncles lacking to 5 mm long, minutely puberulous. Heads 8-9 mm high, heterogamous, disciform; ca¬ lycular bracts 2 or 3, narrowly subulate, 2 3 mm long, glabrous; involucral bracts ca. 8, glabrous, 5 6 mm long, 1-2 mm wide, short-acute. Female florets ca. 4; corollas yellow , tubular, glabrous, tube ca. 4 mm long, lobes ca. 1 mm long, ca. 0.4 mm wide at base. Disk florets ca. 16; corollas yellow, glabrous, tube ca. 3 mm long, throat ca. 2 mm long, lobes ca. 1.5 mm long, 0.5 mm wide, tips scarcely thickened with sclerified mamillose cells outside; an¬ ther collar with cells of the short, broadened base poorly thickened in material seen; anther thecae ca. 1 .8 mm long, basal tails half to two-thirds as long as collar, apical appendage narrowly oblong, 0.28 mm long, ca. 0.01 mm wide. Achenes ca. 1.5 nun long, glabrous; pappus bristles 5-6 mm long, tips not broadened. Pollen grains 27-30 gm diam. The new species keys out close to Pentacalia sevillana and seems most closely related to that species. The latter differs most obviously by its arched leaf blades that are commonly conduplicate when pressed, by the more ascending secondary veins that reach near the leaf margin, and by the much longer, mostly narrowly thyrsoid inflorescences with strictly sessile heads. Pentacalia (subg. Pentacalia) palaciosii H. Rob¬ inson & .[. Cuatrecasas, sp. nov. TYPE: Ec¬ uador. Napo: Canto El Chaco, margen derecha del Rio Quijos, Finca “La Ave Brava” de Se- gundo Pacheco, 77°39'W, (XP^'S, 1,800- 1.900 m, 7-10 Sep. 1990, Palacios 5808 (holotype, US; isotype, MO). 290 Novon In foliis herbaceis vel tenuiter coriaceis in inflorescentiis pyramidaliter thyrsoideis et in capitulis heterogamis dis- ciformis distincta. Plants scandent or shrubby, often indefinitely elongate; stems roughened by persistent bases of evanescent hairs, narrowly fistulose. Leaves alter¬ nate, petioles ca. 1.5 cm long, with rough surface; blades oblong-ovate, firmly herbaceous to thinly co¬ riaceous, glabrous, 1.0-1. 3 cm long, 4-5 cm wide, base obtuse to rounded with short acumination, mar¬ gins entire, apex short-acuminate, upper surface dark, glabrous, with veins and veinlets scarcely raised above surface, lower surface somewhat paler, sparsely puberulous, with veins and veinlets distinctly raised above surface, included veinlets slightly raised above surface; secondary veins ca. 10 on each side, spread¬ ing at 60° or more. Inflorescence lateral from axils of leaves, nearly twice as long as leaves, bearing only reduced bracts, broadly thyrsoid with branches ascending at 30-60°, basal bracts elliptical, 3-4 cm long, 0.9- 1 .5 cm wide, upper bracts papery, mostly 0.5- 1.5 cm long, 0. 1-0.3 cm wide, more sharply acute; peduncles 0.7- 1.3 cm long, minutely pu¬ berulous. Heads 9-10 mm high, heterogamous, dis¬ ciform; subcalycular and calycular bracts 1 or 2, narrowly subulate, 3-4 mm long, glabrous; invo- lucral bracts ca. 8, glabrous, ca. 8 mm long, 1 .0- 1.5 mm wide, apices short-acute. Female florets ca. 4; corollas yellow, tubular, tube ca. 6 mm long, lobes ca. 1.3 mm long, ca. 0.5 mm long at base. Disk florets ca. 20; corollas yellow, glabrous, tube 4-5 mm long, throat ca. 2 mm long, lobes ca. 1.3 mm long, 0.5-0. 6 mm wide, tips not thickened, with nearly plain shield paved with large cells bearing central papillae outside; anther thecae ca. 1.5 mm long, basal tails half as long as collar, apical ap¬ pendage narrowly oblong, ca. 0.3 mm long, 0.1 mm wide. Achenes ca. 1.8 mm long, glabrous; pappus bristles ca. 6 mm long, tips not broadened. Pollen grains ca. 30 /am diam. In general aspect of leaf venation and position of inflorescences, the new species seems closest to Pen¬ tacalia hurtadoi described above, also from Napo. The leaves of the latter differ by the fewer secondary veins, the veinlets that are not prominent, the ho- mogamous heads, and the shorter, more racemiform inflorescences without small leafy bracts near the base. Pentacalia (subg. Pentacalia ) zakii H. Robinson & ,1. Cuatrecasas, sp. nov. TYPE: Ecuador. Pichincha: Reserva Kloristica — Ecologico “Rio Cuajalito,” km 59 de la carretera antigua Qui- to-Sto. Domingo de los Colorados, a 3.5 km al NE de la carretera estribaciones occidentalus del Volcan Pichincha, 00°13'53"S, 78°48'10"W, 1,800-2,200 m, 18 Sep. 1986, Zak 1213 (holotype, US; isotype, MO). In foliis late oblongis integris glabris in inflorescentiis lateralibus axillaribus foliis longioribus distaliter corym- bosis in capitulis subsessilibus heterogamis disciformibus distincta. Plants scandent, indefinitely elongate; stems stri¬ ated, sparsely and finely puberulous, producing many short adventitious roots, with solid pith. Leaves al¬ ternate, petioles ca. 1.8 cm long; blades herbaceous to thinly subcoriaceous, broadly oblong, 12-13 cm long, 5. 5-6.0 cm wide, base obtuse with slight acu¬ mination, margins entire, apex subacute and short- acuminate, upper and lower surfaces with veins and veinlets slightly raised above surface, essentially gla¬ brous, slightly paler below, with ca. 6 widely spread¬ ing, strongly arching secondary veins on each side. Inflorescences lateral from axils of vegetative leaves, half again as long as leaves, with long, unbranched, leafless basal part 10-12 cm long, corymbosely branched distally, branches ending with 2 or 3 sub- sessile heads; peduncles 0. 5-1.0 mm long, finely puberulous. Heads 9-10 mm high, heterogamous, disciform; calycular bracts 2 or 3, subulate, ca. 1.5 mm long, glabrous; involucral bracts 7 or 8, gla¬ brous, ca. 7 mm long, 1.0- 1.3 mm wide, apices narrowly acute. Female florets ca. 4, tubular; co¬ rollas white, glabrous, ca. 6 mm long, tube ca. 4 mm long, lobes ca. 1.7 mm long, 0.4 mm wide. Disk florets ca. 10; corollas white, glabrous, 6.0- 6.5 mm long, tube 3. 0-3. 5 mm long, throat ca. 1 .5 mm long, lobes ca. 2 mm long, 0.5-0. 6 mm wide, tips minimally thickened with subsclerified shield of large, somewhat bulging cells outside; anther the¬ cae ca. 1.5 mm long, basal tails ca. half as long as collar, apical appendage narrowly oblong, ca. 0.35 mm long, 0.13 mm wide. Achenes 2. 0-2. 5 mm long, glabrous, usually 5-ribbed; pappus bristles 4.0 4.5 mm long, tips not or scarcely broadened. Pollen grains ca. 27 /am diam. Pentacalia zakii seems closest to P. huilensis (J. Cuatrecasas) J. Cuatrecasas of western Colombia and cited here from Ecuador: Pichincha: Parroquia Nanegal, Webster & Castro 28943 (US). The latter species has leaf blades of similar texture and indu- ment and lateral inflorescences with long, un¬ branched bases. Pentacalia huilensis differs pri¬ marily by having distinctly ligulate rays, but also has thicker and more ovate leaves with more as¬ cending secondary veins. The presence of adventi¬ tious roots on the stem of the new species may be Volume 3, Number 3 1993 Robinson & Cuatrecasas New species of Pentacalia 291 variable, but no such roots have been seen in P. huilensis. The inflorescence of P. huilensis has a terminal, nearly sessile, corymbose inflorescence on the tapering tips of the stems in addition to the axillary lateral inflorescences, and the same form may be seen in the new species when more material is examined. Pentacalia (subg. Pentacalia) zamorana H. Rob¬ inson & J. Cuatrecasas, sp. nov. TYPE: Ec¬ uador. Zamora: road from Loja to Zamora, km 12-14, elev. 2,800 m, 18 Nov. 1961. Dodson cY Thien 1375 (holotype, US; isotype, SET). In foliis acutissimis argute dentatis glabris in inflores- centiis terminalibus dense pyramidaliter paniculatis et in capitulis sessilibus heterogamis disciformis distincta. Plants scandent, indefinitely elongate; stems gla¬ brous, striate, not or narrowly fistulose. Leaves al¬ ternate, petioles 0.5-0. 7 cm long; blades coriaceous, ovate, 3-6 cm long, 1 .7-3.5 cm wide, base rounded, margins regularly and sharply dentate, apex apic- ulate, surfaces with veins and veinlets strongly to moderately raised above surface, glabrous or rarely with traces of evanescent arachnoid tomentum near midvein; secondary veins ca. 6 on each side, spread¬ ing at 75-85° angles. Inflorescences terminal, densely pyramidal with widely spreading lower branches, with only small subulate bracts. Heads sessile in mostly small clusters, ca. 6 mm high, heterogamous, disciform; calycular bracts 3 or 4, subulate, 1 .0- 1.5 mm long, with few hairs; involucral bracts ca. 8, glabrous, ca. 4 mm long, 1.0- 1.5 mm wide, apices obtuse to short-acute. Female florets 3 or 4; corollas white, tubular, glabrous, tube ca. 2.3 mm long, lobes ca. 0.8 mm long. Disk florets ca. 10; corollas white, glabrous, ca. 4.5 mm long, tube ca. 1.8 mm long, throat ca. 1.8 mm long, lobes ca. 1 mm long, 0.5 mm wide, tips somewhat thickened w ith sclerified shield ol many narrow papillae out¬ side; anther thecae ca. 1 mm long, with basal tails half as long as collar; apical anther appendage ob¬ long, ca. 0.25 mm long, 0.17 mm wide. Achenes ca. 1.5 mm long, glabrous, 5-ribbed; pappus bristles ca. 4 mm long, without enlarged apical cells. Pollen grains ca. 30 /am diam. The new species seems closest to l*. millei from the same general area of southern Ecuador. The latter has only small teeth on the leaf margins and evanescent arachnoid tomentum on the leal under¬ surface and the surface ol the involucral bracts. Paratype. ECUADOR. Zamora: road from Loja to Zamora, km 12-14, elev. 2,800 m, 18 Nov. 1961, Dodson <£ Thien 1375 (SEL). Key to the Species of Pentacalia subg. Pentacalia in Ecuador la. Heads with distinct rays. 2a. Stem becoming slightly but distinctly inflated near base of the terminal inflorescence; inflorescence with primary bracts large, foliose, oblong; rays white . P. luteynorum H. Robinson & J. Cuatrecasas 2b. Stem not or scarcely swollen at base of inflorescence, usually without large primary bracts in terminal inflorescences; rays yellow. 3a. Inflorescences mostly or completely lateral from axils of lull-sized vegetative leaves. 4a. Stems and leaf undersurfaces with numerous crisped hairs; lateral inflorescences with foliose bracts on basal part . P carmelana H. Robinson & J. Cuatrecasas 4b. Stems and leaf surfaces essentially glabrous; lateral inflorescences without foliose bracts on basal part. 5a. Involucral bracts ca. 8; veinlets of leaf blade often slightly raised above surface . . P. huilensis (J. Cuatrecasas) J. Cuatrecasas 5b. Involucral bracts 5-7; leaf surface plane, with veinlets completely immersed . . P. riot intis (J. Cuatrecasas) J. Cuatrecasas 3b. Inflorescences mostly or wholly terminal on leafy branches. 6a. Leaves with tomentum spread over entire undersurface. 7a. Leaves small, 2-5 cm long, ovate, blunt; stems and leaf undersurfaces rufotomentose . P. ruficaulis (Greenman & J. Cuatrecasas) J. Cuatrecasas 7b. Leaves large, mostly 6-14 cm long, ovate-oblong, blunt; stems and leaf undersurfaces thinly arachnoid tomentose . P. campii (J. Cuatrecasas) J. Cuatrecasas 6b. Leaves glabrous or with tomentum only on midrib beneath. 8a. Leaf blades oblong or obovate, with apices rounded or obtuse to short-acute. 9a. Leaf blades oblong, with bases obtuse to short-acute, apices often rounded; veinlets distinctly raised above the surface . P. corazonensis (Hieronymus) J. Cuatrecasas 9b. Leaf blades obovate, with cuneate bases, apices obtuse to short-acute; veinlets scarcely raised above surface . P ■ dorrii H. Robinson & J. Cuatrecasas 8b. Leaf blades ovate, with apices acute. 10a. Leaf with trinervate venation; surface smooth, with veinlets not pellucid; margins entire . P. lanceolifolia (J. Cuatrecasas) J. Cuatrecasas 292 Novon 10b. Leaf venation pinnate; surface with pellucid veinlets raised above surface; margins often with a few teeth . P. andrei (Greenman) J. Cuatrecasas lb. Heads disciform or homogamous, without obvious rays. 11a. Heads homogamous, all florets bisexual. 12a. Leaf margins entire; inflorescences mostly or wholly lateral from axils of full-sized leaves. 13a. Leaf blades elliptic or oblong, fleshy with secondary veins obscure; inflorescences loosely racemiform-thyrsoid; corollas yellow . P. moronensis H. Robinson & J. Cuatrecasas 13b. Leaf blades elliptical, with prominent secondary veins; inflorescences mostly racemiform; corollas white. 14a. Leaf blades smooth between secondary veins; pith of stems solid; secondary veins arching and joining well inside of margin . . P. kurtadoi H. Robinson & J. Cuatrecasas 14b. Leaf blades with veinlets raised above surface; stems fistulose; secondary veins rather evenly ascending almost to margin . . P. napoensis H. Robinson & J. Cuatrecasas 12b. Leaf margins with teeth; inflorescences terminal or subterminal from axils of reduced leaves. 15a. Leaf blades densely tomentose below; heads sessile; corollas yellow . . P. gibbiflora (J. Cuatrecasas) J. Cuatrecasas 15b. Leaf blades glabrous or sparsely pubescent below; heads pedunculate; corollas white or greenish white. 16a. Leaves narrowly ovate to elliptical . . . . P. hitchcockii (J. Cuatrecasas) J. Cuatrecasas 16b. Leaves ovate. 17a. Axis of inflorescence mostly straight; leaves with secondary veins spreading at nearly 90° . P. arborea (HBK) H. Robinson & J. Cuatrecasas 17b. Axis of inflorescence ± zigzagged; leaves with secondary veins spreading at 70-80° angles . P. theae folia (Bentham) J. Cuatrecasas lib. Heads heterogamous, with tubular peripheral female florets. 18a. Leaf margins denticulate; inflorescences terminal. 19a. Leaf blades large, to 20 cm long, with closely and acutely denticulate margins; heads pedunculate . P. foribunda J. Cuatrecasas 19b. Leaf blades mostly less than 10 cm long, with less than 20 teeth on each margin; heads sessile. 20a. Undersurfaces of leaves with dense tomentum; branches of inflorescences often retroflexed, directed backward . P. oronocensis (DC.) J. Cuatrecasas 20b. Undersurfaces of leaves glabrous or with evanescent arachnoid tomentum; branches of inflorescences not retroflexed. 21a. Margins of leaf blades with small mucronate-denticulations; stems, leaf un¬ dersurfaces, and inflorescences with distinct evanescent arachnoid tomentum . P. millei (Greenman) J. Cuatrecasas 21b. Margins of leaf blades with numerous sharp teeth and intervening rounded sinuses; stems, leaves, and most of each inflorescence essentially glabrous . P. zamorana H. Robinson & J. Cuatrecasas 18b. Leaf margins entire; inflorescences partially or wholly lateral from axils of full-sized vegetative leaves. 22a. Undersurfaces of leaf blades and involucral bracts pubescent. 23a. Heads 10-13 mm high from base of receptacle to tips of disk florets, with ca. 40 disk florets and 10-12 short-ligulate, 3-5-lobed peripheral female florets . . P. hillii (Greenman) J. Cuatrecasas 23b. Heads mostly 9-10 mm high, with 16-30 disk florets and 4-7 peripheral female florets. 24a. Peripheral female florets 6-7, tubular, 5-lobed; bisexual disk florets 25-30 . P. disciformis (Hieronymus) J. Cuatrecasas 24b. Peripheral female florets ca. 4, subradiate, with deeply 3-lobed limb; bisexual disk florets 17-19 . P. carchiensis (J. Cuatrecasas) J. Cuatrecasas 22b. Leaf blades and involucral bracts glabrous. 25a. Lateral inflorescences with naked, long, unbranched basal parts that are longer than distal branching part, corymbiform distally; leaves broadly oblong . . P. zakii H. Robinson & J. Cuatrecasas 25b. Lateral inflorescences with basal parts short or closely bractiferous, thyrsoid or racemiform distally; leaves ovate or elliptical. 26a. Leaf blades smooth except for midvein, secondary and tertiary veins obscure; corollas white . P. cazaletii H. Robinson & J. Cuatrecasas 26b. Leaf blades with prominent secondary veins and weakly to strongly raised veinlets; corollas yellow or pale yellow. 27a. Heads loosely racemose, disposed on slender peduncles; bases of lateral inflorescences with foliose bracts . . P. palaciosii H. Robinson & J. Cuatrecasas Volume 3, Number 3 1993 Robinson & Cuatrecasas New species of Pentacalia 293 27b. Heads sessile or subsessile; bases of lateral inflorescences often with only small subulate bracts. 28a. Leaf blades somewhat arched, often conduplicate when pressed; secondary veins reaching near leaf margin; inflorescence longer than subtending leaves, distinctly thyrsoid with branches bearing many sessile heads ... P. sevillnna (J. Cuatrecasas) J. Cuatrecasas 28b. Leaf blades pressing flat; secondary veins strongly upwardly curved and joining well inside of margin; inflorescence not longer than subtending leaves, mostly racemiform with shortly pedun¬ culate heads . P. pailasensis H. Robinson & J. Cuatrecasas New Species of Pentacai.ia (subg. Pentacalia) in Peru Pentacalia (subg. Pentacalia) balsasana J. Cua¬ trecasas & H. Robinson, sp. nov. TYPE: Peru. Cajamarca: Celendin, Rio Maranon, canyon above Balsas, 3-4 km below summit, rd. to Celendin, alt. 2,950 m, 21 May 1964, Hutch¬ ison & It right 5212 (holotype, I S; isotypes, F, UC, USM). In nodis prominentibus in foliis oppositis in laminis oblongis base rotundatis vel breviter acutis subtus cinereo- tomentosis in capitulis radiatis et in bracteis involucri ca. 1 3 distincta. Shrubs ca. 2 m high; stems somewhat hexagonal, thinly white-lanuginose, nodes appearing swollen with persistent enlarged bases of petioles, with solid pith. Leaves opposite, petioles 1 .0-1.5 cm long, 0.8- 1.3 cm wide above enlarged persistent bases; blades subcoriaceous, 2. 5-8.0 cm long, 1.0-2. 8 cm wide, base rounded to short-acute, margins with numerous minute, close denticulations, sinuses as wide as deep, apex obtuse to short-acute, upper surface shiny green with slight evanescent, whitish, arachnoid wool, low¬ er surface covered with dense whitish tomentum; secondary veins ca. 8 on each side, widely spreading, reticulum of veinlets prominent below. Inflorescence terminal, and from axils of uppermost reduced leaves, broadly cymose with corymbose branches; peduncles 0.3-1 .5(— 2.7) cm long, thinly whitish tomentose. Heads 10 13 mm high, heterogainous, radiate; 8- 10 calycular bracts and few subinvolucral bracteoles 3-4 mm long, 0.6-0. 7 mm wide, with few arachnoid hairs, with tips acute and brownish, margins mi¬ nutely ciliate; involucral bracts ca. 13, ca. 6 mm long, mostly oblong, 1.7 1.9 mm wide, outermost linear, ca. 1 mm wide, all glabrous with acute ciliate apices. Ray florets 10; corollas yellow, glabrous, 10- 12 mm long, tube 3.0 -3.5 mm long, limb oblong- elliptical, ca. 8 mm long, 2. 2-2. 4 mm wide. Disk florets ca. 29; corollas yellow, ca. 8 mm long, gla¬ brous, tube ca. 3 mm long, throat ca. 5 mm long, lobes ca. 1 mm long, 0.8-0. 9 mm wide, lobes scarce¬ ly thickened at tips with scarcely bulging cells out¬ side; anther thecae ca. 2.5 mm long, basal tails a third and a half as long as collar, apical appendage oblong-ovate, ca. 0.4 mm long, 0.25 mm wide. Achenes submature, 2. 5-3.0 mm long, glabrous, weakly 5-ribbed; pappus bristles ca. 6 mm long, with long-projecting but not broadened apical cells. Pollen 37-40 /am diam. The type is described on the label as fairly com¬ mon, fragrant, a shrub to 2 m, coarse, with leaves glossy above, woolly below, and leathery. Pentacalia balsasana is one of three species of this genus in Peru with opposite leaves. Of the others, P. wur- dackii differs by being obviously a vine, by its lack of persistent petiole bases, its glabrous leaves, fewer involucral bracts, its lack of rays, and its more deeply cut corolla lobes. Pentacalia vargasiana from southern Peru is described as a shrub, but has cor¬ date bases and sparse pubescence on I lie undersur¬ faces of the leaves, lacks the persistent petiole bases, has the shortest anther tails seen in the subgenus, and has longer corolla lobes with more thickened and roughened tips. The three species do not seem closely related. Paratype. PERU. Cajamarca: Celendin, Rio Mara¬ non canyon, above Balsas, 3-4 km below summit on road to Celendin, alt. 2,950 m, 21 May 1964, Hutchison & Wright 52 12 A (F, UC, US, USM). Pentacalia (subg. Pentacalia) cutervonis II. Robinson & J. Cuatrecasas, sp. nov. TYPE: Peru. Cajamarca: Cutervo National Park, 12 km NE of San Andres de Cutervo, transect 4, 06°10'S, 78°40'W, 2,230 m, 11 Sep. 1991, Gentry, Diaz 1.5 cm diam., the current shoot(s) to about 0.8 cm. Leaves 4-6-foliolate, the base extending across the twig, sheathing, blunt above with no significant free portion; petiolules 1-2.8 cm long. Blades elliptic, glabrous, green and somewhat shiny above, darkly glaucous and minutely pubes¬ cent to glabrescent below (brightly sericeous-pubes¬ cent in young growth), 6.3-10 cm long, 3-6(-6.7) cm wide, stiff, coriaceous; apex rounded, the tip absent or retuse, base obtuse to rounded, ± decur¬ rent into the petiolule, margins entire, slightly in¬ curved when dry, curving ± uniformly; surfaces relatively smooth, the veins and reticulum above very slightly recessed when dry, ± slightly but dis¬ tinctly raised below; midrib beneath prominent, ± glaucous; primary veins 11 or 12, slender, spread¬ ing-ascending, gently rising, sometimes divaricating, abruptly curving and anastomosing just inside mar¬ gin to form an inframarginal vein; secondary veins variously developed, some extending almost to mar¬ gin; reticulation fairly fine, lacking in organized cross¬ veins. Inflorescence initially terminal, later pseu¬ dolateral, umbellate, basically twice compound (but some coflorescences with lateral single reiterations), the axes largely glabrescent, drying relatively dark; main axis absent; primary rays 1 (or perhaps 2), stout-looking, 24 cm long, 5.5 mm diam.; secondary rays all terminal, 1.5-6 cm long, essentially of two lengths; ultimate coflorescences umbellate, com¬ prising, if fully manifest, a central cluster of 6-10 or more bisexual, fertile flowers subtended by one or more umbellules of probably unisexual (male) flowers (evidently mostly absent or undeveloped or abortive in the material studied), the peduncle 0.8 cm long, sericeous-pubescent; ovary usually 3- or 4-locular. Maturing fruit gray-green, 3- or 4-angled, globose, about 5x5 mm; calyx rim about 1 mm wide, wall-like, the teeth relatively obscure; disk < 2 mm across, recessed; stylar column ca. 3 mm long, 3- or 4-fid at apex, the arms radiating at right angles, about 0.5 mm long. Schefflera dissidens is superficially similar to the apparently sympatric S. umbellata , but differs from it in the leaves having less strongly marked venation and margins not evidently incurved or revolute, inflorescences with 1(2) branch(es) and secondary rays of widely varying lengths with the longest ex¬ ceeding those in S. umbellata, and fruits 3- or 4-angled. In ,S. umbellata the secondary rays are mostly of similar length. The morphology of the leaflets also resembles ,S. acaro punctata, but in that species the leaflets are generally smaller and the inflorescence branches rather shorter; it usually also has one empty node. Schefflera dissidens is known only from the type collection and occurs on the summit of Mt. Roraima in open rocky areas with ravines and a small lake. 386 Novon Schefflera duidae Steyermark, Fieldiana, Bot. 28: 444. 1952. Contrary to Maguire et al. (1984), I believe that Schefflera quinquestylorum should retain specific rank. The arguments are presented under that spe¬ cies. Schefflera duidae subsp. duidae then becomes a plant with an extensive range in the western Gua- yana, extending also to Serra Araca in northern Brazil (Prance et al., 1992). Schefflera duidae is a shrub or small tree, 5-6 m tall, or a liana. It is found along streams or on forested sandstone ledges or declivities, at 1,250-1,800 rn in Estado Bolivar (Sierra Pacaraima, Cerro Sarisarinama) and Estado Amazonas (Cerro Cuao, Cerro Duida, Cerro Hu- achamacari, Cerro Neblina, Sierra Parima) in Ven¬ ezuela, and northwestern Amazonian Brazil (Cerro Neblina, Serra Pirapucu, Serra Surucucus). Schefflera eximia Frodin, sp. nov. TYPE: Brazil. Amazonas: Serra da Neblina [S rim], Camp 12 [in the headwaters of the Bio Ariobo drainage, 2 km SW of Pico da Neblina], 1,950 m, 26- 27 Feb. 1985, Hoorn, Buck & Brewer-Carias 5969 (holotype, K; isotype, NY). Figure 13. Arbor parva montium nebulosorum Schefflerae cdri- aceae affinis, at ab ea per folia minor elliptica etiam inflorescentias semel compositas statim diagnoscenda. Per- icarpium fructus in statu sicco ut videtur non sulcatum sed tantum rugosum. Tree 4 m tall, 12 cm diam.; twigs glabrous, ca. 8 mm across below apex, thickening gradually. Leaves 5- or 6-foliolate; petioles 16.5-22 cm long, base sheathing, not or only slightly pulling away from stem on drying, the free portion scarcely de¬ veloped; petiolules 1-5 cm long. Blades narrowly oblong-elliptic, glabrous above, rather finely puber- ulent below, 11-15 cm long, 5-6.2 cm wide, co¬ riaceous; apex tapering, shortly rounded, the tip retuse, base obtuse to rounded, margin flat or slightly incurved; upper surface fairly glossy, drying olive green, the veins and reticulum slightly recessed; undersurface initially rusty hairy, later turning patchily gray, somewhat veiny, the midrib strongly expressed; primary lateral veins 10-13 on each side, somewhat differentiated from the secondary veins, ascending, sometimes divaricating, gradually curv¬ ing and merging into a consolidated marginal vein; secondary veins partially to almost completely de¬ veloped; reticulum faintly expressed, in dry state more readily discernable above. Inflorescence ter¬ minal, umbellate, once compound, bracts fugacious; primary rays about 7, unbroken, glabrous, 15-17 cm long, 2.5 mm diam., drying dark; coflorescences simply umbellate, 20-25-flowered, the pedicels 18- 21 mm long, ± finely rusty sericeous; flowers not known. Fruit remaining inferior, globose, green be¬ fore maturity, to 7 x 7 mm, 5-angled when dry; calyx rim narrow, with 5 shallow teeth; disk about 3 mm across; stylar column 3 mm long, only the stigmata slightly discrete. Schefflera eximia is without apparent immediate relatives; outstanding features include the once com¬ pound umbellate inflorescence, comparable to those in the Plerandra group in the western Pacific, and, among species of the Guayana Highland, the rela¬ tively large, five-pyrenate fruits. The fine, rusty indumentum on younger leaflets is also distinctive. Somewhat coarse leaflet venation and five-pyrened fruits are also seen in S. coriacea and S. japurensis ; in the former, the primary rays of the inflorescence each have a subsidiary verticil of secondary rays, while in the latter the styles are free. Schefflera gracillima Steyermark & Maguire has 4- or 5-locular fruits, but in that species the leaves are smaller and the inflorescence much more delicate. The type collection contains another, possibly extraneous element: a twice umbellate inflorescence of about 6 primary rays 17-18 cm long, each with a naked intermediate node and about 25 terminal secondary rays or peduncles 3. 7-4. 5 cm long; on each of the latter are 10-12 fruits similar to but smaller than those described in the protologue, with the end of the stylar column a little more divided. No leaves are attached. The spread and proportions of the secondary rays and the naked nodes recall S. huachamacarii Maguire, Steyermark & Frodin, but in that species the fruits are 3-pyrened and the leaf has 3-5 leaflets. Schefflera eximia is known only from the type collection and is described as occurring in Bonnetia- dominated scrub forest. This is part of the tepui vegetation formation (Huber & Alarcon, 1988). Camp 1 2 was situated by a waterlogged savanna, but the environs, close to Pico da Neblina, also included a thick, stunted, heavily mossed cloud for¬ est (Brewer-Carias, 1988: 150). Schefflera guanayensis Maguire, Steyermark & f rodin, Mem. New York Bot. Card. 38: 66. 1984. TYPE: Venezuela. Amazonas: Cerro Guanay [summit area], 1,800 m, 2 Fell. 1951, Maguire et al. 31740 (holotype, NY). The original description of S. guanayensis con¬ tained elements of what are now S. clausa, S. eras- silimba, and S. paruana. The brief description of the nominate subspecies is likewise drawn from dis¬ cordant elements; the cited paratype, Steyermark et al. 108065, is S. clausa. A revised description, Volume 3, Number 4 1993 Frodin Schefflera in the Venezuelan Guayana 387 which incorporates one collection made since 1984, is accordingly presented below. Treelet or small tree 2-5 m, glabrous except for thinly appressed hairs on inflorescence axes and peduncles; twigs relatively thick and pithy, 1 . 1 cm diam. below tbe apex. Leaves 1 0- 1 2-foliolate; pet¬ ioles 13-27 cm, drying dark brown, the base ex¬ tending into a stipular ligule for ca. 1 cm long and widening below; petiolules unequal, 1.2-5 cm. Blades in a single plane, flattened, green above, pale green below, oblong to oblong-ovate to elliptic-ovate, the widest point at or somewhat above middle, 6-12 cm long, 3-6.7 cm wide, coriaceous; surfaces some¬ what glossy above, dull below, somewhat contrasting when dry but not markedly so; apex obtuse, tip acuminate or (in leaves from younger trees) shortly caudate, base broadly obtuse to subrotund, margins flat, unbroken; midrib somewhat prominent beneath; primary lateral veins ca. 10-14, not sharply con¬ trasting with mesophyll when dry except for being paler on the upper surface, spreading-ascending, extending to and intersecting with margin, hardly curving but partially breaking up distally into per- current tertiary cross-veins; secondary lateral veins divergent at a higher angle, partially developed, weakening into the reticulation. Inflorescences ter¬ minal, twice compound, paniculate, drying dark, the flowers in umbellules; primary branches up to 6, 20-27.5 cm long, 3 mm diam. near base, ascending to almost erect, thinly to sparsely appressed-hairy, basal bracts lanceolate, ca. 1.1 cm long; peduncles 28 50, scattered, slender, 0.9 3 cm long, their subtending bracts 2.5-3 mm long; umbellules ca. 10-flowered or so, pedicels about 2 mm long, the basal bracteoles 1 each, reduced or early caducous. Stamens 5. Immature ovary 2.5-3 mm long; disk gently sloping, then sharply curving into a stylar column 2 mm long, bifid at the apex, the stigmata ± spreading; calyx almost entirely adnate, the rim nearly flush with disk, with 5 small acute lobes. Fruit not seen, but probably remaining entirely inferior. Among related species, Schefflera guanayensis sensu stricto has no readily discernable features, except perhaps the comparatively small, fairly close¬ ly veined leaflets with unbroken margins and the slightly tapering partial panicles. Only ,S. crassilim- ba and clausa share this combination of relatively small leaflets and flowers in umbellules. Schefflera crassilimba, previously described as S. guanay¬ ensis subsp. sipapoensis Maguire, Steyermark & Frodin, may be distinguished by somewhat thicker, relatively narrower leaflets with ± narrowed bases, less contrast between the leaflet surfaces when dry, and somewhat longer inflorescence branches. Schef¬ flera clausa has a more slender appearance, foliage drying pale with distinctive purplish margins, and usually conduplicately folded leaflets. In addition, S. crassilimba is usually on lower slopes and in valleys, whereas S. guanayensis is found in more exposed situations, comparable to those of ,S. clausa. Small leaflets are also found in S. argophylla, S. contracta, and S. meiurophylla, but all these have their flowers in heads. The somewhat larger- foliaged but also umbelluliferous S. maguireorum (S. reticulata subsp. yutajensis) bas leaflets with coarser-looking venation and flowers almost in heads, the pedicels being only ca. 0.75 mm long. Schefflera paruana , formerly associated with S. guanayensis, has indented margins like those in S. pedicelligera Maguire, Steyermark & Frodin, and as presently known cannot be confused with the Cerro Guanay plants. The relatively thick twigs of this species resemble two other umbelluliferous species, both of the Neb- lina massif: 5. asymmetrica and S. concolor. These, however, have larger, differently veined leaflets and longer inflorescence branches and pedicels; more¬ over, the peduncles are not as widely ranging in length, thus causing the partial panicles to be less tapering. Also, the twigs of S. concolor are as much as 2.5 cm across when dry. Schefflera guanayensis inhabits scrub and dwarf forests in the summit area and on ridges. This is comparable to the situation of S. clausa. The only other known ridge and summit dwellers are the very distinct 5. tremuloidea and ,S. marahuacensis. Additional specimen examined. VENEZUELA. Amazonas: Serrania Guanay, sector NE, toward Rio Parguaza, ± 1,700 m, 20-28 Oct. 1985, Huber 11059 (MO, MYF not seen, NY). Schefflera huberi Frodin, sp. nov. TYPE: Ven¬ ezuela. Amazonas: Sierra de Maigualida, sector nor-occidental, cabeceras del Rio Iguana, af- luente del Rio Asita, 1,720 m, 25 Mar. 1988, Huber 12701 (holotype, K; isotype, MYF not seen). Figure 14. Arbor parva Schefflerae ulocephalae affinis, sed ab ea foliolorum venis lateralibus modice ascendentibus et ova- riis brevissime pedicellatis differt. Folia interdum 4-foliolata foliolis pagina infera minus tomentosa. Tree 5-7 m, with broad, dense crown; twigs pithy, 1.4 cm across, the surfaces, including the leaf bases, brown pubescent. Leaves 3- or 4-foliolate; petiole 13.5-16 cm long, the base broad, sheathing, with a free portion to 3 mm long; petiolules 0.9-2. 4 cm long. Blades oblong-elliptic, dark green and glabrous above, dark grayish maroon and tomentulose be- 388 Novon neath, 9.2-15.4 cm long, 3.8 8.1 cm wide, cori¬ aceous; apex rounded, tip absent, base broadly acute to rounded, margins entire, distinctly incurved; up¬ per surface somewhat glossy, moderately rugose, the venation recessed; undersurface strongly veined, the midrib prominent, decreasing distally; primary lateral veins 11-15, spreading-ascending, gently curving upward before passing into a semi-consoli¬ dated marginal vein formed from the anastomoses; secondary veins almost as strongly developed; cross¬ veins few to several on each side but variously ori¬ ented and passing only between primary and sec¬ ondary veins. Inflorescences initially terminal, later pseudolateral, umbellate, three times compound, at least the secondary bracts persistent; main axis not developed; primary rays ca. 26.5 cm long, 4 mm diarn., glabrescent, drying dark, their number un¬ known but presumably not more than 5 or 6; sec¬ ondary rays 2. 3-3. 5 cm long, dark-tomentulose, radiating hemispherically at ends of primary rays and outwards in a subsidiary pseudowhorl of 6 at ca. 6 cm below, subtended by bracts 4-5 mm long; ultimate coflorescences tertiary, with a cluster of central bisexual, fertile flowers on pedicels 2 mm long at ends of secondary rays and (0)1 or 2 lateral, tertiary, ca. 7-flowered umbellules arising immedi¬ ately below, the peduncles and pedicels of the latter respectively 8-12 mm and 2-3 mm long, tomen- tulose, the pedicels each subtended by a minute bracteole; calyx rim 1 mm wide, 5-toothed; ovary obconic, 1 mm long, the disk flat, the stylar column 2.5-3 mm long, the tip bifid or trifid. Immature fruit 4-5 mm long, bright green, 4 mm across, compressed or 3-angled when dry, hardly hetero- chronous. Among related species, S. huberi has the strong¬ est-looking venation and the least consolidated mar¬ ginal veins. It shares primarily trifoliolate leaves with S. baculosa and S. ulocephala, but differs from the latter in the flowers being pedicellate and the leaflets having thinner pubescence and more ascending ve¬ nation; in addition, the stylar column is distinctly longer. From the former it differs in the leaflet margins being incurved when dry, the undersurface more finely tomentulose with coarser, more strongly expressed and ascending venation, more slender pri¬ mary inflorescence rays, and pedicellate flowers. Schefflera huberi is known only from the type collection and is reported as being of sparse occur¬ rence in scrub vegetation of upper headwaters, at least some of it reported as secondary. Associated species may include ,S. longistyla , of similar habit and aspect, and the more readily distinguishable S. umbraculifera. Paratypes. VENEZUELA. Amazonas: Sierra Maig- ualida, NW sector, small valley along upper tributary of Cano Iguana, 2,000 m, 28 Feb. -3 Mar. 1991, Berry et al. 4833 (MO). Schefflera japurensis (Martius & Zuccarini ex E. Marchal) Harms, in Engler & Prantl, Nat. Pflanzenfam., Ill 8: 38. 1894; Maguire, Stey- ermark & Frodin, Mem. New York Bot. Card. 38: 71. 1984. TYPE: Colombia. Caqueta: Ar- aracoara, Japura River, Martius s.n. (holotype, M). This is one of the two most widely distributed species in the “Crepinella” group, though seemingly relatively scattered in occurrence. Knowledge of its range remains imperfect. Since 1984 it has been collected in two parts of Peru: Dept. San Martin, Rioja Province, logging road to N of Carretera Mar¬ ginal (km 431), 56 km W of Rioja, 2 km E of Naranjos, 980 m, 1 June 1986, Knapp et al. 7445 (MO); Dept. Huanuco, Pachitea Province, W part of Sira Mountains, above “Campamento Pato Rojo,” l, 200 m, 22 Dec. 1987, W’aUndfer 17-221287 (K, WU not seen), and 1,300 m, 23 Dec. 1987, Wall- nofer 12-231287 (K, WU not seen). These are ca. 700-1,000 km southwest and south of the southern edge of the Guayana Shield, where the otherwise most westerly record occurs (Araracuara on Rio Caqueta, southeastern Colombia). The several south¬ eastern Colombian records are in turn distinctly separated geographically from those in or near the Neblina massif, cited below. In addition, Cardona 1765 (US; see below) extends the range into the Gran Sabana uplands. Changes to the list of exsiccatae presented in Maguire et al. (1984) are also necessary, due to redeterminations. Steyermark 90225 belongs here, not under S. coriacea ; Cardona 1765 likewise be¬ longs here and not under S. umbellata ; and Maguire et al. 36904 is a distinct species, S. clusietorum. In Brazil, Froes 12638 likewise is distinct; it will be described elsewhere. Known Venezuelan records now include: Boli¬ var: Bio CaronI, Gran Salto de Eutobarima [as “Eutonarima”], orillas, 720 m, 8 Oct. 1946, Car¬ dona 1765 (US, VEN not seen); base central del Guaiquinima-tepui, 85 km S of La Paragua, 500 m, 1 1 May 1987, Aymard 5821 (MO, PORT not seen); Cerro Guaiquinima, Rio Paragua, Camp 3, 1,200 m, 19 Dec. 1951, Maguire et al. 32745 (K, NY); Guaiquinima, cumbre, Rio Szczerbauari (Carapo), 1-2 km from falls, 750 m, 20 Jan. 1977, Steyermark et al. 1 13241 (NY, US, VEN not seen); Guaiquinima, cumbre, Rio Szczerbanari (Carapo), Volume 3, Number 4 1993 Frodin Schefflera in the Venezuelan Guayana 389 1-2 km from falls, 750 m, 22 Jan. 1977, Stey- errnctrk et al. 113476 (NY, VEN not seen); Sierra Ichun, N de Salto Maria Espuma (Salto Ichun), 625-725 m, 27 Dec. 1961, Steyermark 9 0225 (NY, US, YEN not seen). Amazonas: Flanco N del Duida, bosque del Rio Negro, afluente del Cun- ucunuma, 210 m, 3°43'N, 65°35'W, Apr. 1990, A. Fernandez 7613 (MO, PORT not seen); flanco N del Duida, 2 km al S de la poblacion Culebra, 700 m, Apr. 1990, A. Fernandez 7773 (MO, PORT not seen); slope of Cerro Huachamacari, 850 m, 6 Mar. 1985, Liesner 18339 (MO, NY); vie. y al norte del Cerro Vinilla, ca. 30 km SSW de Ocamo, 440-600 m, 1 Mar. 1984, Steyermark et al. 130362 (MO, VEN not seen); [Neblina massif], 15 km NNE of Pico Phelps, Camp IV [in Canon Gran¬ de], 780 m, 15-18 Mar. 1984, Liesner 16708 (MO). For convenience, the following Brazilian rec¬ ord is also included: BRAZIL. Amazonas: Rio Ma- turaca, between Missao [de Maturaca] and Serra Pirapucu [SW of Serra da Neblina], 400 800 m, 13 Jan. 1966, Silva & Brazao 60814 (K, NY). The considerable contrast in the leaflet surfaces of S. japurensis is further enhanced by retention of the sericeous indumentum on the undersurface; this is reflected in descriptions ol it as “rusty” by more than one collector. Schefflera jauaensis (Maguire, Steyermark & Frodin) Frodin, comb, et stat. nov. Basionym: Schefflera reticulata subsp. jauaensis Ma¬ guire, Steyermark & Frodin, Mem. New York Bot. Card. 38: 59. 1984. TYPE: Venezuela. Bolivar: Cerro Jaua, cumbre de la porcion cen¬ tral-occidental, 1,922-2,100 m, 22-27 Mar. 1967, Steyermark 98078 (holotype, NY; is¬ otypes, K (imperfect), VEN not seen). An amended description is presented because of the exclusion of the original paratype of this taxon as well as the change of rank. I ree 3 m tall. Leaves 5-foliolate; petiole 40 cm long, 6 mm diam. near base, the base extending into a glabrous stipular ligule 2.4 cm long, its apex acute and body only gradually broadening below the mid¬ dle; petiolules 2-7.5 cm long, relatively stout in the manner of the petioles. Blades oblong, coriaceous, glabrous, 16.5-20.5 cm long, 8 9 cm wide, showing some evidence of conduplicate folding when lresh, the undersurface reported as silvery green; apex broadly rounded with a short, incongruously small acumen to 0.7 cm long, base shallowly cordate, margin wavy, tucked in at endings of primary veins; midrib fairly prominent; lateral veins about 16 on each side, conspicuous above and below, spreading, not strongly raised when dry but on both surfaces distinct, oriented straight toward margin and ending only at the marginal vein; secondary venation hardly developed and finer reticulation obscure. Inflores¬ cence presumably twice compound, paniculate, slightly whitish appressed-hairy all over, the length of the main axis and number of primary branches not known; primary branches about 60—61 cm long, subtended by a basal bract to 2.5 cm long and bearing numerous capitula; peduncles spreading, 2. 2-2. 9 cm long with persistent subtending bracts 4-8 mm long; capitula 1.1 1.4 cm diam., about 20-flowered. Ovary 2-locular; stylar column thick¬ ened, less than 1 mm long. Maturing fruit about 4 mm long, entirely inferior, distinctly fleshy, some¬ what compressed. Schefflera jauaensis most nearly resembles S. sessiliflora, but differs from that species in having longer peduncles and more congested heads with more flowers; the stylar column is also shorter. From 5. contracta and its relatives it differs in larger leaflets, inflorescences, and heads, although the mor¬ phology of the leaflets is most like S. contracta. As originally described, S. jauaensis encom¬ passed its type collection and Steyermark et al. 124075 from Cerro Marutani close to the Brazilian border. The latter, however, is S. sessiliflora, a species not reported from the Meseta de Jaua. Schefflera jauaensis is known only from the type collection and is described as inhabiting dense dwarf recumbent plateau forest bordering streams, of mixed woody growth with Bonnetia and Clusia. No habitat or association data are available for S. contracta, but the related S. meiurophylla does occur in com¬ parable situations on Serrania Uasadi. Schefflera longistyla Frodin, sp. nov. TYPE: Venezuela. Amazonas: Sierra de Maigualida, sector NW, cabeceras del Rio Iguana, afluente del Rio Asita, 1,720 m, 25 Mar. 1988, Huber 12692 (holotype, K; isotype, MYF not seen). Figure 15. Arbor parva Schefflerae ulocephalae cognata, sed haec species nova foliis quinquefoliolatis, foliolis latitudine 6 cm non excedentibus etiam inflorescentiis radiorum se- cundariorum verticillos subsidarios carentibus notata. Ovaria brevissime stipitata atque columna stylaris 3 mm longa. Tree 4-5 m, with broad, dense crown; twigs pithy, 1 cm across, the surfaces, including the leaf bases, pale brown pubescent. Leaves usually 5-foliolate (3- foliolate in young plants); petiole 10-15.5 cm long, the base broad, sheathing; petiolules 1-2.9 cm long. Blades narrowly elliptic to oblong-elliptic, green and 390 Novon Figures 15-18. — 15 (Top left). Schefflera longistyla Frodin. Huber 12692 (holotype, K). — 16 (Top right). Schefflera meiurophylla Frodin. Huber 12875 (holotype, K). — 17 (Bottom left). Schefflera suaveolens Frodin. Huber 12837 (paratype, K). — 18 (Bottom right). Schefflera umbraculifera Frodin. Huber 12682 (holotype, K). glabrous above, maroon to grayish maroon and fer- rugineous-tomentulose beneath, 6.6-14.6 cm long, 3. 2-5. 7 cm wide, coriaceous; apex rounded, tip absent or slightly refuse, base broadly acute to ob¬ tuse, margins entire, distinctly incurved; upper sur¬ face glossy, rugose, the venation and reticulum re¬ cessed; undersurface somewhat strongly veined, the midrib prominent, decreasing distally; primary lat¬ eral veins 12-15, somewhat spreading-ascending, curving slightly upward before passing into a con¬ solidated marginal vein formed from the anasto¬ moses; secondary veins extending to or nearly to Volume 3, Number 4 1993 Frodin Schefflera in the Venezuelan Guayana 391 the margin; cross-veins not regularly developed. In¬ florescences initially terminal, later pseudolateral, umbellate, three times compound, at least the sec¬ ondary hracts persistent; main axis not developed; primary rays ca. 32 cm long, 5 mm diam. in lower part, decreasing to 4.5 mm, initially (along with all other reproductive parts) pale pubescent, later gla- brescent, drying dark, their number unknown; sec¬ ondary rays about 15, 3.2-4 cm long, pale-tomen- tulose, radiating mostly hemispherically at ends of primary rays (with an occasional ray further down), subtended by bracts 4-5 mm long; ultimate coflo¬ rescences tertiary, with an almost headlike cluster of central bisexual, fertile flowers on pedicels 1.5 mm long at ends of secondary rays and about 3 lateral, tertiary, ca. 5- or 6-flowered headlike um- bellules arising immediately below, the peduncles and pedicels ol the latter respectively 10-13 mm and 1 mm long, pale-tomentulose, the pedicels each subtended by a minute bracteole; calyx rim 1 mm wide, 5-toothed; corolla green; stamens whitish. Im¬ mature fruit 4-5 mm long, bright green, 4 mm across, compressed or 3-angled when dry, hardly heterochronous; stylar column 3 mm long, barely bifid or trifid at the apex. Distinguishing features of Schefflera longistyla among related species are the narrowly elliptic to oblong-elliptic, somewhat rugose leaflets, the loca¬ tion of the secondary inflorescence rays usually all at or near the ends of the primary rays, the barely pedicellate flowers (and fruits), reiteration of inflo¬ rescences to no more than three times, and, in the fruits, the almost entirely united stylar column. The presence of pedicels separates it from S. baculosa, S. brachypoda, and ,S. ulocephala; the more evi¬ dently united styles, absence of a distinct subsidiary whorl of secondary rays and narrower leaflets sep¬ arates it from S. huberi ; and differently colored indumentum and lesser inflorescence reiteration (as well as the lack of a subsidiary verticil of secondary rays) distinguishes it from S. pallens (with which the type collection was originally identified). Schefflera longistyla is one of a partly geograph¬ ically concentrated series of species, apparently the result of localized diversification on Sierra de Maig- ualida and Serrania Uasadi. Elsewhere, members include S. ulocephala (Cerro Duida, Cerro da la Neblina) and S. pallens (Cerro de la Neblina). Paratypes. VENEZUELA. Amazonas: Sierra de Maigualida, sector nor-occidental [locality as for 12692], 1,720 m, 25 Mar. 1988, Huber 12696 (K, MYF not seen); Sierra Maigualida, NW sector, small valley along upper tributary of Cano Iguana, 2,000 m, 5°30'N, 65°15'W, 28 Feb. -3 Mar. 1991, Berry et al. 4834 (MO). Schefflera maguireorum Frodin, nom. et stat. nov. Replaced name: Schefflera reticulata subsp. yutajensis Maguire, Steyermark & Fro¬ din, Mem. New York Bot. Card. 38: 59. 1984, not Steyermark & B. Holst, 1988. TYPE: Ven¬ ezuela. Amazonas: Cerro Yutaje, summit, NW Ridge, Rio Manapiare, 1,400 m, 1 1 Feb. 1953, Maguire & Maguire 35154 (holotype, NY; isotype, K). An amended description is presented because of the transfer in this paper of one of the two original paratypes of this taxon to S. asymmetrica and the change of rank. Free 4 m tall; twigs initially 12 13 mm diam., increasing to 15 mm. Leaves 7- or 8-foliolate, the leaflets in a single plane; petioles 30-33 cm long, remaining sparsely gray-woolly through anthesis, later glabrescent, the axis emerging about halfway up the base, the free portion of the stipular ligule about 1.2 cm long; petiolules 2-4 cm long, initially thinly pubescent. Blades elliptic to narrowly elliptic, coriaceous, tending to be conduplicately folded, 12.5-15 cm long, 4. 8-5. 9 cm wide, glabrous with some scattered hairs near midrib on undersurface; apex obtuse, the tip shortly acuminate (not caudate), base ± smoothly rounded, margin slightly wavy; upper surface ± glossy; undersurface dull, slightly paler than the upper; midrib fairly prominent be¬ neath; primary lateral veins 9-12 on each side, only slightly raised below, spreading and curving upward slightly, extending to and uniting with the marginal vein; secondary lateral veins scarcely developed, with reticulation usually including 2 or 3 irregular cross-veins on each side toward the margins. Inflo¬ rescences terminal, twice compound, paniculate, with (6)7 or 8 radiating primary branches; main axis short; branches 25-33 cm long, finely but ± densely buff-pubescent at anthesis, subtended by basal bracts 2 cm long; headlike flower clusters numerous, crowded along most of the length of the branches, irregularly spaced, with some naked secondary bracts below them; peduncles 1-2.5 cm long, buff-pubes¬ cent, their subtending bracts 5-7 mm long. Flowers yellowish; pedicels about 0.75 mm long, densely buff-pubescent as the ovaries, each subtended by one bracteole; calyx rim very narrow, with 5 minute teeth; petals 5, coherent in a calyptra, the calyptra opening basally as the stamens expand with the segments separating upward; stamens ca. 3 mm long; anthers reddish; ovary about 1.25 mm long, the disk flat, the stylar column 2 mm long, its lower Vi pale and upper % dark when dry. Fruit as yet unknown. 392 Novon Schefflera maguireorum was originally described as a subspecies of S. reticulata (Gleason) Maguire, Steyermark & Frodin, but here the flowers are less clearly in heads; rather they are in congested clus¬ ters. The species thus links capituliferous (such as S. jauaensis and S. sessiliflora) and umbelluliferous (such as S. concolor and S. paruana) members of what appears otherwise to be a rather natural alli¬ ance — the “Cheilodromi”— all hut confined to Gua- yana, characterized most notably by marginally veined leaflets, paniculate inflorescences with elon¬ gate primary branches, and 2(3)-pyrened, wholly inferior fruits. The weakness as a character of the distinction between heads and umbellules has been discussed elsewhere (Frodin, 1975). Floral distinctions aside, the leaflets in S. ma¬ guireorum and S. paruana are superficially similar. However, those in the latter are somewhat larger and more thinly coriaceous, and the subsidiary lat¬ eral veins are somewhat more consistently devel¬ oped, with no evident cross-veins; in the former there are usually two or three such veins, any subsidiary venation usually being undeveloped or with occa¬ sional elongate secondary veins. In addition, in S. paruana the basal veins are more discrete, the un¬ dersurface somewhat glossier, and the midrib there less prominent. From Schefflera asymmetric a, the first collection of which was originally included here, S. magui¬ reorum can be distinguished by its smaller, more oblong leaflets with lesser development of secondary lateral venation, and flowers in almost headlike clus¬ ters with very short pedicels. Similarly, S. concolor has larger leaflets and flowers in distinct umbellules. Schefflera maguireorum is known only from the type collection and has been described as occurring in scrub on ridge summits. Its reported altitude is below that of the summit area and falls within the known local range of ,S. suaveolens (1,375-1,760 m) as well as the ranges, on other mountains, of more closely related species such as S. nigrescens, S. paruana, and S. pedicelligera. Schefflera marahuacensis (Maguire, Steyermark & Frodin) Frodin, comb, et stat. nov. Basion- ym: Schefflera tremuloidea var. marahuacen¬ sis Maguire, Steyermark & Frodin, Mem. New York Bot. Gard. 38: 63. 1984. TYPE: Ven¬ ezuela. Amazonas: Cerro Marahuaca, cumbre, seccion NW, 2,500 m, 16 Feb. 1981, Stey¬ ermark, Brewer-Cartas & Liesner 124406 (holotype, NY; isotypes, MO, US, VEN not seen). Trees 3-4 m tall; twigs pithy, 11-14 mm diam. Leaves 6-foliolate, tightly clustered toward twig ends; petioles glabrous, 16.5-19 cm long, emerging half¬ way up their associated stipular ligules; ligules 2- 2.5 cm long, 2 cm broad at base, with pale, scarious margins 1 mm wide; petiolules 4 6 cm long. Blades glabrous, ovate-elliptic, 5.8-11.2 cm long, 3. 4-7. 8 cm wide, coriaceous; apex acute, passing into an acumen 8-10 mm long (on new shoots the acumen to 25 mm), base obtuse to rounded, margin entire; both surfaces ± glossy, the upper more strongly shining; midrib on undersurface prominent, raised; primary vasculature comprising a pair of prominent, spreading basal veins and 9 or 10 additional veins on each side, the latter somewhat irregularly spaced, spreading-ascending, extending to margin, some¬ what prominent on undersurface but hardly raised; secondary veins partly developed; reticulation coarse. Inflorescence terminal, twice compound, paniculate; main axis absent; primary branches few, 25-46.5 cm long, drying dark purple or blackish, subtended at base by bracts 2 cm long; flowers in numerous racemosely arranged, fairly congested umbellules arising along upper % of primary branches, their peduncles 10-21 mm long and subtended by per¬ sistent secondary bracts ca. 2 mm long; pedicels 1 . 5(— 2) mm long, each subtended by 1 minute brac- teole; calyx rim scarious, with 5 small teeth; ovary obconic., about 1 mm long. The large stipular ligules (“auricles” in the pro- tologue), broader leaf bases, and larger leaflets were the character states originally used to distinguish these plants at varietal rank. Reexamination of the type, along with a subsequent collection, suggests that the twigs are also larger and thicker than in any of the material of S. tremuloidea Maguire, Steyermark & Frodin var. tremuloidea, the leaflets are thicker and more oblong, the umbellules are more uniformly and extensively distributed along the inflorescence branches, and the peduncles are shorter. Indeed, these plants are perhaps less closely allied to variety tremuloidea than that taxon is to S. pedicelligera, originally ranked as a species. Variety marahuacensis seems, therefore, worthy of specific rank. I am not able at present to postulate any close allies for S. marahuacensis. Apart from S. tremu¬ loidea, it is altitudinally the highest known member of its series. All collections to date come from Cerro Marahuaca, where it occurs in forests along creeks on the tepui summit at 2,500-2,600 m elevation. Additional specimen examined. VENEZUELA. Amazonas: Cerro Marahuaca, cumbre, parte central de la meseta sud-oriental, Quebrada Yekuana, 2,560 m, 10- 12 Oct. 1983, Steyermark 129431 (MO, NY, VEN not seen). Volume 3, Number 4 1993 Frodin Schefflera in the Venezuelan Guayana 393 Schefflera meiurophylla Frodin, sp. nov. TYPE: Venezuela. Amazonas: Serrania Uasadi, sector NW, cumbres montanosas ubicadas en las ca- beceras orientales del Rio Asita, afluente de- recho del Rio Ventuari, ± 1,850 m, 22 Nov. 1988, Huber 12875 (holotype, K; isotype, MYF not seen). Figure 16. Arbor Schefflerae contractae valde affinis, seel ab ea foliis fere truncato-obovatis nec conduplicatis paginis su- perna infernaque minus discrepantibus recedit; vere inter duas omnino in effectu exiguior. Tree 4-6 m tall, sparsely foliaged and somewhat flat-topped, with a slight smell when cut. Leaves gathered toward twig ends, 5- or 6-foliolate; petiole glabrous, somewhat darkened when dry, 6.5-14 cm long, less than 3 mm thick, the base extending into a stipular ligule > 3 mm long, acute; petiolules 1- 4.1 cm long, ca. 1.5 mm thick. Blades almost trun- cate-obovate, widest well above the middle, stiff when dry, green above, pale green below, 5.7-11.8 x 3-6.7 cm, not conduplicately folded; apex rounded- truncate, occasionally slightly emarginte, the tip cus¬ pidate-acuminate, 5-6 mm long, base broadly cu- neate, and margins somewhat incurved; upper surface when dry ± shining, not much darker than lower surface, the veins conspicuous; lower surface with midrib only somewhat prominent, the veins ± distinct from the mesophyll; primary lateral veins closely spaced, 12 or 13 on a side, ascending, ex¬ tending straight to margin and little forking; sec¬ ondary veins fairly well developed, breaking into irregular fine reticulation hut without evident cross¬ veins. Inflorescences terminal, semi-erect, twice compound, paniculate, the axes rather dark maroon; length of main axis or number of branches not known; primary branches (from different inflores¬ cences) 14.5 39.5 cm long, 2-4 mm diam., patch¬ ily or thinly pale hairy; bearing racemosely arranged capitula in the upper % and subtended by a basal bract to 2 cm long; peduncles pale hairy, 6-12 mm long, ca. 1 mm diam., subtended by persistent bracts 2-3 mm long; capitula ca. 5 mm across after an- thesis, later up to 10 mm, ca. 15-flowered; calyx rim narrow, 5-toothed; ovary 2-locular, the disk flat or slightlv sloping, small, passing into a short hut distinct stylar column 0.5 mm long with free parts to another 0.25 mm long. Fruit obovoid-ellipsoid, to 5 mm long and 4 mm across, wholly inferior, initially green, later red; stylar column persistent. Schefflera meiurophylla is most closely related to S. contracta and also shows affinities with S. argophylla. It differs from S. contracta in the pet¬ ioles longer in relation to the blade length and thus less congested; the petiolules as well as the petioles less stout and not obviously purplish; the blades not conduplicate and distinctly obovate with the widest point well above the middle; the surfaces of the blades with less contrast; and the midrib on the undersurface less prominent. Schefflera meiurophylla is known only from the type collection and occurs in low, mixed montane forests, at least sometimes along creeks. On the Sierra de Maigualida it seems to occur in moister habitats than S. argophylla. Schefflera monosperma Maguire, Steyermark & Frodin, Mem. New York Bot. Card. 38: 51. 1984. Replaced name: Crepinella gracilis E. Marchal ex Im Thurn, Timehri 5: 195. 1886; I'rans. Linn. Soc. ser. 2, Bot. 2: 275. 1887, not Schefflera gracilis (Miquel) Viguier 1909. TYPE: Venezuela [as “Guyana”]. Mt. Rorai- ma, upper slope, above 5,000 ft., 10 Dec. 1884, Im Thurn 162 (lectotype, K; isotype, BM). When Maguire et al. (1984) proposed the new name Schefflera monosperma for this species, the reference for the replaced name was incomplete and is given in full here. With the segregation of Schef¬ flera cracens, discussed more fully under that spe¬ cies, S. monosperma becomes limited to the Ro- raima-tepui and its outliers. Further collections have become available or have been rediscovered in the last decade and are cited below. Maguire & Ma¬ guire 40837, doubtfully identified in Maguire et al. (1984), is confirmed as S. monosperma. Observa¬ tion of fruiting material supports the contention of Baumann-Bodenheim (1955) that the ovary is pseu- domonomerous. Additional specimens examined. GUYANA. Mt. Ro- raima, 6,900 ft., 26 Mar. 1978, A. lledley & P. J. Edwards KER.081 (K); Mt. Rorainia, escarpment, 7,600 ft., 1 Nov. 1973, Persaud 144 (BRG not seen, K). VENEZUELA. Bolivar: Uei-tepui (Serra do Sol), on Brazil /Venezuela frontier, 6,500 ft., 28 Dec. 1954, Ma¬ guire & Maguire 40837 (K, MG not seen, NY, YEN not seen). BRAZIL. Mt. Roraima, Abhang, 2,200 m, Jan. 1910, Ule 8706 (K, L). Schefflera morototoni (Aublet) Maguire, Stey¬ ermark & Frodin, Mem. New York Bot. Card. 38: 51. 1984. TYPE: illustration of Panax undulata [sic] Aublet, Hist. I’l. Guian. 2: pi. 360. 1775; supporting herbarium collection yet to be selected, hut at BM there is an Aublet specimen seen by both Seemann (1868) and the author. Panax splendens Kunth in HBK, Nov. Gen. Sp. 5:11 (ed. fol. 9). 1821. Schefflera splendens (Kunth) 394 Novon Frodin ex Lindeman in A. Pulle et al., FI. of Suriname 3(2): 352. 1986. Didymopanax splendens (Kunth) J. Decaisne & Planchon ex Seemann, J. Bot. 6: 132. 1868. TYPE: Colombia. Cauca: Popayan, Hum¬ boldt & Bonpland s.n. (lectotype, selected by See¬ mann, 1868, P not seen). Panax splendens Kunth was first reduced to the slightly earlier name Aralia rnicans Humboldt & Bonpland ex Schultes by Krug & Urban (1899). They retained that taxon, however, as a distinct species under the name Didymopanax rnicans (Humholdt & Bonpland) Krug & Urban. It is per¬ haps not yet fully recognized that both of these names, which may be homotypic (Jan Lindeman, pers. comm.), represent the juvenile form of S. rno- rolotoni. The capitula included with the type of Aralia rnicans (B-WILLD) are extraneous. Prance et al. 4700 should he omitted from the list of exsiccatae given under this species in Maguire et al. (1984: 51-52); it is a distinct species, which will be described elsewhere. The following records from the Venezuelan Gua- yana are assigned here to Schefflera morototoni: Amazonas: Alto Orinoco, Isla del Esfuerzo, ca. 200 m, 5 Sep. 1951, Croizat 562 (F); bosque ribereiio del Rio Asisa, 100 m, 4°30'N, 65°48'W, Oct. 1989, L. Delgado 887; Rio Puruname, 40 km above mouth, ca. 100 m, 29 May-5 June 1982, Huber Tillett 06390 (K, MYF not seen); Rio Orinoco, Cano Tama-Tama, 150 m, 12 Sep. 1954, J. S. Level 142 (MO, NY); Cerro Sipapo, Cano Grande, slopes above lower escarpment, 4,500 ft., 21 Jan. 1949, Maguire & Politi 28528 (MO, NY, VEN not seen); Sierra Parima, near Simarawochi, 795-830 m, 18 Apr. -25 May 1973, Steyermark 107383 (NY, VEN not seen); 1’ rente No. 1, Piedra Sapo, Rio Atacavi, 140 m, Nov. 1989, J. Velazco 0997 (MO, POR I' not seen). Bolivar: Campamento “El Paraiso,” 48 km NE del caserio Los Rosos, 17 km de Upata, altitude not given, Apr. -June 1965, Blanco 039 (US, VEN not seen); Cerro Curichapo, 25 km al NE de la Paragua, 450 m. Mar. 1987, Fernandez 4083 (MO, POR I' not seen); bosque en lomerio 4-5 km al N del Cerro Camaron, Macizo Guaiquinima, 240 m, Oct. 1988, Fernandez Aymard 4813 (MO, POR I' not seen); alrededores de San Ignacio, 240 m, 27 Apr. 1987, L. Her¬ nandez 486 (MO); SW slope of Amuray-tepui, E of Auyan-tepui, W of Aparaman-tepui, 600-700 m, 28 Apr. 1986, Holst & Liesner 2742 (MO); 20-35 km SW of Manteco, road to San Pedro de las Dos Bocas, ± 200 m, 1-3 Aug. 1978, Liesner & Gonzalez 5862 (MO, VEN not seen); Chinese Slide, Cerro Bolivar, 300-400 m, 24 Oct. 1953, Maguire et al. 35966 (NY, VEN not seen); Mina “Los Pijiguaos,” Bajo Rio Suapure, altiplanicie de la Mina, ± 600 m, Apr. -May 1987, Ramirez <§: Paredes 77 (MO, MYF & TFAV not seen); Imataca, altitude not given, July 1983, Stergios et al. 6032 (MO, PORT & UEC not seen); Gran Sabana, be¬ tween Kun and Lladuara-paru [S of Mt. RoraimaJ, l, 065-1,220 m, 1 Oct. 1944, Steyermark 59085 (F); between Santa Teresita de Kavanayen and Rio Pacairao (tributary of Rio Mouak), 1,065-1,220 m, 20 Nov. 1944, Steyermark 60436 (F, NY); Cerro La Reforma, above junction of Rio Reforma with Rio Toro, 200-250 m, 15 Dec. 1960, Stey¬ ermark 88099 (F, GH, NY, US); Rio Paramichi entre la boca del Rio Paramichi y el Salto de Chal- imano, 525-625 m, 8 Jan. 1962, Steyermark 90762 (NY, US, VEN not seen); bosque a 2 km al NE de Santa Maria de Erebato, 370 m, Feb. 1989, M. Sucre 2562 (MO, PORT not seen); Medio Caura, Salto de Para, 280 m, 9 Mar. 1939, LI. Williams 1 1455 (F, LJS, VEN not seen); at E base of Cerro Coroba, Hato la Vergarena, 400 m, 19 Oct. 1954, Wurdack & Guppy 124 (F, NY, US); Cerro Mar- imarota, 100-250 ft., 26 Jan. 1956, Wurdack & Monachino 41384 (MO, NY, VEN not seen). Delta Amacuro: 5- 1 4 km ESE of Los Castillos de Gua- yana, 50-200 m, 28 Mar.-2 Apr. 1979, Davidse & Gonzalez 16279 (MO, VEN not seen); mountain area ca. 13 km by road ESE of town of Sierra Imataca, 400 m, 4-6 Apr. 1979, Davidse & Gon¬ zalez 16582 (MO, VEN not seen); vicinity of Cano Jotajana, tributary of Cano Cuiniquina, NW of Eparia, 50 m, 20 Oct. 1977, Steyermark et al. 1 15094 (MO, VEN not seen). Schefflera morototoni var. august ipetala (E. Marchal) Frodin, comb. nov. Basionym: Di¬ dymopanax morototoni f3 angustipetalum [sic] E. Marchal in Martius, FI. Bras. 11(1): 241. 1878. TYPE: Venezuela. Amazonas: Pacimoni River, banks, low altitude, [11-12] Feb. 1854, Spruce 3456 (K, NY; lectotype not selected at this time). I bis taxon has evidently not been recorded before for Venezuela. Marchal, in his protologue, mistak¬ enly attributed the type locality to Brazil; but the Rio Pacimoni, a tributary of Brazo Casiquiare, lies wholly within Venezuela (Huber & Wurdack, 1984). Marchal distingushed this variety by its petals being three times as long as broad and by the pe¬ duncles of the umbellules being reflexed. To this may be added that the leaves are only 5-foliolate, the primary veins of the leaflets curve continuously, the inflorescence appears only to be twice com¬ pound, with primary branches to 38 cm long, and Volume 3, Number 4 1993 Frodin Schefflera in the Venezuelan Guayana 395 the peduncles are 3-4 cm long, with the longer ones toward the ends of the branches. However, with the imperfect collections available (neither of them fruit¬ ing) I hesitate to recognize it at any higher rank. The reflexed peduncles resemble another species, as yet undescrihed, which occurs outside the Ven¬ ezuelan Guayana in a broken arc from eastern Co¬ lombia to northeastern Peru. Representative collec¬ tions include Schultes & Cabrera 16010 (GH. US) and Croat 18642 (MO, NA). The foliage may be compared with S. tamatamaensis Maguire, Stey- errnark & Frodin, but without knowledge of the fruits in the present taxon no further comparisons are attempted. Additional specimen examined. Venezuela. Ama¬ zonas: Cano Tama-Tama, 2 km al E de Tama-Tama, 140 m, 3°07'N, 65°50'W, 3 Mar. 1990, Aymard & Delgado 8430 (MO, PORT not seen). Schefflera myrioneura Frodin, sp. nov. TYPE: Venezuela. Amazonas: Cerro Sipapo, Phelps Camp to North Savanna, 1,400 m, 17 Dec. 1948, Maguire & Politi 27738 (holotype, K; isotypes, MO, NY, FIS). Arbor mediocris ad 1 5 m alta, Schefflerae psilophyllae ut videtur affinis sed ab ea foliolorum venis numerosissimis (primariis cum secundariis 35-60) atque arte patentibus et fructu in statu sicco non sulcato differt. Bene notabilis. Tree 12-15 m tall, the trunk 12-15 cm diam. Leaves (4-)6 8-foliolate; petiole ca. 24 cm long, 4 mm diam.; petiolules 2.4-7 cm. Blades usually ob- long-obovate but sometimes obovate, glabrous, 9.5- 25.5 cm long, 6. 7-8. 6 cm wide, chartaceous when dry; apex broadly rotund or truncate, ± distinctly emarginate, base acute to more usually broadly ob¬ tuse, the edges sometimes recurving into the peti- olule, margins entire, in the dry state slightly in¬ curved; surfaces when dry about the same color or the upper slightly lighter; midrib moderately prom¬ inent beneath, longitudinally collapsing on drying; lateral venation very fine, the primary and second¬ ary veins together 35-60 on each side, spreading, extending to a continuous marginal vein, the veins sometimes narrowly once divaricating toward the margin; tertiary lateral venation ± elongate, emerg¬ ing from the midrib or primary or secondary lateral veins. Inflorescence initially terminal but later pseu¬ dolateral, umbellate, compound, the primary and secondary branches stiff, straight, and stout; main axis hardly developed; primary rays 3 or 4, glabrous, 11.5-27 cm long, 4-5 mm diam., with or without a subsidiary whorl of secondary rays (when present, the upper internode > 4.5 6 cm long); secondary rays flattened, 2. 5-6. 5 cm long, up to 10 at ends of primary branches and 2-4 in subsidiary whorls, those at the ends oriented in all directions; ultimate coflorescences umbellate, with 3-5 central, bisexual flowers on pedicels 5-7 mm long and lateral tertiary reiterations with flattened peduncles (tertiary rays) 1-2.7 cm long and congested clusters of flowers on pedicels 2—3 mm long; flowers small, 4x2 mm in bud, the calyx rim poorly developed with minute teeth, the corolla calyptrate, pale when dry; ovary drying dark, 3- or 4-locular, the stylar column ca. 1 mm long. Fruit 9-10 mm long, 7-8 mm across, the widest point above the middle during maturation, remaining inferior, the disk depressed, ca. 2 mm across, the stylar column 1.5 mm long without free portions. Schefflera myrioneura differs from S. psilo- [ >hylla , where it was previously included by Maguire et al. (1984), in having fewer primary rays in the inflorescence as well as emarginate leaf apices. It also differs from S. suaveolens in the presence of fewer terminal secondary rays, larger fruits, and united styles. From both these species it differs in its very fine lateral venation, although in this respect it is closer to ,S. suaveolens. The inflorescence has an angular appearance, which among presumably related species in the Guayana can best be compared with S. pallens rather than either S. psilophylla or S. suaveolens. This new species is an occasional to frequent tree of wet mixed montane forests, some at least of a mossy aspect. Like many other araliads in forest zones, it may be an opportunist, taking advantage of gaps created by rockfalls, windthrows, or fires, or dwelling on forest or scrub edges. The structure of the vegetative parts suggests that a considerable amount of water may be lost upon drying. By con¬ trast, the fruits when dry do not shrink around the pyrenes as is common in most species of Schefflera; in this they may be compared with S. gabriellae Baillon of New Caledonia. Schefflera myrioneura is one of the distinctive endemics of Cerro Sipapo, along with ,S. crassilimba, S. pauciradiata Maguire, Steyermark & Frodin, and S. sipapoensis Maguire, Steyermark & Frodin. This westernmost of the Venezuelan Guayana tepuis, however, is “merely the highest point in a vast area of gently rising contour” (Paynter, 1982: 189). Shortly to the east are two patches depicted on the vegetation map of Huber & Alarcon (1988) as hav¬ ing “vegetacion tepuyana”; these areas are appar¬ ently unexplored botanically. This suggests a pos¬ sible larger “support area” for four such distinctive endemics — two of them initially described in 1984 — rather than merely the 55 km2 summit area (95 396 Novon km* if the talus slopes are included) of Cerro Sipapo alone. Paratypes. VENEZUELA. Amazonas: Cerro Sipapo, Savanna Camp to North Escarpment, 1,400 m, 23 Dec. 1948, Maguire & Politi 27880 (K, NY); Cerro Sipapo, above Cano Grande, 1 km NW of Savanna Camp, 1,500 m, 28 Dec. 1948, Maguire & Politi 28015 (K, NY); Cerro Sipapo, above lower escarpment, above Cano Gran¬ de, 4,500 ft., 21 Jan. 1949, Maguire & Politi 28539 (K, NY). Schefflera neblinae (Maguire, Steyermark & Frodin) Frodin, comb, et stat. nov. Basionym: Schefflera coriacea subsp. neblinae Maguire, Steyermark & Frodin, Mem. New York Bot. Card. 38: 79. 1984. TYPE: Brazil. Amazonas: slope of Serra Pirapucu [NE of] Rio Maturaca, [SW of] Serra da Neblina, 1,300 m, 26 Jan. 1966, Silva Brazao 60903 (holotype, NY; isotype, K). Tree 3 m high, 10 cm diam. Leaves 5- or 6-foliolate; petioles 10-13.5 cm long, the sheathing base small; petiolules 1. 3-2.1 cm. Blades oblong- elliptic to oblong-obovate, usually broadest above middle, the surfaces contrasting, 9 13.1 cm long, 4-5 cm wide, coriaceous; apex broadly obtuse, usu¬ ally ending in a broad point, base obtuse, rounded at insertion and not decurrent, margin only narrow¬ ing about !^-%-way down from apex, incurved but not noticeably inrolled; upper surface glabrous, the venation hardly impressed, undersurface grayish hairy, the indumentum tightly appressed; midrib prominent below; primary lateral veins about 12 14 on a side, ± closely parallel, just showing as thin lines on undersurface, near margin suddenly anastomosing and each joining with the primary vein below; secondary venation not well developed, usu¬ ally breaking and toward margin joining with pri¬ mary vein below. Inflorescence terminal, compound- umbellate; primary rays ca. 3-5, 28-30 cm long, drying very dark; secondary rays both at apex and in a single subsidiary whorl, some 20-23 at the former, far fewer at the latter; all axes dark seri¬ ceous-pubescent, the hairs much appressed, but later glabrescent; umbellules 13-15-flowered, the pedi¬ cels fairly slender, 7-9 mm long, sericeous-hairy; buds 3 mm long; petals 5, in bud sericeous without, at anthesis spreading outwards before falling; ovary 3-locular, the stylar column united, 0.5 mm long. I'his species is known only from the type collec¬ tion. For Prance, Steward, Ramos <£• Farias 979 6 from the Serra Parima, formerly included with the subspecies (Maguire et ah, 1984: 79), see Schefflera varisiana; the remaining records included by Ma¬ guire et al. (1984: 79-80) are S. acaro punctata. The generally smaller foliage and inflorescences, the absence of a distinctively reddish indumentum, and a shorter stylar column along with the trilocular ovary all indicate that specific rank is merited for this taxon. The only known locality of ,S. neblinae is relatively remote from the reported range of S. coriacea, although S. varisiana, clearly a sister species of S. neblinae, occurs closer. Schefflera neblinae is also allied to S. pauciradiata Maguire, Steyermark & Frodin. Schefflera nigrescens Frodin, sp. nov. TYPE: Venezuela. Amazonas: 1 km N of Camp XI, 6.5 mi. NNE of Pico Phelps [Cerro Neblina], 1,300 m, 28 Feb. 1985, Nee 31 186 (holotype, NY; isotypes, MO, US). Species habitu formaque Scheffleras varisianam et neblinam in memoriam redigit, sed haec species nova foliolis anguste obovatis, ad basin cuneatis vel attenuatis, in pagina super minus rugulosis, ad apicem late obtusatis vel rotundatis notata. Foliola valde in colore discrepantes, inter paginas supernam infernamque. Tree 8 m tall, trunk to 20 cm diam. Twigs fairly slender, 6 mm across in upper portion when dry. Leaves 5- or 6-foliolate; petioles 15-16 cm long, 2 mm diam.; basal sheath not extending into a stipular ligule; petiolules 1-2.5 cm long. Blades in a single plane, narrowly obovate, 9.2 13 cm long, 3. 3-4. 4 cm wide, thinly coriaceous; surfaces contrasting, the upper quite glossy, drying quite dark, glabrous with distinctly sunken primary and secondary lateral veins, the under finely golden tan hairy, with only the midrib prominent; apex rounded or broadly obtuse, passing into a short acumen 2-3 mm long, base cuneate or attenuate, not constricted at insertion with the petiole, margins when dry curving slightly upward; lateral veins many, spreading-ascending, less hairy below and so contrasting with the pubes¬ cent mesophyll, visibly camptodromous, arching near but not evidently reaching the margin, the intra¬ marginal loops obscure. Inflorescence compound- umbellate, probably terminal and without a main axis; primary rays 2(3?), about 25 cm long, bearing at some 8-10 cm below the end a subsidiary whorl of 2-5 secondary rays 2.6-4 cm long, the remaining secondary rays all at the end; terminal sprays com¬ prising 19-21 secondary rays (or peduncles) 2.3- 3.4 cm long, these somewhat golden sericeous at anthesis and ending in 10- 1 5-flowered umbellules; pedicels 7-9 mm long. Flowers small; buds 3.5 mm long, petals 5, ca. 1 mm or so long, ± sericeous outside, their base hidden by the calyx rim, opening Volume 3, Number 4 1993 Frodin Schefflera in the Venezuelan Guayana 397 from the top and spreading before falling; disk slight¬ ly sloping from a sunken base, to 2 mm across; free portion of calyx rim obscure; ovary usually 3-locular, ± straight, 1-1.5 mm long, then suddenly widening below calyx rim; stylar column shorl . F ruit not known. Related species include Schefflera varisiana and S. neblinae, both of which have compound-umbel- late inflorescences with a single subsidiary whorl of secondary rays and ± sericeous indumentum. Schef¬ flera nigrescens differs from these in the more dis¬ tinctly obovate leaflets, the greater contrast of their surfaces, and finer venation. Schefflera neblinae has more oblong-obovate to oblong-elliptic leaflets with bases rounded at the insertion of the petiole; it is presently known only from Serra Pirapucu, an iso¬ lated range near the southwestern corner of the Neblina massif but mostly cut off from it by the Rio Ariaba. Schefflera nigrescens is known only from the type collection and is found in cloud forest on ridges and slopes north of the southern rim of the Neblina massif, falling toward Canon Grande and the Rio Baria. Nearby Camp XI was in the upper reaches of the most northern tributary of the Canon Menor (maps in Brewer-Carias, 1988: 11, 39). Schefflera pallens Maguire, Steyermark & Kro- din, Mem. New York Bot. Gard. 38: 72. 1984. TYPE: Brazil. Amazonas: trail to Pico Phelps, Serra da Neblina, 2,425 m, 3 Dec. 1965, Ma¬ guire el al. 60503 (holotype, NY; isotype, NY). Nee reported that this tree did not occur much below the site of Camp VII (1,730 1,850 m; Brew- er-Carias, 1988: 149). This suggests that it is prob¬ ably confined to the higher parts of the Neblina massif, perhaps in relatively open places above the main montane forest zone. More collections are needed, however, to ascertain the relative distri¬ bution, frequency, and abundance of this and two other species apparently characteristic of the tepui vegetation zone on Neblina, S. eximia (described above) and S. simplex Steyermark & B. Holst. Additional specimen examined. VENEZUELA. Amazonas: [Cerro de la Neblina] Camp VII, 4 km NE of Pico Phelps, 1,950 m, 4 Feb. 1985, Nee 30749 (MO, NY, US). Schefflera paruana Frodin, nom. nov. Basionym: Schefflera guanayensis subsp. paruensis Ma¬ guire, Steyermark & Frodin, Mem. New York Bot. Card. 38: 67. 1984. TYPFh Venezuela. Amazonas: Cerro Paru, Rio Paru, Cano Asisa, 1,800 m, 12 Feb. 1951, Cowan & II urdack 31405 (holotype, NY; isotype, k). The epithet “ paruensis ” is not taken up here for the species because of the previously existing name Schefflera paraensis Huber, which might cause con¬ fusion. An emended description is furnished below, given the change in status proposed and the exclusion of two of the paratypes (now referred to S. clausa) from the account in Maguire et al. (1984). Slender, probably sparingly branched tree 3 8 m tall; twigs to 1 .5 cm diam. Leaves 4-1 1 -loliolate; petioles to 37 cm long and decreasing from 5 to 2.5 mm diam., emerging from the sheathing base at 2 mm above its insertion with the node, the free portion of the base, or ligule, to more than 2.5 cm long; petiolules 3. 5-8. 5 cm long, fairly slender. Blades oblong-elliptic to oblong-obovate, glabrous, 7-18 cm long, 5. 4-7. 4 cm wide, thinly coriaceous; apex obtuse, the tip acuminate, to 1 cm long, base rounded, margins flat, vertically a little wavy, slight¬ ly tucked in at vein endings and thickened by a marginal vein; upper surface dull, undersurface somewhat glossy; venation of coarse appearance, primary lateral veins 8- 1 0, spreading-ascending, joining with the marginal vein, secondary veins part¬ ly developed, organized cross-veins not evident. In¬ florescence terminal, paniculate, twice compound; main axis short, thickened; primary branches 20 30, 47-68 cm long, ca. 5 mm diam., thinly grayish hairy; peduncles racemosely arranged, 65-85 in upper % to % of the branch, 0.4- 1.1 cm long, subtended by persistent small puberulent bracts, the umbellules 7-9-flowered; pedicels 1.5 3 mm long; flowers after anthesis 1-1.5 mm long, with flattened disk and stylar column about 1 mm long, the ovary 2-locular. Schefflera paruana was originally distinguished from S. guanayensis at the subspecific level on account of its larger, oblong-elliptic leaflets, and greater number of peduncles on each primary branch. However, when viewed in a wider context and with the use of additional characters, each of the non- nominate subspecies described with S. guanayensis merits specific rank. Schefflera paruana is most closely related to S. marahuacensis and S. pedicelligera Maguire, Stey¬ ermark & Frodin, both endemics of the central- southern tepui region, and S. crassilirnba and S. maguireorum in the northwestern tepui region (Hu¬ ber, 1988). It differs from S. pedicelligera mainly in its more densely crowded and numerous umbel- 398 Novon lules with shorter peduncles and pedicels, obtuse to rounded leaflet bases, and petiolar insertion; from S. marahuacensis it differs by its longer, differently shaped, less strongly veined leaflets. The partial union of petiole and base above the insertion of the latter is a character state shared by this species with ,S. marahuacensis. A longer acumen, along with leaflet size and color (when dry), distinguishes S. paruana from S. crassilimba. From S. maguireo- rum, with somewhat similar leaflets, it differs in the umbellules not being headlike and with very short pedicels. Schefflera paruana is characterized by up to 20-30 primary branches in the inflorescence, a state apparently paralleled by S. steyermarkii but more than in the four aforementioned species. Schefflera paruana is a tree of summit streams and the bases of tepui escarpments; it appears to be part of the tepui vegetation zone and in the upper part of the montane zone (Huber & Alarcon, 1988). Along with S. asymmetrica and S. concolor in the south and ,S. ayangannensis Maguire, Steyermark & Frodin in Guyana, it appears to he one of the more generalized species in its series. Additional specimen examined. VENEZUELA. Amazonas: Cerro Paru, from camp cano to Point 2 (just N of Point 1), 2,000 m, 31 Jan. 1951, Cowan & Wur- dack 31072 (K, NY). Schefflera psilophylla (Harms) Maguire, Stey¬ ermark & f rodin, Mem. New York Bot. Gard. 38: 68. 1984. TYPE: Venezuela. Bolivar: Mt. Roraima, 1,900 m, Ule 8702 (lectotype, here designated, G; isolectotypes, K, L). The exclusion of most records included here by Maguire et al. (1984) — either to S. rnyrioneura or •$. suaveolens- — indicates that S. psilophylla is lim¬ ited to mid-elevations on Roraima and Ilu-tepui. Three collections are known: Ule 8702 (G, K, L); Maguire et al. 33364 (K, NY), and 33484 (K, NY). Schefflera quinquecarinata Steyermark, Bol. Soc. Venez. Ci. Nat. 25(106): 44, fig. 2. 1963. TYPE: Venezuela. Bolivar: vicinity of Km 134, bordering headwaters of Rio Cuyuni, NE of Luepa, Steyermark & Nilsson 483 (holotype, VEN). Tillett et al. 45736 should be excluded from the list of records in Maguire et al. (1984: 5). It is a form of S. morototoni, but its status has yet to be resolved. In addition, Prance et al. 10159 from Serra dos Surucucus in Roraima Territory, Brazil, distributed as this species hut not listed in Maguire el al. (1984), appears to be S. yutajensis Steyer¬ mark & B. Holst, a vicariant species. Schefflera quinquecarinata thus remains limited to a relatively small area on the Venezuela-Guyana border. Schefflera quinquestylorum Steyermark, Bol. Soc. Venez. Ci. Nat. 25: 46. fig. 3. 1963. Schefflera duidae subsp. quinquestylorum (Steyermark) Maguire, Steyermark & Frodin, Mem. New York Bot. Gard. 38: 54. 1984. TYPE: Venezuela. Bolivar: Cerro Venamo, NW slopes, SE of road camp 125, to line of NE- facing sandstone bluffs, 1,100-1,140 m, 21 Apr. 1960, Steyermark & Nilsson 429 (ho¬ lotype, VEN not seen; isotypes, K, NY). This taxon is not simply a vicariant of S. duidae but part of a small series of apparently sibling taxa, mainly in eastern Colombia and with links to S. sprucei (Seemann) Harms. Moreover, the leaflets are more acute, with a caudate-acuminate tip, and when dry exhibit greater surface contrast; the pu¬ bescence on the undersurface of the leaflets, when present, is also differently colored. There is also no evidence, as in 5. duidae, that this is a facultative liana. I thus propose renewing its recognition as a species. Geographically, the area of S. quinquestylorum is centered on Cerro Venamo and is limited by comparison with 5. duidae, which is now known to range from Cerro Marutani to the central-southern tepuis (Huber, 1988) and south to Serra Araca and Serra Pirapucu (the latter near the Neblina massif) in northern Brazil. Schefflera sessili flora Splithof-Heerschop ex Fro¬ din, nom. nov. Replaced name: Schefflera re¬ ticulata (Gleason) Maguire, Steyermark & Fro¬ din, Mem. New York Bot. Gard. 38: 57. 1984, not Philipson, 1951. Basionym: Didymopanax reticulatus Gleason, Bull. Torrey Bot. Club 58: 434. 1931. TYPE: Venezuela. Amazonas: Cer¬ ro Duida, summit of Mt. Duida, Peak No. 7, 7,100 ft., Aug. 1928-Apr. 1929, Tate 606 (holotype, NY; isotype, K). Because the epithet reticulata was validly pub¬ lished in 1951 for a New Guinean species, a new name is needed. The new epithet relates to the position of the flowers and was introduced by Mrs. Splithof-Heerschop in her unpublished study of Ar- aliaceae in the Guianas, carried out at Utrecht. Additional material of S. sessilijlora has been collected in the last decade, and its variability can now be more effectively assessed. Two collections should, however, be excluded from the list of exsic- Volume 3, Number 4 1993 Frodin Schefflera in the Venezuelan Guayana 399 catae presented under 5. reticulata in Maguire et al. (1984): Steyermark 98138 and Steyermark et al. 109690. Both of these are S. steyermarkii. To be added is Steyermark et al. 124075 from Cerro Marutani, previously cited under S. reticulata subsp. jauaensis. The resulting distribution pattern appears irreg¬ ular, but may reflect reliction. Two populations are disjunct from the main center in the central-southern tepui region (Huber, 1988), which ranges from Cer¬ ro Paru to the Duida group. One of these is on or near Cerro Marutani in the Sierra Pakaraima, in the far south of the central-eastern region; and the other is at Auyan-tepui in the eastern region. The leaflets in the first are comparatively large, while the latter differs in fruit. The variability in the central area is phenotypic, most evidently in vegetative fea¬ tures. Schefflera steyermarkii Frodin, sp. nov. TYPE: Venezuela. Bolivar: Cerro Jaua, cumbre, at base of second waterfall, 1,922-2,100 m, 22-27 Mar. 1967, Steyermark 098138 (holotype, NY; isotypes, K, VEN not seen). Arbores ad 10 rn altae; haec species cum Schefflera sessiliflora hactenus confusa sed ab ea foliolis angustior- ibus apicibus acutis venis tenuibus et capitulis minoribus (ad 5 mm diametro) differt. Foliola pagina inferna argen- tea. Tree 8-10 m tall, mostly glabrous; branchlets stout, 1.3 cm diam. Leaves around 1 0-foliolate; petiole ca. 26 cm long, arising from base 0.5-1 cm above its insertion, the free ligular portion ca. 1.5 cm long, with papery margins; petiolules around 2.4 cm long. Blades narrowly elliptic, glabrous, 7-13 cm long, 2-4.5 cm wide, thinly coriaceous; apex narrowly acute, passing ± insensibly into a caudate- acuminate tip to 1.2 cm long, lower part narrowing, base abruptly rounded but not recurving into peti- olule, margins incorporating a marginal vein, ver¬ tically slightly wavy and sometimes slightly tucked in at vein endings; upper surface dull, undersurface minutely glandular-papillate, furnishing when dry a dull metallic reflection; midrib somewhat prominent below; lateral veins closely spaced, the primary veins 12-14 or more, spreading and extending to margin, the secondary veins closely resembling them hut only reaching some %-way to margin, visible retic¬ ulation coarse. Inflorescence terminal, twice com¬ pound, paniculate; main axis 6. 5-7. 5 cm long, 0.8- 1 cm diam., the zone of proliferation in the upper 3-4 cm; primary branches tightly clustered, 10 or more, rather straight and rigid, 35-43 cm long, 3.5 mm diam. near base, dull purple, subtended by bracts 3.5 4 cm long; capitula numerous, ca. 5 mm diam. after anthesis, 10- 12 -flowered, racemosely ar¬ ranged, their peduncles 10-14 mm long, appressed silvery hairy, subtended by bracts 1.5-3 mm long (but where empty in lower part of branches to 6- 7 mm long); buds dull purple with green; calyx silvery hairy below, the rim with 5 small teeth; petals and stamens 5; ovary 2(3)-locular, tawny yellow, disk ca. 1.3 mm across, stylar column ca. 1 mm long, tapering. Fruit not known. The narrow leaflets of Steyermark 98138 were noted by Maguire et al. (1984: 57) in introducing the records listed for S. reticulata (= S. sessiliflora ). An earlier judgment by the author that a distinct species might be involved has, on the other hand, been given support by a second collection, Steyer¬ mark 109690. The two collections are thus here described as newT. Other features that support the distinction of S. steyermarkii from S. sessiliflora include the finer leaflet venation and heads only about 5 mm diam. after anthesis. The relatively large number of primary branches in the inflores¬ cence also appears to be distinctive; only S. paruana has more, and that species differs in leaflet form and morphology and in having flowers in umbellules. Schefflera steyermarkii is limited to the Meseta de Jaua, which, with neighboring Cerro Sarisarinama and Cerro Guanacoco, is home to a number of other endemics in its series (,S. clausa, S. contractu, and ,S. jauaensis) as well as, from among compound- umbellate species, the distinctive ,S. gracillima Stey¬ ermark & Maguire. All have been found in the summit zone. When compared with other tepui regions, however, endemism in the group seems 1owt; other species are S. coriacea, here at its western limit, and S. montana (Gleason) Maguire, Steyer¬ mark & Frodin, otherwise occurring further west. Schefflera sessiliflora, too, is apparently absent, unless it has been overlooked. Paratype. VENEZUELA. Bolivar: Cerro Jaua, cumbre, E tributary Rio Marajano, 1,810-1,880 in, 28 Feb. -3 Mar. 1974, Steyermark et al. 109690 (NY, US, VEN not seen). Schefflera suaveolens Frodin, sp. nov. TYPE: Venezuela. Amazonas: Serrania Yutaje, Cerro Coro-Coro, Rio Manapiare, 4,500 ft., 2 Mar. 1953, Maguire & Maguire 35483 (holotype, NY). Figure 17. Arbor sylvarum submontanarum usque ad 1 8 m alta, Schefflerae psilophyllae cognata autem ab ea ovario pler- umque 4-5-loculari, foliolis tenuiter coriaceis (vel etiam chartaceis), et infiorescentiae radiis priinariis fere semper verticillos subsidiaros radiorum secundariorum infra api- cem 4-20 cm ferentibus recedit. 400 Novon Tree 7-18 m tall, when young with as few as 3 orthotropic branches, in maturity with a rounded crown; adult twigs to 1.4 cm diam. below growing point. Adult leaves (3-)5-or 6(-8)-foliolate; petiole 1 2-28(-38) cm long (in ultimate leaves of a shoot relatively short), generally shorter than inflorescence branches, to 5 mm diam.; petiolules 1~6 cm long, widely varying within a given leaf. Blades oblong- elliptic to oblong-obovate, the widest point at or above middle, glabrous, 8-22 cm long, 3. 4-8. 4 cm wide, chartaceous to thinly coriaceous (when young at least sometimes membraneous), drying greenish with only slightly contrasting surfaces; apex acute to obtuse or rounded, tip absent or retuse, base acute to narrowly obtuse, with a mid-range of ca. 90° and shortly decurrent into the petiolule; margins ± wavy, not or only slightly inrolled; venation thin but distinct, spreading, ± ascending; primary and secondary lateral veins virtually indistinguishable, up to 40 on a side; subsidiary lateral venation var¬ iously developed. Blades in leaves from young plants smaller, narrowly elliptic or ovate, 7.5-10 cm long, 2. 4-2. 8 cm wide, the apex acute, the tip acuminate. Inflorescences compound, umbellate, all parts gla¬ brous, initially terminal, later pseudolateral as new shoots develop from beneath and grow alongside; primary rays 3 or 4(5), 23-67 cm long, 3-7 mm diam., always or nearly always with a subsidiary whorl (occasionally 2 whorls) of secondary rays, the internode between apex and the whorl below it 6.5- 13 cm; secondary rays 3-6(-7) cm long, the sub¬ tending bracts 4 mm long but largely becoming lost during fruit maturation, 10-20( 30) in terminal umbel (or just below it), ca. 6 in subsidiary whorl(s), often gently curving; ultimate coflorescences um¬ bellate, comprising a central cluster of 10-12 bi¬ sexual, fertile flowers and about 2 or 3 irregularly spaced lateral umbellules arising at 2- 1 2 mm below; pedicels in central cluster 8-1 1 mm long (length¬ ening slightly in fruit), each subtended by a persistent acicular bracteole; lateral peduncles 9-16 mm long with up to 10 (probably functionally male) flowers on pedicels 5-7 mm long; flowers glabrous, green in bud; calyx rim 0.75 mm wide, projecting outward above the ovary, the 5 teeth quite small; corolla pale, the petals 5, opening and spreading before falling; filaments 2 mm long; ovary (3-or 4-) 5-locular, oblong, 1 mm long, 0.8-0. 9 mm across; stylar col¬ umn 1 mm long, undivided. F ruit remaining wholly inferior, initially green, maturing red-purple and ul¬ timately purple-black, ovoid to globose, 4-5.5 mm long, 4-5 mm across (but up to 8 mm across when fresh); calyx rim slightly projecting, the disk ca. 1.75 mm across; stylar column ca. 2 mm long, divided at the tip, its base filling most of the disk; pyrenes usually 5, sometimes fewer. The first collection of this species was initially and tentatively identified with Didymopanax spru- ceanus by Gleason (1931); later it and other col¬ lections were placed hy Maguire et al. (1984) in Scheffiera psilophylla. The abundant material now available shows S. suaveolens not to represent either of these species. It can be distinguished from S. spruceana by the (3-)4-or 5-celled ovary and from S. psilophylla sensu stricto by the presence in the inflorescences of fewer primary rays, each with 1(2) subsidiary whorls of secondary rays in addition to those at the apex. The venation in S. psilophylla is also somewhat coarser and the leaflet apices are usually ± acute; moreover, the ovary is only 3-locular. From S. myrioneura, described above, the present species differs primarily in its less closely spaced lateral venation and more graceful, less an¬ gular-appearing inflorescences; from S. urnbracu- lifera it differs mainly in having thinner, wholly glabrous leaflets with ± flat margins and finer, less strongly contrasting lateral venation. Scheffiera suaveolens has a relatively wide dis¬ tribution from Serrania Yutaje and Serannia Uasadi south to the Neblina massif and its outliers and east to Auaris in the Serra Parima, and also a wide altitudinal range; it is described as occurring in mon¬ tane forest, mixed mossy-forest, scrub forest and Bonnetia forest (sometimes along watercourses), and is occasional to frequent. Ecologically, it probably favors nutrient-rich soils with a steady water supply, and it may be opportunistic. When cut, it may yield an aromatic, transparent and sticky resinous exu¬ date. Paratypes. VENEZUELA. Amazonas: Serrania LJasadi, sector nor-occidental, cumbres montanosas ubi- cadas en las cabeceras orientales del Rio Asita, afluente derecho del Rio Ventuari, ± 1,850 m, 22 Nov. 1988, Huber 12837 (K, MYF not seen); 8 km NW of Yutaje settlement, W of Serrania de Yutaje, 1,500-1,760 m, 4 Mar. 1987, Liesner & Holst 21637 (MO); Agiiita, slopes of Cerro Duida, 3,100 ft., Dec. 1928, Tate 876 (NY); Cerro Marahuaca, “Sima Camp,” S-C portion of forested slopes, 1,140 m, 22 Feb. 1985, Steyermark & Holst 130506 (MO, NY); Cerro Marahuaca, summit on undulating plateau, 2,520-2,650 m, 26 Feb. 1985, Stey¬ ermark & Holst 130750 (MO, NY); [Cerro de la Nebli¬ na:] Rio Yatua, below NW escarpment between Camp 4 and Cumbre Camp, 1,700-2,000 m, 13 Jan. 1954, Ma¬ guire et al. 37274 (K, NY); Canon Grande, NW head, 2,000 m, 8 9 Dec. 1957, Maguire et al. 42326 (K, MO, NY, US); Canon Grande, NW head, 2,000 m, 8- 9 Dec. 1957, Maguire et al. 42327 (MO, NY); E Es¬ carpment, upper Canon Grande, 1,900 m, 14 Dec. 1957, Maguire et al. 42406 (K, MO, NY, US); Cano[n] Grande Volume 3, Number 4 1993 Frodin Schefflera in the Venezuelan Guayana 401 SSE of Cumbre Camp, 1,100-1,150 m, 26 Dec. 1957, Maguire et al. 42520 (K, MO, NY, US); Pico Phelps, S face, S of Camp V, 1,550-1,650 m, 12 Apr. 1984, Gentry & Stein 46579 (MICH, MO); Camp VII, 1,850 m, 3 Dec. 1984, W. Anderson 13475 (MICH); Camp VII, 5 km NE of Pico Phelps, 1,800 m, 2 Feb. 1985, Nee 30706 (MO, NY); Camp XI, 6 km NNE of Pico Phelps, 1,550-1,650 m, 27 Feb. 1985, Nee 31161 (MO, NY); 26 km ENE of Base Camp [on S rim], ridge on border divide, 2,000 m, 15 Apr. 1984, Plowman & Thomas 13612 (F, K, NY). BRAZIL. Roraima: upper slopes of Serra Parima, S of Auaris [Uavaris Valley], 4°03'N, 64°22'W, 1,400-1,520 m, 10 Feb. 1969, Prance et al. 9801 (CGE, INPA not seen, NY, S, US). Ama¬ zonas: between Missao Salesiana and Serra Pirapucu, Rio Maturaca [SW of Neblina massif], 800-1,000 m, 23 Jan. 1966, Silva & Brazdo 60866 (K, NY). Schefflera ulocephala Frodin, nom. nov. Re¬ placed name: Schefflera globulifera Maguire, Steyermark & Frodin, Mem. New York Bot. Card. 38: 69. 1984, not Grushvitzky & N. T. Skvortsova, 1969. TYPE: Venezuela. Ama¬ zonas: Cerro de la Neblina [N rim], in Cumbre Camp, 1,700-2,000 m, 23 Jan. 1954, Ma¬ guire, Wurdack & Hunting 37354 (holotype, NY; isotype, K). Because the epithet globulifera was validly pub¬ lished in 1969 for a Southeast Asian species, a new name is needed. The new epithet is based on the Greek oulos, curly or woolly, and kephalos , head. Schefflera ulocephala is part of a series of six species, which on Neblina includes S. pallens; the other members are on Sierra de Maigualida and Serrania Uasadi, where there has been some local diversification, as yet little studied or documented. The species include S. brachypoda, S. huberi, and S. longistyla on Maigualida and S. baculosa on Uasadi. Of these, the last named most resembles S. ulocephala. Schefflera umbellata (N. E. Brown) Viguier, Ann. Sci. Nat., Bot., ser. 9, 9: 367. 1909. Scio- daphyllum umbellatum N. E. Brown, Trans. Linn. Soc. Bot. ser. 2, 6: 32. 1901. TYPE: Venezuela [as “Guyana”]. Mt. Roraima, sum¬ mit, Oct. 1898, McConnell & Quelch 666 (lectotype, K; isotype, BM). Schefflera umbellata subsp. duidana of Maguire et al. (1984) has been accorded specific rank in this paper and renamed Schefflera acaropunctata. Re¬ cords from the western Gran Sabana, including Au- yan-tepui and the Chimanta massif, are also de¬ scribed as a distinct, vicariant species, S. clavigera. Cardona 1765 is also transferred to S. japurensis. As a result, Schefflera umbellata is now limited to the cluster of tepuis in the southeastern Gran Sabana centering on Roraima-tepui. Some variability exists among presently available material, hut no attempt is made here to analyze it in more detail. Additional specimens examined (not included in Ma¬ guire et al., 1984). VENEZUELA. Bolivar: Matahui- tepui [Kukenan-tepui], 2,700-3,000 m, 22 Aug. 1982, Castillo 2716 (K); Kukenan-tepui, cumbre meridional, 2,700 m, 15 June 1985, Huber & Alarcon 10552 (CM, MYF not seen, NY); Kukenan-tepui, summit, 2,550 m, 11 Apr. 1988, Liesner 23139 (MO), summit, 2,550 m, 12 Apr. 1988, Liesner 23164 (MO); Yuruani-tepui, 12 km NNE of Kukenan-tepui, 2,200 m, 29 Feb. 1984, Huber 9092 (MO, MYF not seen, NY, US); Mt. Roraima, SW slopes, 7,400 ft., 11 Jan. 1939, Pinkus 153 (GH, NA, US); Mt. Roraima, on slopes among rift, headwaters of Great Central Rift, 2,700-2,740 m, 28 Sep. 1944, Steyermark 58884 (F); Mt. Roraima, cumbre, NE part, 2,750-2,800 m, 26 Aug. -2 Sep. 1976, Steyermark et al. 112445 (MO, NY, U); Roraima Peak, “am Abhang,” Dec. 1909, Ule 8704 (G, K, L). Schefflera umbraculifera Frodin, sp. nov. TYPE: Venezuela. Amazonas: Sierra de Maigualida, sector NW, cabeceras del Rio Iguana, afluente del Rio Asita, on a SE-facing slope, 1,720 m, 25 Mar. 1988, Huber 12682 (holotype, K; isotype, MYF not seen). Figure 18. Arbor sylvarum humilium exigua cujus foliola Schef- flerae suaveolentis paginam supernam habent sed ab ea foliolis crassioribus ad marginem incurvatis ac ad paginam infernam molliter pubescentibus valde recedit. Foliola ob- longo-obovata et late emarginata. Tree 4-7 m, with broad, dense crown; twigs pithy, 1.4 cm across just below uppermost leaf and inflo¬ rescence. Leaves 5-foliolate; petiole ca. 21.5 cm long, the base as wide as the twig and forming flanges but not extending into a ligule; petiolules 2. 3-3. 8 cm long. Blades ohlong-obovate, glabrous above, pale green and minutely brownish to grayish seri- ceous-tomentose below (except on the paler primary and, to a lesser extent, secondary veins), 1 1.3-14.2 cm long, 6—6.6 cm wide, coriaceous, little varying in size within a given leaf; apex emarginate, base broadly acute, slightly recurved into the petiolule, margin entire, noticeably incurved when dry; upper surface glossy, the venation slightly recessed; un¬ dersurface with prominent hut rapidly fading midrib and slightly raised venation; primary lateral veins 9-12 on each side, spreading, sometimes divari¬ cating in outer part of blade, curving sharply near margin and passing up just inside it to anastomose with the vein above; secondary veins ± distinctly developed; cross-veins usually 1 or 2 on each side, passing from one primary or secondary vein to an¬ other at an angle of ca. 90° with the midrib. Inflo- 402 Novon rescence initially terminal, later pseudolateral, um¬ bellate, three times compound, without persistent bracts; main axis not developed; primary rays ca. 48.5 cm long, minutely and thinly sericeous-puber- ulent, their number unknown but presumably not more than 5 or 6; secondary rays 6.2-8 cm long, radiating hemispherically at ends of primary rays and outwards in a subsidiary whorl of 4 at ca. 12 cm below, with an occasional extra in between; ul¬ timate coflorescences tertiary, with central bisexual, fertile flowers on pedicels 8-9 mm long at ends of secondary rays and 6-8 lateral, tertiary, 6-8-flow- ered umbellules arising from 2 to 23 mm below (the interval much more pronounced in the subsidiary whorl, relatively small among the terminal secondary rays), the peduncles and pedicels of the latter re¬ spectively 1.4-2 cm and 4-5 mm long, minutely sericeous-puberulent, the pedicels each subtended by a minute bracteole; ovary obconic, 1 mm long, the disk flat, the stylar column 1 mm long. Maturing fruit at ends of secondary rays becoming black, globose, 5-5.5 mm long, 5 mm across, angular when dry, the calyx rim minute, the disk 2-2.5 mm across, narrow, soon rising into a slender stylar column 2- 2.5 mm long, the uppermost part 3-5-fid; fruit on lateral peduncles green, immature, 2.5-3 mm long. Schefflera umbraculifera is superficially closest to 5. suaveolens but differs most markedly in its leaflets, which are glabrous above but entirely pu¬ bescent beneath except for the main venation. Also, the secondary venation is not as strongly developed, the leaflets are thicker, and the inflorescence inter¬ nodes are longer. Schefflera neblinae, with which the type collection was initially identified, and S. v arisiana both have smaller, narrower leaflets with more or less whitish undersurfaces and smaller in¬ florescences with trilocular ovaries. Schefflera umbraculifera is known only from the type collection and has been noted as frequent in shrubby vegetation of upper headwaters, at least some of it secondary. Schefflera varisiana Frodin, sp. nov. TYPE: Bra¬ zil. Roraima: upper slopes of Serra Parima, S of Auaris [Uavaris Valley], 4°03'N, 64°22'W, 1,400-1,520 m, 10 Feb. 1969, Prance , Stew¬ ard , Ramos & Farias 09796 (holotype, K; isotypes, CGE, F, NY, US; A, INPA, MG, R, S, U, & VEN, not seen). Arbor parva Scheffleram neblinae simulans, sed haec species nova foliis, 6-7-foliolatis, foliolis anguste obovatis vel obovato-ellipticis ad basin plus minusve attenuatis, paginae infernae venis prominulis a mesophyllo discre- pantibus notabilis. Tree to 8 m high, 12 cm diam. Leaves 6- or 7-foliolate; petioles 15.5-16.5 cm long, the sheath¬ ing base small; petiolules 1.8-2. 5 cm. Blades nar¬ rowly obovate to obovate-elliptic, usually broadest above middle, the surfaces contrasting, 9-12.3 cm long, 3.2-5 cm wide, coriaceous, brittle when dry; apex obtuse to rounded, tip not present, base at¬ tenuate to acute, slightly decurrent at insertion of petiolule, margin narrowing from %-V3 of the way down from apex, incurved but not noticeably inrolled when dry; upper surface glabrous, the venation ± impressed, undersurface whitish or grayish seri¬ ceous, the indumentum tightly appressed, the ve¬ nation somewhat raised in dry state and contrasting with the mesophyll; midrib prominent below; pri¬ mary lateral veins ca. 12-16 on a side, ± closely parallel, near margin suddenly anastomosing and forming a submarginal vein; secondary venation ± well developed; reticulation fine, irregular, visible mainly on upper surface, with a few cross-veins. Inflorescence terminal, umbellate, twice compound; primary rays ca. 16.5 cm long, tbcir number not known; secondary rays both at apex and in a single subsidiary whorl, ca. 20 at the former, far fewer in the latter; all axes sericeous-puberulent, the hairs much appressed, later glahrescent; umbellules 10- 15-flowered, the pedicels slender, 5-6 mm long, sericeous-puberulent; flowers brown, in bud 3 mm long; petals 5, in bud sericeous without, at anthesis spreading outwards before falling; ovary 3-locular, the stylar column united, 0.5 mm long. Fruit not known. Schefflera varisiana is closely related to S. neb¬ linae and appears to be a sister species. From its sibling it differs in its narrowly obovate to obovate- elliptic leaflets, attenuate to acute leaflet base, better developed secondary venation, and particularly the leaflet surfaces: here the upper surface is more ev¬ idently rugose and the venation below is more prom¬ inent and shows greater contrast. In 5. neblinae the leaflets are oblong-obovate to oblong-elliptic, the base of the blade is more or less rounded at its insertion with the petiolule, and the blades have less well developed secondary venation and relatively fea¬ tureless surfaces. The pattern of differences of S. varisiana with S. coriacea is discussed under S. neblinae. Schefflera varisiana is known only from the type collection and occurs in higher parts of the northern Serra Parima, in scrub forest. The locality is further described in Huber et al. (1984). It is about 375 km from Pirapucu, the type locality of S. neblinae, where on its upper slopes at 1,300 m similar forest might be expected. Volume 3, Number 4 1993 Frodin Schefflera in the Venezuelan Guayana 403 The epithet is derived from the name of the valley in which Auaris, the type locality, is situated. The initial “ua” has been omitted in the interests of euphony. Acknowledgments. I thank the late Bassett Ma¬ guire and, in more recent years, Patricia Holmgren for making available the collections of The New York Botanical Garden, essential to this study; and Benjamin Stone for the use of facilities at the Acad¬ emy of Natural Sciences of Philadelphia and for obtaining specimens on loan between 1986 and 1 990. Thanks are also due to Otto Huber for timely ship¬ ments of new collections; to the keepers of the her¬ baria at the Royal Botanic Gardens, Kew, and the Natural History Museum, London, for permission to examine the key Roraima collections by Irn Thurn and McConnell and Quelch; to the director of the Rijksherbarium, Leiden, for making available the Ule material; to John Pruski of the Smithsonian Institution for assistance during the period of prep¬ aration of this paper; and to Paul Berry and Bruce Holst of the Missouri Botanical Garden for extensive help in the preparation and final editing of the manu¬ script. I also thank Jan Lindeman for kindly making available a copy of Mrs. Splithofs paper; I use her new epithet, Schefflera sessiliflora, in acknowledg¬ ment of her work. Literature Cited Baumann-Bodenheim, M. G. 1955. Ableitung und bau bicarpellat-monospermer und pseudomonocarpellater Araliaceen und Umbelliferen-Friichte. Ber. Schweiz. Bot. Ges. 65: 481-510, illus. Brewer-Carias, C. (editor). 1988. Cerro de la Neblina: Resultados de la expedicion 1983-1987. Fundacion para el Desarrollo de las Ciencias Fisicas, Mateina- ticas y Naturales, Caracas. Frodin, D. G. 1975. Studies in Schefflera (Araliaceae): The Cephaloschefflera complex. J. Arnold Arbor. 56: 427-448, illus. - . 1989. Studies in Schefflera (Araliaceae), IV. Synopsis of the Formenkreis comprised of Didy- mopanax attenuatus (Sw.) E. Marchal and allied species, with nomenclatural changes. Proc. Acad. Nat. Sci. Philadelphia 141: 313-319. Gleason, H. 1931. Botanical results of the Tyler-Duida Expedition. Bull. Torrey Bot. Club 58: 277-506. Greuter, W., H. M. Burdet, W. G. Chaloner, V. Deinoulin, R. Grolle, I). L.. Hawksworth, D. H. Nicholson, P. C. Silva, F. A. Stafleu, E. G. Voss & J. McNeill. 1988. International Code of Botanical Nomencla¬ ture. Regnum Veg. 118. Huber, 0. 1988. Guayana highlands vs. Guayana low¬ lands, a reappraisal. Taxon 37: 595-614. - & C. Alarcon. 1988. Mapa de vegetacion de Venezuela. 1:2,000,000. Caracas, Ministerio del Ambiente y de los Recursos Naturales Renovables, Venezuela (distributed by BIOMA). - & D. Frame. 1989. Venezuela. Pp. 362-374 in D. G. Campbell & H. D. Hammond (editors), Floristic Inventory of Tropical Countries. The New York Bot. Card., New York. - & J. J. Wurdack. 1984. History of botanical exploration in Territorio Federal Amazonas, Vene¬ zuela. Smithsonian Contr. Bot. 56: 1-83, 1 map. - , J. A. Steyermark, G. T. Prance & C. Ales. 1984. The vegetation of the Sierra Parima, Ven¬ ezuela- Brazil: Some results of recent exploration. Brittonia 36: 104-139. Maguire, B. 1979. Guayana, region of the Roraima sandstone formation. Pp. 223-238 in K. Larsen & L. B. Holm-Nielsen (editors), Tropical Botany. Ac¬ ademic Press, London. - , J. A. Steyermark & D. G. Frodin. 1984. Ar¬ aliaceae. In: B. Maguire & Collaborators, The Botany of the Guayana Highland — Part XII. Mem. New York Bot. Gard. 38: 46-82. Nelson, B. W . , C. A. C. Ferreira, M. F. da Silva & M. L. Kawasaki. 1990. Endemism centres, refugia and botanical collection density in Brazilian Amazo¬ nia. Nature 345: 714-716. Paynter, R. A., Jr. 1982. Ornithological gazetteer of Venezuela. Bird Department, Museum of Compara¬ tive Zoology, Harvard University, Cambridge, Mas¬ sachusetts. Prance, G. T. & D. M. Johnson. 1992. Plant collections from the plateau of Serra do Araca (Amazonas, Bra¬ zil), and their phytogeographic affinities. Kew Bull. 47: 1-24. Seemann, B. 1865. Revision of the natural order Hed- eraceae [continuation]. J. Bot. 3: 265-276. - . 1868. Revision of the natural order Heder- aceae. J. Bot. 6: 129 142. Steyermark, J. A. 1957. Araliaceae. In: Contributions to the Flora of Venezuela, 4. Fieldiana, Bot. 28: 1043-1045. - . 1967. Araliaceae. In: Flora del Auyan-tepui. Acta Bot. Venez. 2(5-8): 271-277. - . 1979. Flora of the Guayana Highland: En- demicity of the generic flora of the summits of the Venezuelan tepuis. Taxon 28: 45-54. - — & B. Holst. 1988. Araliaceae. In: J. A. Stey¬ ermark, Flora of the Venezuelan Guayana, V. Ann. Missouri Bot. Gard. 75: 1082-1083. A New Species of Galaxia (Iridaceae subfamily Iridoideae) from Namaqualand, South Africa Peter Goldblatt B. A. Krukoff Curator of African Botany, Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A. E. G. H. Oliver Stellenbosch Herbarium, National Botanical Institute, P.0. Box 471, Stellenbosch 7600, South Africa ABSTRACT. Galaxia fenestralis is new a species from central interior Namaqualand, Cape Province, South Africa. Apparently most closely related to another Namaqualand endemic, G. kamiesmontana Goldblatt, G. fenestralis is distinguished by its terete, succulent leaves, and small light pink to mauve or whitish flowers with a yellow center and narrowly clawed tepals. It grows on exposed granite outcrops in virtually no soil, and flowers early in the rainy season, in June and July, a few weeks after the first of the season’s rains have fallen. Comprising some 1 5 species, including the one described here, Galaxia Thunberg is restricted to the winter rainfall region of the southern and western Cape Province, South Africa (Goldblatt 1979a; 1984). A member of subfamily Iridoideae tribe Ir- ideae (Goldblatt, 1991), Galaxia is distinguished from the related genera Moraea Miller and Homeria Ventenat by the absence of an aerial stem and by the cup-shaped flowers borne above the basal cluster of leaves on a short closed perianth tube. Two sub¬ genera are currently recognized in Galaxia (Gold¬ blatt, 1979a). Subgenus Galaxia comprises species with yellow or white flowers and apomorphic style lobes that are deeply and regularly fringed. In sub¬ genus Eurystigma the flowers may be yellow but are more often pink to purple or reddish (also some¬ times white), and the style lobes have entire margins, the plesiomorphic state. The new species, G. fenes¬ tralis (Figure 1 ), clearly accords with subgenus Eu¬ rystigma both in flower color and stigma features. Its flowers resemble most closely those of the only other Namaqualand member of the subgenus, G. kamiesmontana Goldblatt, in their pale pink to mauve color with a yellow center, small size, and in the rather narrowly clawed tepals. In other species of the suhgenus the distinction between tepal limb and claw is obscure if present at all, and the flowers are considerably larger. Cytologically, subgenus Eurystigma is notable for a descending dysploid series. The presumed ances¬ tral basic chromosome number for Galaxia is x = 9 (Goldblatt, 1979b, 1984), also the base number for all species of subgenus Galaxia and for G. ka¬ miesmontana. Other species of subgenus Eurystig¬ ma have n = 8, 7, or 6. Unfortunately, chromosome number has not been determined for the new species, the only member of the genus for which this infor¬ mation is unknown. Galaxia fenestralis is distinguished by apomor¬ phic nearly terete and succulent leaves with an ad- axial band of transparent tissue (Fig. 2A). It is also unique in the genus in having the inner whorl of tepals connate for 2 mm with the filament column (Fig. IB). Nectaries are present at the base of the outer tepals and can be recognized by their crescent shape and maroon edges (Figure 1 B). Nectaries have not been described in other species of Galaxia (Gold¬ blatt, 1979a), but perigonal nectaries are typical of tribe Irideae (Goldblatt, 1990) where they com¬ monly occur at the base of the outer tepals, as in Moraea and Iris (Goldblatt, 1991), or also at the base of the inner tepals. The short-lived flowers, which last from mid-morning when they open until about 3:30 PM, are typical of the genus. Such fugacious flowers are plesiomorphic for subtribe Homeriinae, to which Moraea and Homeria also belong (Goldblatt, 1990, 1991). The unusual leaves of Galaxia fenestralis (Figure 2A) are interesting anatomically. Although nearly terete, they are not centric. The peculiar transparent band on the upper leaf surface lacks chlorenchyma and is continuous with the cortical mesophyll, which is compact and lacks intercellular air spaces, which renders the leaf succulent. The “window” is bor¬ dered by single strands of subepidermal sclerenchy- ma fibers three cell layers thick. Elsewhere the epi¬ dermis overlies a palisade mostly of 3 cell layers. The outer walls of the epidermal cells are micro- Novon 3: 404-407. 1993. Volume 3, Number 4 1993 Goldblatt & Oliver Galaxia fenestralis 405 Figure 1. Morphology and distribution of Galaxia fenestralis. — A. Whole plant (scale bar = 2.5 mm). — B. Detail of flower with outer tepal bent forward to show basal nectary surrounded by dark pigment and entire filament column (note inner tepals united for a short distance with the filament column) (scale bar = 5 mm). (Drawn by I. Oliver.) scopically striate, with two striae per cell. The outer walls and cuticle are otherwise unremarkable. Sto¬ mata are abundant above the chlorenchyma (absent from the “window”), are not sunken, and commu¬ nicate with the cortex via substomatal chambers. The vascular bundles are arranged in a ring along the inner edge of the palisade with the xylem in¬ ternal. A complete parenchymatous outer bundle sheath is present. The cells at the poles, especially at the phloem pole, each contain a single large rhomboidal crystal. The inner sheath is partially sclerified. A sclerenchymatous phloem cap is present on all but the most minor bundles, and an incomplete xylem cap is found on only the major bundles. The leaf of Galaxia kamiesmontana, putatively the clos¬ est relative of G. fenestralis is similar in its general anatomical features (Figure 2B), but it is much more slender, lacks the adaxial window, and is not suc¬ culent. The leaves of Galaxia kamiesmontana and G. fenestralis are typical of malacophylous xeromorphs in their more or less terete shape with vascular bundles arranged in a cylinder, their compact me- sophyll, and relatively greater amount of photosyn¬ thetic tissue in relation to the amount of exposed leaf surface (Eames & McDaniels, 1925). The abun- 406 Novon Figure 2. Leaf anatomy of Galaxia fenestralis (A) and closely related G. kamiesmonlana (B) (scale bar 0.1 mm). (Drawn by J. C. Manning.) dant surface stomata, unremarkable epidermis, and thin cuticle argue against significant drought toler¬ ance. In G. fenestralis the strategy of maximizing photosynthetic tissue while minimizing surface area is apparently developed further through enhancing light utilization by the photosynthetic window which permits light to penetrate into the center of the leaf. This is presumably a strategy to enable rapid growth in the short period when water is available each growing season, followed by drought avoidance through geophytism. Galaxia fenestralis Goldblatt & E. G. H. Oliver, sp. nov. TYPE: South Africa. Cape Province: 3018 (Kamiesberg) Kliprand District, valley northwest of Matjesfontein towards Middelpos, sandy seep on low granite outcrop, 960 m, 27 June 1991 (DA), Oliver 9 817 (holotype, STE; isotypes, K, MO, PRE) (Figure 1). Plantae parvae, ad 2.5 cm altae, foliis productis 3-4, laminis teretibus falcatis succulentis 10-35 x 0.6-1 mm, floribus cupuliformibus pallidis carneis ad malvinis vel albescentibus centribus luteis, tubo perianthii ca. 8 mm longo, tepalis unguiculatis ca. 15 mm longis, unguibus tepalorum interiorum ad columnam filamentorum con- natis, limbis patentibus, filamentis ca. 7 mm longis in columnam connatis supra divergentibus, antheris ca. 2 mm longis, stylo diviso prope medium antherorum, mar- ginibus stigmatum integribus undulatis. Plants small, to 2.5 cm high. Conn ± ovoid, 6- 8 mm diam., the tunics light brown, composed of reticulate fibers with prominent vertical ribs. Foliage leaves 3-4 per plant, blades terete, falcate, suc¬ culent, 10-35 x 0.6-1 mm, with a transparent adaxial band, the sheaths broad and transparent. Stem subterranean hut in fruit reaching shortly above the ground. Flowers actinomorphic, cuplike, pale pink to lilac (or whitish) with a yellow center; peri¬ anth tube ca. 8 mm long, cylindric; tepals ca. 15 mm long, unguiculate, the inner 3 tepals joined to the filament column for ca. 2 mm, tepal claws as¬ cending, ca. 6 mm long, the limbs spreading, ca. 6 mm long. Filaments ca. 7 mm long, united for the lower 6 mm in a cylindrical column (connate for 2 mm with the inner tepals), free and diverging above; anthers ca. 2 mm long, erect, yellow. Ovary ca. 4 mm long; style slender, reaching to about mid anther level, there diverging into 3 lobes, these with entire, undulate margins. Capsules ovoid -ellipsoid, ca. 6 mm long; seeds ± globose, ca. 0.6 mm diam., dark brown. Flowering June but depending on the advent of rain (probably until mid July); flowers normally open¬ ing ca. 10:00 hr. and collapsing ca. 15:30 hr. Distribution and habitat. Galaxia fenestralis is endemic to the eastern Kamiesberg Mountains of central Namaqualand and to the lower hills of the interior in the Kliprand District (Figure 1). It was discovered only in 1 990 and its entire range remains to be demonstrated. Like the apparently closely re¬ lated G. kamiesmontana, G. fenestralis favors rocky habitats where it grows in shallow sandy soil on exposed granite outcrops. It flowers at the beginning Volume 3, Number 4 1993 Goldblatt & Oliver G ataxia fenestralis 407 of the wet season when these sites remain damp for a few weeks and are sometimes waterlogged. In this basically very extreme habitat, which may dry out rapidly in warm weather, we surmise that the suc¬ culent leaves store enough water to permit plants to survive periodic desiccation of their habitat. The transparent adaxial window may serve as a means to increase light penetration into the inner tissues of the leaf so that photosynthesis is enhanced during the short periods when moisture conditions permit this to occur. Pnrntypes. SOUTH AFRICA. Cape: 3018 (Kamies- berg) Leliefontein Communal Area, valley southeast of Gooirnanskraal between Paulshoek and Nourivier, shallow- sand in seep on granite sheets, 1,130 m, 28 June 1991 (AC), Oliver 9858 (STE); Platbakkies, plateau northwest of Banke, seeps and damp sand on granite sheets, 1,100 m, 28 June 1991 (AD), Oliver 9852 (PRE, STE); Kli- prand District, between Wabreek and Middelpos, 1,080 m, 27 June 1991 (BC), Oliver 9849 (STE); hill slopes 4 km southwest of Kliprand, on Rietmond road, south-facing granite slabs, 960 m, 27 June 1991, Oliver 9846 (STE); 7.5 km southeast of Kliprand on Kamas road, south¬ facing slopes on granite, 840 m, 5 July 1990 (fr), Oliver 9510 (STE), 27 June 1991 (DB), Oliver 9844 (MO, PRE, STE). Acknowledgments. We thank John Manning for assistance with the leaf anatomical study, and Inga Oliver and John Manning for the illustrations used here. Literature Cited Eames, A. J. & L. H. McDaniels. 1925. An Introduction to Plant Anatomy. McGraw Hill, New York. Goldblatt, P. 1979a. Biology and systematics of Gal- axia (Iridaceae). J. S. African Bot. 45: 385-423. - . 1979b. Chromosome cytology and karyotype change in Galaxia (Iridaceae). PI. Syst. Evol. 133: 61-69. - . 1984. New species of Galaxia (Iridaceae) and notes on cytology and evolution in the genus. Ann. Missouri Bot. Card. 71: 1082-1087. - . 1990. Phylogeny and classification of Irida¬ ceae. Ann. Missouri Bot. Gard. 77: 607-627. - . 1991. An overview of the systematics, phy¬ logeny and biology of the southern African Iridaceae. Contrib. Bolus Herb. 13: 1-74. Ruptiliocarpon (Lepidobotryaceae): A New Arborescent Genus and Tropical American Link to Africa, with a Reconsideration of the Family1 Barry E. Hammel Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A. Nelson A. Zamora Instituto Nacional de Biodiversidad, Apdo. 22-3100, Santo Domingo de Heredia, Costa Rica and Escuela de Ciencias Ambientales, Universidad Nacional Autonoma, Apdo. 86-3000, Heredia ABSTRACT. A new genus and species of trees from Costa Rica and northern South America, Ruptilio¬ carpon caracolito, is described and compared to the African Lepidobotrys. It is distinguished from Lepidobotrys primarily by its much shorter fila¬ ments, basifixed rather than versatile anthers, its lack of styles, its two-rather than three-locular ova¬ ry, and by its irregularly dehiscent fruits with a woody exocarp and cartilagineous endocarp. In wood anatomy, apart from the presence of vestured pits in Ruptiliocarpon , the two genera are remarkably similar. They match in important floral and fruit characteristics (dioecious; 5 + 5 stamens of unequal length with filaments fused at base; two apical, col¬ lateral ovules per locule; one-seeded fruit; black seed with a large red aril and no endosperm), and both have unifoliolate leaves with very fugaceous stipules and stipels, and leaf-opposed inflorescences. The familial placement of Lepidobotrys has been con¬ troversial: the genus was placed first in the Linaceae by Engler, then in the Oxalidaceae by Hallier, then in its own family (between Linaceae and Erythrox- ylaceae) by Leonard, and then again in the Oxali¬ daceae by Cronquist. Reviewing evidence, old and new, we maintain Ruptiliocarpon and Lepidobo¬ trys (only) in the Lepidobotryaceae and suggest that they relate more to Sapindales or possibly Euphor- biaceae than to Oxalidaceae. Populations of Cedro caracolito, a local name for the new taxon described below, lay hidden and pro¬ tected in Costa Rica’s Osa Peninsula until recently built roads exposed the area to easy access for botanical exploration and the process of deforesta- 'The usual editorial policy of Novon only to allow papers that present new taxa has been relaxed for the two fol¬ lowing papers (Mennega, Tobe & Hammel); each was written in concert with and designed to be published with the core paper in which a new genus is described. tion. There, caracolito is a large, locally common tree with light wood having good qualities for cabinet work, but it is not generally known or sought by wood harvesters. Ongoing investigations of bark ex¬ tract show that it has promise as a biocide (natural- product agricultural pesticide, Arnason, pers. comm.). Several earlier collections of Ruptiliocarpon from South America (most of them filed among the Fa- baceae) have come to light since its discovery in Costa Rica. Study of the flowers led us to search among the Sapindales (Cronquist, 1981) and, fol¬ lowing submission of an earlier draft of this paper, we distributed numerous Costa Rican collections (in¬ cluding the type) with the new name but placed in Meliaceae. Bringing together information on wood anatomy, floral anatomy, embryology, and pollen morphology, we were later convinced to describe it as a new family. Finally, we saw it to its proper home as a distinct new genus of the hitherto mono- typic Lepidobotryacaeae, itself of controversial re¬ lationships. The novelty, problematic placement, and economic potential of Ruptiliocarpon underscore the urgency of continued exploration, study, and protection of tropical floras. Ruptiliocarpon caracolito Hammel & N. Za¬ mora, gen. et sp. nov. TYPE: Costa Rica. Li- mon: Cordillera de Talamanca, Canton de Ma- tina, cuenca media del Rio Barbilla, margen izquierda, sendero entre Cerro Amu siguiendo la fila hacia el este hasta estribaciones del Cerro Tigre, 200 m, 9 Nov. 1988 (fr), Herrera & Martinez 2310 (holotype, CR; isotypes, AAU, BM, CAS, COL, DUKE, F, G, GB, K, KYO, LE, MEXU, MICH, MO, NY, PM A, QCA, QCNE, RSA, S, TEX, UC, US, USJ, VEN, WIS). Lepidobotrys similis sed inflorescentiis et floribus mas- culinis femineis similibus, antheris basifixis antisepalis ses- Novon 3: 408-417. 1993. Volume 3, Number 4 1993 Hammel & Zamora Ruptiliocarpon 409 silibus, antipetalis in filainentis parvis, ovario biloculari, stylis quasi nullis, stigmatibus 2, capsulis exocarpio lignoso ruptili differt. Dioecious, evergreen trees 20-30(-40) rri tall, (20-)50-90 cm DBH; bole straight, smooth and branch-free for the lower ca. 15 m, light gray with large scattered lighter and darker patches; bark with very shallow, narrowly lanceolate, longitudinal fis¬ sures, bitter; wood pinkish white, porous and light¬ weight. Leaves unifoliolate; leaflets elliptic, (5-)6.5- 16 cm long, (2.7-)4-7.5 cm wide, entire, without pellucid lines or dots, without obvious fragrance, chartaceous, sparsely appressed-pubescent at the base with small simple trichomes, obtuse at the base, the apex acute to acuminate, the acumen ca. 1 cm long; lateral veins 5-7 pairs; reticulate venation obvious on the lower leaf surface; petiolule pulvin- ulate (thickened) for its entire length, 2-4 mm long with a distinct articulation al juncture with petiole and on emerging leaves, subtended by a slender clasping, deciduous stipel 4-5 mm long; petiole (0.4-)0.6-1.5 cm long, pulvinate at the base; stip¬ ules paired, ensiform, 1-1.5 mm long, soon decid¬ uous, visible on emerging leaves and leaving minute, scarcely visible scars on the twigs. Inflorescence a lax, mostly leaf-opposed (terminal) panicle of spikes; peduncle 1-2 cm long; 1 — 3(— 4) branches 2.3-8(- 10) cm long, each with a small bracteole at the base; rachis puberulous. Flowers cryptically unisexual (the staminate with pistil but only rudimentary ovules, the pistillate with stamens but no pollen), globose to ovoid, 4-4.5 mm long, 3.5-4 mm wide, at anthesis opening only slightly, green, subtended by 3 round¬ ed, abaxially sparsely puberulous, filiate bracts 0.5- 0.8 mm long; sepals 5, free, imbricate, ciliate, 1.6 2 mm long and wide; petals 5, free, imbricate, api- cally ciliate, 3-4 mm long, 2.5 mm wide. Stamens ca. 3 mm long (including filament tube), slightly shorter in female flowers than in male; filaments fused into a tube/nectary ca. 0.5 mm long; anthers 10, narrowly cordate, ca. 1 (— 1-2) mm long, inserted in 2 alternating series, the antipetalous at apex of filaments (ca. 0.5 mm long), the antisepalous ± sessile on the margin of the tube, rarely with a small, simple appendage produced on the margin of the tube between the anthers; connective produced to form a small pubescent appendage. Pollen tricol- porate with no thickenings at apertures, subspher- oidal, 10.5-15 gm (polar axis) x 11-16 /am (equa¬ torial axis); exine variable (di-or polymorphous?), psilate or verrucate to fossulate/foveolate; amb tri¬ angular; colpi rather short with thin costae ca. 0.8- 1 .0 /am thick; pores sometimes slightly protruding, elongated along polar axis, ca. 2. 5-4.0 /am; exine 0.8- 1.0 /am thick, tectate, columellae hardly visible. Intrastaminal or gynophoreal nectary lacking. Ova¬ ry ± ovoid, ca. 1.5-4 mm, smaller in the male flowers than in the female, glabrous, 2 -locular; each locule with 2 collateral ovules, pendulous from near the summit of the partition; obturator from funicular tissue present. Style essentially lacking, the summit of the ovary ± directly produced into an obscurely 2-lohed stigma. Fruit an ovoid 1 (rarely 2 (-seeded capsule, 2. 5-3. 5 cm long, 1.5-2. 5 cm wide; exo¬ carp coriaceous to woody, irregularly rupturing and falling away to expose 2 horny endocarps, one nearly completely surrounding the seed, the other usually empty and smaller, these also falling away, the larger taking on the shape of a snail shell. Seed globose, shiny black, % covered with a red-orange aril, pen¬ dulous (by the aril) at the end of a coriaceous strip (the partition) attached to the pedicel. Figures 1-3. Pnratypes. COSTA RICA. Puntarenas: Canton de Golfito, steep forested slopes above Golfito Airstrip, 1- 200 in, 19 Jan. 1984 (st), Pennington et al. J 1398 (CR); Reserva Nacional de Vida Silvestre Golfito, en fila entre Golfito y Villa Briceno, 200 m, 27 Jan. 1992 (fr), U. Chavarria et al. 511 (CR, F, MO); Canton de Osa. Reserva Forestal Golfo Dulce, entre Chacarita & Rincon, ca. 15 km de Chacarita, Alto los Mogos, 100 m, 27 Mar. 1991 (fl), Aguilar & Hammel 101 (AAU, BM, CAS, COL, CR, DUKE, F, G, GB, K, KYO, LE, MEXU, MICH, MO, NY, PMA, QCA, QCNE, RSA, S, TEX, UC, US, USJ, VEN, WIS); ca. 4 km de Rincon, 230 m, 9 Nov. 1990 (st), Hammel & M. M. Chavarria 17965 (CR, MO); antiguo campo de aterrizaje de Rincon, en fila al N de la Estacion Agua Buena de Boscosa, 300 m, 28 Nov. 1990 (fr), Hammel 17983 (CAS, CR, F, MO, NY, K, US, USJ, WIS); entre Rancho Quemado — por camino nuevo — y Drake, 50 m, 29 Mar. 1991 (imm fr), Aguilar et al. 103 (CR, F, MO); 300 m, 20 Mar. 1991 (fl), Hammel et al. 18154 (CR, F, MO); 300 m, 17 June 1990 (fr), Herrera 4198 (CR, F, MO); 21 Mar. 1989 (fl), Jimenez et al. 672 (CR); Fila Ganado, between Ran¬ cho Quemado — along old road — and Drake, 400 m, 5 June 1988 (fr), Hammel et al. 17034 (CR, F, MO); entre Rancho Quemado — por camino nuevo de madereros — y Guerra, 300 m, 28 Mar. 1991 (fl), Hammel et al. 18166 (CR, F, MO); Finca de Juan Marin, cerca a Guerra, 250 m, 6 Aug. 1991 (fr), Marin 87 (CR, MO); Reserva Indigena Guaymi, ca. 2 km noreste de la union del Rio Pavon con Rio Rincon, 100 m, 20 Oct. 1990 (fr), Ham¬ mel et al. 17911 (CR, F, MO); San Pedrillo, Playa Cam- panario, 10 m, 27 Mar. 1991 (fl), Harmon 210 (CR, F, MO). COLOMBIA. Valle: Bajo Calima, 15 km N of Buenaventura, 50 m, 26 Mar. 1986 (fr), Gentry et al. 53632 (MO). Etymology. The essence of caracolito’s most characteristic feature is captured by combination of the Latin “ruptilis,” irregularly splitting, with the Greek “carpon,” fruit. We explicitly choose this hybrid word “Ruptiliocarpon,” against recommen¬ dation of the Code, because we consider the purely Greek or Latin options decidedly inelegant (e.g.. 410 Novon Figure 1. Ruptiliocarpon caracolito Hammel & N. Zamora. — A. Seedling with insets: 1-stipel, 2-stipule. — B. Fruiting and flowering branches. — C. Flower showing longitudinal section and androecium. — D. Articulation of leaflet. — E. Fruit dehiscence series. Flowering material from Aguilar & Hammel 101, other material from Hammel 17983. “Ruptilifructum” or “Klastocarpon”) by compari¬ son. Likewise, we intentionally take “io” as the combining form rather than either “i” or “o” alone because it effectively and smoothly bridges the Latin/ Greek chasm. The epithet “caracolito,” meaning small snail, as in the shape of shell macaroni (also called “caracolito” in Spanish), is in reference to the shape of the fallen, horny endocarp and is taken Volume 3, Number 4 1993 Hammel & Zamora Ruptiliocarpon 411 Figures 2, 3. Flower and pollen of Ruptiliocarpon caracolito. — 2. Mature flower bud with perianth and seven anthers removed, showing staminal tube and ovary. Bar = 0.5 mm; Hammel 18154. — 3. Pollen. Bar = 5 /xm; Harmon 210. directly from the common name (Cedro caracolito). As such, it assumes the gender of its genus, in this case neuter, without change. We clarify these ori¬ gins so that well-meaning Latinists will resist cor¬ recting our spellings. In Costa Rica the common name “cedro” is used mostly — often in combination with a clarifying epithet — for species of Cedrela, Carapa (Meliaceae), Calophyllum (Clusiaceae), and Tapirira (Anacardiaceae). A common name tor Ruptiliocarpon in Peru is Cedro masha (R. Vas- quez, pers. comm.). Phenology. In Costa Rica, Ruptiliocarpon car¬ acolito flowers in late March to early April, directly following a flush of new leaves. The fruits are nearly mature by late December and last into February. By the time flowering begins, the previous year’s crop of fallen seeds has germinated and grown to the third or fourth leaf. Apparently not all mature individuals flower in a particular year. However, we have found flowering individuals that had obviously flowered and fruited the previous season. Germination. Seeds fallen below trees germinate readily in situ, and those taken Irom ripe fruits also germinate readily in pots; the radicle begins to emerge within ca. 2 weeks of planting. Germination is epi- geal; the thick cotyledons are green and the first leaves are opposite. The first few seedling leaves are often reddish below and, in general aspect, are strik¬ ingly similar to those of Protium aracouchini (Au- blet) Marchand (Burseraceae) and Pterocarpus spp. (Fabaceae). Habitat and Distribution. In Costa Rica this often large tree is common on slopes and hills of the Osa Peninsula and nearby Golfito. We have found it near sea level but most often between 100 and 400 m elevation and always in well-drained primary forest, typically with red clay soils. It has also been collected from near Barbilla (the type locality) on the Atlantic slope. This transmontane, distribution between the Caribbean lowlands and the Osa Peninsula on the Pacific slope, is exhibited in Costa Rica by numerous other wet forest, basically South American taxa such as Dendrobangia boli¬ viano Rusby (Icacinaceae), Hirtella tubiflora Prance (Chrysobalanaceae), Humiriastrum diguense Cua- trecasas (Humiriaceae), Pleurothyrium trianae (Mez) Rohwer (Lauraceae), Qualea paraensis Ducke ( Vochysiaceae ), and Thoracocarpus bissectus (Ve 1- lozo) Harling (Cyclanthaceae). On the basis of that pattern we had predicted the presence of the genus in South America. Now that it is known from Co¬ lombia, Peru, and Suriname, Ruptiliocarpon seems conspicuously absent from Panama and Ecuador. In addition to the Colombian specimen, included among the paratypes, numerous other South Amer¬ ican, mostly Peruvian, collections have also come 412 Novon Table 1. Characters shared by Ruptiliocarpon and Lepidobotrys. Leaves unifoliolate with stipules, stipels and disarticulation at upper pulvinus Inflorescences leaf opposed Flowers unisexual; 5 imbricate, free se¬ pals; 5 imbricate, free petals; 10 stamens with 5 longer and 5 shorter filaments; filaments fused at base into nectary/tube Ovules apical axial placentation, 2 collat¬ eral ovules per locule; placental obturator Fruits exocarp and endocarp separate; dehiscence septicidal Seeds black, one per fruit; orange aril at apex of fruit attached to tip of partition; endosperm lacking to our attention. No flowering material from South America has been seen, but the observed variation suggests that more than one species may be involved. Therefore, we explicitly isolate the following South American collections of Ruptiliocarpon from those cited as R. caracolito: PERU. Loreto: 7 km SW of Iquitos, 31 July 1972 (fr), Croat 18606 (MO); Maynas, Pucacuro, Rio Chambira, 160 m, 20 Apr. 1986 (fr), Vdsquez et al. 7452 (MO); Maynas, Nauta, Carretera a Iquitos, 150 m, 8 Dec. 1986 (lr), Vdsquez & Jaramillo 8475 (MO); Maynas, Iquitos, Quebrada de Aucaya hasta Union, 22 June 1976 (fr), McDaniel & Rimachi \. 20801 (MO); Maynas, Iquitos, Allpahuayo-IIAP, 150 m, Nov. 1990 (st), Vdsquez & Jaramillo 14820 (MO), Dec. 1990 (st), Vdsquez & Jaramillo 15543 (MO), (bud), Vdsquez & Jaramillo 15740 (MO), 22 May 1991 (fr), Vdsquez & Jaramillo 1 6314 (MO), 11 July 1991 (st), Vdsquez & Jaramillo 17366 (MO); Sa- puena, Rio Ucayali, 170 m, 2 July 1991 (fr), Gran- dez et al. 2732 (MO); Yanamono “Explorama Lodge,” 120 m, 19 May 1979 (fr), Diaz et al. 1140 (MO); 130 m, 26 June 1983 (st). Gentry et al. 42185 (MO), 10 July 1983 (st). Gentry et al. 42866 (MO). SURINAME. Nassau Mts., 530 m, 23 Mar. 1949 (st), Lanjouw & Lindeman 2877 (U), 1955 (st), Lindeman & Cowan 7020 (U). The fact that Ruptiliocarpon (in spite of the large population at Golfito) was neither reported by Allen (1956) nor apparently collected in Costa Rica before 1984 may result, in part, from the inconspicuous nature (small green flowers) of the species. However, as with Ticodendron (see Hammel & Burger, 1991) the discovery of Ruptiliocarpon surely has more to do with a recent general upsurge in collecting efforts and, especially in Costa Rica, an intensive focus on collecting and classifying the large trees. We should also expect that more specimens from earlier collections, throughout the Neotropics, will now come to light. Relationships. The section on relationships in earlier drafts of this paper focused on a search for a family for Ruptiliocarpon among the Sapindales. Due, in part, to a strong resemblance between the wood of Ruptiliocarpon and Trichilia (Mennega, 1993; R. Miller, pers. comm.; C. Morton, pers. comm.), but also because of floral similarities, most importantly, the filament tube, one earlier draft de¬ scribed Ruptiliocarpon as a new genus of Melia- ceae. Another, bringing together information on wood anatomy, floral anatomy, embryology, and pollen morphology, presented it as a new family. Curiously, some 40 years earlier Leonard (1950) had come to this same conclusion after a similar, family-by-fam¬ ily, search (in part among Sapindales) focused on the African genus we now believe to be Ruptilio¬ carpon ’s nearest relative. The African (Gabon-Cameroon region) Lepido¬ botrys is a monotypic genus that has been variously compared or assigned to Linaceae (Engler, 1902), Oxalidaceae (Hallier, 1923; Cronquist, 1981), Lep- idobotryaceae, Erythroxylaceae, and Sapindales (Leonard, 1950). Ruptiliocarpon and Lepidobo¬ trys are nearly identical in their wood and leaves. Although floral and seed anatomy of Lepidobotrys have not been analyzed to the same detail as Rup¬ tiliocarpon, the two genera coincide unambiguously in important floral and fruit characters, as well as in the unusual leaf-opposed position of inflorescences (Table 1). Specimens of them compared side by side immediately proclaim kinship from across the At¬ lantic Ocean. Here follows the history of why it took at least 40 years (after many millions) to make that comparison and get these two back together. We first saw and recognized Ruptiliocarpon as problematic after collecting fruiting material in Costa Rica in 1988. As in nearly all examples of earlier collections and identifications from elsewhere that have come to light, the plant was soon determined to be a legume. Convergence among workers in identifying collections of Ruptiliocarpon as Faba- ceae is remarkable. The earliest known collection (Suriname, 1949 (st), Lanjouw & Lindeman 2877) was placed provisionally at U in the “Papilionaceae” (Mennega, pers. comm.). The first known fertile collection (Peru, 1972 (fr), Croat 18606) was iden¬ tified as Swartzia simplex (Swartz) Sprengel. Most other collections that have come to our attention had been identified as Bocoa, a papilionoid legume. Volume 3, Number 4 1993 Hammel & Zamora Ruptiliocarpon 413 Although the fruits (immature) of our early col¬ lections, outwardly similar to those of species of Cynometra, were initially interpreted as drupaceous, the arillate seed was contradictory and suggested Swartzia. Furthermore, a scan through the Faba- ceae material at MO revealed that the double pulvini, articulation of the leaflet, and overall appearance of leaf venation make caracolito look very much like a unifoliolate legume such as Bocoa prouacensis Aublet (we now know that some specimens of this species at MO were actually sterile, misidentihed collections of Ruptiliocarpon), Dalbergia mone- taria L.f., or Swartzia hostmannii Bentham. The discovery of stipules and stipels on seedling leaves further supported Fabaceae. As noted below, many of the wood characters of Ruptiliocarpon, including vestured pits, are also in accordance with that fam- ily. However, once phenology and logistics Anally co¬ incided (in 1990), numerous flowering individuals were seen in the held, and flowers from five different collections were examined microscopically, it be¬ came clear that Ruptiliocarpon could not be a legume. The flowers, with a staminal tube/nectary, a compound, 2-locular ovary with 2 collateral ovules per locule and apical axial placentation, decisively eliminate the Fabaceae and would seem to place the new taxon within the Sapindales of Cronquist (1981). These and other floral characters give Ruptilio¬ carpon a resemblence to Meliaceae, although the secretory nature of the staminal tube would appear to be discordant in that family (Tobe & Hammel, 1993). Pollen morphology characters of Ruptilio¬ carpon are also discordant in Meliaceae because the characteristic (for Meliaceae) apertural thickenings are not present in Ruptiliocarpon (Hooghiemstra, pers. comm.). Although the wood of Ruptiliocarpon is very similar to Trichilia (Mennega, 1993) one important feature, vesturement of the vessel pits, which was discovered late in this analysis and reconfirmed with SEM, does not coincide with Trichilia nor with any other Sapindales and adds to the list of characters that isolate Ruptiliocarpon from Meliaceae. It is remarkable, here, that just as overall vegetative appearances of Ruptiliocarpon suggest Fabaceae, many of the wood characters, including vestured pits, are also in accordance with that family (Men¬ nega, 1993). In any case, as shown elsewhere in this paper, floral and fruit characters of Ruptilio¬ carpon must eliminate Fabaceae from consideration and suggest Sapindales. Other families of Sapindales such as Rutaceae and Simaroubaceae are similar to Ruptiliocarpon in many reproductive characters, but none coincides in convincing detail. In the Simaroubaceae stipules do occur (rarely) but arils are not known. Details of the flowers (free stamens, presence of a disk and usually only partially united carpels) tend to elimi¬ nate the family from consideration for Ruptilio¬ carpon. The capsular or follicular Rutaceae have fruits with similar loose, horny endocarps, but many other characters combine against placing caracolito in that family. In Rutaceae the exocarp remains with the infructescence rather than falling from the seed, the seeds lack arils, the flowers are bisexual, and stipules are wanting. Gland dots are present in the leaves of Rutaceae but lacking in Ruptiliocar¬ pon. In Burseraceae (where stipules are rare but known) fruit dehiscence in certain genera (e.g., Bur- sera) is somewhat similar to caracolito in that the leathery exocarp falls away leaving the arillate di- aspore attached to the pedicel, but the “arif’ (pseu- daril, fide Daly, 1989) is attached to the endocarp, which is stony and sealed until germination. In the genus Dacryodes the fruit is indehiscent with an oily and resinous mesocarp and an exarillate seed (Daley, pers comm.), but the cartilaginous endocarp is very similar to that of Ruptiliocarpon . The en¬ docarp is separable into two pieces, one smaller and empty with the margin folded over, the other larger and covering the seed, just as in Ruptiliocarpon. Nevertheless, in contrast to Ruptiliocarpon, the tribe Canarieae, which includes Dacryodes, is char¬ acterized by 3-merous flowers, fused sepals, and valvate petals. Furthermore, in contrast to typical Burseraceae, caracolito lacks resin ducts in the bark, lacks a floral disk, has two (rather than four or five) carpels and has the stamens fused in a single whorl rather than free in two whorls. The Burseraceae were eliminated early on in the wood anatomy anal¬ ysis because, in distinction to Meliaceae, Sapinda- ceae, and Rutaceae, the Burseraceae are rather uniform in their anatomy and not at all like Rup¬ tiliocarpon (Mennega, 1993). The Sapindaceae are known to have stipules in some of the lianas and agree with caracolito in many floral details, although they always have free stamens and, typically, a nectary disk and an ovary with one or two ovules in each of three locules. In fruit, the only similarity between caracolito and Sapindaceae (other than the presence of an aril) is that in certain genera (e.g., Paullinia and Thouinia) the mature carpels often fall away (entire) from the central axis, and in car¬ acolito the exocarp and endocarp split off (in pieces) leaving the seed dangling by the central axis. But for a casual glance at the excellent revised edition of Thonner’s key to families (Geesink et al., 1981), we might have published Ruptiliocarpon in its own new family, leaving Lepidobotrys stranded 414 Novon Table 2. Differences between Ruptiliocarpon and Lepidobotrys. Ruptiliocarpon Lepidobotrys Vestured pits + - Dioecy cryptic + obvious Fibrous exotegmen + - Gynoecium 2-merous 3-merous Styles - + Exocarp woody leathery Endocarp cartilaginous chartaceous Filaments short long Anthers basifixed versatile Connective apiculate truncate Inflorescences panicle of spikes fasciculate spikes in Africa a while longer. The cryptic dioecy of Rup- tiliocarpon, which was only revealed in 1992 by anatomical studies (Tobe & Hamrnel, 1993), is per¬ haps the one major clue that led to this late success with Thonner. Lepidobotrys staudtii Engler has unifoliolate leaves with fugaceous stipules and sti- pels, and an articulation at the upper pulvinus. It has leaf-opposed inflorescences; small green flowers with five imbricate sepals and petals; ten stamens of two different lengths with the filaments fused at the base into a nectary-tube; two collateral, apical ovules per locule; obturators; capsular fruits splitting septicidally to reveal a single, black, exendosperm- ous seed with a red aril attached to the tip of the partition, which is in turn attached to the pedicel (Table 1 and Fig. 1). Rank. Although cedro caracolito is very similar to Lepidobotrys, we distinguish Ruptiliocarpon be¬ cause the two are different in ways that can be regarded as generic: Ruptiliocarpon has vestured pits, Lepidobotrys does not (Mennega, 1993); Rup¬ tiliocarpon has nearly identical male and female inflorescences and flowers, in Lepidobotrys dioecy is more obvious; pedicels are lacking in Ruptilio¬ carpon, 5-8 mm long (male flowers) in Lepidobo¬ trys ; anthers are basifixed and apiculate in Rupti¬ liocarpon, versatile and truncate in Lepidobotrys; the fused part of the filaments (nectary-tube) is much longer than the free part in Ruptiliocarpon, much shorter in Lepidobotrys; Ruptiliocarpon has a ses¬ sile, bilobed stigma and 2-locular ovary, Lepido¬ botrys has 3 styles and 3-locular ovary; the exocarp is woody and irregularly dehiscent with a cartila- genous endocarp in Ruptiliocarpon but leathery and 3-parted and with a papery endocarp in Lep¬ idobotrys; the persistent partition is much narrower than the seed in Ruptiliocarpon, nearly as wide as or wider than the seed in Lepidobotrys; the mature seed of Ruptiliocarpon has a fibrous exotegmen lacking in Lepidobotrys (Tobe & Hamrnel, 1993) (Table 2). Lepidobotryaceae Reconsidered Although Leonard (1950) reviewed the literature concerning Lepidobotrys and described the family Lepidobotryaceae, the issue was not settled. Hutch¬ inson (1959, 1967, 1973) included the primarily tropical Asian Dapania and Sarcotheca in Lepi¬ dobotryaceae. However, as pointed out by Veldkamp (1967) in his revision of these two genera and also by Willis (1973), Dapania and Sarcotheca are de¬ cidedly members of Oxalidaceae and closely related to Averrhoa, whereas their relationship to Lepi¬ dobotrys is questionable. Oltmann (1971) stated that the pollen of Lepidobotryaceae (including Da¬ pania and Sarcotheca) stands next to Oxalidaceae but also shows affinity to the broadly circumscribed Linaceae complex where the floral morphology is more concordant, particularly with that of Lepi¬ dobotrys (e.g., presence of obturators). Huynh (1969) considered that the pollen of Lepidobotrys can be distinguished from all others of the family (Oxalidaceae) by its very long apertures. Oltmann (1971) also felt that Lepidobotrys has pollen distinct enough to support status as a separate family. Leaf anatomy (cf. discussion in Mennega, 1 993) is un¬ informative on the question and wood anatomy, it¬ self, does not presently resolve the issue. The most striking similarities between Lepido¬ botrys and Oxalidaceae (i.e., Dapania and Sar¬ cotheca) lie in the unifoliolate, articulate leaves and the woody habit. According to Leonard (1950), this similarity to leaves of Oxalidaceae, which generally have compound leaves with articulate leaflets, was Hallier's (1923) and later Knuth's (1931) principal reason for removing Lepidobotrys from Linaceae and placing it in Oxalidaceae. They might just as Volume 3, Number 4 1993 Hammel & Zamora Ruptiliocarpon 415 well have placed it in the Fabaceae, where groups with not only compound leaves and articulated leaf¬ lets exist, but also with stipules and stipels, features lacking in Oxalidaceae including Dapania and Sar- cotheca. The filaments of two lengths are also sug¬ gestive of Oxalidaceae, but that condition is present, as well, in some Sapindales, e.g., Trichilia spp. (Pennington, 1981). The main new evidence presented here in support of Leonard s recognition ol Lepidobotryaceae as a lamily distinct from Oxalidaceae has to do with obturators and endosperm. Engler (1902) described the ovules of Lepidobotrys as having the placenta dilated into a caruncle that covers the micropyle. As pointed out by Hallier (1923) and Oltmann (1971) this structure is an obturator, which presumably serves to guide pollen tubes to the micropyle (Cron- quist, 1981). Placental obturators are also present in Ruptiliocarpon but lacking in Oxalidaceae. Both Lepidobotrys and Ruptiliocarpon lack endosperm in mature seeds (Tobe & Hammel, 1993). Although endosperm is often scanty or absent in some mem¬ bers of CronquisFs Geraniales (Geraniaceae and Bal- saminaceae), it is nearly always copious in Oxali¬ daceae. Furthermore, Leonard did not point out that the petals are clawed in Oxalidaceae but unclawed in Lepidobotrys (and Ruptiliocarpon). Leaves and bark of Ruptiliocarpon have bitter substances while those of Oxalidaceae are sour or acidic. The Ox¬ alidaceae are said to be tenuinucellate (Cronquist, 1981), whereas at least Ruptiliocarpon is crassin- ucellate. However, Averrhoa (Oxalidaceae) is also crassinucellate (Thathachar, 1942). In addition to the above, and as pointed out by Leonard, Lepidobotryaceae differ importantly from Oxalidaceae in having septicidal (or irregular) rather than loculicidal (or no) dehiscence, collateral rather than superposed ovules, two or three carpels rather than five, and a disk (nectar-producing fused portion of filaments) (Table 3). With regard to the disk, a similarity to Oxalidaceae should be examined fur¬ ther; Cronquist stated that the outer filaments are often thickened nectariferous below in Oxalidaceae. Also the nature of the so-called disk in Lepidobotrys needs to be studied histologically. In contrast to published descriptions and drawings (e.g., Knuth, 1931; Tisserant, 1949; Leonard, 1950; Hutchin¬ son, 1959; Badre, 1973), our examination of re¬ constituted dried male flowers of L. staudtii revealed no discontinuity (other than fusion) between free parts of filaments and the point where the androecial structure attaches to the base of the petals. The filaments simply expand gradually from tip to base and the “disk” seems quite obviously to be the fused basal portion of the filaments. We had this same problem with Ruptiliocarpon , interpreting the structure first as a disk, then as a filament tube until finally it was shown, histologically, to be both, sta- minal tube and nectary (Tobe & Hammel, 1993). All evidence considered, placing Lepidobotrys in Oxalidaceae seems no more defensible than placing Ruptiliocarpon in Fabaceae. The leaves, outwardly so similar to Dapania and Sarcotheca in the former and Swartzia and Bocoa in the latter, have been the principal culprit in both these errors and are homologous, we believe, only in the case of uniting Ruptiliocarpon with Lepidobotrys. Where, then, do the broader affinities of Lepidobotryaceae lie? Leonard (1950) also compared Lepidobotrys with Linaceae and Erythroxylaceae, hut none of the new evidence we have accumulated for Ruptiliocarpon suggests a particularly close relationship to either of those two families (Mennega, pers. comm.; Tobe, pers. comm.). Our preliminary studies of Ruptilio¬ carpon focused almost entirely on comparison with families of Sapindales, and that comparison, al¬ though discarded for Lepidobotrys by Leonard (1950), is still viable on the basis of floral, seed coat, and wood anatomy. Ruptiliocarpon seemed partic¬ ularly close to Meliaceae, but its pollen, while similar to that of Sapindaceae (Hooghiemstra, pers. comm.; Pennington, pers. comm.), is not consistent with placement in the Meliaceae as it lacks the thick¬ enings at the apertures, characteristic of that family (Hooghiemstra, pers. comm.). Lepidobotryaceae, apparently, has never been compared to the Euphorbiaceae. However, its dioecy and obturators (both revealed very late in the in¬ vestigation of Ruptiliocarpon) give credence to that possibility. The paired, apical, collateral ovules co¬ incide directly with the primitive subfamily of eu- phorbs, Phyllanthoideae (Webster, 1975). Fibrous exotegmen, a character emphasized by Tobe & Hammel in comparing Ruptiliocarpon to Melia¬ ceae, is also found among the Phyllanthoideae (Cor¬ ner, 1976). Other important characters consistent with Euphorbiaceae include: two or three carpels; septicidal dehiscence; differentiation and separation of exocarp and endocarp; shape of the endocarp; and attachment of the seed via the aril (as in e.g., Aporosa and Richeria) to the persistent axis, which is suggestive of the characteristic euphorb columella (Cronquist, 1981). Although all of the wood ana¬ tomical characters manifested by Lepidobotryaceae may be found among the Phyllanthoideae, its par¬ ticular set of characters is not found in any one genus (Mennega, pers. comm.) Furthermore, the androecium of Lepidobotryaceae, with its filament- tube disk, and the unifoliolate, articulate leaves are discordant in Euphorbiaceae. If Lepidobotyraceae 416 Novon Table 3. Comparison of Lepidobotry aceae to Meliaceae, Oxalidaceae, and Euphorbiaceae. Lepidobotryaceae Meliaceae Oxalidaceae Euphorbiaceae Leaves unifoliolate compound, rarely unifoliolate compound, rarely unifoliolate simple, rarely compound or unifoliolate (?) Petiole articulate with the petiolule continuous articulate with petiolule in unifoliate spp. continuous Stipules + - - + Wood ± - - — (+ in one genus) (vestured pits) Inflorescence leaf-opposed axillary, terminal or rarely extra- axillary axillary axillary, terminal or very rarely leaf-opposed Flowers unisexual uni-bisexual bisexual unisexual Stamens 10 10 (-25) 10 (l-)5-many Filaments fused, bicydic fused, sometimes bicyclic fused, bicyclic free-fused, not bicyclic Nectary staminal tube separate or lacking ? (as scales) or lacking separate or lacking Carpels 2 or 3 ( 1 — )2— 5(— 20) 5 2 or 3(-4-many) Ovules/locule 2 collateral (l-)2(-12) collateral or superposed (l-)2-several superposed 1 or 2 collateral Obturator + ± - * Placentation apical-axial axial (often apical) axial apical-axial Fruit septicidal septi-loculicidal loculicidal septicidal capsule capsule capsule (schizocarp)- various Seed 1 1 - many numerous (l-)3-several Seed coat ± ± - ± (fibrous exotegrnen) Endosperm - + (-) + + (-) were found to be a close outgroup to Euphorbiaceae, the generally accepted, simple-leaved origin of the Euphorbiaceae might be challenged. We have found a family for Ruptiliocarpon but the exact placement of Lepidobotryaceae among other Rosidae is not yet clear. Biogeography Apparent sister genera separated by the Atlantic, Ruptiliocarpon and Lepidobotrys keep company with such taxa as Cecropia Musanga and Pour- ouma-Myrianthus (Cecropiaceae — Berg, 1978), Duguetia Pachypodanthium (Annonaceae — Schatz, pers. comm.) and numerous others cited by Thorne (1973). These pairs, endemic to their re¬ spective continents, are examples suggesting that vicariance of the original population via plate tec¬ tonics resulted in the taxonomic structure we see. The substantial differentiation between Ruptilio¬ carpon and Lepidobotrys, as well as the wide dis¬ tribution of Ruptiliocarpon within the Neotropics, is consistent with a division in the range of the ancestor population somewhere near or relatively Volume 3, Number 4 1993 Hammel & Zamora Ruptiliocarpon 417 soon after the South American and African [dates began to separate about, 100 million years ago (Raven & Axelrod, 1974). In contrast to the re¬ cently described A 'yssa (Cornaceae) and Ticoden- dron (Ticodendraceae) from Costa Rica (cf. Hammel & Burger, 1991; Hammel & Zamora, 1990) we have no evidence here to support a boreotropical origin of Ruptiliocarpon (cf. Lavin & Luckow, 1993). Rather, the wide distribution and variability of Ruptiliocarpon in South America and its restric¬ tion to Costa Rica otherwise, suggest dispersal into Central America from a South American origin. Within this context, Lepidobotryaceae may he quite old, as is also suggested by its similarity to primitive members of various more or less disparate groups in the Rosidae. Acknowledgments. Supported by grants from the National Science Foundation (BSR-9006449) and the National Geographic Society (M. Grayum, 3317- 86) to the collaborative project “A Manual to the Plants of Costa Rica.” We thank Francisco Hodgson for the line drawings, Mary Merello (flower and pollen) for SEM photos, and D. Daley, G. Davidse, M. Grayum, T. Pennington, D. Stevens, B. Styles, and W . Burger, as well as two anonymous reviewers, for critical comments on earlier drafts ot this paper. We also thank our colleagues and collaborators at the Museo Nacional/Herbario Nacional de Costa Rica and the Instituto Nacional de Biodiversidad for continuing support of facilities and scientific inter¬ action. This work has benefitted greatly through our long and fruitful correspondence with Alberta Men- nega and Hiroshi Tobe. Literature Cited Allen, P. H. 1956. The Rainforests of Golfo Dulce. Univ. Florida Press, Gainsville. Badre, F. 1973. Lepidobotryaceae. In: Flore du Gabon 2 1 : 40-42. Museum National d'Histoire Naturelle, Paris. Berg, C. C. 1978. Cecropiaceae, a new family of the Urticales. Taxon 27: 39-44. Corner, E. J. H. 1976. The Seeds of Dicotyledons, 2 vols. Cambridge Univ. Press, Cambridge. Cronquist, A. 1981. An Integrated System of Classifi¬ cation of Flowering Plants. Columbia Univ. Press, New York. Daly, D. C. 1989. Studies in neotropical Burseraceae II. Generic limits in New World Protieae and Can- arieae. Brittonia 41: 1 7 — 27. Engler, A. 1902. Linaceae Africanae. Bot. Jahrb. 32: 108. Geesink, R., A. J. M. Leeuwenberg, C. E. Ridsdale & J. F. Veldkamp. 1981. Thonner’s Analytical Key to the Families of Flowering Plants. Leiden Univ. Press, Boston. Hallier, H. 1923. Lepidobotrys Engl., die Oxalidaceen und die Geraniaceen. Beih. Bot. Central!). 39(2): 163. Hammel, B. & W. G. Burger. 1991. Neither oak nor alder, but nearly: The history of Ticodendraceae. Ann. Missouri Bot. Card. 78: 89 95. - & N. Zamora. 1990. Nyssa talamancana (Cornaceae), an addition to the remnant Laurasian Tertiary flora of southern Central America. Brittonia 52: 165-170. Hutchinson, J. 1959. The Families of Flowering Plants, 2nd ed. Oxford Univ. Press, Oxford. - . 1973. The Families of Flowering Plants, 3rd ed. Oxford Univ. Press, Oxford. - . 1967. The Genera of Flowering Plants. Ox¬ ford LIniv. Press, Oxford. Huynh, K.-L. 1969. Etude du pollen des Oxalidaceae I. Morphologie generate - Palynotaxonomie des Ox- alis americains. Bot. Jahrb. 89: 271-303. Knuth, R. 1931. Oxalidaceae. In: Engler & Prantl, Die Natiirlichen Pflanzenfamilien, ed. 2, 19a: 40-41. Lavin, M. & M. Luckow. 1993. Origins and relation¬ ships of tropical North America in the context of the boreotropics hypothesis. Amer. J. Bot. 80: 1-14. Leonard, J. 1950. Lepidobotrys Engl. Type d’une fam- ille nouvelle de spermatophytes: Les Lepidobotry¬ aceae. Bull. Jard. Bot. Bruxelles 20: 31-40. Mennega, A. M. W. 1993. Comparative wood anatomy of Ruptiliocarpon caracolito (Lepidobotryaceae). Novon 3: 418-422. Oltmann, O. 1971. Pollenmorphologisch — Systema- tische Untersuchungen innerhalb der Geraniales. Dis- sertationes botanicae, 11. Pennington, T. D. 1981. A Monograph of Neotropical Meliaceae. FI. Neotrop., Monogr. 28. Raven, P. H. & D. I. Axelrod. 1974. Angiosperm biogeography and past continental movements. Ann. Missouri Bot. Card. 61: 539 673. Thathachar, T. 1942. Studies in Oxalidaceae (Bioph- ytum sensitivum DC., Averrhoa carambola L. and Averrhoa bilimbi L.). J. Indian Bot. Soc. 21: 21. Thorne, R. 1973. Floristic relationships between trop¬ ical Africa and tropical America. Pp. 27-47 in B. Eggers et al. (editors), Tropical Forest Ecosystems in Africa and South America. Smithsonian Institution Press, Washington, D.C. Tisserant, P. C. 1949. Sur le Lepidobotrys staudtii Engl. Bull. Soc. Bot. France 96: 214. Tobe, H. & B. Hammel. 1993. Floral morphology, embryology and seed anatomy of Ruptiliocarpon caracolito (Lepidobotryaceae). Novon 3: 423-428. Veldkamp, J. F. 1967. A revision of Sarcotheca Bl. and Dapania Korth. (Oxalidaceae). Blumea 15: 519 543. Webster, G. 1975. Conspectus of a new classification of the Euphorbiaceae. Taxon 24: 593-601. Willis, J. C. 1973. A Dictionary of the Flowering Plants and Ferns, 8th ed. Cambridge Univ. Press, Cam¬ bridge. Note added in proof. We have now found flowering material from South America from among the family indets. at the Field Museum of Natural History: PERLJ. Loreto: Mishuyacu, near Iquitos, 100 m, Jan. 1930 (fl), King 749 { F). Annotations indicate that Erythroxylaceae, Meliaceae, and Burseraceae had been successively con¬ sidered. The flowers are somewhat larger than those of the Costa Rican material. Comparative Wood Anatomy of Ruptiliocarpon caracolito (Lepidobotryaceae) Alberta M. If . Mennega University of Utrecht, Institute of Systematic Botany, Heidelberglaan 2, P.0. Box 80.102, 3508 TC Utrecht, The Netherlands ABSTRACT. Wood anatomy of Ruptiliocarpon caracolito is described and compared in general to Sapindales, and specifically to Trichilia (Meliaceae) and the monotypic, African Lepidobotrys (Lepi¬ dobotryaceae). It is aberrant in all groups compared for having vestured pits. Otherwise, it is most similar to both Trichilia and Lepidobotrys. Wood anatomy does not conflict with the recognition of Ruptilio¬ carpon as a second genus of Lepidobotryaceae; how¬ ever, the question of affinities of that family needs further investigation. Analysis of wood anatomy is often crucial for the elucidation of relationships of taxa problematic al the generic or higher levels (cf. Hayden & Brandt, 1984; Mennega, 1984; Pennington & Styles, 1975). During the course of an investigation (Hammel & Zamora, 1993) into the affinities of a Costa Rican tree that could not be placed to family, an analysis of its wood anatomy became essential. Materials and Methods Microtome sections and macerations of wood of Ruptiliocarpon caracolito Hammel & N. Zamora from the trunk ( Hammel & Chavarria 17965, bole ca. 17 cm diarn.; MO) and from a branch ( Hammel 17983, 4.5 cm diam.; MO) both from near Rincon de Osa on the Osa Peninsula of Costa Rica, of Lepidobotrys staudtii Engler (Breteler 2087, Cam¬ eroon near village Zende, tree 15 m; Uw, WAGw), and of Trichilia lepidota C. Martius subsp. leu- caste ra (Sandwitb) Pennington (Maas 10841, Su¬ riname, Maratakka, tree 13 cm diam.; Uw) were prepared according to standard methods (Mennega, 1982). Descriptions, counts, and measurements fol¬ low recommendations of the International Associa¬ tion of Wood Anatomists Committee (I AW A, 1989). Wood Anatomy of Ruptiliocarpon caracolito GENERAL ASPECT A straight-grained wood apparently without dif¬ ferentiation in sapwood and heartwood, color uni- Novon 3: 418-422. 1993. lormly light, pinkish cream; moderately light, vol¬ ume weight ca. 0.40. MICROSCOPIC CHARACTERS (FIG. 1 ) Growth rings faint, formed by a narrow zone of flattened fibers and occasionally by a 1- or 2-celled band of parenchyma. Vessels solitary for about 45%, the remainder in radial multiples of 2 or 3(-8) and a few clusters, the latter mainly on the border of the growth ring; number 5(0-12) per sq. mm, dis¬ tribution somewhat irregular; perforations simple, perforation plates oblique; cross section oval to round, diameter (70— ) 1 00— 140 yum, average vessel mem¬ ber length 860(600-1,300) yum, mostly with long, narrow tails; intervascular pits alternate, crowded, vestured, diameter 5-6.5 yum, the slits enclosed or locally confluent; vessel/ray pitting similar; resinous contents occasionally present. Fibers regularly dis¬ tributed, angular in cross section, thin-walled, di¬ ameter 22-28 yam, the walls 2. 5-3. 5 yam wide; nonseptate; minute bordered pits restricted to the radial walls; length 1,000(750-1,290) yum. Fiber/ vessel ratio 1.16. Rays uniseriate, homogeneous or nearly so, cells procumbent, except for a marginal row of slightly higher and shorter cells, which re¬ semble square cells; number 5-7 per mm; width 15 yum, height 130-500 yum, up to 21 cells high; no contents. Parenchyma as scattered strands and in fine, often interrupted, rather straight bands 1 or 2 cells wide, also paratracheal, narrow vasicentric, occasionally aliform; terminal parenchyma as a band 1 or 2(-4) cells wide. Number of bands 6-8 per mm; strands of (2 -)4-8 cells. Rhombic crystals nu¬ merous in subdivided cells of the isolated strands. Discussion Preliminary analysis of wood samples of Rupti¬ liocarpon suggested a relationship to Meliaceae by way of a very close match to the genus Trichilia. This was in agreement with Hammel and Zamora’s independent conclusion that Ruptiliocarpon seemed to belong to Sapindales. Although they found that Volume 3, Number 4 1993 Mennega Wood Anatomy of Ruptiliocarpon caracolito 419 Figure 1. Ruptiliocarpon caracolito Hammel & N. Zamora. — A. Transverse section, x45. Vessel and parenchyma distribution. — B. Tangential longitudinal section, x 1 12. Uniseriate rays and parenchyma strands, partly with crystals. — C. Tangential longitudinal section, x 1 12. A vessel member with a lump of gum. — D. Radial longitudinal section, x 1 12. Homogeneous to weakly heterogeneous rays. All from Hammel & Chavarria 17965. — E. Vestured inter- vascular pitting; Hammel 17983. 420 Novon Figure 2. A, B. Trichilia lepidota C. Martius subsp. leucastera (Sandwith) Pennington. — A. Transverse section, x45. Vessel and parenchyma distribution. — B. Tangential longitudinal section, xll2. Intervascular pitting and uniseriate rays. Both from Maas 10841. C, D. Lepidobotrys slaudtii Engler. — C. Transverse section, x47. Vessel and parenchyma distribution. — D. Tangential longitudinal section, x 120. Uniseriate rays, intervascular pitting, crystal-bearing parenchyma strand (left side). Both from Breteler 2087. Volume 3, Number 4 1993 Mennega Wood Anatomy of Ruptiliocarpon caracolito 421 certain features of the fruits of Burseraceae resemble those of Ruptiliocarpon, vessels with large pits, septate fibers, and scarce parenchyma are salient features of Burseraceae that characterize their dif¬ ference from Ruptiliocarpon and eliminate that family from consideration as a close relative. Three Sapindalean families, Sapindaceae, Hippocastana- ceae, and Meliaceae, have members with wood that superficially resembles that of Ruptiliocarpon. In Sapindaceae, which like Meliaceae has great diver¬ sity in wood structure, the genus Talisia shows a remarkable similarity to Trichilia (Mennega, 1972) and therefore with Ruptiliocarpon (see below). The main difference is the occurrence of septate fibers in Talisia , in particular in the species with thin- walled fibers. Billia (Hippocastanaceae) is compa¬ rable to Ruptiliocarpon but here more numerous dissimilarities occur; these include a different pa¬ renchyma distribution with mainly rather wide con¬ spicuous terminal bands and hardly any diffuse strands, random occurrence of septate fibers, and shorter vessel members. Among Sapindales, the closest match to Rupti¬ liocarpon is Trichilia of the Meliaceae. Superficial examination of the branch sample suggested a sim¬ ilarity to Trichilia (Fig. 2A, B), and that impression was confirmed in detail on closer study of cross sections. Wood characteristics of Ruptiliocarpon such as nonseptate fibers, small intervascular pits, uniseriate wood rays that are weakly heterogeneous and not over 20 cells high, parenchyma in narrow wavy bands and partly aliform-confluent, several strands with rhombic crystals, and normal strands of 4 8 cells are features present in Trichilia ac¬ cording to older and more recent literature (e.g., Pennington & Styles, 1975). In a study of wood samples from the Guianas, Klaassen (1988) con¬ firmed the above features for nine species of Tri¬ chilia and also recorded a vessel member length of 550-760 fan, fiber length of 870-1,300 /am, very similar to those for Ruptiliocarpon. One important feature, vesturement of the vessel pits, that was discovered later in this analysis and reconfirmed with SEM (Fig. IE) does not coincide with Trichilia or with any other Meliaceae (Kribs, 1930; Record & Hess, 1943; Metcalfe & Chalk, 1950). A reference in Metcalfe & Chalk (1983) to a report of vestured pits in Meliaceae is an error; the original paper (Kanazawa, 1968) does not make that claim. It is remarkable, here, that just as overall vegetative appearances of Ruptiliocarpon suggest Leguminosae (see discussion in Hammel & Zamora, 1993), many of the wood characters, including ves¬ tured pits, are also in accordance with that family. In fact, in Record’s (1944) key one is led to a choice between Trichilia with vascular pits less than 4 /am wide and Leguminosae with pits more than 4 /xm; Ruptiliocarpon, with pits 5-6.5 ^nn in diameter, should key to Leguminosae. On the other hand, vessel members in legumes are seldom over 500 /am (Baretta-Kuipers, 1981; Metcalfe & Chalk, 1950; Reinders-Gouwentak & Rijsdijk, 1968), whereas in Ruptiliocarpon the length ranges from 600 to 1,300 /um. In any case, floral and fruit characters of Rup¬ tiliocarpon must eliminate Leguminosae from con¬ sideration (Hammel & Zamora, 1993;Tobe& Ham¬ mel, 1993). Ruptiliocarpon differs from Trichilia on the ba¬ sis of wood anatomy primarily because of tbe ves¬ turing of the vascular pits, a feature not reported for Meliaceae (nor any other Sapindales). Presence or absence of this feature has long been considered constant for a given family or genus. The few ex¬ ceptions include the tribe Bauhinieae in Legumi¬ nosae, which lacks vestured pits, otherwise present in the family; Bridelia, the only Euphorbiaceae with vestured pits (Mennega, 1987); and certain species of Prunus from China (Zhang & Baas, 1992), a genus that otherwise lacks vestured pits. It may be that the exceptions are too many and that we should no longer attribute such great value to this feature as a condition sine qua non in assigning a given taxon to a family or genus, but it does add, impor¬ tantly, to the list of characters suggesting that Rup¬ tiliocarpon does not belong in Trichilia or even in Meliaceae. On the eve of describing Ruptiliocarpon as a monotypic genus in its own family within Sapindales, congruence in a majority of vegetative, floral, and fruit characters with Lepidobotrys, an African monotypic genus in its own family, was discovered (Hammel & Zamora, 1993; Hammel, pers. comm.). Examination of wood of Lepidobotrys staudtii En- gler (Fig. 2C, D) also revealed a close conformity to the wood of Ruptiliocarpon. Apart from the absence of vestured pits in Lepidobotrys, the main difference is found in the more diffuse parenchyma in the latter species. In wood anatomy Trichilia and Lepidobotrys appear to differ and agree with Rup¬ tiliocarpon in similar ways, but the preponderance of other evidence favors a relationship with Lepi¬ dobotrys (Hammel & Zamora, 1993). The question of where the affinities of Lepidobotryaceae (Lepi¬ dobotrys and Ruptiliocarpon ) lie remains open. The historical alignment of Lepidobotryaceae in the Linaceae complex or in Oxalidaceae was discussed by Van Welzen & Baas (1984) based on a leaf anatomical study. Since leaf anatomy proved neutral 422 Novon with respect to the question, they saw no reason to differ from the generally accepted preference for its placement in Oxalidaceae. However, the wood struc¬ ture of Lepidobotrys, poorly known at the time Van Welzen & Baas published their analysis (cf. Metcalfe & Chalk, 1950: 272), is in several respects entirely different from that of Averrhoa, one of the few woody members of Oxalidaceae. The Linaceae com¬ plex comprises several mainly woody families with a rather great diversity of structure. From literature and personal knowledge of the wood structure of genera belonging to these families, a close relation¬ ship of Lepidobotryaceae with that complex seems improbable. For a well-founded statement, compar¬ ative research of a much broader scope than this paper would be necessary. The present study, along with accumulated knowledge (cf. Mennega, 1987), suggests that a relationship to the Sapindales or to Euphorbiaceae, as proposed by Hammel & Zamora (1993), should also be considered. Wood anatomy alone does not support or eliminate the possibility that Lepidobotryaceae may lie close to Euphorbi¬ aceae. Although most of the wood anatomical char¬ acters present in Lepidobotryaceae are manifested among the genera of Euphorbiaceae in subfamily Phyllanthoideae, they are not found together in any one genus. Also, rays in Phyllanthoideae woods are decidedly heterogeneous, whereas those of Lepido¬ botryaceae are homogeneous or nearly so. Acknowledgments. It was a great pleasure to have the opportunity to discuss the problematic wood of Ruptiliocarpon with Regis B. Miller during his recent brief visit to Utrecht. I am also greatly in¬ debted to him for the kind offer to have the SEM photographs made in Madison at the U.S. Forestry Products Laboratory. I thank my colleagues J. Koek- Noorman, L. Y. Th. Westra and B. J. H. ter Welle for critical remarks and for technical advice and as¬ sistance. Thanks are also due to B. Hammel for his much appreciated suggestions for textual improve¬ ments. To I). Makhan I am indebted for the prep¬ aration of slides. Literature Cited Baretta-Kuipers, T. 1981. Wood anatomy of Legu- minosae: Its relevance to taxonomy. Pp. 677-705 in R. M. Polhill & P. H. Raven (editors), Advances in Legume Systematics. Part II. Royal Botanic Gar¬ dens, Kew. Hammel, B. & N. Zamora. 1993. Ruptiliocarpon (Lep- idobotryaceae): A new arborescent genus and tropical American link to Africa, with a reconsideration of the family. Novon 3: 408-417. Hayden, W. & D. Brandt. 1984. Wood anatomy and relationships of Neowawraea (Euphorbiaceae). Syst. Bot. 9: 458-466. IAW'A Committee. 1989. I AW A list of microscopic features for hardwood identification. IAWA Bull. 10: 219-323. Kanazawa, K. 1968. Electronmicroscopic investigation on the vestured pit. [Japanese: English summary.] Bull. Fac. Agric. Shizuoka Univ. No. 18, 75-83. For. Abstr. 31/1278. Klaassen, R. 1988. Wood anatomy of the Meliaceae in the Guianas. Unpublished manuscript, Utrecht. Kribs, D. A. 1930. Comparative anatomy of the woods of the Meliaceae. Amer. J. Bot. 17: 724-738. Leonard, J. 1950. Lepidobotrys Engl. Type d’une fain- ille nouvelle de spermatophytes: Les Lepidobotry¬ aceae. Bull. Jard. Bot. Bruxelles 20: 31-40. Mennega, A. M. W. 1972. Wood structure of the genus Talisia (Sapindaceae). Acta Bot. Neerl. 21: 578- 586. - . 1982. Stem structure of the New World Men- ispermaceae. J. Arnold Arbor. 63: 145-171. - . 1984. Wood structure of Jablonskia con- gesta Euphorbiaceae. Syst. Bot. 9: 236-239. - . 1987. Wood anatomy of the Euphorbiaceae, in particular of the subfamily Phyllanthoideae. Bot. J. Linn. Soc. 94: 111-126. Metcalfe, C. R. & L. Chalk. 1950. Anatomy of the Dicotyledons. Clarendon Press, Oxford. - . 1983. Anatomy of the Dicotyledons, 2nd ed., vol. II. Clarendon Press, Oxford. Pennington, T. D. & B. T. Styles. 1975. A generic monograph of the Meliaceae. Blumea 22: 419-540. Record, S. J. 1944. Keys to American woods. XIV. Dicotyledonous woods with xylem rays virtually all uniseriate. Trop. Woods no. 79: 25-34. - & R. W. Hess. 1943. Timbers of the New World. New Haven. Reinders-Gouwentak, C. A. & J. F. Rijsdijk. 1968. Hout van Leguminosae uit Suriname. Veenman & Zonen, Wageningen. Tobe, H. & B. Hammel. 1993. Floral morphology, embryology, and seed anatomy of Ruptiliocarpon caracolito (Lepidobotryaceae). Novon 3: 423-428. Welzen, P. C. Van & P. Baas. 1984. A leaf anatomical contribution to the classification of the Linaceae com¬ plex. Blumea 29: 453-479. Zhang, Shu-Yin & P. Baas. 1992. Wood Anatomy of trees and shrubs from China. III. Rosaceae. IAWA Bull. n.s. 13: 21-91. Floral Morphology, Embryology, and Seed Anatomy of Ruptiliocarpon caracolito (Lepidobotryaceae) Hiroshi Tobe Department of Biology, College of Liberal Arts and Sciences, Kyoto University, Kyoto 606, Japan Barry E. Hammel Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A. ABSTRACT. Floral and seed anatomy of Ruptilio¬ carpon caracolito , based on histological studies, are described in detail. Cryptic dioecy, a nectariferous starninal tube, and ovules with obturators comprise tbe characters most crucial to elucidation of rela¬ tionships of this taxon. Originally compared among Sapindales, Ruptiliocarpon seemed close to but dis¬ tinct from Meliaceae; floral anatomy suggested that it be recognized in its own separate family, a move obviated by the discovery of its close relationship to the monotypic African Lepidobotryaceae. However, the floral and seed anatomy of Lepidobotrys have not been studied in the same detail as those of Ruptiliocarpon , and the relationships of Lepido¬ botryaceae ( Ruptiliocarpon and Lepidobotrys ) re¬ main controversial. A preliminary survey of tbe literature supports the suggestion that Lepidobotry¬ aceae may be closer to Sapindales or perhaps Eu- phorbiaceae than to other groups with which it has been compared. Histological studies of floral and seed anatomy often provide critical information for discovering relationships of problematic taxa (cf. Tobe, 1991; Hammel & Wilder, 1989). Material from collections of yet another Costa Rican tree that could not be placed to family was analyzed to that end. Materials and Methods Buds ( Aguilar & Hammel 101, Aguilar et al. 103 , Hammel et al. 18154, 18166), young fruits and mature seeds ( Aguilar et al. 103, Hammel 17983) were fixed in FAA for this analysis. Buds and young fruits were softened as described in Tobe & Raven ( 1 984) and thin sectioned with a rotary microtome following standard paraffin methods. Sec¬ tions were stained with Heidenhain’s hematoxylin, safranin, and fastgreen FCF. Pieces of mature seed coat were dehydrated through an ethanol series, embedded with Technovit 7100 (Kulzer & Co., Ger¬ many) resin, sectioned on a rotary microtome and stained with 0.02% Toluidineblue 0 (Tobe et ah, 1992). Floral Morphology As indicated by anatomical sections, flowers are unisexual and the plants apparently dioecious. All flowers have stamens and a pistil, but the pistil of male flowers ( Aguilar & Hammel 101, Hammel et al. 18154, 18166) is smaller than that of female flowers ( Aguilar et al. 103) and bears only rudi¬ mentary ovules (Figs. 1, 2). Female flowers have stamens with shorter filaments and lack pollen grains in anthers (Figs. 4, 5). Flowers are pentamerous, comprising five sepals, five petals, ten stamens, and one pistil (Figs. 2, 4, 7). Both sepals and petals are free and imbricate in aestivation and alternately arranged. The ten sta¬ mens are inserted at the same level of the receptacle and fused along nearly the entire length of their filaments, forming a starninal tube (Fig. 2). The starninal tube is particularly conspicuous in the male flowers, where filaments are longer. There, the five antisepal stamens have a short length of free filament extending beyond the starninal tube, while the five antipetal stamens lack free filament (Fig. 7 A. H, J). In the male flowers the nectariferous starninal tube is densely stained and even has vascular tissue that is profusely branched on the adaxial side (Fig. 3). The starninal tube is massive and histologically ap¬ pears to represent a nectary. Neither an intrastam- inal nor a gynophoreal nectary is present. In female flowers, however, the starninal tube is less conspic¬ uous (Fig. 6) and may not actually exude nectar. Vascular tissue just below the level oi the recep¬ tacle in male flowers forms a vascular cylinder of ca. 10 discrete vascular bundles (Fig. 7B). At and above the level of the calyx, the vascular bundles divide and successively supply vascular tissue to sepals, petals, stamens, and pistil (Fig. 7C-F). Each Novon 3: 423-428. 1993. 424 Novon Figures 1-6. Floral anatomy of Ruptiliocarpon. — 1. Longitudinal section of male flower. — 2. Transverse section of male flower. — 3. Transverse section of nectariferous staminal tube of male flower. Arrowheads indicate vascular tissue of the nectary. — 4. Transverse section of female flower. — 5. Longitudinal section of female flower. — 6. Longitudinal section of stamen base of female flower. Arrowhead points to adaxial side of filament. Abbreviations: nc, nectary (nectariferous staminal tube); ov, ovule; pe, petal; ps, pistil; se, sepal; st, stamen. Scales equal 10 jun in Figures 3 and 6, 1 mm in Figures 1, 2, 4, 5. Volume 3, Number 4 1993 Tobe & Hammel 425 Floral and Seed Anatomy of Ruptiliocarpon caracolito Figure 7. Diagrams illustrating the vascular anatomy of male flower of Ruptiliocarpon. — A. Median longitudinal section through the line presented in J. — B-J. Transverse sections at levels marked b-j in A. Sepals and petals are not drawn in H and only stigmas are presented in I. Abbreviations: nc, (nectariferous staminal tube); pe, petal; ps, pistil; se, sepal; st, stamen. 426 Novon Figures 8-16. Anatomy of ovules and seeds of Ruptiliocarpon. — 8. Longitudinal section of mature ovule. — 9. Longitudinal section of an upper half of mature ovule. — 10. Longitudinal section of younger ovule at 4-nucleate embryo sac stage. — 11. Transverse section of young seed. — 12. Longitudinal section of an upper part of young seed. — 13. Longitudinal section of young seed. — 14. Half of longitudinally dissected mature seed. — 15. Longitudinal section of mature seed coat on raphal side. —16. Transverse section of mature seed coat. Abbreviations: ar, aril; ch, chalaza; cot, cotyledon; cp, nucellar cap; ext, exotegmen; ii, inner integument; mic, micropyle; pc, parietal cell(s); ob, obturator; oi, outer integument; r, raphe; sc, seed coat; tg, tegmen; ts, testa; vs, vascular bundle. Scales equal 10 mui in Figure 7, 50 //m in Figures 8, 9, 100 gm in Figures 11, 15, and 16, 200 gm in Figures 10, 12, and 14, and 5 mm in Figure 13. Volume 3, Number 4 1993 Tobe & Hammel 427 Floral and Seed Anatomy of Ruptiliocarpon caracolito sepal generally receives three or more vascular bun¬ dles (Fig. 7C, D). while each petal and stamen re¬ ceives a single vascular bundle (Fig. 7D, E). Vascular bundles enter the base of the ovary arranged in a ring and divide upwardly into ovary wall bundles and axial placental bundles (Fig. 7E-G). In female flowers the ovule supply is provided from the axial bundles to nearly the top of the locule (Fig. 5). Ovule Morphology and Embryology Ovules are anatropous, bitegmic and crassinu- cellate (Figs. 8 10). In the youngest ovules avail¬ able, a 10-cell-layered parietal tissue is present above the eight-nucleate embryo sac (Fig. 10). A thick, three-cell-layered nucellar cap derived from apical dermal cells of the nucellus is formed (Fig. 9). Both the inner (ii) and outer (oi) integuments are multi¬ plicative; at the mature ovule stage the ii is about four cells thick and oi 7-10 cells thick. A micropyle is formed by the ii and oi together (Fig. 12). Ovules and young seeds are more or less pachv- chalazal. A funicular vascular bundle divides at the chalaza, forming a vascular plexus, but none of the vascular branches enters the integuments (Fig. 13). An obturator is formed from funicular tissue near the micropyle (Fig. 9). No hypostase is differenti¬ ated. Seed Anatomy Mature seeds are ellipsoid, about 17-19 mm long and 11-12 mm in diameter with a funicular aril. They are exalbuminous and contain a straight em¬ bryo with massive cotyledons (Fig. 14). The mature seed coat has both testa (developed oi) and tegmen (developed ii) (Fig. 16). Early in development only exotegmic cells are enlarged, while all remaining cells are not specialized (Figs. 11, 12). All consti¬ tuting cells of the young testa are persistent; at maturity the testa is 8-10 cells thick, comprising small, flattened, tanniniferous cells. Compared to other cells, exotestal cells may be more or less enlarged and have thick outer cell walls. On the other hand, the tegmen of the mature seed is much more spe¬ cialized, comprising a well-developed fibrous exo- tegmen and a collapsed but persistent meso-and endotegmen. Cells of the exotegmen are longitudi¬ nally elongate, thick-walled, pit ted, and up to 70- 90 jam thick (Figs. 15, 16). Discussion Prior to discovery of the similarity between Rup¬ tiliocarpon and the African Lepidobotrys search for affinities of Ruptiliocarpon concentrated on Sapindales (Hammel & Zamora, 1993; Mennega, 1993). Among Sapindales, Ruptiliocarpon coin¬ cides with Meliaceae in numerous reproductive char¬ acters. In both, flowers are pentamerous with five sepals, five petals, and ten stamens. They have a compound superior ovary with no distinct styles and each of the two locules has two ovules attached near the top of the partition. Filaments are connate to form a staminal tube; ovules are anatropous, bi¬ tegmic, and crassinucellate; an obturator and nu¬ cellar cap are present; seeds are pachychalazal (less conspicuous in Ruptiliocarpon ), exalbuminous, ar- illate (pro parte in Meliaceae), and exotegmen of mature seed coats is fibrous (pro parte in Meliaceae, including some Trichilia spp.). Ruptiliocarpon, however, appears to differ importantly from Meli¬ aceae with respect to the position of the floral nec¬ tary. In Ruptiliocarpon the floral nectary is the staminal tube itself, while in Meliaceae it is usually (always?) intrastaminal or gynophoreal and present as a separate disk (Boesewinkel, 1981; Corner, 1976; Nair, 1958, 1959a, b. 1963, 1970; Nair & Kanta, 1961; Narayana, 1958; Pennington, 1981). This and a number of other characters were considered sufficient to exclude Ruptiliocarpon from Melia¬ ceae, and describing it in its own family was con¬ sidered (see Hammel & Zamora, 1993). However, the formerly monotypic Lepidobotryaceae has sur¬ faced as the more logical disposition of Ruptilio¬ carpon. Since no floral material was available for anatom¬ ical analysis of Lepidobotrys , the close relationship between that genus and Ruptiliocarpon rests on the similarities apparent from examination of her¬ barium material and the literature, as discussed by Hammel & Zamora (1993). As far as is known from the literature, Ruptiliocarpon agrees with Lepi¬ dobotrys in floral anatomy in much the same way it agrees with Meliaceae. In addition, the floral disk (cf. Leonard, 1950) would appear to be more like the nectariferous staminal tube of Ruptiliocarpon than what is known for the Meliaceae. One mature seed of Lepidobotrys staudtii Engler ( Etuge & Tho¬ mas 478, MO) was available and examined for com¬ parison. Contrary to some literature reports (e.g., Tisserant, 1949), the mature seeds of Lepidobotrys certainly lack endosperm as do those of Ruptilio¬ carpon. The seed coat of Lepidobotrys does lack a fibrous exotegmen, present in Ruptiliocarpon and thought to be important in its comparison to Meli¬ aceae. However, this character is variable within a number of families, including Meliaceae, and does not necessarily exclude association of Ruptiliocar¬ pon with Lepidobotrys at the familial level. The suggestion of a relationship to (but not within) 428 Novon Euphorbiaceae (Hammel & Zamora, 1993) deserves attention. In terms of ovule and seed morphology and anatomy, Ruptiliocarpon agrees well with the primitive subfamily Phyllanthoideae hut not with advanced members of the Euphorbiaceae. A fibrous exotegmen, which has sometimes been considered significant for determining relationships, is found in quite a few genera of Phyllanthoideae (Corner, 1976). Nevertheless, a more definite resolution of the af¬ finities of Lepidobotryaceae ( Lepidobotrys and Ruptiliocarpon) must await examination of floral and young fruiting material of Lepidobotrys. Literature Cited Boesewinkel, F. D. 1981. Development of the seed of Trichilia grandifolia Oliv. (Meliaceae). Acta Bot. Neerl. 30: 459-464. Corner, E. J. H. 1976. The Seeds of Dicotyledons, 2 vols. Cambridge Univ. Press, Cambridge. Hammel, B. & G. Wilder. 1989. Dianthoveus: A new genus of Cyclanthaceae. Ann. Missouri Bot. Gard. 76: 112-123. - & N. Zamora. 1993. Ruptiliocarpon (Lepi¬ dobotryaceae): A new arborescent genus and tropical American link to Africa, with a reconsideration of the family. Novon. 3: 408-417. Leonard, J. 1950. Lepidobotrys Engl. Type d'une fam- ille nouvelle de spermatophytes: Les Lepidobotry¬ aceae. Bull. Jard. Bot. Bruxelles 20: 31-40. Mennega, A. 1993. Comparative wood anatomy of Ruptiliocarpon caracolito (Lepidobotryaceae). No¬ von 3: 418-422. Nair, N. C. 1958. Studies on Meliaceae. III. Floral morphology and embryology of Sandoricum indicum Cav. Phyton 10: 145-151. - . 1959a. Studies on Meliaceae. I. Floral mor¬ phology and embryology of Naregamia alata W. & A. J. Indian Bot. Soc. 38: 353-366. - . 1959b. Studies on Meliaceae. II. Floral mor¬ phology and embryology of Melia azedarach L. — A reinvestigation. J. Indian Bot. Soc. 38: 367-378. - . 1963. Studies on Meliaceae. VI. Morphology and floral anatomy of the tribe Cedrelieae and dis¬ cussion on the floral anatomy of the family. J. Indian Bot. Soc. 42: 177-189. - . 1970. Meliaceae. In: Proceedings of the Sym¬ posium on Comparative Embryology of Angiosperms. Bull. Indian Natl. Soc. Acad. 41: 151-155. - & K. Kanta. 1961. Studies in Meliaceae. IV. Floral morphology and embryology of Azadirachta indica A. Juss. — A reinvestigation. J. Indian Bot. Soc. 40: 382-396. Narayana, L. L. 1958. Floral anatomy and embryology of Cipadessa baccifera Miq. J. Indian Bot. Soc. 37: 147-154. Pennington, T. D. 1981. A Monograph of Neotropical Meliaceae. Flora Neotropica, Monograph No. 28. New York Bot. Card., New York. Tisserant, P. C. 1949. Sur le Lepidobotrys staudtii Engl. Bull. Soc. Bot. France 96: 214. Tobe, H. 1991. Reproductive morphology, anatomy, and relationships of Ticodendron. Ann. Missouri Bot. Gard. 78: 135-142. - & P. H. Raven. 1984. An embryological con¬ tribution to systematics of the Chrysobalanaceae. I. Tribe Chrysobalaneae. Bot. Mag. (Tokyo) 97: 397- 411. - , S. Yasuda & K. Oginuma. 1992. Seed coat anatomy, karyomorphology, and relationships of Simmondsia (Simmondsiaceae). Bot. Mag. (Tokyo) 105: 529-538. Rahowardiana globifera (Solanaceae), a New Species from Colombia Sandra Knapp The Natural History Museum, Cromwell Road, London, SW7 5BD, United Kingdom William G. D’Arcy Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. Abstract. Rahowardiana globifera, newly de¬ scribed from Antioquia, Colombia, is a second spe¬ cies for this striking, high-climbing solanaceous ge¬ nus. Rahowardiana is a small group of relatively rare epiphytic shrubs first described from Panama (D’Arcy, 1973: 670). It is a member of tribe Juan- ulloeae, whose members are all neotropical woody epiphytes. Rahowardiana is most closely related to Juanulloa, sharing with that genus narrow tubular flowers and highly branched inflorescences. Work in progress by V. Persson, S. Knapp & S. Blackmore suggests the genera share several pollen characters. Rahowardiana is distinctive, however, in its con¬ gested globose inflorescence with highly contrasting corollas and calyces. Rahowardiana globifera Knapp & D’Arcy, sp. now TYPE: Colombia. Antioquia: Parque Na- cional Natural “Las Orquideas,” Sector Ven- ados, 890 m, 6°33'N, 76°19'W, 1 Apr. 1988 (fl, fr), Cogollo, Ramirez & Alvarez 2895 (holotype, JAUM-018167; isotypes, COL not seen, FMB not seen, MO). Figure 1. Frutex scandens, foliis obovatis coriaceis magnis pe- tiolatis. Inflorescentia congesta globosa, pedicellis brevis. Flores calycibus tubulosis, coriaceis purpurea, corollis al- bis, antheris linearis. Bacca anguste turbinata. High-climbing, unarmed, scandent shrub, twigs stout, drying strongly angled; pubescence (seen only on leaf undersides) of slender, erect, weak, collaps¬ ing, simple, uniseriate hairs 3-5 cells long, the distal cells smaller. Leaves obovate, 15-30 x 10-21 cm, apically rounded, basally obtuse, veins obscure above, prominently elevated beneath, the lateral veins 4 on each side, looping and anastomosing 5-10 mm from the margin, the minor venation mostly per¬ pendicular, the margins slightly revolute, subcor- iaceous, concolorous, glabrous above, beneath softly and evenly pubescent; petioles drying dark, 15-25 mm long, ca. 5 mm thick, drying longitudinally ridged and crossed with numerous transverse fis¬ sures. Inflorescence a condensed, crowded, globose panicle or raceme ca. 15 cm long, forming spheres ca. 35 cm diam.; peduncle not evident, pedicels stout, ca. 5 mm long, glabrous. Flowers with calyx purple, tubular, strongly angled, 7-8 cm long, ca. 15 mm wide apically, coriaceous, irregularly sinuate lobed, the tips of the lobes green; corolla white, the tips of the lobes purple dorsally, ca. 6.5 cm long (immature), the lobes 0.5-1 cm long; stamens 5, the filaments straight, glabrous, inserted at or near the base of the corolla tube, anthers linear, ca. 15 mm long, 2-3 mm wide, yellow, the connective drying dark, the thecae confluent at the apex, de¬ hiscing longitudinally; ovary pale brown, conical, ca. 1 cm long, narrowly beaked, glabrous, surmounting a distinct, slightly lighter colored, slightly undulating annular nectary, style white, markedly different in color from the ovary, straight, 5-6 cm long, stigma strongly bifurcate, the lobes ca. 1 mm long, minutely papillate. Fruit a pale yellow, narrowly ovoid berry, ca. 9.5 cm long, basally 1 cm wide, expanding to ca. 2 cm wide about 4 mm from the base, then narrowing into a slender beak 4-5 cm long; seed fabiform, ca. 6x3.5 mm, tan colored, the surface minutely pitted. Rahowardiana globifera resembles Rahowar¬ diana wardiana D’Arcy, from Panama. It differs in its stouter twigs; larger, more coriaceous leaves, which are puberulent beneath; in its short pedicels; larger calyces; white instead of yellow flowers; and larger fruit. The inflorescences of this species are twice or three times as large as those of R. war¬ diana. Recent collecting in northwestern Colombia has uncovered many interesting new plants, one ot which is described here. The collecting area, Parque Na- cional “Las Orquideas,” is on the Pacific slope of the Western Cordillera, an area of high rainfall and high relief. This species was found in premontane rainforest, climbing in the overstory within hand’s reach. It is locally abundant, and the large, unusual inflorescences can be seen at a distance (Cogollo, pers. comm.). Novon 3: 429-430. 1993. 430 Novon E o If) CO A Figure 1. Rahowardiana globifera Knapp & D’Arcy ( Cogollo , Ramirez & Alvarez 2895, MO). —A. Twig with leaves and pendent inflorescence. — B. Twig with leaves. — D. Fruit, opened to show seeds. — E. Corolla opened ti Acknowledgments. Gratitude is noted for the help of Alvaro Cogollo, Jardin Botanico Joaquin Antonio Uribe, Medellin, and to John J. Pipoly, III, Missouri Botanical Garden. The illustration was prepared by John Myers. — C. Portion of inflorescence showing flowers and fruit, show stamens. Literature Cited D'Arcy, W. G. 1973. [1974]. Solanaceae. In: R. E. Woodson et ah, Flora of Panama. Ann. Missouri Bot. Card. 60: 573-870. A New Species of Cyanea (Campanulaceae: Lobelioideae) from KauaT, and the Resurrection of C. rernyi Thomas G. hammers Department of Botany, Field Museum of Natural History, Chicago, Illinois 60605-2496, U.S.A. David H. Lorence National Tropical Botanical Garden, Lawa‘i, Hawaii 96765, U.S.A. ABSTRACT. Two species are added to Cyanea , both endemic to Kaua‘i. One is new, based on a recently collected specimen, l be other was described pre¬ viously but excluded from the most recent account of the genus due to the inadequacy of the available material. It is reinstated on the basis of data obtained from several newly collected specimens. The genus now includes 56 species and nine heteronymic sub¬ species. Cyanea Gaudichaud (Campanulaceae: Lobelioi¬ deae) is a genus of trees and shrubs endemic to the Hawaiian Islands. Lammers’s (1990) treatment of the genus for the Manual of the Flowering Plants of HawaPi included 52 species and nine heteron¬ ymic subspecies. Recently, hammers (1992) added two species from Kaua‘i to the genus. In the present [taper, Cyanea is further enlarged by the addition of two more species from Kauah. One is newly described; the other is a previously described species that was not recognized in the Manual because of the inadequacy of the available material. As such, Cyanea now includes a total of 56 species and nine heteronymic subspecies. Both of these species were collected at a relatively inaccessible locality known as “The Blue Hole.” This site, located east of Kaua‘i’s center at the headwaters of the Wailua River, is a deep and extremely wet valley head surrounded on three sides by vertical cliffs 900 m high and laced with waterfalls. The vegetation here is an unusual mix of shrubland and low rainforest on saturated rocky ground. The can¬ opy is just 1-3 m tall, and the ground layer is rich in pteridophytes. A brief comment on the spelling of Hawaiian place names in the specimen citations is in order. The spellings adopted here are those given by Pukui et al. (1974), in which the macron (— ) is used to indicate long vowels and the hamza (‘) to indicate glottal stops. Although non-Polynesian authors are often tempted to omit these diacritical marks, doing so is a grammatical gaffe comparable to omitting the umlauts from German words or the tildes from Spanish ones. New Species During an excursion into the Blue Hole in June 1990, Steve Perlman collected a Cyanea with very unusual leaves. The lamina was less than three times as long as wide, cordate at base, and only half as long as the petiole. Nearly all species of Cyanea have leaves in which the lamina is several to many times longer than wide; attenuate, cuneate, obtuse, truncate, or rounded at base; and much longer than the petiole. The sole exception is the aptly named C. asarifolia H. St. John, which is endemic to east¬ ern Kaua‘i and also occurs in the Blue Hole. How¬ ever, the unidentified Cyanea differed from C. asar¬ ifolia by having the lamina ovate or oblong (vs. cordiform), 3. 7-5. 8 (vs. 7-8) cm wide, twice as long as wide (vs. almost as wide as long), with scat¬ tered hairs along the midrib on the lower surface (vs. glabrous), the margin callose-serrulate (vs. mi¬ nutely callose-crenulate), the base often markedly asymmetric (vs. symmetric); and by having the in¬ florescences 8- 1 1 -flowered (vs. 30-40-flowered), sparsely pubescent (vs. glabrous), the peduncle 4 4.5 (vs. 2.5-3) cm long, the pedicels 12-15 (vs. 7-10) mm long, and the bracts 5-7 (vs. 1.5) mm long. Perlman was able to locate only a single large individual just beginning to flower plus three juve¬ niles. Despite the lack of mature flowers and fruit, it is clear that these plants cannot be referred to any previously described member of the Lobelioideae and that they represent an undescribed species. We realize that publishing a novon on the basis of a single immature specimen is a less than commend¬ able practice. However, extenuating circumstances dictate timely publication in this case. First, fertile material is not likely to become available anytime in the foreseeable future. A survey of the type lo¬ cality in October 1992, six weeks after Hurricane Novon 3: 431-436. 1993. 432 Novon Iniki struck Kaua‘i, revealed that all known indi¬ viduals of this species had been destroyed by the hurricane. Although it is likely that additional plants survive on the cliffs surrounding the Blue Hole, the vertical aspect and great height of these cliffs ren¬ ders them virtually impossible to explore. Second, because of its apparent rarity, this species merits protection under the Endangered Species Act of 1973. However, to be listed, it must first “exist,” i.e., be described and have a name. Cyanea dolichopoda Lammers & Lorence, sp. nov. TYPE: Hawaii. Kaua‘i: Blue Hole, N fork of Wailua River, S side of Blue Hole, Metros- ideros polymorpha lowland wet shrubland with Pipturus, Cyrtandra, Touchardia, Cyanea asarifolia, Lysimachia filifolia, Hedyotis ela- tior, 1 June 1990, S. Perlman 1 1070 (holo- type, PTBG). Figure l . A Cyanea asarifolia laminis ovatis, duplo longioribus quam latioribus, 3.7 5.8 cm latis, pubescentibus in pagina inferna secus costam, margine calloso-serrulato, basi saepe asymmetrica; et inflorescentiis 8-1 1-floris, pubescentibus, pedunculis 4-4.5 cm longis, pedicellis 12-15 mm longis, bracteis 5-7 mm longis differt. Shrub, 1 m tall; stem unbranched, erect, un¬ armed, glabrous. Lamina ovate or oblong, 6.5-10 cm long, 3. 7-5. 8 cm wide; upper surface green, glabrous; lower surface light green, with scattered hairs along the midrib; margin callose-serrulate; apex acute or obtuse; base cordate, often markedly asym¬ metric; petiole 9-16 cm long, slender, glabrous. Inflorescence (not fully expanded) spreading or pen¬ dent, 8 1 1 -flowered, sparsely pubescent; peduncle 4-4.5 cm long, slender; rachis 2-2.7 cm long; bracts linear, 5-7 mm long; pedicels 12-15 mm long, spreading, bibracteolate at the middle; brac- teoles linear, 7.5-8 mm long. Corolla (in bud) pink¬ ish, glabrous. Fully expanded flowers, fruits, and seeds not seen. Distribution , Habitat and Phenology. Cyanea dolichopoda is known only from the relatively in¬ accessible type locality on Kaua'i, where it grew on a cliff face at ca. 700 m elevation (S. Perlman, pers. comm.). Judging from the size of the flower buds, anthesis might be expected to commence in middle to late June. Etymology. The specific epithet was coined by combining the Greek words dolichos, long, and po- dion , foot, in reference to both the long peduncles and the long petioles. Relationships. Both Cyanea dolichopoda and its close relative C. asarifolia are members of section Delisseoideae (Hillebrand) Rock. This group in¬ cludes about a dozen species and is found on all of the main Hawaiian Islands but Hawai‘i. The Delis¬ seoideae are characterized by their shrubby habit, relatively slender unarmed stems, entire or minutely toothed petiolate leaves, typically glabrous flowers, hypanthium 2-6(-8) mm long, calyx lobes typically dentiform, 0.5-1 mm long, and corollas 15-30(- 40) mm long, with lobes about as long as the tube. In the treatment of Cyanea in the Manual , C. dolichopoda would key with difficulty to C. asari¬ folia, from which it may be distinguished by the characters given above. Resurrection of Cyanea remyi Rock In addition to C. dolichopoda, botanists exploring the Blue Hole collected a second unusual species of Cyanea. This plant was present in larger numbers, with a total of three healthy populations numbering approximately 200 individuals scattered in the un¬ derstory at 600-730 m elevation (S. Perlman and K. Wood, pers. comm.). Similar plants were encountered in south-central Kaua‘i, some 9 10 km south of the Blue Hole, in the Wahiawa Mountains and the drainage of Wah- iawa Stream, during a botanical inventory of that region. The rugged mountainous terrain of this bowl¬ shaped region is dissected by numerous stream sys¬ tems and covered by lowland rainforest and boggy shrubland. Six populations of the unidentified plant, numbering 110 juveniles and adults in all, were encountered here at elevations of 660—780 m in forest understory (Lorence & Flynn, 1991). Most recently, this same unidentified species was encountered in northern Kaua'i at the headwaters of Wai‘oli Stream. Here, two sterile plants were discovered in the understory of lowland rainforest at 370 m elevation (S. Perlman and K. Wood, pers. comm.). This third site is located approximately 1 1 km north of the Blue Hole and 20 km north of the Wahiawa Mountains. The plants from these three localities could not be referred to any species of Cyanea described in the Manual and were thought by their collectors to represent a species new to science. However, when specimens were shown to the senior author in 1989, he suggested that they might represent C. remyi, which had not been included in the Manual. This species had been described by Rock (1919) on the basis of a single specimen deposited at Paris, which had been collected on “Kauai ou Nihau” (i.e., Kaua'i or Nifihau) sometime during the early 1850s by Jules Remy. 'This specimen, which the senior author had examined at Paris in 1987, consisted of a single stem bearing a few large leaves, two inflorescences Volume 3, Number 4 1993 Lammers & Lorence Cyanea dolichopoda 433 Figure 1. Cyanea dolichopoda Lammers & Lorence, apex of plant; inset shows portion of leaf margin, enlarged (from the holotype). 434 Novon with immature flower buds, plus a detached inflo¬ rescence apex (cf. Rock, 1917: pi. 12; 1919: pi. 158). In the 130 years following Remy’s visit, C. remyi was collected just once more. In 1916, C. N. Forbes gathered sterile specimens of an unknown Cyanea in the Hi‘i (Wahiawa) Mountains of Kaua'i. It was subsequently identified as C. remyi by Wimmer (1943). In preparing the treatment for the Manual, the senior author of the present paper came to essen¬ tially the same conclusion regarding C. remyi as Wimmer (1943: 78), who commented “Species non satis nota et eius positio dubia.” With only the incomplete holotype and Forbes’s sterile gathering at hand, it was not possible to characterize the spe¬ cies adequately and to include it in the dichotomous key. Instead, C. remyi was merely mentioned in the discussion under C. truncata (Rock) Rock, a species with leaves that are somewhat similar in size and shape. In order to determine whether the unidentified plants were referable to C. remyi, the holotype was borrowed and examined by both authors in 1991. After a thorough comparison, there was no doubt that the recent collections from all three localities on Kaua‘i, as well as Forbes’s 1916 collection, were in fact all referable to that species. The holotype and Forbes’s specimen fell well within the range of variation seen within and among the three extant populations. As a result, C. remyi is here resurrected and described more fully on the basis of the flowering and fruiting material now available. Cyanea remyi Rock, Bull. Torrey Bot. Club 44: 233. 1917. Delissea remyi (Rock) H. St. John, Phytologia 63: 87. 1987. TYPE: Hawaiian Islands, “Kauai ou Nihau,’’ 1851-55,7. Remy 302bis (holotype, P). Figure 2. Shrub, 0.9-2 m tall; stem unbranched (occa¬ sionally with 1 or 2 branches from base), 1-2.5 cm diam., erect, unarmed, dark purple and pubescent toward apex, brown and glabrous below; latex light yellow (occasionally white), thin. Lamina broadly elliptic, ovate, or broadly oblong, 16-40 cm long, 9.5-19.5 cm wide; upper surface green, glossy, glabrous, the midrib maroon or dark purple at least in the lower %; lower surface whitish green, glossy, with scattered short white hairs on the midrib and veins; margin minutely callose-denticulate; apex ob¬ tuse, acute, acuminate, or cuspidate; base subcor- date, truncate, rounded, or obtuse; petiole 8-20 cm long, 2-5 mm diam., maroon or dark purple, gla¬ brous or pubescent with short white hairs. Inflores¬ cence ascending, 6- 1 3 -flowered, pubescent with short white hairs; peduncle 4-8 cm long, 2-4 mm diam., maroon or dark purple; rachis 0.5-2 cm long, maroon or dark purple; bracts linear, 3 mm long, 0.6 mm wide, deciduous; pedicels 10-21 mm long, bibracteolate at or below the middle, maroon or dark purple; bracteoles linear, 1-3 mm long. Hypanthium obovoid or obconic, 9-12 mm long, 5-9 mm diam., dark purple, pubescent with short white hairs. Calyx lobes triangular or narrowly tri¬ angular, spreading or ascending, 4-6 mm long, 1- 2 mm wide, dark maroon; apex acute. Corolla bi¬ labiate, 40-53 mm long, dark purple, shading to purplish white at the apex of the lobes on their inner surface, densely pubescent with short white hairs; tube curved, 30-31 mm long, 5-5.5 mm diam., cleft dorsally for ca. V2 its length; dorsal lobes linear, spreading and slightly recurved, 19-22 mm long, 2-2.5 mm wide, acute at apex; ventral lobes linear, spreading and slightly recurved, 15-20 mm long, 2-2.5 mm wide, acute at apex. Staminal column slightly exserted; filaments 32-40 mm long, purplish white, glabrous; anther tube 1011 mm long, 2 mm diam., purplish white, the lower 2 anthers with tufts of white hairs at apex, otherwise glabrous. Berry spherical, 1013 mm long, 10-13 mm diam., ma¬ roon or dark purple with orange flesh, tuberculate. Seeds minute, black or dark brown, smooth, shining. Chromosome number (M. Kiehn, pers. comm.) 2 n = 28. Distribution, Habitat and Phenology. Extant populations are known from three localities on Kaua‘i, in the upper reaches of the Wailua River, Wahiawa Stream, and Wai‘oli Stream. The plants occur on wet seeping or saturated rocky substrates in areas of rainforest and shrubland at 370-770 m. Flow¬ ering specimens were collected in June, July, and September, and fruiting ones in September, Novem¬ ber, and April. Although the holotype is labeled “Kauai ou Ni¬ hau,” C. remyi is here considered to be endemic to the former. Certainly, the extreme wetness of the known localities, as well as their elevational range, suggests that the holotype could not have been col¬ lected on the much drier island of Ni'ihau, which is just 390 m above sea level at its highest point. Etymology. This species commemorates French naturalist and traveler Ezechiel Jules Remy ( 1 826— 1893), who collected several hundred botanical specimens in the Hawaiian Islands during the early 1850s. Additional specimens examined. HAWAII. Kaua4i: Hanalei District, Wai'oli Stream, back of valley, Wood <£ Perlman 1494 (PTBG); boundary of Llhu'e and Ka- Volume 3, Number 4 1993 Lammers & Lorence Cyanea dolichopoda 435 Figure 2. Cyanea remyi Rock, apex of plant (based on Wood et al. 0098); inset shows single flower, enlarged slightly (based on Lorence et al. 6592 and on a color transparency by D. H. Lorence). waihau districts, headwaters of N fork of Wailua River, area called “The Blue Hole” or “The Crater,” Lorence et al. 5327 (F, PTBC), Wood et al. 0098 (F, PTBG), Wood et al. 0352 (PTBG), Lorence et al. 6561 (F, PTBG), Lorence et al. 6565 (F, PTBG), Flynn et al. 3990 (PTBG), Lorence et al. 6592 (PTBG), [voucher for chromosome count] Kiehn 900823-1/5 (WU not seen), Wood et al. 1125 (PTBG), Wood et al. 1114 (PTBG), Wood & Perlman 1353 (PTBG); Hi‘i Mts., Forbes 660. K (B, BISH); Llhu‘e-Koloa F. R., Wahiawa Mts., just NE of Wahiawa Bog, along N fork of Wahiawa Stream, NW of Mt. Kahili, Lorence et al. 5949 (BISH, PTBG), Wag¬ ner et al. 6061 (BISH not seen), Wood et al. 0544 (PTBG); Llhu‘e-K5loa F. R., Wahiawa Mts., W ridge, below rope trail, W ood et al. 1206 (PTBG); Llhu'e-Koloa F. R., Wahiawa Mts., NW of Wahiawa Bog, along trib¬ utary of Wahiawa Stream, NW of stream and SE of Hulua, Flynn & Wood 3237 (PTBG), Flynn & Wood 3238 (PTBG); Llhu4e-K5loa F. R., W ahiawa drainage, Flynn et al. 4493 (PTBG); Llhu'e-Koloa F. R., W side of Wahiawa drainage, below ridge between Hulua and Pu'uau'uka, Wood et al. 1365 (PTBG). Relationships. Cyanea remyi is referable to sec¬ tion Hirtellae Rock, a group of eight species en¬ demic to Kaua‘i, which is characterized by shrubby habit, relatively slender unarmed stems, entire or minutely toothed petiolate leaves, densely pubescent flowers, hypanthium 6-11 mm long, calyx lobes triangular, l-5(-16) mm long, and corollas 25-53 mm long, with lobes about as long as the tube. Cyanea remyi may be incorporated into Lammers’s (1992) key to the species of Hirtellae by appending the following couplet to the beginning of that key: 436 Novon Oa. Lamina ovate, broadly elliptic, or broadly ob¬ long, the base subcordate, truncate, rounded, or obtuse; petiole 8-20 cm long . C. remyi Ob. Lamina narrowly elliptic, obovate, oblanceolate, or oblong, the base attenuate or cuneate (rarely obtuse); petiole 0.5-9 cm long . . (remainder of Hirtellae). In the Manual, C. remyi would key with some difficulty to C. sylvestris A. Heller, another member of the Hirtellae. The two species are similar in their relatively wide leaves (typically more than 6 cm wide), ascending peduncles, and large flowers (co¬ rolla more than 4 cm long). However, C. rernyi differs from C. sylvestris in its shorter stature (0.9- 2 m vs. 1.5-6 m tall) and in having the lamina ovate, broadly elliptic, or broadly oblong (vs. obovate or oblanceolate) and subcordate, truncate, rounded, or obtuse (vs. cuneate) at base; petioles 8-20 (vs. 2.5-8) cm long; peduncles 4-6.5 (vs. 1-4) cm long; corolla dark purple (vs. white with longitudinal pur¬ ple stripes); and berries spherical (vs. obovoid), 10- 13 (vs. 7-10) mm in diameter, maroon or dark purple (vs. orange or yellow). Cyanea truncata, the species under which C. remyi was previously discussed (Lammers, 1990), is a member of section Cyanea. It differs from C. remyi by having its stems and petioles muricate; lamina obovate, with a cuneate or truncate base; inflorescences horizontal or pendent; bracts narrowly oblong, 9-20 mm long; hypanthium 5-6 mm long; calyx lobes narrowly oblong, obtuse at apex; corolla 32-42 mm long, the lobes only VA~Vi as long as the tube; and berries orange, 8-10 mm long, 8-10 mm in diameter. Additionally, it is known only from 0‘ahu. Acknowledgments. The curators of BISH and P are acknowledged for the loan of specimens. Lam- mers’s visit to B and P in 1987 was supported by a Graduate Student Alumni Research Award from The Ohio State University. We are grateful to Tim Flynn, Steve Perlman, and Ken Wood for providing information on population sizes and other field ob¬ servations; Zorica Dabich for preparing the illustra¬ tions; and Michael Kiehn for permission to publish his chromosome number determination for C. rem yi. Literature Cited Lammers, T. G. 1990. Campanulaceae. Pp. 420 489 in W. L. Wagner, D. R. Herbst & S. H. Sohmer, Manual of the Flowering Plants of Hawai'i. Univ. Hawai‘i Press, Honolulu. - . 1992. Two new combinations in the endemic Hawaiian genus Cyanea (Campanulaceae: Lobelioi- deae). Novon 2: 129-131. Lorence, D. H. & T. Flynn. 1991. Botanical survey of the Wahiawa Drainage, Kaua'i. Final report sub¬ mitted to State of Hawai'i, Department of Land and Natural Resources, Division of Forestry and Wildlife. [Unpublished.] Pukui, M. K., S. H. Elbert & E. T. Mookini. 1974. Place Names of Hawaii, rev. ed. Univ. Hawai'i Press, Honolulu. Rock, J. F. 1917. Notes on Hawaiian Lobelioideae, with descriptions of new species and varieties. Bull. Torrey Bot. Club 44: 229-239 + pi. 9-16. - . 1919. A monographic study of the Hawaiian species of the tribe Lobelioideae family Campanula¬ ceae. Mem. Bernice Pauahi Bishop Mus. 7(2): i xvi, 1-395. Wimmer, F. E. 1943. Campanulaceae-Lobelioideae. I. Teil. Pp. i-iv, 1-260 in R. Mansfeld (editor), Das Pflanzenreich, IV. 276b. Wilhelm Engelmann, Leip¬ zig. Merger of the Endemic Hawaiian Genera Cyanea and Rollandia (Campanulaceae: Lobelioideae) Th omas G. hammers Department of Botany, Field Museum of Natural History, Chicago, Illinois 60605-2496, U.S.A. Thomas J. Givnish and Kenneth J . Sytsma Department of Botany, University of Wisconsin, Madison, Wisconsin 53706-1381, U.S.A. ABSTRACT. Phylogenetic analyses based on mo¬ lecular data indicate that the species of Rollandia are embedded within the genus Cyanea and that together these genera form a inonophyletic group. Cyanea and Rollandia are therefore merged under the former name. A revised key to the baccate genera of Hawaiian Fobelioideae and an expanded description of the genus are provided; eight new' combinations and one new name are proposed; and lectotypes are selected for Rollandia humboldtiana and R. longi flora var. angustifolia. This expanded circumscription makes Cyanea the largest genus in the Hawaiian flora, with 64 species and 10 heter- onymic subspecies. The flora of the Hawaiian Islands includes 114 endemic species of woody Lobelioideae (Campanu¬ laceae), 95 of which have baccate fruits (hammers, 1990, 1992a; Lammers & Lorence, 1993). Tra¬ ditionally (Gaudichaud, 1826; Bentham & Hooker, 1876; Hillebrand, 1888; Schonland, 1889; Hock, 1919; Wimmer, 1943), these baccate species have been divided among four genera: Clermontia Gau¬ dichaud, Cyanea Gaudichaud, Delissea Gaudi¬ chaud, and Rollandia Gaudichaud. However, var¬ ious alternative classifications have been proposed. Presl (1836) and de Candolle (1839) recognized a fifth genus, Macrochilus C. Presl. Gray (1861), however, recognized only three; Macrochilus was included in Cyanea and the species of Rollandia were divided between Cyanea and Delissea. En- dlicher (1836) and Dietrich (1839), taking a very broad view of generic circumscription, assigned all Hawaiian lobelioids to cosmopolitan Lobelia L. Bail- Ion (1885), on the other hand, treated the baccate Hawaiian species as a single genus, Delissea, with the four traditional genera as sections. The classi¬ fication of von Post and Kuntze (1903) was similar, except Rollandia was maintained as distinct. St. John (1987) merged Cyanea and Delissea under the latter name, but maintained Clermontia and Rollandia as distinct genera. The traditional four- genus classification was employed in the most recent taxonomic summary of the Hawaiian Campanula¬ ceae (Lammers, 1990). Recent phylogenetic analyses based on restriction site variation in the chloroplast DNA of Hawaiian lobelioids (Givnish et al., in press) have shown that the species comprising Rollandia are embedded within Cyanea as traditionally circumscribed, that Cyanea without Rollandia is paraphyletic, and that the two genera together form a monophyletic group. Specifically, Rollandia is part of a major clade that is defined by 10 synapomorphic mutations and gen¬ erally characterized by orange fruits and an un¬ branched habit. Within this major clade, Rollandia forms an unresolved clade defined by three muta¬ tions. Morphologically, the two genera are very similar. Every author for over 100 years has distinguished Rollandia solely on the basis of a single feature, i.e., the adnation of the stamina! column to the corolla (Hillebrand, 1888; Schonland, 1889; von Post & Kuntze, 1903; Hock, 1919; Wimmer, 1943; Lammers, 1990). The traditional classification of Campanulaceae includes several such cases of close¬ ly related genera distinguished by a single morpho¬ logical feature, e.g., Campanumoea Blume and Co- donopsis Wallich, Lobelia and Pratia Gaudichaud. The circumscription of such genera must be revised if a natural classification of the family is ever to be achieved (Lammers, 1992b, c). In order to provide a more natural classification of the baccate Hawaiian lobelioids, Cyanea and Rol¬ landia are here merged. Although Gray (1861) transferred some species of Rollandia to Cyanea . the species that was later designated as the lectotype of the genus (R. lanceolata) was assigned to De¬ lissea. Because no previous author has effectively merged just Cyanea and Rollandia , and because the two generic names are of equal priority, we have the prerogative of choosing the name to be retained. Novon 3: 437-441. 1993. 438 Novon We here choose Cyanea, as doing so will require far fewer nomenclatural innovations. The eight species of Rollandia (one comprising two subspecies) recognized by hammers ( 1 990) are here transferred to Cyanea with no change in their circumscription or rank. In addition, lectotypes are designated for two inadequately typified names. De¬ scriptions, distributional and habitat data, and lists of heterotypic synonyms are found in hammers’s (1990) treatment and will be amplified in the forth¬ coming monograph of Cyanea (hammers, in prep.). Assignment of these species to an infrageneric taxon within Cyanea is problematic. As noted previously (hammers, 1990), the infrageneric classification of Cyanea is badly in need of revision. The genus has been divided into either three (Hillebrand, 1888; Wimmer, 1943; St. John, 1969) or five sections (Rock, 1919; Stone, 1967). Neither classification maps cleanly onto the molecular clades defined by Givnish et al. (in press); for example, Cyanea sect. Palmiformes (Hillebrand) Rock comprises species belonging to at least three different clades. A definite infrageneric assignment is deferred, therefore, pend¬ ing completion of a comprehensive reevaluation of relationships within the enlarged Cyanea. This merger makes Cyanea the largest genus in the Hawaiian flora, with 64 species and 10 heter- onymic subspecies (Wagner et ah, 1990; hammers & horence, 1993). A morphological description of the newly enlarged genus is provided below, as is a revised key to all of the Hawaiian genera of hobe- lioideae with baccate fruit. Both are adapted from hammers (1990). A brief comment on the spelling of Hawaiian place names in the specimen citations is in order. The spellings adopted here are those given by Pukui et ah (1974), in which the macron (— ) is used to indicate long vowels and the hamza (4) to indicate glottal stops. Although non-Polynesian authors are often tempted to omit these diacritical marks, doing so is a grammatical gaffe comparable to omitting the um¬ lauts from German words or the tildes from Spanish ones. Revised Key to the Baccate Hawaiian Genera of Lobe- LIOIDEAE la. Stems repeatedly branched, spreading; inflores¬ cences (l-)2-3(-10)-flowered; corolla cleft dor- sally nearly to base . Clermontia lb. Steins unbranched or only sparingly branched, erect or ascending; inflorescences ( 3 — )5 — 2 5 ( 40)-flowered; corolla cleft to about the middle. 2a. Corolla with a small knob at the terminus of the dorsal cleft, sometimes with two ad¬ ditional lateral knobs; seeds 0.7- 1.5 mm long, white, dull, conspicuously transverse¬ ly rugose . Delissea 2b. Corolla smooth or rarely muricate, without definite dorsal or lateral knobs; seeds 0.2 0.5(— 2) mm long, dark brown to black, shiny, smooth or microscopically foveate- reticulate . Cyanea Cyanea Gaudichaud, Voy. Uranie 457. 1829. De¬ lissea sect. Cyanea (Gaudichaud) Baillon, Hist. PI. 8: 364. 1885. Kittelia Ih G. h. Reichen- bach, Handb. Nat. Pfl.-Syst. 186. 1837, nom. illeg. TYPE: Cyanea grimesiana Gaudichaud. Rollandia Gaudichaud, Voy. Uranie 458. 1829, syn. nov. Delissea sect. Rollandia (Gaudichaud) Baillon, Hist. PI. 8: 364. 1885. TYPE: Rollandia lanceo- lata Gaudichaud (selected by Rock (1919: 363)). Macrochilus C. Presl, Prodr. Monogr. Lobel. 47. 1836. TYPE: Macrochilus superbus (Chamisso) C. Presl (based on Lobelia superba Chamisso). Shrubs, treelets, or trees, rarely lianas, terrestrial or rarely epiphytic; stems unbranched or sparingly branched, erect or ascending, unarmed, muricate, or aculeate (more so in juveniles), with numerous helically arranged leaf scars near apex; pith solid or chambered; latex white, yellow, or tan, viscous. Leaves alternate, petiolate or sessile, glabrous or pubescent, sometimes muricate or aculeate, es¬ pecially on the petiole; margin entire, callose-toothed, or pinnately lobed, cleft, parted, or divided (more so in juveniles), the lobes or segments then entire, callose-toothed, lobed, or divided. Inflorescences ax¬ illary, racemose, (3-)5-25(-40)-flowered, the ra- chis sometimes obsolete and the raceme thus ap¬ pearing subumbellate. Flowers epigynous, resupinate, protandrous. Calyx synsepalous; tube adnate to the ovary, forming a hypanthium, obconic or obovoid, rarely ovoid, oblong, cylindrical, or depressed ob¬ ovoid; lobes 5, valvate, persistent, dentiform, tri¬ angular, or oblong, rarely linear, distinct or rarely connate, much shorter to somewhat longer that the hypanthium, rarely as long as the corolla. Corolla zygomorphic, bilabiate or unilabiate, white or various shades of magenta, purple, or pink, rarely greenish or yellowish, sometimes longitudinally striped, gla¬ brous or pubescent, rarely muricate; tube suberect, curved, arcuate, or sigmoid, dorsally cleft to about the middle; lobes 5, valvate, spreading, deflexed, or rarely erect. Stamens 5, syngenesious, included or exserted; filaments connate, free from the corolla or dorsally adnate for %-% its length; anthers con¬ nate, dithecal, opening introrsely by longitudinal slits, the 3 dorsal ones a little longer than the 2 ventral ones, the latter or rarely all 5 with apical tufts of white hairs; pollen tricolporate, prolate, ellipsoidal. Ovary 2-loculed; placentae large, axile; style slen- Volume 3, Number 4 1993 Lammers et al. Merger of Cyanea and Rollandia 439 der, terete, with a ring of stiff white hairs near the apex; stigma 2-lobed, the lobes appressed and non- receptive as the style grows through the anther tube, pushing out pollen, after which the stigmas spread and become receptive. Fruit a yellow, orange, or purple berry. Seeds numerous, small, dark brown to black, shiny, smooth or microscopically foveate- reticulate. Chromosome number n = 14 (Lammers, 1988). Cyanea crispa (Gaudichaud) Lammers, Givnish & Sytsma, comb. nov. Basionym: Rollandia cris¬ pa Gaudichaud, Voy. Uranie 459. 1829. Lo¬ belia crispa (Gaudichaud) Endlicher, Ann. Wiener Mus. Naturgesch. 1: 170. 1836; not Graham, Edinburgh New Philos. J. 1: 173. 1826. Cyanea rollandia A. Gray, Proc. Amer. Acad. Arts 5: 149. 1861, nom. illeg. TYPE: ‘Ties Sandwich.,” Gaudichaud s.n. (holotype, P). Based on Gaudichaud’s itinerary (St. John & Fit- comb, 1983) and the known range of the species (Lammers, 1990), the holotype probably was col¬ lected on 0‘ahu, in the Ko‘olau Mountains above Honolulu, during 26 29 August 1819. Cyanea huinboldtiana (Gaudichaud) Lammers, Givnish & Sytsma, comb. nov. Basionym: Rol¬ landia humboldtiana Gaudichaud, Voy. Bon- ite, Bot., pi. 76. 1842. TYPE: “lies Sand¬ wich.,” Oct. 1836, Gaudichaud s.n. (lectotype, selected here, W). Both Rock (1919) and St. John (1940) failed to locate type material of this species among Gaudi¬ chaud’s specimens at P; Lammers likewise was un¬ able to find any during a visit in September 1987. However, he did discover one sheet ot original ma¬ terial at W, which is here designated as the lecto¬ type. Based on Gaudichaud’s itinerary (Lasegue, 1845) and the known range of the species (Lam¬ mers, 1990), the type probably was collected on O'ahu, in the Ko‘olau Mountains above Honolulu. Cyanea koolauensis Lammers, Givnish & Syts¬ ma, nom. nov. Replaced name: Rollandia an- gustifolia (Hillebrand) Rock, Bull. Torrey Bot. Club 45: 136. 1918; not Cyanea angustifolia (Chamisso) Hillebrand, FI. Hawaiian Isl. 253. 1888. Basionym of replaced name: Rollandia longiflora var. angustifolia Hillebrand, FI. Ha¬ waiian Isl. 246. 1888. TYPE: Hawaii. O'ahu: Palolo Valley, Hillebrand s.n. (lectotype, se¬ lected here, BISH). Only one other syntype, a sheet at GH labeled “Oahu from Manoa Eastward,” has been seen by Lammers. A specimen at B cited by Rock (1918, 1919) could not be located; presumably, it was among those destroyed when the herbarium burned in 1943 (Hiepko, 1987). Cyanea lanceolata (Gaudichaud) Lammers, Giv¬ nish & Sytsma, comb. nov. Basionym: Rollan¬ dia lanceolata Gaudichaud, Voy. Uranie 458. 1829. Lobelia lanceolata (Gaudichaud) Hook¬ er & Arnott, Bot. Beechey Voy. 88. 1832; not Hooker & Arnott, Bot. Beechey Voy. 301. 1838; nor Kuntze, Revis. Gen. PI. 378. 1891. Lobelia rollandia Gaudichaud ex I). Dietrich, Syn. PI. 1: 728. 1839, nom. illeg. Delissea lanceolata (Gaudichaud) A. Gray, Proc. Amer. Acad. Arts 5: 147. 1861. TYPE: “lies Sand¬ wich.,” Gaudichaud s.n. (holotype, P; isotype, G). Based on Gaudichaud’s itinerary (St. John & Tit- comb, 1983) and the known range of the species (Lammers, 1990), the type probably was collected on 0‘ahu, in the Ko‘olau Mountains above Honolulu, during 26-29 August 1819. Cyanea lanceolata subsp. calycina (Chamisso) Lammers, Givnish & Sytsma, comb. nov. Bas¬ ionym: Lobelia calycina Chamisso, Linnaea 8: 222. 1833. Rollandia calycina (Chamisso) G. Don, Gen. Hist. 3: 699. 1834. Delissea ca¬ lycina (Chamisso) C. Presl, Prodr. Monogr. Lobel. 47. 1836. Rollandia lanceolata suhsp. calycina (Chamisso) Lammers, Syst. Bot. 13: 507. 1988. TYPE: Hawaii. “0 Wahu” [i.e., 0‘ahu]: Chamisso s.n. (holotype, LE). Cyanea longiflora (Wawra) Lammers, Givnish & Sytsma, comb. nov. Basionym: Rollandia lon¬ giflora Wawra, Flora 56: 44. 1873. TYPE: Hawaii. 0‘ahu: leg. Hillebrand, If awra 2285 (holotype, W). Cyanea parvifolia (C. Forbes) Lammers, Givnish & Sytsma, comb. nov. Basionym: Rollandia parvifolia C. F orbes, Occas. Pap. Bernice Pau- ahi Bishop Mus. 5: 10. 1911. TYPE: Hawaii. Kaua‘i: Wai‘oli Valley, 1 Aug. 1909, Forbes 103. K (holotype, BISH). Cyanea purpurellifolia (Rock) Lammers, Giv¬ nish & Sytsma, comb. nov. Basionym: Rollan¬ dia purpurellifolia Rock, Coll. Hawaii Publ. Bull. 2: 44. 1913. TYPE: Hawaii. 0‘ahu: Hau‘ula ridge, Punalu‘u Mts., 2,000 ft., Aug. 440 Novon 1911, Rock 8821 (holotype, BISH; isotypes, BISH, GH, K, W). Cyanea st.-johnii (llosaka) Lammers, Givnish & Sytsma, comb. nov. Basionym: Rollandia st.- johnii Hosaka, Occas. Pap. Bernice Pauahi Bishop Mus. 11(13): 14. 1935. TYPE: Hawaii. 0‘ahu: Ko‘olau Range, Waipi‘o, Klpapa Gulch, wet windswept main divide, 2,800 ft., 6 Aug. 1933, Hosaka 1 159 (holotype, BISH). Acknowledgments. Fieldwork by TGL was suit- ported by National Science Foundation (NSF) Dis¬ sertation Research Grant BSR-83 13285 (T. F. Stuessy, principal investigator) and a Sigma \i Grant- in-Aid of Research. Travel by TGL to American and European herbaria, respectively, was made possible by an Herbarium Travel Award from the American Society of Plant Taxonomists and a Graduate Stu¬ dent Alumni Research Award from The Ohio State University. The staff's of B, BISH, G, GH, k. LE, P, PTBG, and W are acknowledged for the loan of specimens and for courtesy extended to TGL during visits to their institutions. Field studies and molecular systematic analyses by IJG and KJS were supported by a grant from the National Geographic Society, and by NSF grants BSR-9007293 and BSR-9020055; additional logis¬ tic support was provided by the National Tropical Botanical Garden. TJG and KJS extend their heart¬ felt thanks to S. Anderson, L. Cuddihy, T. Flynn, R. Hobdy, J. Jacobi, I). Lorence, A. Medeiros, E. Misaki, J. Obata, S. Perlman, and K. Woods, who accompanied them in the field and helped locate many rare species of Hawaiian lobelioids. Literature Cited Baillon, H. 1885. Lobelieae. Pp. 362-367 in Histoire des Plantes, vol. 8. Librairie Hatchette et Cie., Paris. Bentham, G. & J. D. Hooker. 1876. Genera Plantarum, vol. 2, part 2. Reeve, London. Candolle, A. L. P. P. de. 1839. Lobeliaceae. Pp. 339- 413, 784-786 in A. P. de Candolle, Prodromus Systematis Naturalis Regni Vegetabilis, vol. 7. Treut- tel & Wiirtz, Paris. Dietrich, D. 1839. Synopsis Plantarum, vol. 1. Voigt, Weimar. Endlicher, S. 1836. Bemerkungen fiber die Flora der Sfidseeinseln. Ann. Wiener Mus. Naturgesch. 1: 1 29 190. Gaudichaud, C. 1826. Botanique. In L. de Freycinet, Voyage autour du Monde, Entrepris par Ordre du Roi . . . Execute sur les Corvettes de S. M. l’Uranie et la Physicienne, pendant les Annees 1817, 1818, 1819 et 1820. Pillet-aine, Paris. Givnish, T. J., K. J. Sytsma, J. F. Smith & W. J. Hahn. In press. Molecular evolution, adaptive radiation, and geographic speciation in Cyanea (Campanula- ceae), the largest plant genus endemic to Hawai'i. In W. L. Wagner & V. A. Funk (editors), Patterns of Speciation and Biogeography in the Hawaiian Biota. Smithsonian Institution Press, Washington. Gray, A. 1861. Notes on Lobeliaceae, Goodeniaceae, &c. of the collections of the U.S. South Pacific Ex¬ ploring Expedition. Proc. Amer. Acad. Arts 5: 146- 152. Hiepko, P. 1987. The collections of the Botanical Mu¬ seum Berlin-Dahlem (B) and their history. Englera 7: 219-252. Hillebrand, W. 1888. Flora of the Hawaiian Islands: A Description of their Phanerogams and Vascular Cryptogams. Williams & Norgate, London. Lammers, T. G. 1988. Chromosome numbers and their systematic implications in Hawaiian Lobelioideae (Campanulaceae). Amer. J. Bot. 75: 1130-1134. - . 1990. Campanulaceae. Pp. 420-489 in W. L. Wagner, D. R. Herbst & S. H. Sohmer, Manual of the Flowering Plants of Hawai‘i. Univ. Hawai'i Press, Honolulu. - . 1992a. Two new combinations in the endemic Hawaiian genus Cyanea (Campanulaceae: Lobelioi¬ deae). Novon 2: 129-131. - . 1992b. New combinations for Asian Cam¬ panulaceae. Bot. Bull. Acad. Sin. 33: 285-287. - . 1992c. Systematics and biogeography of the Campanulaceae of Taiwan. Pp. 43-61 in C.-I Peng (editor). Phytogeography and Botanical Inventory of Taiwan. Institute of Botany, Academia Sinica, Taipei. - & D. H. Lorence. 1993. A new species of Cyanea (Campanulaceae: Lobelioideae) from Kaua'i, and the resurrection of C. remyi, Novon 3: 431- 436. Lasegue, A. 1845. Musee Botanique de M. Benjamin Delessert. Notices sur les Collections de Plantes et la Bibliotheque. Librairie de Fortin, Masson et Cie., Paris. Presl, C. B. 1836. Prodromus Monographiae Lobeli- acearum. Theophilus Haase, Prague. Pukui, M. K., S. H. Elbert & E. T. Mookini. 1974. Place Names of Hawaii, rev. ed. Univ. Hawai'i Press, Honolulu. Rock, J. F. 1918. New species of Hawaiian plants. Bull. Torrey Bot. Club 45: 133-139 + pi. 6. - . 1919. A monographic study of the Hawaiian species of the tribe Lobelioideae family Campanula¬ ceae. Mem. Bernice Pauahi Bishop Mus. 7(2): i-xvi, 1-395. St. John, H. 1940. Ophioglossum, Rollandia, and Scaevola. Hawaiian plant studies 9. Occas. Pap. Bernice Pauahi Bishop Mus. 15: 351-359. - . 1969. Types of sections in Clermontia, Cy¬ anea, and Delissea (Lobeliaceae). Taxon 18: 483. - . 1987. Enlargement of Delissea (Lobeliaceae). Hawaiian plant studies 138. Phytologia 63: 79-90. - & M. Titcomb. 1983. The vegetation of the Sandwich Islands as seen by Charles Gaudichaud in 1819. A translation, with notes, of Gaudichaud’s “lies Sandwich." Bishop Mus. Occas. Pap. 25(9): 1-16. Schonland, S. 1889. Campanulaceae. Pp. 40-70 in A. Engler, Die Natfirlichen Pflanzenfamilien, Teil IV, Abteilung 5. Wilhelm Engelmann, Leipzig. Stone, B. C. 1967. A review of the endemic genera of Hawaiian plants. Bot. Rev. 33: 216-259. von Post, T. & O. Kuntze. 1903. Lexikon Generum Volume 3, Number 4 1993 Lammers et al. Merger of Cyanea and Rollandia 441 Phanerogamarum. Deutsche Verlags-Anstalt, Stutt- gart. Wagner, W. I,., D. R. Herbst & S. H. Sohrrier. 1990. Manual of the Flowering Plants of Hawai'i. Univ. Hawai'i Press, Honolulu. Wimmer, F. E. 1943. Campanulaceae-Lobelioideae. I. Teil. Pp. i-vi, 1-260 in R. Mansfeld (editor). Das Pflanzenreich, Teil IV, Abteilung 276b. Wilhelm En- gelmann, Leipzig. New Series and Species of Lepanthes (Orchidaceae) from Ecuador Carlyle A. Luer Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, LJ.S.A. Abstract. Three new series in Lepanthes sect. Lepanthes are described, and sixteen new species of Lepanthes are described from Ecuador. Prior to 1983, 30 species of Lepanthes Swartz had been attributed to Ecuador, mostly by Lindley, Reichenbach, and Schlechter from collections most¬ ly by Jameson and Sodiro. Other better known col¬ lectors never bothered to collect a Lepanthes. To¬ day, 295 species are known from Ecuador, but many of these have not yet been published. In order to handle this large number of basically similar species, series based on key-characters are employed. Some individuals of a few species fall between series. Therefore, a few species may be listed in more than one series. The following 16 previously undescribed species are all members of section Lepanthes. Fourteen are in Lepanthes subsect. Elongatae (Luer, 1987) and two are in Lepanthes subsect. Lepanthes. Four series, two in each subsection, are recognized. Lepanthes subsect. Lepanthes ser. Lepanthes is characterized by a congested inflorescence that re¬ mains shorter than the leaf. It contains by far the greatest number of species in the genus. Lepanthes subsect. Lepanthes ser. Filamentosae is character¬ ized by a densely flowered inflorescence that is borne beyond the leaf by an elongated peduncle. Lepanthes subsect. Elongatae ser. Elongatae is characterized by a “lax” or “loose” (distantly flow¬ ered) inflorescence that is longer, or becomes longer than the leaf. Lepanthes subsect. Elongatae ser. Breves is characterized by a loose inflorescence that remains shorter than the leaf. In the Pleurothalli- dinae a distantly flowered inflorescence is charac¬ terized by the exposed portion of the rachis between floral bracts being longer than the pedicels. Lepanthes ser. Lepanthes. TYPE: Epidendrum ovale Swartz, Prodr. 125, 1788 = Lepanthes ovalis (Swartz) Fawcett & Rendle. Lepanthes ser. Filamentosae Luer, series nov. TYPE: Lepanthes Jilamentosa Luer & Hirtz, Novon 3: ???, 1993. Racemus congestus quam folio longiore. Lepanthes ser. Elongatae Luer, series nov. TYPE: Lepanthes elongata Luer & Hirtz, Lindleyana 2: 99, 1987. Racemus laxus quam folio longiore. Lepanthes ser. Breves Luer, series nov. TYPE: Lepanthes monoptera Lindley, J. Bot. 1: 10, 1834. Racemus laxus quam folio breviore. The vast majority of the species of Lepanthes from Ecuador that have come to my attention since 1978 have been discovered by Alexander C. Hirtz, a mining engineer in Quito, Ecuador. Not only does he have the enthusiasm to hunt them and an un¬ canny ability to find them, but he knows them by name, habitat, and locality, far better than I. He has also had the perseverance to photograph nearly every one. For these reasons I do not consider him merely the collector. I am pleased to include his name with mine in the following new descriptions. Lepanthes andreettae Luer, sp. nov. TYPE: Ec¬ uador. Morona-Santiago: without locality, col¬ lected by A. Andreetta, probably in the vicinity E of Paute, ca. 1 ,800 m, flowered in cultivation at Paute by A. Andreetta, C. Luer 13855 (ho- lotype, MO). Figure 1. Planta parva; inflorescentia racemosa laxa folio elliptico multilongiore; Boris sepalis purpureis late luteomarginatis, sepalo dorsali synsepaloque anguste ovatis acuminatis, synsepalo supra medium inciso, petalis transverse bilobis, lobis triangularibus, superiore ter majore, labelli laminis duabus anguste ovatis acuminatis, corpore angusto, ap- pendice digitiformi pubescenti. Plant small, epiphytic, caespitose; roots slender. Ramicauls slender, erect, 5-8 mm long, enclosed by 2-3 loose, microscopically ciliate, lepanthiform sheaths. Leaf erect, coriaceous, elliptical, obtuse, 12-16 mm long, 6-8 mm wide, the base cuneate into a petiole 2-3 mm long. Inflorescence a loose, distichous, successively several-to many-flowered ra¬ ceme up to 9 cm long, borne by a slender peduncle 15-20 mm long; floral bract 2 mm long; pedicel 1.5-2 mm long; ovary 1 mm long; sepals purple Novon 3: 442-454. 1993. Volume 3, Number 4 1993 Luer Lepanthes from Ecuador 443 Figures 1-4. — 1 (Top left). Lepanthes andreettae Luer. — 2 (Top right). Lepanthes campodostele Luer & Hirtz. — 3 (Bottom left). Lepanthes corkyae Luer & Hirtz. — 4 (Bottom right). Lepanthes delhierroi Luer & Hirtz. 444 Novon with hroad, yellow margins, the dorsal sepal narrowly ovate, acute, acuminate, 7 mm long, 2 mm wide, connate to the lateral sepals for 0.5 mm, the lateral sepals microscopically denticulate, narrowly ovate, acute, acuminate, oblique, 8 mm long, 3 mm wide together, connate 3 mm, the apices in apposition; petals pale rose, microscopically pubescent, trans¬ versely bilobed, 0.75 mm long, 2.6 mm wide, the lobes triangular, the upper lobe larger; lip pale rose, bilaminate, the laminae narrowly ovate, 2 mm long, the bases rounded, the apices acute, long-acuminate, microscopically pubescent, the connectives narrow, from the base of the blades, the body narrow, con¬ nate to the base of the column, the appendix pu¬ bescent, narrowly linear, incurved in the natural position and in contact with the stigma; column 1 mm long, the anther dorsal, the stigma ventral. Etymology. Named in honor of Father Angel Andreetta, an Italian-born Salesian Padre at Paute, Ecuador, who discovered this species. This species was collected by Father Andreetta, probably east of Paute, and cultivated by him at the mission near Paute. It is distinguished by the small habit and a successively flowered raceme much lon¬ ger than the leaves. The flowers are comparatively large hut narrow with acuminate sepals, the laterals in apposition. The blades of the lip are very narrowly ovate with the connectives arising at the base. The appendix is narrow and curved up to he intimately associated with the stigma. Lepanthes campodostele I.uer & Hirtz, sp. nov. TYPE: Ecuador. Zamora-Chinchipe: Cordil¬ lera del Condor, epiphytic in cloud forest E of Los Encuentros, 1,550 m, 18 May 1988, C. Luer, A. Hirtz, If’. Flores, A. Andreetta & If . Teague 13472 (holotype, MO). Figure 2. Planta parva caespitosa; inflorescentia racemosa laxa foliis orbicularibus longiore; floris sepalis ovatis longicilia- tis, petalis pubescentibus transverse lobatis oblongis, la- bello pubescenti bilobo, lobis oblongis erectis, appendice externa pubescenti, columna grandi cylindrica apice trun- cato bicornuto. Plant very small, epiphytic, caespitose; roots com¬ paratively fleshy. Ramicauls erect, slender, 12-15 mm long, enclosed by 4 long-ciliate, lepanthiform sheaths. Leaf erect, coriaceous, orbicular, 10 mm long, 10 mm wide, the rounded base contracted into a petiole 1.5 mm long. Inflorescence a loose, dis¬ tichous, successively few-flowered raceme up to 30 mm long including the filiform peduncle 1 5 mm long, sparsely pubescent; floral bracts 1.5 mm long, spiculate; pedicels 1 .5 mm long; ovary 1 .5 mm long; sepals carinate-spicidate, the margins erose-ciliate, the dorsal sepal translucent, suffused with purple, ovate, deeply concave, acuminate, acute, 5 mm long, 4 mm wide unexpanded, connate to the lateral sepals for 1 mm, the lateral sepals purple, ovate, oblique acute, 6.5 mm long, 2.25 mm wide, connate 3.5 mm, deeply concave medial to the midveins, convex lateral to the midveins, convex along the medial margins in apposition; petals green pubes¬ cent, transversely bilobed, 0.5 mm long, 3 mm wide, the lobes subequal, oblong with rounded ends, the apex of the upper lobe incurved; lip green, ciliate, bilobed, the lobes oblong with rounded apices, 1 mm long, erect to either side of the column, the body cleft anteriorly, connate to the base of the column, the appendix external, oblong, recurved, pubescent; column comparatively large, cylindrical, 2 mm long, the apex truncate and flat with a pair of erect angles on the upper margin, the clinandrium on the lower half of the apical surface, separated from the sub- apical stigma by the rostellar flap along the lower margin. Etymology. From the Greek campodes, “like a caterpillar,” and stele, “the column,” referring to the unusual shape of the column. The flower of this species is large for the tiny plant with orbicular leaves. The sepals are carinate spiculate externally with ciliate-erose margins. The dorsal sepal is deeply concave, while the lateral sepals are concave centrally between convex mar¬ gins. The convex medial margins are in apposition except basally, where the lateral sepals are connate and deeply concave. This unusual configuration is seen in the Bolivian L. nycteris Luer & Vasquez. Most unusual is the comparatively large column. The oblong lobes of the lip are held erect to either side. The column is cylindrical with the apex flat. Two erect angles are present on the upper margin of the flat apical end. The clinandrium with the anther occupies most of the lower half of the end. The stigma is located along the lower margin. Al¬ though unrelated, the apex of the column is remi¬ niscent of Salpistele Dressier. Lepanthes corkyae Luer & Hirtz, sp. nov. TYPE: Ecuador. Carchi: epiphytic in cloud forest be¬ tween Tulcan and Maldonado, 2,300 m, 2 Apr. 1984, C. Luer, S. Dalstrom & T. Hdijer 9934 (holotype, MO). Figure 3. Species haec Lepanthis schizicis Luer similis, sed se¬ palis glabris, synsepalo concavo cum apicibus brevibus acutis differt. Plant small, epiphytic, caespitose; roots slender. Ramicauls erect, slender, 20-30 mm long, enclosed Volume 3, Number 4 1993 Luer Lepanthes from Ecuador 445 by 5-6 close, minutely ciliate, lepanthiform sheaths. Leaf erect, coriaceous, elliptical, obtuse, 10 12 mm long, 5-8 mm wide, the base cuneate into a petiole 1 mm long. Inflorescence a loose, flexuous, succes¬ sively several -flowered raceme up to 6 cm long, including the slender peduncle 1-2 cm long; floral bract 1 mm long; pedicel 2-3 mm long; ovary 1.5 mm long; sepals orange to red-brown, carinate, gla¬ brous, the dorsal sepal ovate, obtuse, concave, 5 mm long, 3.5 mm wide unexpanded, the apex abruptly acuminate into a slender tail 2 3 mm long, the lateral sepals 11-12 mm long including the slender tails 5-6 mm long, the blades connate 4.5 mm into an oblong, bifid lamina 6 mm long, 4 mm wide, the free portions acute, contracted into non- approximate tails; petals yellow-orange to brown, transversely bilobed, 0.8 mm long, 4 mm wide, the upper lobe oblong with the apex rounded, the lower lobe narrowly triangular with the apex acute, lightly recurved; lip yellow-orange to brown, bilaminate, the blades oblong, lightly concave, glabrous, 1 .5 mm long, with the apices subtruncate, the connec¬ tives cuneate, the body narrow, connate to the col¬ umn above the base, the appendix small, triangular, concave, with an uncinate, apical segment; column 1 mm long, the anther dorsal, the stigma ventral. Etymology. Named for Corky, Skeezix’s sister, a character from Frank King’s Gasoline Alley. This species is very similar to L. schizix, and for years was considered to be the same. However, a recent collection and re-examination of pickled (low¬ ers reveal subtle differences. The sepals of L. cork- yae are not ciliate; the synsepal is proportionately shorter and broadly concave with short, acute apices contracted into longer tails. Lepanthes corkyae oc¬ curs on the western slopes of northern Ecuador in the provinces of Carchi and Imbabura. Lepanthes schizix occurs on the western slopes of Pichincha. Paratypes. ECUADOK. Carchi: cloud forest between Tulcan and Maldonado, 2,050 m, 2 Apr. 1984, C. Luer , S. Dalstrom & T. Hoijer 9919 (MO), 2,300 rn, 17 Mar. 1991, C. Luer , J. Luer , A. Hirtz, X. Hirtz & J. del Hierro 15147 (MO). Imbabura: Selva Alegre, 1,400 m, May 1 989, A. Hirtz & X. Hirtz 4250 (MO). Lepanthes delhierroi Luer & Ilirtz, sp. nov. TYPE: Ecuador. Napo: epiphytic in cloud lor- est between El Carmelo and La Bonita, 2,400 m, Aug. 1990, A. Hirtz, X. Hirtz, J. del Hierro & F. Sarrniento 4965 (MO), C. Luer illustr. 15282. Figure 4. Planta parva caespitosa; racemo paucifloro foliis ellip- ticis crassis multilongiore; floris grandis sepalis late ovatis concavis obtusis acuminatis, petalis transverse bilobis, la- belli laminis duabus oblongis longitudinaliter callosis, con- nectivis cuneatis, appendice cupulata processum stigma- ticum amplectenti. Plant small, epiphytic, caespitose, roots slender. Ramicauls slender, erect, 2-4 cm long, enclosed by 5 7 minutely ciliate-scabrous, lepanthiform sheaths. Leaf erect, thickly coriaceous, ± suffused with pur¬ ple beneath, elliptical, subacute to obtuse, 17-28 mm long including a 2-4-mm-long petiole, 7-12 mm wide, the base cuneate into the petiole. Inflo¬ rescence a loose, flexible, successively few-to sev¬ eral-flowered raceme, up to 6 cm long including the slender peduncle 2-4 mm long; floral hract ciliate, 1 .5-2 mm long; pedicel 2-3 mm long; ovary cos¬ tate, 1 -2 mm long; sepals red, sometimes with yellow stripes along the veins, glabrous, the dorsal sepal ovate, broadly ovate to suborbicular, concave, the blade 8-10 mm long, 5-8 mm wide, unexpanded, connate to the lateral sepals for 1 mm, the apex subacute to obtuse, acuminate into a slender tail 3- 5 mm long, the lateral sepals connate 6 mm into an ovate to broadly ovate, bifid lamina, 10-11 mm long, 7-12 mm wide, with the subacute apices con¬ tracted into tails 2-4 mm long; petals red to orange- brown, minutely pubescent, transversely bilobed, 1 mm long, 4.5 mm wide, the upper lobe oblong, obtuse, the lower lobe smaller, narrowly triangular; lip red to orange-brown, bilaminate, the laminae oblong with obtuse ends, minutely pubescent, lon¬ gitudinally callous below the middle, 2 mm long, the connectives short, cuneate, from near the base, the body connate to the base of the column, with a membranous, ciliate, slinglike appendix with a knob¬ like apex that holds an elongate process from the stigma; column terete, 1 mm long, the anther apical, the stigma ventral with a process grasped by the appendix. Etymology. Named in honor of Juan del Hierro of Quito, Ecuador, co-discoverer of this species. This showy species is distinguished by a loose, flexible, successively few-flowered raceme much lon¬ ger than the elliptical leaf. The large, red flower with yellow veins is characterized by broad, concave sepals that are abruptly acuminate into short tails. The red, bilobed petals are not remarkable. The red blades of the lip are longitudinally callous toward the base. Most remarkable is the membranous, sling¬ like appendix with a knoblike apex that holds a long, curved process that descends from the stigma. A form with narrower sepals occurs farther south. Paratypes. ECUADOR. Morona-Santiago: epi¬ phytic in cloud forest, Valle del Paute, 2,200-2,400 m, collected by M. Portilla, flowered in cultivation by A. Andreetta at Paute, 16 May 1988, C. Luer 13366 (MO). 446 Novon Figures 5 8. — 5 (Top left). Lepanthes drymocharis l.uer & Hirtz. — 6 (Top right). Lepanthes echo Luer & Hirtz. — 7 (Bottom left). Lepanthes exogena Luer & Hirtz. — 8 (Bottom right). Lepanthes Jilamentosa Luer & Hirtz. Volume 3, Number 4 1993 Luer Lepanthes from Ecuador 447 Lepanthes drymocharis Luer & Hirtz, sp. nov. TYPE: Ecuador. Carchi: epiphytic in cloud for¬ est E of Maldonado, 2,300 m, 17 Mar. 1991, C. Luer, J. Luer, A. Hirtz, X. Hirtz & J. del Hierro 15152 (holotype, MO). Figure 5. Species haec Lepanthis schizicis Luer similis, sed fio- ribus duplomajoribus, sepali dorsalis vix concavi cauda recurva, synsepalo proportione breviore, et labelli laminis duabus lateribus deflexis differt. Plant small, epiphytic, caespitose; roots slender. Kamicauls erect, slender, 20-30 mm long, enclosed by 5-6 ciliate, lepanthiform sheaths. Leaf erect, coriaceous, elliptical, obtuse, 10-20 mm long, 6 9 mm wide, the base cuneate into a petiole 1 mm long. Inflorescence a loose, flexuous, successively several-flowered raceme up to 9 cm long, including the slender peduncle 2-3 cm long, 2 flowers often produced simultaneously; floral bract 1.5 mm long; pedicel 1.5 mm long; ovary 1 mm long; sepals red- brown, striped with yellow along the veins, carinate, minutely ciliate, the dorsal sepal broadly ohovate, obtuse, 8-9 mm long, 6.5 mm wide, the apex acu¬ minate into a slender, recurved tail 4-5 mm long, the lateral sepals 15 mm long including the slender tails 5-6 mm long, the blades connate 7 mm into an ohovate, bifid lamina 5 mm wide, the free portions acute, tapered into nonapproximate tails; petals yel¬ low, minutely pubescent, transversely bilohed, 1 mm long, 4.5 mm wide, the upper lobe oblong with the apex rounded, the lower lobe narrowly triangular; lip white, suffused with brown medially, bilaminate, the blades oblong, lightly concave on the upper surface, with deflexed sides, microscopically pubes¬ cent, 2 mm long, with the apices rounded, the con¬ nectives broad, short, the body broad, connate to the column at the base, the appendix pubescent, oblong, with an incurved, truncate apical segment; column 1 mm long, the anther dorsal, the stigma ventral. Etymology. From the Creek drymocharis, “a woodland grace,” referring to the beauty of the flowers. This handsome species is related to L. corkyae, with which it is sympatric, and L. schizix from farther south. Lepanthes drymocharis is distin¬ guished by the larger flowers with a broad, minimally concave dorsal sepal with a recurved tail. I he sepals are minutely ciliate. The blades of the lip differ from those of the two related species in the broad, deflexed sides. Lepanthes echo Luer & Hirtz, sp. nov. TYPE: Ecuador. Zamora-Chinchipe: epiphytic in for¬ est near Nambija, 1,300 m, Feb. 1987, A. Hirtz 3315 (holotype, MO), C. Luer illustr. 14712. Figure 6. Planta mediocris, inflorescentia cum racemo grandiffo- ro subcongesto disticho cum pedunculo gracili folio an- guste elliptico duplolongiore, sepalo dorsali synsepaloque triangularibus acutis leviter acuminatis, petalis transverse bilobis, lobo superiore ovato obliquo, lobo inferiore su- boblongo incurvato apice incrassato, labello bilobo, lobis flabellatis marginis incrassatis, appendice parva pubes- centi. Plant medium in size, epiphytic, caespitose; roots slender. Kamicauls slender, erect, 3-7 cm long, enclosed by 4-7 ciliate, lepanthiform sheaths. Leaf erect, coriaceous, narrowly elliptical, acute, 2.5- 4.5 cm long, 0.8-1 cm wide, the base cuneate into a petiole 2-4 mm long. Inflorescence a subcong¬ ested, distichous, successively few-flowered raceme up to 10 mm long, borne beyond the leaf by a slender peduncle 4 5 cm long; floral bract 1.5-2 mm long, lightly muriculate; pedicel 1 .5-2 mm long; ovary 2.5 mm long; sepals glabrous, the dorsal sepal triangular, acute, lightly acuminate, 17 mm long, 1 1 mm wide, connate to the lateral sepals for 4 mm, the lateral sepals connate 12 nun into a triangular synsepal 18 mm long, 11 mm wide, the apices in apposition, acute, lightly acuminate, free for 4 mm; petals minutely pubescent, transversely bilohed, 1 mm long, 4 mm wide, the upper lobe ovate, oblique, acute, the lower lobe suboblong, incurved near the middle with the incurved apical portion thickened, the lower lobe thrice longer than the upper lobe; lip red, bilohed, pubescent, the lobes flabellate with rounded, thickened margins as rudimentary blades, 1.9 mm long, 3.6 mm wide expanded, the lobes (connectives) cuneate to form the narrow body, con¬ nate to the base of the column, the appendix small, oblong, pubescent; column 1 .5 mm long, the anther dorsal, the stigma apical. Etymology. Named for the mythological nymph Echo, fhe large, gaping flower is reminiscent of /.. dalessandroi Luer. This species is known only from the type collec¬ tion by Alexander Hirtz. It is similar to L. dales¬ sandroi, known from nearby localities, but it is easily distinguished by the short, more or less congested, instead of loose, raceme (suhsect. Lepanthes) borne by a peduncle beyond the leaf (ser. Elongatae). fhe flower is large, with the triangular, noncaudate dor¬ sal sepal and synsepal spreading widely in opposite directions. The incurved, apical portion of the lower lobe of the petal is thickened. The blades of the lip consist merely of the thickened margins of fan¬ shaped lobes of the lip. 448 Novon Lepanthes exogena Luer & Hirtz, sp. nov. TYPE: Ecuador. Zamora Chinchipe: epiphytic in cloud forest S of Yangana above Valladolid, 2,450 m, 23 Mar. 1985, C. Luer, J. Luer, A. Hirtz & ff . Flores 10944 (holotype, MO). Figure 7. Planta parva; inflorescentia laxe multiflora folio late ovato multilongiore; floris sepalis acutis minute denticu- latis, petalis transverse bilobis, lobo superiore oblongo quain lobo inferiore obtuse triangulari majore, labelli lam- inis duabus anguste ovatis, connectivis brevibus cuneatis, corpore angusto, appendice stipitata pubescenti ex pagina externa corporis orienti. Plant small, epiphytic, caespitose; roots slender. Ramicauls erect, slender, 10-30 mm long, enclosed by 5-8 ciliate, lepanthiform sheaths with dilated ostia. Leaf erect, coriaceous, broadly ovate to or¬ bicular, 7-13 mm long, 7-12 mm wide, the apex subacute, obtuse to rounded, the base broadly cu- neate into a petiole 1.5-2 mm long. Inflorescence a loose, flexuous, successively several-flowered ra¬ ceme up to 12 cm long, including the slender pe¬ duncle 1.5 cm long; floral bract 1.25-2 mm long; pedicel 1 mm long; ovary 1 mm long; sepals dark red with narrow, green margins, carinate, minutely denticulate, the dorsal sepal ovate, acute, 4-5 mm long, 2.5-3.25 mm wide, connate to the lateral sepals for 0.5 mm, the lateral sepals ovate, acute, oblique, 4-5 mm long, 1.5-2 mm wide, connate 1 mm; petals orange, microscopically pubescent to glabrous, transversely bilobed, 0.5-1 mm long, 3- 3.5 mm wide, the upper lobe oblong, oblique, the apex obtuse, the lower lobe smaller, triangular, with the apex rounded; lip white or yellow, suffused with red medially, bilaminate, the blades ovate, micro¬ scopically pubescent to glabrous, 2 mm long, with the apices acute and bases rounded, the connectives short, cuneate, the body narrow, connate to the column at the base, the appendix ovoid, pubescent, stipitate, originating from the external surface of the body at the attachment to the column; column 2 mm long, the anther dorsal, the stigma ventral. Etymology. From the Latin exogenus, “formed on the outside,” referring to the position of the appendix. This little species is distinguished by the flexuous raceme that becomes much longer than the broadly ovate to orbicular leaves. The red sepals with thin, green margins are acute and minutely denticulate. The petals are bilobed and broad. The blades of the lip are narrowly ovate with short connectives that form a narrow body. The stipitate appendix origi¬ nates from the external surface. Pnratypes. ECUADOR. Zamora-Chinchipe : cloud forest S of Yangana above Valladolid, C. Luer, J. Luer, A. Hirtz & l r: Flores 10852 (MO). Lepanthes filamentosa Luer & Hirtz, sp. nov. TYPE: Ecuador. Esmeraldas: epiphytic in cloud forest W of Lita, 750 m, 18 Jan. 1987, C. Luer, J. Luer, C. //. Dodson, A. Hirtz, D. Benzing & D. Bermudes 12378 (holotype, MO). Figure 8. Planta minutissima caespitosa; inflorescentia racemosa subdensa folio elliptico microscopice ciliato longiore; floris sepalis ovatis apice longissime filamentosis, petalis bifur- catis apicibus filamentosis, labello bilobo, lobis auriculatis superne laminas indistinctas formantibus, inferne apicibus incurvatis, appendice nulla. Plant minute, epiphytic, caespitose; roots com¬ paratively thick. Ramicauls erect, slender, 4-10 mm long, enclosed by 3-5 microscopically ciliate, lepanthiform sheaths. Leaf erect, coriaceous, ellip¬ tical, 3-7 mm long, 2.5-5 mm wide, the apex obtuse to subacute, prominently apiculate, the margins mi¬ croscopically cellular-ciliate, the base cuneate into a petiole up to 1 mm long. Inflorescence a subdense, distichous, successively several-flowered raceme up to 7 mm long, borne by a filiform peduncle 10-18 mm long; floral bracts 1-1.5 mm long, muriculate; pedicels 1.5-2. 5 mm long; ovary 1 mm long, with the ribs cellular-papular; sepals yellow to white suf¬ fused with purple centrally, carinate, ovate, with the acute apices extremely long-attenuate, the dorsal sepal 10 17 mm long, 1.25-1.5 mm wide, connate to the lateral sepals for 0.5 mm, the lateral sepals oblique, 10-16 mm long, 1-1.25 mm wide, connate 0.5 mm; petals yellow, bifurcate 1 mm above the base, the divisions filiform, 3-5.5 mm long; lip yel¬ low, glabrous, bilobed, the lobes auricular, 1.25 mm long, with the half above the column thickened to form an ill-defined lamina, the half below the middle falcate with broadly rounded, incurved, overlapping apices, without an appendix, the body connate to the base of the column; column 1 mm long, the anther and the stigma apical. Etymology. From the Latin filamentosus, “fila¬ mentous,” referring to the long-acuminate sepals and lobes of the petals. Mature plants of this species are some of the smallest known in the genus. The flowers are also minute, but the lengths of the exceedingly long, filamentous tails of the sepals and the filamentous lobes of the petals create dimensions comparable to flowers of much larger species. Lepanthes filamentosa grows relatively frequent¬ ly in the wet, lowland forest of northwestern Ecuador where it is nevertheless difficult to find. It grows buried in thick layers of moss on small branches of trees. Sometimes only the minute, spiderlike flowers are visible at a distance of 2 or 3 cm from the hidden plant. Although the racemes are congested (subsect. Volume 3, Number 4 1993 Luer Lepanthes from Ecuador 449 Lepanthes), they are borne far beyond the leaves by hairlike peduncles (ser. Elongatae). Paratypes. ECUADOR. Esmeraldas: cloud forest W of Lita, color forms, C. Luer et al. 12377, 12408 (MO). Lepanthes grossiradix Luer & Hirtz, sp. nov. TYPE: Ecuador. El Oro: epiphytic in forest remnant W of Pacha, 2,250 m, 23 May 1988, C. Luer, A. Hirtz, IT. Flores, A. Andreetta & W. Teague 13613 (holotype, MO). Figure 9. Planta mediocris caespitosa; radicibus crassissimis; in- florescentia racemosa laxe multiflora foliis ellipticis mul- tilongiore; floris sepalis ovatis acutis sparse ciliatis, petalis transverse lobatis, lobo superiore oblongo, lobo inferiore triangulari, labelli laminis duabus pubescentibus lunatis, connectivis late cuneatis, corpore lato, appendice rninuta fissa. Plant medium in size, epiphytic, caespitose; roots purple, coarse, thick. Ramicauls erect, slender, 3- 6 cm long, enclosed by 6-8 ciliate, lepanthiform sheaths dilated at the ostia. Leaf erect, coriaceous, elliptical-ovate, acute, slightly acuminate, 2.5 4 cm long, 0.9- 1.2 cm wide, the base cuneate into a petiole 2-4 mm long. Inflorescence a loose, disti¬ chous, successively many-flowered raceme up to 6 cm long including the slender peduncle ca. 1 cm long; floral bracts 1.5 mm long; pedicels 1.5 mm long; ovary 0.5 mm long; sepals brown with yellow margins, sparsely ciliate, the dorsal sepal ovate, concave, acute, 3 mm long, 2 mm wide expanded, connate to the lateral sepals for 0.5 mm, the lateral sepals ovate, oblique, acute, 3 mm long, 1 .2 mm wide, 1 -veined, connate 1 mm; petals brown, mi¬ nutely pubescent, transversely bilobed, 0.5 mm long, 1 .9 mm wide, the upper lobe oblong, obtuse, the lower lobe triangular; lip brown, minutely pubescent, bilaminate, the laminae lunate with indistinct blades, 1.5 mm long, the connectives broadly cuneate, the body broad, connate to the base of the column, the appendix minute, oblong, cleft down the middle; column terete, 1.5 mm long, the anther dorsal, the stigma apical. Etymology'- From the Latin grossiradix, “a thick root,” referring to the thick, coarse, purplish roots. Most remarkable of this species are the roots that are extremely thick for the size of the plant. The racemes are loosely flowered, beginning much short¬ er than the leaf, but eventually lengthening to about twice the length of the leaf. The sepals are acute and sparsely ciliate, the laterals with only one vein. The lobes of the lip are lunate, surrounding the apex of the column with a dorsal anther and apical stigma. The appendix is a minute, oblong structure with a cleft down the center. Lepanthes illinizae Luer & Hirtz, sp. nov. TYPE: Ecuador. Pichincha: epiphytic in wet forest, W flank of Volcan Illiniza, Cerro Azul, 2,900 m, 23 Jan. 1987, C. Luer, J. Luer & A. Hirtz 12478 (holotype, MO). Figure 10. Species haec L. bilobae Findley affinis, sed sepalis denticulatis et petalorum lobis apice rotundatis differt. Plant medium in size, epiphytic, caespitose; roots slender. Ramicauls erect, stout, 5-12.5 cm long, enclosed by 10-14 dark, loose, ciliate, lepanthiform sheaths. Leaf erect, coriaceous, elliptical-ovate, acute, acuminate, 4-6.5 cm long, 1.5-2 cm wide, the base cuneate to rounded, contracted into a pet¬ iole ca. 4 mm long. Inflorescence an elongating, loose, distichous, lightly flexuous, successively many- flowered raceme up to 14 cm long including the pedicel 1.5-2. 5 cm long; floral bracts 3 mm long, pedicels 1.5 mm long; ovary 2 mm long; flowers yellow to orange-brown; sepals ovate, acute, lightly acuminate, carinate, denticulate, the dorsal sepal 7 mm long, 3.5 mm wide, connate to the lateral sepals for 1 mm, the lateral sepals oblique, 7 mm long, 2.25 mm wide, connate 2.5 mm; petals microscop¬ ically pubescent, transversely bilobed. 1 mm long, 3 mm wide, the upper lobe oblong with the end rounded, the lower lobe triangular with the end rounded, shorter than the upper lobe; lip micro¬ scopically pubescent, bilaminate, the blades narrow¬ ly oblong with rounded ends, 2.25 mm long, the connectives cuneate, the body narrow, connate to the base of the column, the appendix small, ovoid, pubescent; column 2 mm long, the anther dorsal, the stigma ventral. Etymology. Named for Volcan Illiniza, on which this species was discovered. This species is another in the hoard belonging to the L. biloba and L. elongata complexes. The loose¬ ly flowered racemes begin flowering shorter than the leaves, but eventually attain lengths exceeding the leaves several times. Lepanthes illinizae is distin¬ guished from the species included in the above com¬ plexes by the following combination of features: pe¬ duncles shorter than the leaves; sepals acute and lightly acuminate, glabrous but denticulate; lobes of the petals rounded; blades of the lip narrowly oblong, and the appendix small and ovoid. Lepanthes nematostele Luer, sp. nov. TYPE: Ecuador. Carchi: epiphytic in wet, mossy forest E of Maldonado, 1,900 m, 15 Feb. 1989, S. Dalstrdm & T. Hoijer 1234 (holotype, MO), C. Luer illustr. 14715. Figure 1 1. Planta parva; racemo laxo successive plurifloro folio elliptico ter longiore; floris sepalo dorsali synsepaloque 450 Novon Figures 9-12. —9 (Top left). Lepanthes grossiradix L.uer & Hirtz. —10 (Top right). Lepanthes illinizae Luer & Hirtz. — 11 (Bottom left). Lepanthes nematostele Luer & Hirtz. —12 (Bottom right). Lepanthes phrixothrix Luer & Hirtz. Volume 3, Number 4 1993 Luer Lepanthes from Ecuador 451 cristatis concavis ovoideis, petalis transverse bilobis, lobo inferiore longissime acurninato hirsuto, labelli minuti con- nectivis corporeque ad columnam circa apicem adnatis, laminis duabus arcuatis inter se agglutinatis apice rotun- datis ciliatis, appendice triangulari retrorsa, columna lon- gissima gracillima. Plant small, epiphytic, caespitose; roots slender. Ramicauls slender, erect, 3-5 cm long, enclosed by 7-8 long-ciliate, lepanthiform sheaths. Leaf erect, coriaceous, elliptical, subacute, 10-15 mm long ex¬ cluding the petiole 1-1.5 mm long, 8-10 mm wide, the base cuneate into the petiole. Inflorescence an erect, loose, successively several-flowered raceme up to 5 cm long including the slender peduncle 2- 2.5 cm long; floral bract 1.5 mm long; pedicel 1.5- 2 mm long; ovary 3.5 mm long, lightly ribbed; sepals dark red, concave, crested along the veins and mar¬ gins, the dorsal sepal ovoid, acute, 6 mm long, 2.5 mm wide unexpanded, connate to the lateral sepals for 1 mm, the lateral sepals connate 5 mm into an ovoid synsepal similar to the dorsal sepal, 6 mm long, 4 mm wide unexpanded, the apices free for less than 1 mm; petals transversely bilobed, long- pubescent on the back surface, 1 mm long, 5 mm wide, the upper lobe short, semicircular, the lower lobe triangular, acute, long-acuminate; lip bilami¬ nate, the laminae glabrous, arcuate, firmly agglu¬ tinated to each other below the middle over the column, 1.25 mm long, the apices everted, rounded, ciliate, the connectives and body short, adnate to the column near the apex, the appendix triangular, retrorse; column weak, proportionately very long and very slender, 3.5 mm long, the anther dorsal, the stigma ventral with a prominent, descending process. Etymology ■ From the Greek nematostele, “a threadlike column,” referring to the column. This remarkable species is characterized by the small vegetative habit with long-ciliate, lepanthiform sheaths; a loose inflorescence longer than the leaves; concave, crested sepals; petals with a long-acumi¬ nate lower lobe and arising from the shaft of the column; and a long, slender column hearing near the tip a tiny lip. The blades of the lip are adherent over the column. A prominent stigmatic process as large as the retrorse appendix descends from be¬ tween the blades. Lepanthes phrixothrix Luer & llirtz, sp. nov. TYPE: Ecuador. Morona-Santiago: Cordillera del Condor, epiphytic in forest E of Guisme, 1,650 m, 20 Jan. 1989, C. Luer , J. Luer , P. Jesup & A. Jesup 14014 (holotype, MO). Fig¬ ure 12. Planta parva; ramicaulium vaginis longipubescentibus; racemo laxo folio suborbiculato multilongiore; Boris sepalis cristatis, synsepalo cymbiformi, petalis transverse bilobis, lobo superiore triangulari, lobo inferiore majore longicau- dato, labelli laminis duabus oblongis, corpore latissimo ad columnae gracilis medium adnato, appendice triangulari pubescenti. Plant small, epiphytic, caespitose; roots slender. Ramicauls slender, erect, 1.5-6 cm long, enclosed by 5-8 densely and coarsely long-ciliate, lepanthi¬ form sheaths. Leaf erect, coriaceous, suborbicular, obtuse to rounded, 7-15 mm long, 7-11 mm wide, the rounded base contracted into a 1-mm-long pet¬ iole. Inflorescence an arching, loose, successively many-flowered raceme, up to 5 cm long including the slender peduncle 10 15 mm long; floral bract 0.5 mm long; pedicel 0.75 mm long; ovary 1 mm long, lightly ribbed; sepals costate-crested, the dorsal sepal red, triangular, acute, concave, 3 mm long, 1.5 mm wide, connate to the lateral sepals for 1 mm, the lateral sepals completely connate into an ovoid, boat-shaped synsepal, 3 mm long, 1.5 mm wide unexpanded, the apex narrowly obtuse; petals yellow, minutely pubescent, transversely bilobed, 0.5 mm long, 3 mm wide, the upper lobe triangular, the lower lobe oblong, the acute apex attenuate into an equally long, slender tail, the lower lobe much longer than the upper lobe; lip red, bilaminate, the laminae glabrous, oblong, 1 mm long, the apices rounded, the connectives broad, quadrate, forming a broad body, adnate to the middle of the slender column, the appendix triangular, protuberant and decurved, pubescent; column 1.5 mm long, the anther apical, the stigma ventral. Etymology. From the Greek phrixothrix, “with bristling hair,” referring to the lepanthiform sheaths. This little species is known only from the Cor- dillera del Condor. It is characterized by the small vegetative habit with long-ciliate, lepanthiform sheaths; a loose, successively flowered raceme even¬ tually reaching much longer than the small, round leaf; the costate-crested sepals, the laterals connate into a boat-shaped synsepal; the petals with the lower lobes long-tailed; the glabrous oblong blades of the lip with a pubescent, protruding, triangular appen¬ dix. Although the inflorescence is “loose” by defi¬ nition, it approaches being “dense.” Paratypes. ECUADOR. Morona-Santiago: F. of Chuchumbletza, 1,650 m, 21 May 1988, C. Luer, A. llirtz, W. Flores, W. Teague & A. Andreetta 13539 (MO). Lepanthes sigsigensis Luer & Hirtz, sp. nov. TYPE: Ecuador. Morona-Santiago: epiphytic in cloud forest E of the pass E of Sigsig, 2,700 m, 15 May 1988, C. Luer, A. llirtz, W. Elores, A. Andreetta & W. Teague 13350 (MO). Fig¬ ure 13. 452 Novon Planta mediocris caespitosa; inflorescentia racemosa laxissima disticha foliis ovatis multilongiore; floris sepalis ovatis acutis minute denticulatis, petalis transverse lobatis, iobis oblongis obtusis, lobo superiore longiore, labelli lam- inis duabus glabris anguste ovatis, connectivis cuneatis, appendice gracili sigmoidea minute pubescenti. Plant medium in size, epiphytic, caespitose; roots slender. Ramicauls erect, slender, 3-7.5 cm long, enclosed by 7-10 ciliate, lepanthiform sheaths di¬ lated at the ostia. Leaf erect, coriaceous, ovate, acute, slightly acuminate, 3-4 cm long, 1 .3-1.7 cm wide, the rounded base contracted into a petiole 2-3 mm long. Inflorescence a loose, distichous, suc¬ cessively several-flowered raceme up to 10 cm long including the slender peduncle 2 2.5 cm long; floral bracts 2.5-3 mm long, crested; pedicels 3. 5-4. 5 mm long; ovary alate, 1 mm long; sepals light yellow, minutely denticulate, the dorsal sepal with a broad, central, purple stripe, triangular-ovate, acute, 8 mm long, 4 mm wide, connate to the lateral sepals for 1.5 mm, the lateral sepals ovate, oblique, acute. 8 mm long, 3 mm wide, connate 2 mm; petals orange, suffused with purple medially, microscopically cel¬ lular-glandular, transversely bilobed, 1.25 mm long, 5.5 mm wide, the lobes oblong, obtuse, the upper lobe longer; lip red-orange, essentially glabrous, bi¬ laminate, the laminae narrowly ovate, with the apex narrowly rounded, 3 mm long, the connectives short¬ ly cuneate, the body narrow, connate to the base of the column, the appendix slender, sigmoid, mi¬ nutely pubescent; column terete, 2 mm long, the anther dorsal, the stigma ventral. Etymology. Named for the town of Sigsig, near the area where this species was discovered. Although superficially similar to other medium¬ sized species with relatively large flowers borne in a long, loose raceme (e.g., L. capitanea Reichen- bach f. and L. effusa Schlechter), this species is distinguished by the ovate leaves, minutely dentic¬ ulate sepals, simple petals and blades of the lip, hut with a slender, sigmoid appendix borne in the sinus. Lepanthes splendida Luer & Ilirtz, sp. nov. TYPE: Ecuador. Morona Santiago: epiphytic in wet forest along the new' road W of Macas toward Guamote, 1,900 m, 16 Jan. 1989, C. Luer , J. Luer , P. Jesup, A. Jesup, A. Ilirtz & S. Ortega 13937 (holotype, MO). Figure 14. Planta parva; racemo laxo simul florenti folio elliptico ter longiore; floris purpurei grandis sepalo dorsali trian- gulari acuminato, synsepalo ovato breviter bifurcato mul- timajore, petalis transverse bilobis, lobis ovoideis, lobo inferiore multiminore, labelli laminis duabus anguste fal- catis acutissimis, connectivis brevibus, appendice bipar¬ tita. Plant small, epiphytic, caespitose; roots slender. Ramicauls slender, erect, 10-15 mm long, enclosed by 3-4 minutely ciliate, lepanthiform sheaths. Leaf erect, coriaceous, elliptical, subacute, 17-22 mm long excluding the petiole 2 mm long, 6-8 mm wide, the base cuneate into the petiole. Inflorescence an erect, loose, simultaneously flowered raceme of 5- 6 large, red-purple flowers, up to 9.5 cm long in¬ cluding the slender peduncle 3 cm long; floral bract 2 mm long; pedicel 3-4 mm long; ovary 1 mm long, lightly ribbed; sepals red-purple, glabrous, the dorsal sepal triangular, acute, acuminate, 9.5 mm long, 3.5 mm wide, connate to the lateral sepals for 1 mm, the lateral sepals connate 8 mm into a broad lamina 13 mm long, 7 mm wide, the apices free for 3 mm, subacute, shortly apiculate; petals yellow- white, microscopically cellular pubescent, trans¬ versely bilobed, 1 mm long, 2.25 mm wide, the upper lobe ovoid, obtuse, the lower lobe triangular, obtuse, much smaller than the upper lobe; lip rose, bilaminate, the laminae glabrous, subfalcate, 1.25 mm long, the bases rounded, the apices narrowly acuminate, the connectives short, forming a narrow body with a two-parted appendix beneath the col¬ umn; column 1 .5 mm long, the anther apical, the stigma ventral. Etymology. From the Latin splendidus, “splen¬ did, elegant,” referring to the beauty of the species. This remarkable species is characterized by the small vegetative habit and an imposing, erect inflo¬ rescence of several large, simultaneous, purple flow¬ ers. Unfortunately, the only plant found failed to survive. It was discovered by Santiago Ortega in the forest along the road under construction between Macas and Guamote. The broad synsepal is considerably larger than the acute dorsal sepal. The lower lobe of the petals is minute. The blades of the lip are falcate, and the short, two-parted appendix is held between the blades beneath the column. Lepanthes synema Luer & Ilirtz, sp. nov. TYPE: Ecuador. Zamora-Chinchipe: epiphytic in elfin forest E of the pass E of Loja, 2,750 m, 21 Mar. 1985, C. Luer, J. Luer, A. Ilirtz <£• W. Flores 10724 (holotype, MO). Figure 15. Planta parvula; racemo laxo folio suborbiculato mul- tilongiore; floris sepalis obtusis abrupte caudatis, petalis transverse bilobis, labelli laminis duabus planis oblongis, corpore angusto appendice oblonga pubescenti munito. Volume 3, Number 4 1993 Luer Lepanthes from Ecuador 453 Figures 13 16. —13 (Top left). Lepanthes sigsigensis Luer & Hirtz. —14 (Top right). Lepanthes splendida Luer & Hirtz. —15 (Bottom left). Lepanthes synema Luer & Hirtz. —16 (Bottom right). Lepanthes vaginans Luer & Hirtz. 454 Novon Plant very small, epiphytic, caespitose; roots slen¬ der. Ramicauls slender, erect, 2-7 mm long, en¬ closed by 2-4 minutely ciliate, lepanthiform sheaths. Leaf erect, coriaceous, suborbicular, obtuse to rounded, 6-8 mm long, 4.5-6 mm wide, the round¬ ed base contracted into a 1-mm-long petiole. Inflo¬ rescence a loose, successively few-flowered raceme, up to 2 cm long including the slender peduncle 8 10 mm long; floral bract 1.5 mm long; pedicel 1.5 mm long; ovary 0.5 mm long; sepals yellow, sub- carinate, the dorsal sepal suffused with red centrally, suborbicular, 4.5 mm long, 4.5 mm wide, with the rounded apex abruptly contracted into a slender tail 2.5 mm long, connate to the lateral sepals for 1 mm, the lateral sepals oblong, oblique, connate 3 mm into an oblong, bifid lamina, 5 mm long, 5 mm wide together, the apices obtuse, contracted into slender tails 3 mm long; petals yellow, cellular-glan¬ dular, transversely bilobed, 0.75 mm long, 4 mm wide, the lobes subequal, oblong, narrowly obtuse; lip yellow, bilaminate, the laminae glabrous, oblong, 1 .8 mm long, the apices rounded, the connectives cuneate, forming a narrow body, connate to the base of the column, the appendix small, oblong, incurved, pubescent; column 1 mm long, the anther apical, the stigma ventral. Etymology. From the Greek synaimos, “a kins¬ man,” referring to the similarity to some other small species. This tiny twig-epiphyte of subparamo, elfin forests is closely related to several other minute, sympatric species, e.g., L. caudata Luer & Escobar, L.Jloresii Luer & Hirtz, L. jimburae Luer & Hirtz, L. par- adoxa Luer & Hirtz, and L. vermicularis. From all of them L. synema is distinguished by the smaller habit; round, abruptly caudate sepals; and flat, ob¬ long blades of the lip with a small, oblong appendix. Lepanthes vaginuns Luer & Hirtz, sp. nov. TYPE: Ecuador. Carchi: epiphytic in cloud forest E of Maldonado, 2,300 m, 17 Mar. 1991, C. Luer, J. Luer , A. Hirtz, X. Hirtz & J. del Hierro 1515 6 (holotype, MO). Figure 16. Planta mediocris; racemo laxo flexuoso folio elliptico breviore ad multilongiore; floris sepalis late ovatis ciliatis, lateralibus uninervibus, petalis minutis angustis, labello bilobo, lobis obcuneatis deflexis columnam vaginantibus. Plant medium in size, epiphytic, caespitose; roots slender. Ramicauls erect, slender, 5-8 cm long. enclosed by 8-11 microscopically scabrous, lepan¬ thiform sheaths with dilated ostia. Leaf erect, co¬ riaceous, elliptical, narrowly obtuse, 3-3.5 cm long, 1-1.3 mm wide, the base cuneate into a petiole ca. 3 mm long. Inflorescence a loose, flexuous, succes¬ sively several-to many-flowered raceme up to 9 cm long, including the slender peduncle ca. 1 cm long, 2 4 flowers often produced simultaneously; floral bracts 1.25 mm long; pedicel 2-2.5 mm long; ovary 0.75 mm long; sepals yellow, suffused with rose medially, carinate, coarsely ciliate, broadly ovate, subacute, lightly acuminate, the dorsal sepal 2- 2.5 mm long, 2 mm wide, 3-veined, adnate to the lateral sepals for 0.5 mm with the sides reflexed, the lateral sepals 2 2.5 mm long, 1.5 mm wide, 1 -veined, connate 0.5 mm; petals rose, cellular glandular, transversely bilobed, 0.25 mm long, 1.4 mm wide, the upper lobe narrowly oblong, obtuse, the lower lobe similar but much shorter; lip rose, bilobed, the lobes obliquely subtruncate, 1 mm long, surrounding the column, with cuneate connectives forming a broad body, adnate to the base of the column, the sinus narrow with a small, oblong, pubescent ap¬ pendix; column 1 mm long, the anther and the stigma apical. Etymology. From the Latin vaginans, “sheath¬ ing,” referring to the column covered by the lip. This species has features that seem to ally it with several diverse groups. The loose, flexuous, succes¬ sively flowered racemes begin flowering while much shorter than the leaf, but continue to flower until much exceeding the leaf. As many as four or five flowers are often produced simultaneously. The small sepals are coarsely dentate, and the laterals are one- veined. The petals are minute and slender. The lip is bilobed without any development of marginal blades. The lobes are obcuneate and sheathe the gynostenium. A rudimentary appendix is present in the sinus between the lobes. The lip suggests section Haplocheilus. Acknowledgment. I thank Dan H. Nicolson for constructive criticism. Literature Cited Luer, C. A. 1987. New Lepanthes from Ecuador — 4. Lindleyana 2: 95. A New Combination in Chonemorpha (Apocynaceae) David J . Middleton Department of Botany, Trinity College, Dublin 2, Ireland ABSTRACT. The new combination Chonemorpha verrucosa is made, and a key to the four species of Chonemorpha in Thailand is given. Chonemorpha verrucosa (Blume) D. J. Middle- ton, comb. nov. Basionym: Tabernaemontana verrucosa Blume, Bijdr. 1029. 1826. Rhyn- chodia verrucosa (Blume) Woodson, Suny- atsenia 3: 102. 1936. TYPE: Blume s.n. (ho- lotype, L; photograph in K). Chonemorpha was first described by G. Don in 1837, although many of the 13 species he described have since been referred to other genera (see Chat- terjee, 1947). The oldest name now referable to Chonemorpha is C. fragrans (Moon) Alston based on a drawing and description by Rheede (1689), which he called “Belutta-kaka-kodi,” and validated as Echites fragrans by Moon (1824) who referred to Rheede. This species encompasses many of the subsequently published names. Chonemorpha is a conserved name against Belutta-kak Adanson, 1763. Rhynchodia was first validly published in Ben- tham & Hooker (1876), although de Candolle (1844) and Miquel (1857) had previously published this genus under the names Rhyncospermum and Cer- cocoma , respectively. These are illegitimate later homonyms of Rhynchospermum Reinwardt (1828) and Cercocoma Wallich ex G. Don (1837). A num¬ ber of species have been described over the large range of the genus, but it appears that they are all referable to one variable species. Pichon (1950) placed both Chonemorpha and R h ynchodia in his subtribe Chonemorphinae of tribe Parsonsieae, while Ly (1986) described a new tribe, Anodendreae, containing both genera. Ly further suggested that Chonemorpha evolved from Rhyn¬ chodia. These two genera are extremely similar in leaf type, calyx and corolla structure, and stamen, disk, and gynoeciurn type. Because the only aspect separating the two genera is the smaller flowers in Rhynchodia , I believe they should be united. Chonemorpha appears to be most closely related to Epigynum and Trachelospermum. Kerr (1939) included two species, under the names C. grandieriana Pierre ex Spire and C. macrantha Pitard, in his list of the species in Thailand. These are synonyms of C. fragrans (Moon) Alston and C. griffithii J. D. Hooker, respectively. A key to the four species of Chonemorpha found in Thailand follows: Key to the Species of Chonemorviia in Thailand la. Calyx 3-4.5 mm long; corolla tube 4.3-7 mm long . C. verrucosa (Blume) D. J. Middleton lb. Calyx 1-3.9 cm long; corolla tube 2. 8-6. 2 cm long. 2a. Sepals connate for about half their length, sparsely puberulent or glabrous; corolla tube 4. 7-6. 2 cm long . . . C. fragrans (Moon) Alston 2b. Sepals connate for most of their length or almost free, densely puberulent or tomen- tose; corolla tube 2. 8-3. 9 cm long. 3a. Sepals almost free to base . . C. griffithii J. D. Hooker 3b. Sepals connate for most of their length . C. megacalyx Pierre ex Spire Acknowledgments. I thank the Royal Society, London, and the Royal Irish Academy, Dublin, for financial support for my work on Thai Apocynaceae, and John Parnell for comments on the manuscript. Literature Cited Bentham, G. & J. D. Hooker. 1876. Apocynaceae. In: Genera Plantarum 2: 681-728. Reeve, London. Candolle, A. L. P. de. 1844. Apocynaceae. In: Prod- romus 8: 317-489. Treuttel & Wiirtz, Paris. Chatterjee, D. 1947. The genus Chonemorpha G. Don (Apocynaceae). Kew Bull. 1947: 47-52. Don, G. 1837. Apocynaceae. In: A General History of the Dichlamydeous Plants 4: 69-105. Rivington, London. Kerr, A. F. G. 1939. Apocynaceae. In: Flora Siainensis Enumeratio 2: 422-476. Siam Society, Bangkok. Ly, T. D. 1986. Die Familie Apocynaceae Juss. in Vietnam. Feddes Repert. 97: 235-273, 405-466, 607-689. Miquel, F. A. W. 1857. Apocynaceae. In: Flora van nederlandsch Indie 2: 384-459. Van de Post, Am¬ sterdam. Moon, A. 1824. Catalogue of the Indigenous and Exotic Plants Growing in Ceylon. Wesleyan Mission, Colom¬ bo. Pichon, M. 1950. Classification des Apocynacees: XXV, Echitoidees. Memoires du Museum Nationale d’His- toire Naturelle, ser. B, Bot. 1: 1 174. Rheede, H. A. 1689. Hortus Indicus Malabaricus 9: 7, t. 5 & 6. Novon 3: 455. 1993. A New Species of Marathrum (Podostemaceae) from Jalisco, Mexico Alejandro l\ovelo R. Departamento de Botanica, Instituto de Biologia, Universidad Nacional Autonoma de Mexico, Mexico, D.F. 04510, Mexico C. Thomas Phi lb rick Department of Biological and Environmental Sciences, Western Connecticut State University, Danbury, Connecticut 06810, U.S.A. ABSTRACT. A new species of Podostemaceae, Mar¬ athrum rubrum, from Jalisco, Mexico, is described and illustrated. Capillaceous leaf divisions and red leaf color distinguish M. rubrum from other species. Resumen. Se describe e ilustra una especie nueva de la familia Podostemaceae ( Marathrum rubrum) del estado de Jalisco, Mexico. Las divisiones capi- liformes y el color rojizo de las hojas distinguen a M. rubrum de las otras especies. The pantropical Podostemaceae are the largest family of aquatic flowering plants. Little taxonomic work has been conducted on New World Podoste¬ maceae since Royen’s treatments (1951, 1953, 1954). Currently most species of Podostemaceae in the New World are poorly collected, taxonomic boundaries at both the specific and generic levels are unclear, and factors relating to the unusually high incidence of endemism (e.g., > 45% among New World species) are unresolved. Marathrum , one of the largest genera of Podo¬ stemaceae, is composed of 25 species occurring in tropical river rapids and waterfalls. The genus rang¬ es from the West Indies and Mexico through Central America, to northwestern portions of South America (Royen, 1951). During field studies aimed at clar¬ ifying the taxonomy of Podostemaceae in Mexico, collections were made from Jalisco that did not cor¬ respond to any described species. These collections serve as the basis for the description of a new species, Marathrum rubrum Novelo & Philbrick. Marathrum rubrum Novelo & Philbrick, sp. nov. TYPE: Mexico. Estado de Jalisco: municipio de Cabo Corrientes, Puente Los Horcones sobre el rio, a 27 km al S de Puerto Vallarta rumbo a Chamela, 6 Mar. 1991, Novelo & Philbrick 1003 (holotype, MEXU; isotype, RSA). Figure 1. Herbae caule longo, prostrato, applanato; lamina it- erum atque iterum pinnata, divisionibus ultimis capillifor- rnibus, 2-4.5 mm longis, 0.02-0.06 mm latis, rubescen- tibus, apice acutis. Flores solitarii, axillares, pedicellati; pedicelli apice expanso, cupulato, cupula 1.5-2 mm dia- metro. Tepala triangularia, filamentis alternantia. Stamina 7-9 in verticillo, libera, pistillum cingentia. Antherae 1 .5- 2.8 mm longae. Styli liberi, conici. Fructus 4.5-5(-6) mm longus, 2 valvis, quaque 3-costata. Aquatic herbs usually with a long prostrate and flattened stem up to 1 cm diam., strongly adhering to rocks. Leaves alternate, up to 20 cm long, petiole 2.5-9 cm long, cylindrical, up to 0.8 mm diam., with a broadened base; blade repeatedly pinnate, the later divisions capillaceous, 2-4.5 mm long, 0.02-0.06 mm wide, apex acute. Flowers her¬ maphroditic, actinomorphic, pedicellate, borne sin¬ gly, axillary, protected by a spathella; spathella 0.7 2 cm long, thin, clavate. Pedicels 3-5.5 cm long, elongating during anthesis, with an expanded apex forming a cuplike process around the capsule base, cup 1.5-2 mm diam., with irregular border. Tepals 0.5-0. 9 mm long, triangular, alternating with the stamens. Stamens 7-9, free, in a ring surrounding the pistil; filaments 2.3-4(-9) mm long, subulate, flattened, elongating during anthesis, deciduous; an¬ thers 1 .5-2.8 mm long, sagittate, basifixed, 2-celled, dehiscing longitudinally by 2 lateral slits. Ovary su¬ perior, ellipsoid, 2-locular; styles 2, 0.6-1. 5 mm long, free, conical, ovules numerous, placenta axile. Fruit 4.5-5(-6) mm long and 1.7-2. 2 mm diam., 2-locular capsule, suture margins thickened; valves 2, subequal, each 3-ribbed, one of them deciduous. Seeds 0.25-0.30 mm long, 0.12-0.18 mm wide, 574 (SD 317, N = 20) per capsule, obovoid. Pollen 14.5 /am (SD 0.8, N = 30) diam., tricolpate, tectum spinulose. Paratypes. MEXICO. Jalisco: Municipio de Cabo Corrientes, Rio Horcones, aprox. 17 km S de Puerto Vallarta rumbo a Chamela, 6 Feb. 1991, Novelo 979 (MEXU, RSA); Puente Los Horcones sobre el rio, a 27 km al S de Puerto Vallarta rumbo a Chamela, 6 Feb. Novon 3: 456-458. 1993. Volume 3, Number 4 1993 Novelo & Philbrick Mara thrum rubrum 457 Figure 1. Marathrum rubrum Novelo & Philbrick. Drawings based on the holotype. —A. General habit of plant with most of the capillar divisions of the largest leaf removed. — B. Leaf base with petiole. — C. Capillar divisions of leaf. — D. Young flowers projecting from between leaf bases; each flower is covered by a spathella. — E. Flower bud as the spathella is ruptured. — F. Flower projecting through the ruptured spathella. — G. Young flower prior to anther dehiscence and stigma receptivity; note tepals. — H. Young flower with three stamens (six have been removed). — I. Mature flower showing orientation of the styles and anthers at the time of dehiscence (three stamens have been removed). — J. Mature styles with receptive stigmas. — K. Cross section of ovary showing placenta and numerous ovules. — L. Capsule showing the expanded apex of the pedicel; the stamens have been shed. — M. Cross section of a capsule showing ribs, dehiscence points of the capsule valves, and the attachment points of the valves to the placenta. — N. Seed. 458 Novon 1991, Novelo 982(MEXU, RSA), 17 Mar. 1992, Novelo & Philbrick 1035 (MEXU). Additional observations. Marathrum rubrum grows submerged in the swift currents of river rapids, attached directly to granitic boulders of various size and sometimes to submerged branches of Salix. Plants have been collected only in the Horcones River, Jalisco, in areas of full sun. Phis species often occurs on the same rock with Vanroyenella plu- mosa Novelo & Philbrick (Novelo & Philbrick, 1993), Tristicha trifaria (Bory ex Willdenow) Sprengel, and Oserya coulteriana Tulasne. Marathrum rubrum belongs to subfamily Podos- temoideae sensu Royen (1951), tribe Podostemeae. This subfamily is characterized by minute, scalelike tepals, and flowers within a single sheathlike spa- thella. Podostemeae is the largest tribe of New World Podostemaceae and is characterized by flowers borne singly, fascicled, or in extra-axillary inflorescences, but not in spiciform monochasia. In addition to Marathrum rubrum, five other species of Marathrum occur in Mexico: M. elegans P. Royen, M. haenkeanum Engler, M. schiedean- um (Chamisso) Tulasne, M. tenue Liebmann, and M. trichophorum P. Royen. Three species (M. ele¬ gans, M. haenkeanum, M. schiedeanum ) share with M. rubrum the expanded apex of the pedicel and posses five or more stamens evenly spaced around the ovary. Marathrum tenue and M. trichophorum lack the expanded pedicel apex, and possess 2 3 stamens that arise on one side of the ovary. The leaves display the most distinctive features of Marathrum rubrum. It is the only species that has capillaceous leaf divisions, which range from 0.02 to 0.06 mm in diameter. In contrast, the leaf divisions of M. haenkeanum (0.075-0.125 mm), M. elegans (0.3- 1.3 mm), and M. schiedeanum (0.1 -0.4 mm) are larger in diameter. The leaves of M. rubrum are distinctively red on both abaxial and adaxial surfaces. In contrast, leaves of the other species of Marathrum in Mexico are green, sometimes with a reddish abaxial surface. Acknowledgments. We thank Fernando Chiang for providing the Latin description, Fernando Chiang, Ron Scogin, and Thomas Elias for helpful comments on the manuscript, and Albino Luna for drawing Figure 1. Support for this study was provided by the Intercambio Academico Office at the National LJniversity of Mexico, the National Geographic So¬ ciety, and a grant to CTP by the National Science Foundation. Literature Cited Novelo R., A. & C. T. Philbrick. 1993. Vanroyenella: A new genus of Podostemaceae from Jalisco, Mexico. Syst. Bot. 18: 64-67. Royen, P. van. 1951. The Podostemaceae of the New World. Part I. Meded. Bot. Mus. Herb. Rijks Univ. Utrecht. 107: 1-151. - . 1953. The Podostemaceae of the New World. II. Acta Bot. Neerl. 2: 1-20. - . 1954. The Podostemaceae of the New World. III. Acta Bot. Neerl. 3: 215-263. Notes on Cybianthus Subgenus Cybianthus (Myrsinaceae) in Southeastern Brazil John J. Pi poly, III Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. ABSTRACT. Cybianthus rupestris, a new species from the Planalto of Brazil, is described and illus¬ trated, and its phylogenetic relationships are dis¬ cussed. Cybianthus glaziovii Mez is placed in syn¬ onymy with C. coriaceus Martius; the former bi¬ nomial is leetotypified, the latter is neotypified, and a complete description is provided for the taxon. The neotropical genus Cybianthus Martius con¬ tains approximately 160 species in 10 subgenera (Pipoly, 1987). Cybianthus subg. Cybianthus con¬ tains 55 species, distributed across the Amazon Ba¬ sin, the Guayana Floristic Province (sensu Maguire, 1979), the Guianas, and central and southeastern Brazil. Members of the Grammadenia-Cybian- thus-Cybianthopsis clade (Pipoly, 1987) possess basifixed anthers with terminal pores, or with pores that open terminally, then for a short length down each side of the thecae. However, all species of subgenus Cybianthus may at once be recognized by the apparently epipetalous stamens, of which the tube is developmentally fused, or entirely adnate to, the corolla tube; the apically free portions of the filaments range from 1 mm to obsolete. Members of the subgenus are only locally common. My field observations indicate that the number of individuals per population is very low, usually less than 20 individuals per hectare in primary forest. I have seen precocious flowering and architectural reiter¬ ation phenomena (sensu Pipoly, 1992), along with sexual variation, in every population thus far ob¬ served. The sexual variation observed consists of monoecy, dioecy, and often polygamy in different populations of the same species. Like the majority of Cybianthus species, the flowers are perfect, but functionally unisexual. Historically, the group’s sys¬ tematic biology has been poorly understood owing to a lack of study in population dynamics. In prep¬ aration for a treatment of the Myrsinaceae for the Flora da Serra do Cipo, located in Minas Gerais, a remarkable new species was found from the nearby Ghapada dos Veadeiros, state of Goias, and is de¬ scribed herewith. Cybianthus rupestris Pipoly, sp. nov. TYPE: Bra¬ zil. Goias: Chapada dos Veadeiros, 14°00'S, 47°00'W, ca. 20 km W of Veadeiros [Alto Paraiso], 1,000 m, 10 Feb. 1966 (stam. fl), H. S. Irwin et al. 12841 (holotype, L’B; iso¬ types, LL-TEX, NY). Figure 1 . Propter ramulos dense stellato-tomentosos, laminam coriaceam revolutamque, corollae lobos suborbiculares dense prominenteque atro punctatos, C. coriaceo valde affinis, sed ab ea petiolis marginatis (non canaliculatis) 0.7-1 (nec 1.5-3) cm longis, perianthio membranaceo (non chartaceo), calyce inequilater (non equilater) diviso, lobis corollinis subapicale incisuris (non integerrimis) secus margines, glandulari-fimbriatos (nec glandulari-granulo- sos), antheris glabris (non minute glandulari-papillatis), denique stigmate capitato (non 4-lobato) praeclare distat. Shrub to small tree to 2 m tall, stems to 3 cm diam.; branchlets terete, 3.5-5 mm diam., densely ferrugineous floccose-tomentose, the tomentuin composed of stiff, arachnoid-stellate trichomes. Leaves coriaceous, the blades linear-oblanceolate to narrowly oblanceolate, (4-)4.5-9 cm long, 0.8 1.9 cm wide, apex acute, base cuneate, nitid and es¬ sentially glabrous above, pallid and densely ferru¬ gineous floccose-tomentose below along the midrib and margin, glabrescent, midrib impressed above, prominently raised below, secondary veins 10-15 pairs, the margin tightly inrolled-revolute, entire; petioles marginate, 0.7-1 cm long, basally pulvi- nate, glabrous above, densely tomentose below, gla¬ brescent. Staminate inflorescence: a simple, erect raceme, 5-7.5(-8) cm long, the peduncle, rachis and pedicels densely tomentose; peduncle 0.8-1 cm long; floral bracts linear-lanceolate, 1.4 1.8 mm long, apex attenuate, densely tomentose above and below, the margin erose, long glandular-ciliate; ped¬ icels cylindrical, thin, 2. 8-4. 6 mm long. Flowers nodding, 2.6 2.9 mm long, dull yellow-green; calyx membranaceous, cotyliform, 1.2- 1.6 mm long, the tube 0. 2-0.4 mm long, unequally divided, the lobes suborbicular, 0.7-0. 9 mm long, 0.7- 1.1 mm wide, apically rounded, hyaline, densely and prominently black punctate-lineate, the margin erose, mostly sub- apically notched, densely glandular-ciliate, the cilia Novon 3: 459-462. 1993. 460 Novon ,0 - ’°°'c^ \«>*f £ it: \%O0 - 1 mm Figure 1. Cybianthus rupestris Pipoly. — A. Habit, showing revolute, tomentose leaves. — B. Staminate flower, showing subapically notched corolla lobes. — C. Staminate calyx and pistillode, showing erose calyx lobe margins and glandular-granulose pistillode. — D. Pistillate flower, showing discoid stigma and translucent glandular-lepidote ovary. A C drawn from isotype. D drawn from H. S. Irwin 32765. long, 3-5-celled, ferrugineous; corolla membrana¬ ceous, subrotate, 2. 5-2. 8 mm long, the tube 0.7- 0.9 mm long, the lobes suborbicular, asymmetric, 1.8-2. 2 mm long and wide, apex broadly obtuse, subapically notched, glabrous without, densely glan¬ dular-granulose throughout within, prominently black punctate medially, the margins glandular-fimbriate, the fimbria ferrugineous; stamens 1.8-2. 2 mm long, the tube inconspicuous, membranaceous, adnate to corolla tube, 0.3-0. 4 mm long, the filaments flat, 0.5 0.6 mm long, densely glandular-granulose, the anthers very widely ovate to suborbicular, 0.6-0. 7 mm long, 0.7-0. 8 mm wide, apex broadly rounded, base cordate, glabrous, the connective prominently punctate dorsally; pistillode conic, ca. 0.9 mm long, 0.4 mm diam., densely and prominently black punc¬ tate and glandular-granulose. Pistillate inflores¬ cence: a simple, stiff, erect raceme, ( 1 ,5-)2-3.5(- 4) cm long, the peduncle, rachis, and pedicels densely tomentose; peduncle 0.4-0. 6 cm long; floral bracts Volume 3, Number 4 1993 Pipoly Cybianthus subgenus Cybianthus 461 linear-lanceolate, 1-1.4 mm long, apex attenuate, densely tomentose above and below, the margin erose, long glandular-ciliate; pedicels cylindrical, thin, 1.5- 1.8 mm long. Flowers nodding, 1.9-2. 3 mm long, dull yellow-green; calyx membranaceous, co- tyliform, 0.9- 1.1 mm long, the tube 0.1 -0.2 mm long, unequally divided, the lobes suborbicular to ovate, 0.7-0. 9 mm long, 0.7- 1 . 1 mm wide, apically rounded to subacute, hyaline, densely and promi¬ nently black punctate-lineate, the margin erose, mostly subapically notched, densely glandular-cili¬ ate, the cilia long, 3-5-celled, ferrugineous; corolla membranaceous, subrotate, 1 .8-2 mm long, the tube 0.7-0. 9 mm long, the lobes suborbicular, asym¬ metric, 1.2- 1.4 mm long and wide, apex broadly obtuse, subapically notched, glabrous without, densely glandular-granulose throughout within, prominently black punctate medially, the margins glandular-fim- briate, the fimbria ferrugineous; staminodes 1.3- 1.5 mm long, the tube inconspicuous, membrana¬ ceous, adnate to corolla tube, 0.6 0.7 mm long, the filaments flat, 0.2-0. 3 mm long, densely glan¬ dular-granulose, the anthers very widely ovate to suborbicular, 0.3-0. 4 mm long and wide, apex broadly rounded, base cordate, glabrous, the con¬ nective prominently punctate dorsally; pistil obtur- binate, 1.3- 1.5 mm long, the ovary 0.8-0. 9 mm long, 0.7-0. 9 mm wide, densely and prominently black punctate and glandular-granulose, the style 0.4-0. 5 mm long, the stigma capitate, flat, discoid, the placenta subglobose, apex apiculate, ovules 3, deeply imbedded in the placenta. Fruit subglobose, 5-7 mm long and in diam., exocarp thin, reddish brown when dried, inconspicuously pellucid punc¬ tate. Distribution. Endemic to the Chapada dos Vead- eiros, Goias, Brazil, at 1,000 1,600 m elevation. Ecology. Cybianthus rupestris grows in gallery forests along rivers, at the margin of cerrado for¬ mations. It appears to be locally common and forms a conspicuous element of the understory. Paratypes. BRAZIL. Goias: Chapada dos Veadeiros, 20 km N of Alto Paraiso, 1,250 rn, 20 Mar. 1966 (pist. fl, fr) H. S. Irwin et al. 32765 (NY, UB), 32765A (MO, NY, UB); ca. 1,600 m, 6 Mar. 1973 (fr), W. H. Anderson 6501 (IAN, NY, UB). Cybianthus rupestris is most closely related to the vicariant C. coriaceus Martius, but is easily recognized by its shorter, marginate petioles, un¬ equally divided calyx, subapically notched corolla lobes with glandular-fimbriate margins, glabrous an¬ thers, and capitate, discoid stigma. Upon study of Cybianthus glaziovii Mez, as the closest congener of C. rupestris, it became apparent that C. coriaceus is the earliest name for the taxon. Mez (1902) separated the two species based on an overlapping character state of acutish to acuminate vs. acute or obscurely acuminate leaf apex, and the prominence of the secondary venation below. How¬ ever, I have examined all material seen by Mez, except the holotype of C. coriaceus and can find no differences. Because the holotype of C. coriaceus has been lost, and no other element was mentioned in the protologue, a neotype is proposed lor C. coriaceus. Cybianthus glaziovii is lectotypified. A complete description, not previously available, is provided below. Cybianthus coriaceus Martius, Flora (Beibl.) 2(2): 19. 1841. TYPES: Brazil. San Sebastian [Hio de Janeiro]: C. Martius s.n. (holotype, M lost). NEOTYPE: Brazil. Hio de Janeiro: Hio de Ja¬ neiro, summit, Morro do Carangola, 22 Oct. 1882 (stain, fl), A. Glaziou 11048 (neotype, here designated, P; isoneotypes. BR, C, G, K, HB). Cybianthus glaziovii Mez in Engler, Pflanzenr. IV. 236 (9): 227. 1902, syn. nov. TYPE: Brazil. Rio de Janeiro: Serra dos Orgaos, 12 Aug. 1888 (stam. fl), A. Glaziou 17121 (lectotype, here designated, C; isolectotypes, K, P, RB). Shrub or small tree to 4 m tall; branchlets an- gulate, 3.5-5 mm diam., densely ferrugineous fur- furaceous stellate-tomentose. Leaves coriaceous, the blades elliptic to oblanceolate, (3-)6-8(- 1 1 .5) cm long, (1.5-)2-3.5 cm wide, apex acute to acumi¬ nate, submucronate, base cuneate, drying grayish, ferrugineous furfuraceous stellate-tomentose, gla- brescent, and scrobiculate above, densely stellate- tomentose below, the tomentum persistent on the midrib and near margin, midrib impressed above, prominently raised below, secondary veins 10-15 pairs, the margin opaque, highly revolute, entire; petioles canaliculate, ( 1 . 5— )2— 2 . 5(— 3) cm long, not pulvinate, densely tomentose, glabrescent. Stami- nate inflorescence: a simple, erect raceme, (3 )5- 8(— 11) cm long, the peduncle, rachis, and pedicels densely tomentose; peduncle 1-1.5 cm long; floral bracts linear, 1.7-2 mm long, 0.2— 0.3 mm wide, apex attenuate, densely tomentose above and below, the margin irregular, somewhat erose, glabrous; ped¬ icels cylindrical, thin, 4 5 mm long. Flowers nod¬ ding, 2. 4-2. 9 mm long, translucent green; calyx chartaceous, cotyliform, 1.4- 1.6 mm long, the tube 0.2-0. 3 mm long, equally divided, the lobes ovate- triangular, 1.1 1.2 mm long, 0.7 -0.8 mm wide, apex acute, densely and prominently black punctate, the margin hyaline, erose, densely glandular-ciliate; 462 Novon corolla chartaceous, subrotate, 2. 3-2. 8 mm long, the tube 0.5-0. 8 mm long, the lobes suborbicular, symmetric, 1. 8-2.1 mm long, 1 .2-1.5 mm wide, apex rounded, densely black punctate, glabrous without, densely glandular-granulose throughout within, the margin entire, glandular-granulose; sta¬ mens 1.3- 1.5 mm long, the tube inconspicuous, 0.5-0. 8 mm long, the filaments flat, 0.4-0. 5 mm long, proximally recurved, glabrous, the anthers su¬ borbicular, 0.5-0. 6 mm long and wide, apex obtuse, base rounded, densely and minutely rufous glan¬ dular-granulose dorsalty, the connective prominent¬ ly black punctate dorsally; pistillode subcylindrical, somewhat tapered apically, 0.7-1 mm long, 0.5- 0.6 mm diam., the stigma truncate, hollow, densely glandular-papillate. Pistillate inflorescence: a sim¬ ple, erect raceme, 3.5-5 cm long, the peduncle, rachis, and pedicels densely tomentose; peduncle 0.4-0. 7 cm long; floral bracts linear, 1.4- 1.8 mm long, 0.2-0. 3 mm wide, apex attenuate, densely tomentose above and below, the margin irregular, somewhat erose, glabrous; pedicels cylindrical, thin, 1 .6-2 mm long. Flowers nodding, 2. 6-2. 9 mm long, translucent green; calyx chartaceous, cotyliform, 1 .1-1 .3 mm long, the tube 0.2-0. 3 mm long, equal¬ ly divided, the lobes deflate to very widely ovate, 0.9- 1 mm long, 1-1.1 mm wide, apex acute, dense¬ ly and prominently black punctate, the margin hy¬ aline, erose, densely glandular-ciliate; corolla char¬ taceous, subrotate, 2. 5-2. 8 mm long, the tube 0.6- 0.8 mm long, the lobes suborbicular, symmetric, 1.8-2 mm long, 2-2.2 mm wide, apex rounded, densely black punctate, glabrous without, densely glandular-granulose throughout within, the margin entire, glandular-granulose; staminodes 1.2- 1.5 mm long, the tube inconspicuous, 0.6-0. 8 mm long, the filaments flat, 0.2-0. 3 mm long, proximally re¬ curved, glabrous, the anthers suborbicular, 0. 5-0.6 mm long and wide, apex obtuse, base rounded, densely and minutely rufous glandular-granulose dorsally, the connective prominently black punctate dorsally; pistil cylindrical, 1.6 1.8 mm long, 1-1.2 mm diam., the ovary densely translucent glandular- lepidote, gradually tapered to the 4-lobed stigma, the placenta deeply cupuliform, the ovules 3, deeply buried in the placenta. Fruit subglobose, 4-5 mm long and in diam., exocarp thin, inconspicuously pellucid punctate. Distribution. Southeastern Brazil, in the states of Minas Gerais and Rio de Janeiro. Representative specimens examined. BRAZIL. Mi¬ nas Gerais: Ouro Preto, near Camasinhos, 6 June 1905 (pist. fl), L. Damay 97 1 (G-BOIS, G-DC); Serra do Ouro Preto, Apr. 1892 (fr), E. Ule 2629 (B destroyed, F, HBG). Rio de Janeiro: Tijuca, 22 July 1864 (stam. fl), A. Glaziou 895 (BR, C, P — 2 sheets); Serra dos Orgaos, 12 Aug. 1888 (pist. fl, fr), A. Glaziou 17120 (P). Cybianthus coriaceus is known only from his¬ torical collections and may be extinct or exceedingly rare. It may be immediately separated from C. ru- pestris by the canaliculate petioles, chartaceous perianth, and minutely glandular-papillate anthers. Acknowledgments. Results presented here were facilitated by my appointment as a Research As¬ sociate of the Department of Botany, U.S. National Museum of Natural History, Smithsonian Institu¬ tion. I thank Peggy Duke for her skillful illustration, which was subsidized by a Smithsonian ROF Grant from the Office of the Assistant Secretary for Re¬ search, No. 1233F089. My work in neotropical plant species diversity is made possible by generous grants from the Andrew W. Mellon Foundation and the John D. and Catherine T. Mac Arthur Founda¬ tion. Literature Cited Maguire, B. 1979. Guayana, region of the Roraima Sandstone Formation. Pp. 223-238 in K. Larsen & L. B. Holm-Nielsen (editors), Tropical Botany. Ac¬ ademic Press, London. Mez, C. 1902. Myrsinaceae. In: A. Engler (editor), Das Pflanzenreich IV. 236: 1 437. Pipoly, J. 1987. A systematic revision of the genus Cybianthus subgenus Grammadenia (Myrsinaceae). Mem. New York Bot. Card. 43: 1-76. The Genus Geissanthus (Myrsinaceae) in the Choco Floristic Province John J. Pi poly. III Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. ABSTRACT. Studies on the plant diversity of Las Orquideas National Park in Antioquia, Colombia, along with preparation of a treatment of the Myr¬ sinaceae for Flora de Colombia, resulted in the discovery of four new taxa and two new combina¬ tions in Geissanthus. Examination of species known from the entire Western Cordillera of Colombia and the Choco Floristic Province, from Panama to Ec¬ uador, resulted in the discovery of 17 species from the area. An artificial key to the species of Geis¬ santhus from the Choco and Western Cordillera is presented, along with descriptions of four new spe¬ cies, G. callejasii, G. betancurii, G. francoae, and G. cogolloi. Three new combinations are made, G. longistamineus, G. per puncticulosus, and G. scrob- iculatus, and revised descriptions based on complete material are presented for G. longistamineus and G. per puncticulosus. Rf.SUMEN. Como resultado de los estudios de la fitodiversidad del Parque Nacional Natural “Las Or¬ quideas,” Antioquia, Colombia, y de actividades pre- liminares al tratamiento de Myrsinaceae para Flora de Colombia, se presentan cuatro taxa nuevos y tres nuevas combinaciones en el genero Geissant¬ hus. Al estudiar los especimenes de la Cordillera Occidental de Colombia y la provincia floristica cho- coana desde Panama hasta Ecuador, se reconocie- ron 17 especies en la zona. Se presenta una clave artificial para separarlos, se describen las cuatro especies nuevas, G. callejasii, G. betancurii, G. francoae, y G. cogolloi, y se publican las tres nuevas combinaciones, G. longistamineus, G. perpuncti- culosus, y G. scrobiculatus; ademas se proveen descripciones revisadas para G. longistamineus y G. per puncticulosus basadas en el estudio de ma¬ terial completo. Determination of collections from an ongoing flo¬ ristic study ol Parque Nacional Natural “Las Or¬ quideas,” jointly conducted by the Fundacion Jardin Botanico, “Joaquin Antonio Uribe” (JAUM) and the Missouri Botanical Garden (MO), concomitant with preparation of a taxonomic treatment of the Myr¬ sinaceae for Flora de Colombia, required a review of the species of Geissanthus J. D. Hooker from the Cordillera Occidental of Colombia and the ad¬ jacent Choco Floristic Province (sensu Gentry, 1982), which extends from the Darien of Panama southward to Los Rios, Ecuador. The genus Geissanthus was circumscribed by Bentham (1876) to include 1 1 taxa from the Andes, which he did not list, nor make the necessary com¬ binations. Bentham described the genus as polyga- mo-dioecious, with a 3-5-lobed calyx, infundibuli- form, 5-lobed corolla, linear-oblong anthers, and terminal panicles with spieate branches. The basis of his opinion that the taxa were polygamo-dioecious was the fact that fruit production per inflorescence branch is extremely low. After examining a number of species, I have concluded that there is a contin¬ uum from wholly staminate to wholly pistillate plants, but the most frequent condition is polygamous; the position of flowers for any particular sex on the inflorescence is apparently random. The flowers of each species appear to be monomorphic, the sta¬ minate ones of normal size but with hollow pistillodes, and the pistillate ones having staminodes slightly shorter, but normal -sized anthers, devoid of pollen. In his worldwide monograph of the family, Mez (1902) distinguished Geissanthus from the other genera of the tribe Myrsineae by its free stamens, dorsifixed anthers, and the calyx closed in bud, open¬ ing later into irregular lobes. In that treatment, 25 species were recognized, of which 17 were new. Subsequent to that treatment, miscellaneous new species were added by Mez (1905, 1920), Macbride (1934), and Cuatrecasas (1951). Agostini (1970) was the first worker since Mez to discuss generic delimitation among taxa assigned to Conomorpha A. DC. and Stylogyne A. DC. Agos¬ tini distinguished Geissanthus from Stylogyne and what was at that time the Cybianthus complex of genera (including Conomorpha), based on a com¬ bination of several characters, including: the ter¬ minal inflorescence; sessile or subsessile flowers; ca¬ lyx lobes at first closed, then rupturing into 2-8 unequal segments; the petals fused '/, or more their length; and capitate stigma. Using those criteria, he transferred three taxa from Conomorpha and one taxon from Stylogyne into Geissanthus. My studies have revealed that Geissanthus may be defined by its unique calyx, which is closed in bud and opens into 2-8 usually unequal lobes, co- Novon 3: 463-474. 1993. 464 Novon rolla with linear or oblong petals that are distally recurved at least 180° in anthesis, and the subver- satile or versatile anthers, which are latrorsely de¬ hiscent by wide or narrow longitudinal slits. Stylo- gyne may easily be separated from Geissanthus by its contorted corolla, with the lobe tips highly twisted in bud. Cybianthus may be separated from Geis¬ santhus by its axillary inflorescences, stamens con¬ nate by their filaments to form a tube, the staminal tube adnate to the corolla tube, and the glandular granules present at least at the junction of corolla lobes and tube. The present study has revealed 17 species of Geissanthus in the region, clearly the most diverse zone for the genus. Synonymies are given where appropriate, and complete descriptions are given only when new data have been revealed as a result of this study. Because there are perhaps an addi¬ tional six undescribed taxa known from fragmentary material, the present key is tentative until a broader survey can be conducted. Morphological terminology follows Lindley (1848) and Pipoly (1987, 1992). The 17 species of Geis¬ santhus occurring in the region may be distinguished by the following artificial key. Key to the Species of Geissaiythvs in the Cordillera Occidental and Adjacent Choc6 Floristic Province la. Leaves coriaceous, the margins revolute. 2a. Inflorescence longer than the leaves. 3a. Leaf blades 4-18.5 cm wide; inflorescence branches spicate. 4a. Leaf blades bullate; petioles deeply canaliculate. 5a. Branchlets and leaves rufous stellate-tomentose; leaf margin entire; pedicels 0.3-0. 5 mm long . G. callejasii Pipoly 5b. Branchlets furfuraceous-lepidote; leaf margin serrate; pedicels 2-3 mm long . . G. serrulatus (Willdenow ex Roerner & Schultes) Mez 4b. Leaf blades smooth; petioles marginate. 6a. Branchlets angulate; corolla membranaceous . G. betancurii Pipoly 6b. Branchlets terete; corolla coriaceous . G. occidentalis Cuatrecasas 3b. Leaf blades 1.5-3. 5 cm wide; inflorescence branches corymbose. 7a. Leaf margins serrate; petioles 1.5-2. 1 cm long; pedicels obsolete . G. goudotianus Mez 7b. Leaf margins entire; petioles 0.5-1 cm long; pedicels 1.5-4 mm long . G. quindiensis Mez 2b. Inflorescence shorter than or subequaling the leaves. 8a. Branchlets angulate; leaf blades decurrent to petiole base; petioles marginate; panicles columnar. 9a. Branchlets and inflorescence rachis glabrous; leaf blades nitid above; petioles 1-1.4 cm long . G. ecuadorensis Mez 9b. Branchlets and inflorescence densely and minutely furfuraceous-lepidote; leaf blades sordid above; petioles 1.7-3. 5 cm long . G. longistylus (Cuatrecasas) Agostini 8b. Branchlets terete; leaf blades only slightly decurrent on the petiole; petioles deeply canaliculate; panicles pyramidal. 10a. Leaf blades nitid above, the bases acute, the margins entire; pedicels obsolete; calyx obconic basally . G. kalbreyeri Mez 10b. Leaf blades sordid above, the bases truncate, the margins serrate; pedicels 2. 5-3. 5 mm long; calyx truncate basally . G. argutus (Kunth) Mez lb. Leaf blades membranaceous to chartaceous, the margins flat. 11a. Leaf blades densely and prominently black perpuncticulose, or punctate and punctate-lineate. 12a. Leaf blades oblong, elliptic, or narrowly oblanceolate, asymmetric, 2-5 cm wide. 13a. Leaf blades nitid above; floral bracts cucullate; pedicels 1.8-2. 5 mm long . . G. cestrifolius (Kunth) Mez 13b. Leaf blades sordid above; floral bracts flat; pedicels 2.5-3 mm long .... G. cogolloi Pipoly 12b. Leaf blades widely oblanceolate to obovate, symmetric, 6-18 cm wide. 14a. Petioles 0.5-1 cm long; calyx membranaceous, 2.7 3.2 mm long, black punctate-lineate . G. angustiflorus Cuatrecasas 14b. Petioles 1-1.5 cm long; calyx chartaceous, 1.2- 1.7 mm long, orange punctate . . G. perpuncticulosus (Lundell) Pipoly lib. Leaf blades pellucid punctate or punctate-lineate, or inconspicuously black punctate and punctate- lineate (the glands not raised). 15a. Branchlets quadrangular; leaf blades densely fufuraceous-lepidote below, the scales overlapping and appearing velutinous, the blade margin serrate; inflorescence from % to subequaling leaf length . G. francoae Pipoly 15b. Branchlets terete; leaf blades sparsely furfuraceous-lepidote, not appearing velutinous, the blade margin entire; inflorescence ca. '/- '/2 leaf length. 16a. Leaf blades scrobiculate above; calyx lobes hyaline, the margins regular, erose . . G. scrobiculatus (Cuatrecasas) Pipoly 16b. Leaf blades smooth above; calyx lobes opaque, the margins irregular, entire . . G. longistamineus (A. C. Smith) Pipoly Volume 3, Number 4 1993 Pipoly Geissanthus in the Choco 465 New Species of Geissanthus Geissanthus callejasii Pipoly, sp. nov. TYPE: Colombia. Antioquia: Mcpio. Jardm, Alto de Ventanas, 15 km SW ol Jardin, on road to Riosucio, 2,400-2,800 m, 05°30'N, 75°50'W, 9 June 1987 (infl. bud, fr), R. Callejas, O. Marulanda, F. Roldan & //. Correa 3916 (holotype, IIUA; isotypes, MO Nos. 3702266, 3702267). Propter laminam coriaceam ellipticam vel oblanceola- tam bullatamque, inflorescentiam quasi latam quam lon- giorem ramulam subspicatamque, lobos calycinos longi- tudine latitudos aequantes, G. bogotensi arete similis, sed ab ea ramulis angulatis (non teretibus) dense rufo- stellati-tomentosis (nec dense adprese lepidoti), petiolis profunde canaliculatis (non late marginatis) 1.2-2 (nec 2-3) cm longis, lobis calycinis suborbicularibus vel ovatis (nec triangularibus), corollis 4. 3-4. 6 (non 3.2-4) mm longis, denique lobis corollinis oblongis (nec ovato-trian- gularibus) statim separabilis. Free to 4 m tall; branchlets strongly ungulate, longitudinally ridged, 5-7 mm diam., densely rufous stellate-tomentose, persistent. Leaves alternate, the blades stiffly coriaceous, elliptic or rarely oblance- olate, (1 1 — ) 1 4.5-1 7(— 2 1 ) cm long, (4-)6-8.5 cm wide, apex acute, base acute, decurrent on the pet¬ iole, midrib and secondary veins deeply impressed above, decurrent to petiole base, prominently raised below, secondary veins 12-17 pairs, rufous stellate- tomentose above at first, early glabrescent, the re¬ sulting remnant pits from fallen trichomes giving a scrobiculate appearance, densely rufous stellate-to¬ mentose below, the hairs at times with all branches pointing upwards from leaf surface, inconspicuously brown punctate and punctate-lineate, the margin entire, somewhat revolute; petiole deeply canalicu¬ late, ( 1 . 2— ) 1 .4- 1 .8(-2) cm long, densely and per¬ sistently rufous stellate-tomentose above and below. Inflorescence terminal, pyramidal bipaniculate, 9- 17 cm long, 8-14 cm wide, the secondary branches 7-9 cm long toward base; peduncle, rachis, bracts, and pedicels densely rufous stellate-tomentose, the trichomes persistent; inflorescence bract unknown; peduncle obsolete to 2 cm; primary branch bracts unknown; secondary branch bracts membranaceous, oblong, 4-4.3 mm long, 2.8 —3 mm wide, apex trun¬ cate, densely stellate-tomentose below, glabrous, densely and prominently black punctate and punc¬ tate-lineate within, the margin stramineous, entire, glabrous; floral bracts membranaceous, 1.3— 1.5 mm long, 0.5 nun wide, apex obtuse, densely stellate- tomentose below, prominently black lineate within, the margin hyaline, glabrous, early caducous; ped¬ icels cylindrical, 0.3-0. 5 mm long. Flowers erect, green; fruiting calyx coriaceous, (2-)3-5-lobed, 2.4- 3.1 mm long, the tube 0.3-0. 7 mm long, extremely unequally divided, the lobes suborbicular to ovate, 2. 1-2.8 mm long, 2. 1-2.8 mm wide, apex acute, asymmetric, densely rufous stellate-tomentose me¬ dially without, glabrous within, inconspicuously black punctate, the margin scarious, hyaline, glabrous, undulate, and at times appearing erose; corolla mem¬ branaceous, campanulate, 4.3 4.6 mm long, the tube 1.2-1. 3 mm long, the lobes oblong, 3. 1-3.4 mm long, 1.4-1. 5 mm wide, apex obtuse, reflexed 1 80° at anthesis, sparsely and prominently red punc¬ tate and punctate-lineate, the margin irregular, somewhat erose; stamens and pistil unknown. Fruit globose, 5-7 mm long and in diam., fruit green; densely and prominently black punctate and punc¬ tate-lineate, the exocarp thin. Distribution. Geissanthus callejasii is known only from the type, from the Western Andean Cordillera, Alto de Ventanas, which is north of Cerro Cara- manta, in an area where the broken terrain yields north-south -facing slopes, at 2,400-2,800 m ele¬ vation. Ecology. This species is known to be occasional, near small watercourses in the montane and upper premontane wet forests. Etymology. It is with great pleasure that I ded¬ icate this species to Ricardo Callejas Posada, pro¬ fessor and curator of the herbarium of the Univ- ersidad de Antioquia; a friend, colleague, and preeminent authority on the morphogenesis, anat¬ omy, and systematics of the Piperaceae. With elliptic or rarely oblanceolate, coriaceous, and bullate leaves, panicles almost as wide as long, and calyx lobes as long as wide, Geissanthus cal¬ lejasii is most similar to G. bogotensis Mez. How¬ ever, the angulate and rufous stellate-tomentose branchlets, shorter, deeply canaliculate petioles, su¬ borbicular or ovate calyx lobes, and oblong corolla lobes easily set Geissanthus callejasii apart. Geissanthus betancurii Pipoly, sp. nov. TYPE: Colombia. Antioquia: Mcpio. Urrao, on trail to Paramo de Frontino, near Finca El Quince, 06°30'N, 76°10'W, 2,900 m, 18 Nov. 1988 (fl), G. McPherson, E. Roldan & J. Betancur 13106 (holotype, HIJA; isotypes, COL, MO, US). Figure 1 . Quoad ramulos subteretos, furfuraceo-lepidotos cortic- emque horizontaliter rimosem, laminas coriaceas obovatas ad apices acutas vel subacuminatas, petiolos marginatos, pedicelos obsoletos vel obconicos, inflorescentiam pyr- amidali-panicutam, G. fragranti valde affinis, sed ab ea ramulis 3.5-6 (non 7-10) mm diametris, laminis secus margines revolutis (non planis), pedicelis 0.2-0. 3 mm longis (nec obsoletis), calyces coriaceis (non chartaceis), 466 Novon antheris ovoideis (non oblongoideis) ad apices truncatis (nec apiculatis), connectivis epunctatis (nec atro-punctatis) praeclare distat. Tree to 12 m tall; branchlets subterete to an- gulate, 3.5-6 mm diam., sparsely ferrugineous, fur- furaceous-lepidote, the bark horizontally checked, glabrescent. Leaves alternate, the blades coriaceous, obovate, ( 1 0.5-)l 2.5- 1 6 cm long, (4.3-)5-6(-7.4) cm wide, apex acute to subacuminate, base cuneate, decurrent on the petiole, midrib impressed above, prominently raised below, secondary veins incon¬ spicuous, ( 1 8-)22-26(-30) pairs, smooth, sordid and glabrous above, pallid, moderately and minutely furfuraceous-lepidote below, sparsely and inconspic¬ uously reddish black punctate, the margin revolute, entire, glabrous; petioles marginate, ( 1 .3 )1 .5-2 cm long, glabrous above, densely and minutely ferru¬ gineous furfuraceous-lepidote below, glabrescent. Inflorescence terminal, a pyramidal pinnate panicle, (10— )13— 1 8(— 2 1 ) cm long, 10-19 cm wide at base, inflorescence bract unknown; peduncle, rachis, sec¬ ondary branches and pedicels moderately and mi¬ nutely rufous glandular-papillate, glabrescent; inflo¬ rescence bract unknown; peduncle ca. 1 cm long; secondary branch bracts unknown; floral bracts un¬ known; pedicels obsolete or obconic to 0.2-0. 3 mm long. Flowers erect, white; calyx coriaceous, obcon¬ ic, (2-3)4-5(-6)4obed, 3. 3-3. 6 mm long, the tube 2. 1- 2.4 mm long, very unequally divided, densely translucent and rufous furfuraceous-lepidote with¬ out, the lobes widely ovate to narrowly lanceolate, 2. 1- 2.2 mm long, 0.7- 1.8 mm wide, apex obtuse on wider lobes, acutish on narrower ones, densely and prominently brown punctate and orange punc- tate-lineate, the margin scarious, irregular, entire to subentire, glabrous; corolla carnose, campanulate, 4-5(-6)-lobed, 4. 9-5. 6 mm long, the tube 2. 1-2.5 mm long, the lobes ovate, 2.5-3. 1 mm long, 1.6- 1 .8 mm wide, apex acute, translucent, prominently brown punctate, glabrous, the margins irregular, entire, glabrous; stamens free, 4. 9-5. 6 mm long, the filaments membranaceous, flat, 3.8-4 mm long, hyaline, glabrous, not widened basally, inserted at corolla tube base, anthers versatile, ovoid, 1.7- 1.9 mm long, 1-1.2 mm wide, apex truncate, base deep¬ ly cordate, longitudinally dehiscent by wide latrorse longitudinal slits, the connective hyaline, epunctate; pistil obturbinate, 3. 8-4. 4 mm long, the ovary 1.6- 1.8 mm long, 0. 7-0.8 mm diam., the style 2-2.2 mm long, the stigma capitate, the placenta subgl- obose, 0.3-0. 4 long and in diam., apiculate, the ovules 4, buried in the placenta. Fruit depressed- globose, 8-12 mm long, 10-13 cm wide, reddish violet at maturity, densely and prominently black punctate and punctate-lineate, the exocarp thin. Distribution. Endemic to the northern sector of the Cordillera Occidental and facing western slopes of the Cordillera Central, in the Department of An- tioquia, Colombia, at 1,500-2,900 m elevation. Ecology. Geissanthus betancurii is a rare forest treelet, growing in areas near creekbeds, on steep slopes. Etymology. It is with great pleasure that I ded¬ icate this species to Julio Betancur (COL), prodigious field botanist and specialist in neotropical Bromeli- aceae. The following characters indicate that Geissan¬ thus betancurii is closely related to G. Jragrans Mez, of the Venezuelan Coastal Cordillera: subter¬ ete, furfuraceous-lepidote branches with horizontally Assuring bark; coriaceous, acute or subacuminate leaves with marginate petioles; and pyramidally pa¬ niculate inflorescences with short, obconic or absent pedicels. However, G. betancurii may be easily rec¬ ognized by its thinner branchlets, revolute leaves, short-pedicellate flowers with coriaceous calyx lobes, and ovoid anthers with truncate apices and epunc¬ tate connectives. Paratypes. COLOMBIA. Antioquia: Mcpio. San Luis, Piedra del Castrillon, Cordillera Central, Ladera Oriental, 06°4'30"N, 74°59'74"W, 1,500-1,700 m, 16 Sep. 1988 (fr), J. Betancur et al. 664 (COL, HUA, MO, US); Cordillera Occidental, 3-4 hours on foot SW of town, 06°01'N, 75°01'W, 1,500 m, 12 Aug. 1987 (fr), D. Daly & J. Betancur 5335 (COL, HUA, MO, US). Geissanthus francoae Pipoly, sp. nov. TYPE: Colombia. Bisaralda: Mcpio. de Pereira, El Ced- ral, old road to Salento, 2,200-2,300 m, 11 June 1989 (fl), G. Galeano, P. Franco, /V. Ladino, E. Eorero & A. Castillo 1955 (holo- type, COL; isotype, MO). Figure 2. Ob ramulos angulatos, adprese-ferrugineo-furfuraceo- lepidotos, folia pseudoverticillata, flores pedicellatos, caly- cem glandulari-papillatum, anthera lanceloideas, subver- satiles necnon pistillo obturbinato, G. perpuncticuloso valde arete affinis, sed ab ea laminis chartaceis (non mem- branaceis) subter squamis lepidotis superpositis sic a vel- veturn similis (nec dissite praeditis), secus margines ser- ratis (nec integerrimis), petiolis (2-)2.5-3 (non 1-1.5) cm longis, calyce membranaceo (non chartaceo) urceolato (nec cupuliforme), corolla membranacea (non chartacea) 3-3.2 (nec 2. 4-2. 7) mm longa, antheris ad apices emar- ginatis (non obtusis), denique in sylvas montanas nebu- losasque (non premontanas) incolens, facile distinguitur. Shrub or small tree to 4 m tall; branchlets qua¬ drangular, the angles suhalate, 7-12 mm diam., densely ferrugineous furfuraceous-lepidote, the scales appressed, the scale margins overlapping, persistent. Leaves pseudoverticillate, chartaceous, the blades oblanceolate, (29-)32-39(-54) cm long, (9-)10 1 3(— 1 6) cm wide, apex short-acuminate, the acu- Volume 3, Number 4 1993 Pipoly Geissanthus in the Choco 467 Figure 1. Geissanthus betancurii Pipoly. — A. Habit. — B. Detail of leaf margin. — C. Calyx. — D. Separated corolla. — E. Fruit, with calyx. AD, drawn from the type. E, drawn from Betancur 664. men 5 10 mm long, gradually tapering to an obtuse base, midrib impressed above, decurrent to petiole base, prominently raised below, secondary veins 19- 29 pairs, scrobiculate and perpuncticulose above, densely ferrugineous furturaceous-lepidote below, the scale margins overlapping and thus appearing ve- lutinous, the margin serrate, the teeth vascularized; petioles deeply canaliculate, (2-)2.5 3 cm long, ta- 468 Novon E CJ Figure 2. Geissanthus francoae Pipoly. — A. Habit. — B. Close up, leaf margin. — C. Calyx. — D. Separated corolla. — E. Fruit, with calyx. A, drawn from Devia 250. B D, drawn from the type. pered to the base, glabrous above, densely lepidote below, glabrescent. Inflorescence pyramidal and bi- pinnately paniculate, (10-) 19-50 cm long, second¬ ary branches (10-)14-25 cm long toward inflores¬ cence base; peduncle, rachis, bracts, and pedicels densely ferrugineous furfuraceous-lepidote, the scales overlapping, persistent; inflorescence bract un¬ known; peduncle 3-5 mm long; primary and sec- Volume 3, Number 4 1993 Pipoly Geissanthus in the Choco 469 ondary branch bracts unknown; floral bracts mem¬ branaceous, linear, 0.4-0. 6 mm long, ca. 0.1 mm wide, hyaline, apex subulate, prominently and dense¬ ly black punctate, the margin entire; pedicels cylin¬ drical, 0.4-0. 8 mm long. Flowers erect, greenish white; calyx membranaceous, urceolate, the base truncate, 4-5(-6)-merous, 1.3 1.8 mm long, the tube unequally divided, 0. 7-0.9 mm long, minutely glandular-papillate, the lobes erect, triangular to subdeltate, 0.6 0.9 mm long, 0.50.8 mm wide, apex acute to subacuminate, densely and promi¬ nently black punctate, glabrous, the margin entire, minutely glandular-ciliolate; corolla membrana¬ ceous, campanulate, 4-5-merous, 3-3.2 mm long, the tube 0.9-1 mm long, the lobes linear, 2. 1-2.2 mm long, 0.6-0. 7 mm wide, apex long-attenuate, reflexed 1 80° at anthesis, hyaline, densely and prom¬ inently black punctate and punctate-lineate, gla¬ brous, the margin entire; stamens free, 2. 1-2.2 mm long, the filaments membranaceous, flat, 1.5- 1.7 mm long, hyaline, glabrous, widened basally, in¬ serted at corolla tube base, anthers subversatile, lanceoloid, 0.9-1 mm long, 0.3-0. 4 mm wide, apex emarginate, base widely subcordate, longitudinally dehiscent by wide latrorse slits, the connective hy¬ aline, epunctate; pistil obturbinate, 2.1 2.4 mm long, the ovary 0.9-1. 1 mm long and in diam., conspic¬ uously black punctate, the style 1.1 -1.2 mm long, the stigma truncate, punctiform, the placenta broad¬ ly cupuliform, ovules 4, buried in the placenta. Fruit globose, 3-5 mm long and in diam., reddish brown at maturity, densely and prominently black punctate and punctate-lineate, the exocarp thin. Distribution. Geissanthus francoae is endemic to the western slopes of the Central and both slopes of the Western Andean Cordillera in the depart¬ ments of Antioquia, Valle del Cauca, and Risaralda, Colombia, at 1,830-2,300 m elevation. Ecology. This species is a common understory component of cloud forests in the region, growing along creek margins at the high water line. Label data from Fonnegra et al. 3202 indicate that Tri- gona bees were observed visiting the flowers, and the flowers are very fragrant. Etymology. It is with great pleasure that I ded¬ icate this species to Pilar Franco, professor at the Herbario Nacional Colombiano, Instituto Nacional de Ciencias, Universidad Nacional de Colombia, col¬ league and specialist in Euphorbiaceae, Moraceae, and Urticaceae. The angulate branchlets of this species, with ad- pressed ferrugineous furfuraceous-lepidote scales, pseudoverticillate leaves, pedicellate flowers with glandular-papillate calyces, subversatile, lanceoloid anthers and obturbinate pistil, clearly indicate a close relationship with Geissanthus perpuncticulosus. Flowever, Geissanthus francoae is easily recognized by its chartaceous, serrate leaves with adaxial scale margins so overlapped as to form a velvety tomen- tum, longer petioles, membranaceous perianth, ur¬ ceolate calyx, and emarginate anthers. It is inter¬ esting to note that the two apparent sister taxa are separated by habitat: Geissanthus perpuncticulosus occurs in premontane pluvial forests, while G. fran¬ coae is restricted to cloud forests. Paratypes. COLOMBIA. Antioquia: Mcpio. Tame- sis, Vereda Rio Frio, ca. 5°40'N, 75°43'W, 2,100 m, 9 Nov. 1989 (fl), R. Fonnegra et al. 3202 (COL, JAUM, MO, US). Valle del Cauea: Mcp io. Versalles, Bocatoma, NW of Versalles, Finca Maribel, 2,000 m, 1 1 Oct. 1983 (fr), W. Devia 250 (COL); Mcpio. Argelia, Vereda Las Brisas, 2,140 m, 21 Jan. 1983 (fr), P. Franco et al. 1696 (COL); Mcpio. Argelia, Vereda La Bella, Finca La Miranda, 1,830 m, 25 Jan. 1983 (fr), P. Franco et al. 1875 (COL). Geissanthus cogolloi Pipoly, sp. nov. TYPE: Co¬ lombia. Antioquia: Mcpio. IJrrao, Parque Na¬ cional Natural “Las Orquideas,” Vereda Calles, Sector Calles, right bank of Rio Calles and Quebrada “El Guaguo,” 06°32'N, 76°19'W, 1,430-1,490 m, 13 Feb. 1989 (fl), A. Cogollo, D. Cardenas & O. Alvarez 3955 (holotype, JAUM; isotypes, COL, FMB, MO). Figure 3. Species haec cum lamina oblonga, elliptica vel oblan- ceolata, asymmetrica glabraque, flores pedicellatos, ca- lycem cupuliformem, anthera epunctata G. lehmannii affinis, sed ab ea laminis membranaceis (non chartaceis), secus margines integerrimis (nec crenulatis), petiolis can- aliculatis (non marginatis) 1.2- 1.5 (nec 0.5-0. 7) cm lon- gis, pedicellis cylindricis (non obconicis) 2.5-3 (non 1.5- 2 ) mm longis, lobis calycinos oblongis (non ovato-trian- gularibus), antheris apiculatis (nec rotundatis) ad bases subcordatis (nec rotundatis) facile cognoscitur. Shrub or small tree to 12 m x 11.5 cm DBII; branchlets terete, 2.5-3 mm diam., red-violet, nitid, densely and prominently black punctate-lineate, gla¬ brous. Leaves alternate, membranaceous, bright red- violet in bud, the blades oblong, elliptic or rarely narrowly oblanceolate, (9.9 )12 l7(-20.5) cm long, (3 .2— )3 .5 — 5(— 6) cm wide, apex rostrate to long- acuminate, the acumen 0.8- 1.5 cm long, base broadly obtuse, slightly asymmetric, decurrent on the petiole, midrib deeply impressed above, promi¬ nently raised and black punctate-lineate below, sec¬ ondary veins 13-18, slightly raised above, promi¬ nently raised below, dark green above and light green below when fresh, nitid and glabrous above, pallid and minutely rufous translucent lepidote be¬ low, densely and prominently black punctate and 470 Novon Figure 3. Geissanthus cogolloi Pipoly. — A. Habit. — B. Close up, leaf margin. — C. Calyx. — D. Separated corolla. A, B, drawn from Pipoly et al. 16689. C, D, drawn from the type. punctate-lineate above and below, the margin hy- neate, glabrous. Inflorescence terminal, pyramidal aline, flat, entire, glabrous; petiole deeply canalic- paniculate, 8-1 0(-28) cm long, 8-1 0( 15) cm wide ulate, 1.2- 1.5 cm long, red-violet when fresh, densely basally, peduncle, rachis, pedicels, and calyces and prominently black punctate and punctate-li- densely rufous glandular-papillate; inflorescence bract Volume 3, Number 4 1993 Pipoly Geissanthus in the Choco 471 unknown; peduncle 3 -5(-7) mm long; secondary branch bracts membranaceous, ovate, 5-7 mm long, 3-4 mm wide, apex acute, base subtruncate, sessile, glabrous, densely and prominently black puntate and punctate-lineate, the margin entire, hyaline, gla¬ brous; floral bract early caducous, membranaceous, linear, 2-2.2 mm long, 0.2-0. 3 mm wide, apex attenuate, glabrous, densely and prominently black punctate and punctate-lineate, the margin entire, glabrous; pedicel cylindrical, 2.5-3 mm long. Flow¬ ers erect, white, 3-3.5 mm long; calyx membra¬ naceous, deeply cupuliform, 4-5(-6)-merous, 2.3- 2.5 mm long, the tube 0.5-0. 8 mm long, unequally divided, the lobes oblong, 1.5-1. 8 mm long, 1-1.1 mm wide, erect, apex obtuse, hyaline throughout, densely and prominently red and black punctate and punctate-lineate, sparsely rufous papillate, the mar¬ gin entire, glabrous; corolla membranaceous, cam- panulate, 5-merous, 2.9 3.1 mm long, the tube 0.4-0. 5 mm long, glabrous, the lobes oblong, 2.2- 2.6 mm long, 1-1.2 mm wide, reflexed 180° at anthesis, hyaline, apex obtuse, densely and promi¬ nently red punctate and punctate-lineate, the margin entire, glabrous; stamens 5, 2.6 3.1 mm long, the filaments free, slightly expanded basally, 1.5-1. 6 mm long, hyaline, epunctate, inserted at corolla tube base, the anthers subversatile, ovoid, 1.1 -1.3 mm long, 0. 5-0.6 mm wide, apex apiculate, base sub- cordate, latrorsely dehiscent by narrow longitudinal slits, the connective hyaline, epunctate; pistil ob- turbinate, 3.9 4.2 mm long, the ovary subglobose, 2-2.2 mm long, 1.8- 1.9 nun diam., densely and prominently red punctate, glabrous, the style 1.9- 2 mm long, epunctate, the stigma subcapitate, the placenta widely cupuliform, 0.7-0. 9 mm long, 1.1- 1.3 mm wide, the ovules 5, uniseriate, deeply im¬ mersed in the placenta. Fruit unknown. Distribution . Endemic to Las Orquideas National Park, western slope of the Western Andean Cor¬ dillera, Urrao Municipality, Antioquia, Colombia, 1,330-1,490 m elevation. Ecology. Geissanthus cogolloi grows on steep slopes in stream canyons throughout the upper pre- montane forest, just above flood line. Individuals of C. cogolloi are found in relatively low numbers (8- 10 per hectare), but are conspicuous because ol the red-violet color of the terminal bud margins, bran- chlet apices, and petioles. The premontane forests surveyed in Parque I, as Orquideas receive approx¬ imately 6,000 8,000 mm of precipitation annually and are frequently surrounded by clouds, despite their low elevation. Geissanthus perpuncticulosus (Lundell) Pipoly, Cybianthus poeppigii Mez, and C. venezuelanus Mez are common elements of the same forest. Etymology. Geissanthus cogolloi is named for Alvaro Cogollo Pacheco, scientific director of the Fundacion Jardin Botanico, “Joaquin Antonio l ribe" (JAUM), Medellin, Colombia. He is a specialist in the systematics of Colombian Myristicaceae, and floristics of the Choco Floristic Province. The oblong, elliptic, or oblanceolate and glabrous leaves of Geissanthus cogolloi, along with its ped¬ icellate flowers, cupuliform calyx, and epunctate anthers, indicate a close relationship with G. leh- mannii. However, the leaf blades with membrana¬ ceous texture and entire margins, canaliculate and longer petioles, long and cylindrical pedicels, oblong corolla lobes, and anthers with an apiculate apex and subcordate base easily distinguish Geissanthus cogolloi from G. lehmannii. Geissanthus lehmannii is reported from an area near Popayan (Department of Cauca) and may be found in the area, but as yet has not been collected. Paratypes. COLOMBIA. Antioquia: Mcpio. Urrao, Parque Nacional Natural Las Orquideas, Quebrada Hon¬ da, 06°29'N, 76°14'W, permanent montane rainforest plot, NW of the Calles INDERENA Cabin, Plot W, subplot W2-W3, 1,340 m, 7 Dec. 1992 (ster.), Pipoly et al. 16689 (COL, FMB, JAUM, MO); Plot W3-W4, Tree #99, 1,330 m, 8 Dec. 1992 (ster.), Pipoly et al. 16789 (COL, FMB, JAUM, MO, US), (ster.), Pipoly et al. 16813 (COL, FMB, JAUM, MO, US), (seedling), Pipoly et al. 16835 (COL, FMB, JAUM, MO), 10 Dec. 1992 (ster.), Pipoly et al. 16916 (COL, FMB, JAUM, MO, US), (ster.), Pipoly et al. 16922 (COL, F, FMB, HUA, JAUM, MO, NY, US); Alto de Cuevas, 10 km W of Blanquita, 12 km W of Nutibara, cloud forest, 1,720 m, 3 Mar. 1992 (fl bud). Gentry et al. 761 1 1A (FMB, MO). New Combinations in Geissanthus Examination of type material, concomitant with recent collections, makes the transfer of Conomor- pha scrobiculata Cuatrecasas, Ardisia longistam- inea A. C. Smith, and Cybianthus perpuncticulosus (Lundell) Pipoly & Lundell to Geissanthus neces¬ sary. The new combinations follow, along with re¬ vised descriptions and citation of specimens exam¬ ined for the latter two species. Geissanthus scrobiculatus (Cuatrecasas) Pipoly, comb. nov. Basionym: Conomorpha scrobi¬ culata Cuatrecasas, Rev. Acad. Colomb. Ci. Ex. 8: 320. 1951. TYPE: Colombia. Valle del Cauca: costa del Pacifico, Rio Yurumangui, El Papayo, 5 Feb. 1944 (fl, fr), J. Cuatrecasas 15998 (holotype, F; isotype, COL). 472 Novon This species is known only from the type, but the calyx development, corolla lobe shape, and anther morphology clearly place this species in Geissan- thus. Geissanthus longistamineus (A. C. Smith) Pi- poly, comb. nov. Basionym: Ardisia longis- taminea A. C. Smith, Amer. J. Bot. 27: 544. 1940. TYPE: Colombia. Narino: Gorgonilla Is¬ land. 130-200 m, 9 Feb. 1939 (fl), E. Killip & //. Garcia 33078 (holotype, NY; isotype, US). Shrub or small tree to 10 m tall; branchlets terete, except angular at apex, 5-8 mm diam., sparsely and minutely rufous glandular-granulose and fur- furaceous-lepidote, glabrescent. Leaves pseudover- ticillate, chartaceous, the blades narrowly to widely oblanceolate, (1 5-) 19-31 (-42.5) cm long, (4— )8— 1 2.5 cm wide, apex acuminate, the acumen (0.7 — )1 — 1 .5(— 2.2) cm long, base abruptly truncate, midrib impressed above, prominently raised below, the sec¬ ondary veins 15-23 pairs, smooth or somewhat impressed above, prominently raised and promi¬ nently black punctate-lineate below, blades smooth and glabrous above, moderately and minutely fur- furaceous-lepidote and black punctate-lineate below, the margin flat, entire, glabrous; petiole canaliculate, (l-)1.5-2.5(-3) cm long, tapered to base, glabrous above, densely and minutely furfuraceous-lepidote below, glabrescent. Inflorescence terminal, pyra¬ midal and bipinnately paniculate, 10-16 cm long, secondary branches 4 6 cm long at base, peduncle, rachis, bracts, and pedicels densely glandular-gran¬ ulose, glabrescent; inflorescence bract unknown; pe¬ duncle obsolete to 3 mm long; primary and second¬ ary branch bracts unknown; floral bracts membranous, linear-lanceolate, 3.2 3.5 mm long, 1-1.1 mm wide, apex attenuate, asymmetric, sparsely and prominently red punctate, the margin hyaline, irregular, with occasional serrulations api- cally, glabrous; pedicel obconic, 0.2- 1.3 mm long, densely glandular-granulose. Flowers erect, 5. 8-6. 3 mm long; calyx membranaceous, campanulate, whit¬ ish, 4-5-merous, 3.4-4 mm long, the tube 1 .3-1.5 mm long, unequally divided, the lobes ovate to widely oblong, 2. 1-2.5 mm long, 1.5 2 mm wide, apex obtuse, medially thickened, densely and prominently brown punctate, and punctate-lineate, densely glan¬ dular-granulose, the margin hyaline, irregular, with occasional papillae, entire, glabrous; corolla mem¬ branaceous, starting out tubular, then the lobes opening to become somewhat campanulate, 5. 5-5. 7 mm long, pinkish cream; early ontogeny: the tube unequally divided, 4.5-5 mm long, the lobes trullate. ca. 0.5-1 mm long, 1-1.1 mm wide, apex acute, erect, hyaline, densely and prominently brown punc¬ tate and punctate-lineate, the margin entire, gla¬ brous; corolla at maturity: tube unequally divided, ca. 2 mm long, the lobes linear, 3. 5-3. 7 mm long, 1-1.4 mm wide, reflexed 180° from tube, hyaline, and as described above; stamens free, 5-5.7 mm long, subequaling the petals, the filaments membra¬ naceous, terete, 4-4.5 mm long, not basally wid¬ ened, inserted at the base of the corolla tube, the anthers oblongoid, 2. 2-2. 4 mm long, 0.6-0. 8 mm wide, apex obtuse, base deeply cordate, versatile, dehiscent by wide latrorse longitudinal slits, the con¬ nective epunctate, hyaline; pistil obturbinate, 3-4 mm long, the ovary 1.2- 1.5 mm long, 1-1.1 mm diam., densely and prominently red punctate, sparsely glandular-granulose, the style 1 .7-2 mm long, erect, epunctate, the stigma ca. 0.2 0.3 mm long, the placenta depressed-conical, ca. 0.2-0. 3 mm long, 0.4-0. 5 mm diam., the ovules 5, uniseriate, deeply imbedded in the placenta. Fruit globose, 6-8 mm long and in diam., densely and prominently black punctate and punctate-lineate, exocarp thin. Distribution. Endemic to the Choco Floristic Province of Colombia and Ecuador, from 80 to 1 , 1 50 m elevation. Ecology. Geissanthus longistamineus is a ridgetop species, occurring at margins of primary forest habitats. Common name. “CapuH.” Specimens examined. COLOMBIA. Choco: Quibdo Tutunendo Rd., ca. 3 km W of Tutunendo, 05°46'N, 76°35'W, 80 m, 8 Jan. 1981 (ster.), A. Gentry et al. 30346 (COL, MO). Narino: Mcpio. de Barbacoas, be¬ tween Barbacoas and 15 km above Rio Telembi, toward Chalchal, 01°40'N, 78°09'W, 170 m, 20 Nov. 1986 (fl), B. Hammel & R. Bernal 15772 (COL, HUA, MO, US); Barbacoas, Corregimiento Santander, Buenavista to Bar¬ bacoas, slopes toward Rio Telembi, 200-840 m, 3-5 Aug. 1948 (fl bud), H. Garcia- Bar riga 13190 (COL); Tumaco, near Pinal Dulce, 7 Oct. 1955 (fr), R. Romero- Castaheda 5342 (COL, MO). ECUADOR. Cotopaxi: Tenefuerste, Rio Pilalo, km 52 53 Quevedo-Latacunga, 750-900 m, 19 July 1992 (fl), C. Dodson et al. 13462 (MO, QCA, SEL). Esmeraldas: Rio Cayapa, Zapallo Grande, 1 km upriver from village, 78°55'W, 0°48'N, 150 m, 3 Aug. 1982 (fr), L. Kvist (ft E. Asanza (AAU, QCA); San Lorenzo, Finca la Chiquita, 01°13'N, 78°49'W, 80 m, 8-15 July 1988 (fl bud), W. Palacios 2652 (MO, QCNE, US). Los Rios: Rio Palenque Biological Station, km 56 Quevedo-Sto. Domingo, 150-220 m, 18 Mar. 1974 (fl), C. Dodson et al. 5506 (MO, QCA, RPSC, SEL), 11 Aug. 1977 (fr), C. Dodson et al. 6731 (MO, QCA, RPSC, SEL), 200 m, 6 Feb. 1974 (ster.), A. Gentry 9684 (MO, RPSC, SEL), 27 Feb. 1974 (fl bud), A. Gentry 10220 ( MO, RPS, SEL), 79°25'W, 00°35'S, 150 m, 19 June 1991 (fr), Palacios & E. Freire 7404 (MO, QCNE); ridge line at El Centinela Ua, on rd. from Patricia Pilar to 24 de Mayo, km 45 on rd. Sto. Domingo to Volume 3, Number 4 1993 Pipoly Geissanthus in the Choco 473 Quevedo, 600 m, 23 May 1983 (fr), C. Dodson & A. Gentry 13822 (MO, SEL, RPSC), 650 m, 26 July 1984 (fr), C. Dodson et nl. 14505 (MO, SEL, RPSC). Manabi: Mahcalilla National Park, zona de San Sebastian, 01°36'S, 80°42'W, 20 Jan. 1991 (fl), A. Gentry & C. Josse 72740 (MO, QCNE). Napo: Huamani, Centro Calluhua Yacu, 31 km E of Tena Baeza Rd., on new rd. to Coca, 0°40'S, 77°40'W, 1,150 m, 24 Dec. 1988 (ster.), Gentry et al. 64134 (MO, QCNE). Geissanthus longistamineus is a relatively well known member of the genus, especially from the fine gatherings noted above from Ecuador. It ap¬ pears to have an as yet undescribed sister species from Pasco, Peru, hut is immediately distinguished from that and all others by its corolla, which is at first tubular, then splits into long lobes, with ob- trullate lobules differentiated at the lobe apices. This unique character allows for rapid recognition in flow¬ er, and in fruit the taxon may be recognized by the spreading, but not recurved, calyx lobes and rela¬ tively large fruits. Geissanthus perpuncticulosus (Lundell) Pipoly, comb. nov. Basionym: Ardisia perpuncticu- losa Lundell, Wrightia 5(6): 194. 1975. C.y- bianthus perpuncticulosus (Lundell) Pipoly & Lundell. Wrightia 7(2): 52. 1982. TYPE: Pan¬ ama. Darien: vie. of Cerro Tacaracuna, summit camp, along N stream of camp, 1 Feb. 1975 (fr), A. Gentry & S. Mori 14049 (holotype, LL-TEX; isotypes, MO — -2 sheets). Shrub or small tree to 15 m tall; branchlets an- gulate, (4.5-)6-8 mm diam., densely furfuraceous- lepidote, glabrescent. Leaves pseudoverticillate, membranaceous, the blades obovate to oblanceolate, or rarely elliptic, ( 1 3.5— )1 6—28( 44.5) cm long, (6-)8 12(-18.5) cm wide, apex subacuminate to long-acuminate, the acumen 0.5 1.5 cm long, base broadly cuneate, decurrent on the petiole, midrib impressed and glabrous above, prominently raised and densely furfuraceous-lepidote below, secondary veins 16—27 pairs, somewhat impressed and gla¬ brous above, prominently raised and densely fur¬ furaceous-lepidote below, densely and minutely scro- biculate and black perpuncticulose above, prominently black punctate and sparsely furfura¬ ceous-lepidote below, the margin flat, entire, gla¬ brous; petioles canaliculate, 1-1.5 cm long, glabrous above, densely furfuraceous-lepidote, glabrescent. Inflorescence a terminal or rarely ( E . Forero et al. 6 869) axillary pyramidal bipinnate panicle, ( 8 — ) 1 3 — 18(-27) cm long, (1 1 . 5 -) 1 3 -2 3(- 34 .5) cm wide at base, inflorescence bract membranaceous, lolia- ceous, oblanceolate, 4.8-5 cm long, 1.3- 1.5 cm wide, apex long-acuminate, the acumen 0.3-0. 5 cm long, base cuneate, sparsely fufuraceous above and below except along midrib and secondary veins, mid¬ rib impressed above, prominently raised below, sec¬ ondary veins 13 15 pairs, scrobiculate and densely black perpuncticulose above, prominently black punctate below, the margin flat, entire, glabrous, caducous, the bract petiole canaliculate, ca. 0.5 cm long, glabrous above, densely furfuraceous-lepidote below; peduncle, rachis, secondary branches and pedicels densely and minutely furfuraceous-lepidote; peduncle obsolete to 1 cm long; secondary branch bracts membranaceous, oblanceolate, 0.3-0. 5 mm long, ca. 0. 1-0.2 mm wide, apex acuminate, base cuneate, furfuraceous-lepidote and densely and prominently black punctate above and below, the margin flat, entire, glabrous; floral bracts membra¬ naceous, linear-lanceolate, 1-1.5 mm long, 0.2- 0.3 mm wide, apex long-attenuate, prominently black punctate, translucent glandular-lepidote and glan¬ dular-papillate, the margin irregular, entire; pedicels cylindrical, 0.9-1. 5 mm long, densely translucent glandular-lepidote and glandular-papillate. Flowers erect, white; calyx cbartaceous, cupuliform, 4 5(- 6)-merous, 1 .2-1.7 mm long, the tube 0.6-0. 7 mm long, very unequally divided, the lobes widely ovate to linear, 0.9-1 . 1 mm long, 0.4- 1 . 1 mm wide, apex obtuse, densely and prominently orange punctate, minutely red glandular-papillate, the margin apically erose, hyaline, glabrous; corolla chartaceous, cam- panulate, 2. 4-2. 7 mm long, the tube 0.3 -0.5 mm long, the lobes linear-lanceolate, 2. 1-2.3 mm long, 0.3-0. 4 mm wide, reflexed 180° at anthesis, apex attenuate, densely and prominently orange punctate, glabrous, the margin entire, glabrous; stamens free, 1.7-2 mm long, the filaments membranaceous, flat, 1 .4-1.7 mm long, widened basally, inserted at co¬ rolla tube base, anthers subversatile, lanceoloid, 0.7- 0.8 mm long, 0.2-0. 3 mm wide, apex obtuse, base cordate, longitudinally dehiscent by narrow, latrorse, longitudinal slits, the connective hyaline, epunctate; pistil obturbinate, 1.4-1. 7 mm long, the ovary 0.7- 0.8 mm long, 0.3-0. 4 mm diam., densely and prom¬ inently red punctate, glabrous, the style 0.6 0.7 mm long, curved, the stigma capitate, the placenta globose, 0.5-0. 6 mm long, 0.2-0. 3 mm diam., ovules 4, buried in the placenta. Fruit globose, 3.5-4 mm long and in diam., densely and prominently black punctate and punctate-lineate, the exocarp thin, red¬ dish purple at maturity. Distribution. Endemic to the Choco Floristic Province, Panama (Darien) and Colombia (Antio- quia, Choco), at 100-1,400 m. Ecology. Geissanthus perpuncticulosus grows in ravines on gravel islands in creekbeds, where fre- 474 Novon quent flash floods occur with onset of heavy rains. Thus far it is known from tall pluvial and premontane pluvial forests, in areas that receive ca. 8,000 mm annual precipitation. Etymology. The specific epithet refers to the numerous small punctations that adorn the leaves of this species. Lundell (1975) described this species in Ardisia because of its furfuraceous-lepidote tomentum, but indicated its generic position was uncertain, and furthermore, that it might belong to Gentlea Lundell. The type specimen is in late fruit and the regularity in merosity of the remaining fruits on it indicated that Cybianthus subg. Weigeltia (A. DC.) Agostini was in order, so Pipoly and Lundell transferred it (Pipoly & Lundell, 1982). However, with additional material, close examination of the inflorescence ves- titure, the campanulate corolla, linear petals, free filaments, and subversatile, latrorsely dehiscent an¬ thers, placement in Geissanthus was indicated. Geissanthus perpuncticulosus is most closely re¬ lated to G. francoae, but is easily distinguished from that taxon by its membranaceous leaves, sparsely placed abaxial lepidote scales, shorter petioles, char- taceous perianth, cupuliform calyx, shorter corolla, obtuse anthers, and premontane forest habitat. Specimens examined. COLOMBIA. Antioquia: Mcpio. de Urrao, Vereda Calles, Parque Nacional Natural “Las Orquideas,” sector Cabana de Calles, Quebrada Agu- delo, 06°31'N, 76°19'W, 1,300-1,400 m, 1 Apr. 1992 (ft, fr), D. Cardenas et al. 3209 (COL, FMB, JAUM, MO); Quebrada Honda, range NW of Cabana de Calles, Plot W, 06°29'N, 76°1 4'W, 1,330 m, 8 Dec. 1992 (ster.), Pipoly et al. 16813 (COL, FMB, JAUM, MO). Choco: Mcpio. San Jose del Palmar, bank of Rio Torito, tributary of Rio Habita, W slopes, Finca Los Guaduales, 630-730 m, 8 Mar. 1980 (fl, fr), E. Forero et al. 6869 (COL, MO), W slopes, 630-680 m, 10 Mar. 1980 (fr), E. Forero et al. 6961 (COL, MO), E slopes, 630 m, 21 Mar. 1980 (fl bud), E. Forero et al. 7554 (COL, MO), 21 Mar. 1980 (fr), E. Forero et al. 7565 (COL, MO); Rio Mutata, tributary of Rio El Valle, between base of Alto del Buey and river mouth, 100 150 m, 9 Aug. 1976 (fr), A. Gentry & M. Fallen 17452 (AAU, COL, LL- TEX, MO). Acknowledgments. Studies on the plant diversity of Parque Nacional Natural “Las Orquideas” are be¬ ing carried out under the auspices of a generous grant from the National Geographic Society, No. 473 1 -92, to the author and Alvaro Cogollo (JAUM). I am most grateful to the staff of the Jardin Botanico and Universidad de Antioquia (JAUM and HUA) for their cooperation and assistance, along with that of the Herbario Nacional Colomhiano (COL). The di¬ versity project has enjoyed the collaboration of: An¬ tonio Jose Lopez de Mesa (Director, Jardin Botan¬ ico), Alvaro Cogollo (Co-PI), Dayron Cardenas, Wilson Rodriguez, Ovidio Alvarez, Luzmarina Velez, Juan Arango, Mara Villa, and Ricardo Callejas, for which I am most grateful. To Felipe Pineda, Antio¬ quia regional director of INDERENA, along with Fabio Ramirez, director of Parque Las Orquideas, I owe my sincere thanks for their collaboration and hospitality during fieldwork. Linda Ellis skillfully pre¬ pared the illustrations. Literature Cited Agostini, G. 1970. Notes on Myrsinaceae. I. Generic assignment of Conomorpha sodiroana Mez, Ardisia ambigua Mart., and related species. Phytologia 20: 401-403. Bentham, G. 1876. Myrsineae. Pp. 639-650 in: G. Bentham & J. D. Hooker (editors), Genera Plantar- um. Lovell Reeve, London. Cuatrecasas, J. 1951. Notas a la flora de Colombia XI. Rev. Acad. Colomb. Ci. Ex. 8: 297-328. Gentry, A. 1982. Phytogeographic patterns as evidence for a Choco Refuge. Pp. 112-136 in G. T. Prance (editor), Biological Diversification in the Tropics. Co¬ lumbia Univ. Press, New York. Lindley, J. 1848. Illustrated Dictionary of Botanical Terms. [Excerpt from Illustrated Dictionary of Bo¬ tanical Terms, by J. Lindley, With an Introduction by A. Eastwood. 1964. School of Earth Sciences, Stanford Univ., Stanford, California.] Lundell, C. L. 1975. Neotropical Myrsinaceae— II. Wrightia 6: 60-100. Macbride, J. F. 1934. New or renamed spermatophytes, mostly Peruvian. Candollea 5: 346-402. Mez, C. 1902. Myrsinaceae. In: A. Engler (editor), Das Pflanzenreich. 9(IV, 236): 1-437. Verlag von Wil¬ helm Englemann, Leipzig. - . 1905. Additamenta monographica. II. Bull. Herb. Boissier ser. 2, 5: 527 538. - . 1920. Additamenta monographica. III. Re- pert. Spec. Nov. Regni. Veg. 16: 410-425. Pipoly, J., III. 1987. A systematic revision of the genus Cybianthus subgenus Grammadenia (Myrsinaceae). Mem. New York Bot. Card. 43: 1-76. - . 1992. The genus Cybianthus subgenus Con¬ omorpha (Myrsinaceae) in Guayana. Ann. Missouri Bot. Gard. 79: 908-957. - & C. L. Lundell. 1982. Contributions toward a monograph of Cybianthus (Myrsinaceae) II. The systematic position of Ardisia perpuncticulosa. Wrightia 7: 52-54. A New Xylopia (Annonaceae) from Panama George E. Sc hat z Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. ABSTRACT. Xylopia panamensis sp. nov. is pro¬ posed based on material from Santa Rita Ridge and El Llano-Carti road formerly referred to Xylopia bocatorena Schery. The new species differs from X. bocatorena by its rounded leaf base, longer in- dument on the bracts and calyx, smaller flowers, fewer seeds per monocarp, and thus shorter mon¬ ocarps. Reexamination of Mesoamerican material of Xy¬ lopia in conjunction with the preparation of a treat¬ ment of the Annonaceae for the Manual to the Plants of Costa Kira has revealed a previously undescribed species from Colon and San Bias Prov¬ inces, Panama. Xylopia panamensis G. E. Schatz, sp. nov. 1 YPE: Panama. San Bias: El Llano-Carti road, km 26.5, 9°19'N, 78°55'W, 200 m, 17 June 1986 (11, fr), de Nevers, Herrera , McPherson , D'. lrcy & Allen 7836 (holotype, MO; isotypes, PM A not seen, U). Figure 1 . Ab Xylopia bocatorena laminis basi rotundatis, brac- teis ut calyces dense aureosericeis, pilis ad 2 mm longis, floribus minoribus, semenibus paucioribus, monocarpiis brevioribus differt. Tree 4-14 m tall, the young branches initially sparsely covered with short, spreading hairs, at length glabrescent. Petiole slender, 2-6 mm long, sparsely covered with short, spreading hairs; lamina char- taceous to slightly membranaceous, broadly lanceo¬ late to narrowly ovate, 5.5-12.5 cm long, 1.7-5. 2 cm broad, the apex long acuminate to cuspidate, the acumen 1-2 cm long, the base rounded to trun¬ cate, rarely obtuse, the upper surface glabrous, the lower surface sparsely covered with appressed, white hairs to 1.5 mm long; venation very weakly bro- chidodromous with 6-10 faint secondary veins per side, the primary vein even with the lamina surface adaxially, somewhat elevated and sparsely covered with appressed, white hairs to 1 .5 mm long abaxiallv. Flowers solitary, axillary. Pedicel 2-4 mm long, borne in the axil of a basal round bracteole and bearing 12 additional distichous, broadly ovate arn- plexicaul bracts, 5 mm long and broad, the apex acute, densely long golden sericeous, the hairs to 2 mm long. Sepals 3, connate at the base for V2 their length, broadly ovate, 6-9 mm long, the lobes 5- 8 mm broad, the apex acute, densely long golden sericeous outside, the hairs to 2 mm long, glabrous inside. Petals 6, in 2 valvate whorls, fleshy, white; outer petals ovate to narrowly elliptic, strongly con¬ cave, 7 mm long, 4-5 mm broad, 1 mm thick, the apex acute, the base obtuse, densely short white sericeous outside, very short tomentellous inside, with a medial keel inside toward the apex marking the line along which adjacent inner petals meet; inner petals rhombic-trullate, 5 mm long, 3 mm broad, 0.5 mm thick, the apex acute, the base cuneate, strongly concave where the petal fits tightly against the androecium, densely short white sericeous out¬ side, glabrous inside. Torus conical, 3.2 mm diam., with a central cavity into which the ovaries are embedded. Stamens ca. 200 (based on scars from the filament bases on the torus), stamens lacking; carpels 3 8 (based on fruiting monocarps present), ovaries lacking. Fruit a cluster of separate stipitate dehiscent monocarps, the stipe 2-3 mm long, the monocarp shortly cylindric to ellipsoid, to 3 cm long, 1.5 cm diam., the apex strongly apiculate, and with 1 to several additional beaklike protuberances along the margin of dehiscence, the surface initially dense¬ ly sericeous, at length puberulent, red; seeds 2 3, interdigitated to form a single row, flattened ellipsoid, 10 mm long, 6 mm diam., shiny black with a fleshy, bilobed, white aril at the base. Much of the material cited has previously been determined as Xylopia bocatorena Schery, to which the new species, X. panamensis, is undoubtedly closely related, and with which it is wholly sympatric. In several respects — leaf length and seed number (and therefore, monocarp length) the new species conforms well to the original description of X. bo¬ catorena (in Woodson & Schery, 1943). However, as numerous collections of X. bocatorena have ac¬ cumulated from Nicaragua to El Llano-Carti Road in Panama, these two features of the type material ( von Wedel 2965 (HT: MO)) from Bocas del Toro, Panama, have proven anomalous. Leaves on the type fall within the lower end of the overall range in variation of leaf length; indeed, A. bocatorena often displays leaves considerably longer (to 22 cm) Novon 3: 475-477. 1993. 476 Novon 5mm D Figure 1. Xylopia pananiensis G. E. Schatz (from de Never* at at. 7836 (flower) and Knapp & Schmalzel 1792 (fruit)). —A. Composite flowering and fruiting branch. — B. Flower with subtending bracts. — C. Adaxial view of outer petal. — D. Adaxial view of inner petal. than those on the type (8-11 cm). Similarly, the number of seeds (2), and hence, minimum number of ovules, described from the type is unusually low, resulting in a concomitantly short monocarp (2 cm). Usually, ca. 6 ovules are present (to 10), resulting in a cylindrical monocarp 5 cm long. Thus, in gen¬ eral V. pananiensis can be characterized by fewer seeds, resulting in a shorter, more flattened ellipsoid monocarp, which, moreover, has one to several cu¬ rious beaklike appendages along the margin of de¬ hiscence, which are lacking on monocarps of X. bocatorena. Xylopia pananiensis differs further from X. bo¬ catorena by a more rounded versus cuneate to ob- Volume 3, Number 4 1993 Schatz Xylopia panamensis 477 fuse leaf base, generally smaller flowers, and es¬ pecially by the markedly longer (to 2 mm long vs. less than 0.5 mm long) golden sericeous indument on the bracts and calyx. Among several other north¬ ern South American species of Xylopia with solitary, relatively large flowers, X. panamensis also exhibits affinities with X. chocoensis R. E. Fries in the size and shape of the monocarps and a rounded leaf base (Fries, 1955). However, leaves of X. chocoensis are nearly always longer (12-15 cm long), and the number of carpels (ca. 25) is significantly greater. Paratypes. PANAMA. Colon: Santa Rita E Ridge, 23 Jan. 1968 (fr), Dwyer 8421 (MO), 300-500 m, 16 Dec. 1972 (fr), Gentry 6571 (MO), 21 km from Tran¬ sisthmian Hwy., 9°26'N, 79°38'W, 400-500 m, 23 Oct. 1981 (fr), Knapp & Schmalzel 1792 (MO, U), ca. 4- 5.5 mi. E of Transisthmian Hwy., 6 Apr. 1969 (fl, fr), Lewis et al. 5275 (MO), 20-25 km from Transisthmian Hwy., 9°24'N, 79°39'W, 30-400 m, 25 Sep. 1980 (fr), Sytsma 1354 (MO), 10 Oct. 1980 (fr), Sytsma 1526 (MO), ca. 3.1 km E of Agua Clara rain gauge, 9 Sep. 1975 (fr), Witherspoon et al. 8316 (MO). Literature Cited Fries, R. E. 1956. Some new contributions to the knowl¬ edge of the Annonaceae in Colombia and Mexico. Ark. Bot., n.s., 3(12): 433-437. Woodson, R. E. & R. W. Schery. 1943. Contributions towards a flora of Panama. VII. Miscellaneous col¬ lections, chiefly by H. von Wedel in Bocas del Toro. Ann. Missouri Bot. Gard. 30: 83-96. Terpsichore , a New Genus of Grammitidaceae (Pteridophyta) Alan R. Smith University Herbarium, University of California, Berkeley, California 94720, U.S.A. Abstract. Terpsichore, a new genus of Gram¬ mitidaceae, is described and combinations are made for the species known to belong to it. Terpsichore differs from other genera in the family by the com¬ bination of the following characters: the presence of usually conspicuous hydathodes that sometimes pro¬ duce calcareous secretions; reddish to atropurpu- reous (less often hyaline) setae mostly 1 -3 mm along the stipe, rachis, costae, and (sometimes) lamina; nonclathrate, usually castaneous to blackish rhizome scales that are usually setulose along the margins; and free, unbranched, pinnate venation in the seg¬ ments of pinnae. The genus comprises about 50 species and is primarily neotropical; one species is known from Africa and offshore islands. A key is provided to distinguish the neotropical genera of Grammitidaceae. This is the last in a series of papers describing new genera of neotropical Grammitidaceae. Earlier papers dealt with the anhydathodous genera Cera- denia (Bishop, 1988), Zygophlebia (Bishop, 1989), and Enterosora (Bishop & Smith, 1992), while more recently the hydathode-bearing genera Lellingeria (Smith et ah, 1991), Melpomene (Smith & Moran, 1992), and Micropoly podium (Smith, 1992) have been delimited. Species of Terpsichore also have hydathodes, sometimes with calcareous secretions. There are approximately 50 known species, making it about equal in size to Lellingeria and Ceradenia. The distinctness of Terpsichore, named for the Greek muse of dance, was first realized hy the late L. Earl Bishop. Unfortunately, he was unable to advance the work beyond the use of the name in labeling some of his own collections. I have made a search of herbaria (UC and MO), recent floristic accounts for Latin American countries (Proctor, 1977, 1985, 1989; Stolze, 1981; Mickel & Beitel, 1988; Lellinger, 1989), Copeland's monograph of Ctenopteris (1956), and other relevant literature to determine those species that should be transferred to the new genus. The following artificial key is provided to distin¬ guish Terpsichore from its New World congeners: Artificial Key to Genera of Neotropical Grammitidaceae 1 . Laminae entire or with margin slightly re- pand, rarely 1 -2-forked with divisions en¬ tire . 2 1 . Laminae shallowly to deeply lobed to pin- natifid or 1 -pinnate, rarely pinnate-pinnat- ifid . 8 2(1). Veins connected by a marginal vascular strand, thus forming a row of areoles along each side of midrib; Greater Antilles .... . Lomaphlebia 2. Veins free, or only casually and infrequent¬ ly anastomosing; widespread in the Neo¬ tropics . 3 3(2). Sori intermixed with whitish, stipitate, spherical glands . . . Ceradenia jungermannioides (Klotzsch) L. E. Bishop 3. Sori lacking glands, or glands, if present, not whitish . 4 4(3). Laminae with a dark brown to black scler- enchymatous border . . Grammitis sect. Grammitis 4. Laminae lacking a strongly differentiated dark border . 5 5(4). Stipes and laminae with numerous dark setae greater than 1 mm . 6 5. Stipes and laminae lacking dark setae, or if such setae present then these usually few and less than 1 mm . 7 6(5). Rhizomes short-creeping; sori round or nearly so; tropical America . Enterosora 6. Rhizomes long-creeping; sori oblong to short- linear; southern Chile and Argentina . . . . Grammitis patagonica (C. Christensen) Parris 7(5). Hydathodes present; tropical America . . . Cochlidium 7. Hydathodes usually absent (sometimes present in G. magellanica Desvaux); southern Chile and Argentina . . Grammitis sect. Grammitastrum 8(1). Hydathodes absent or obscure on adaxial surface of lamina . 9 8. Hydathodes present at the ends of veins on adaxial surface of the laminae . 12 9(8). Laminae simple, a single blade differenti¬ ated into serrate, sterile, proximal part and an entire, usually fertile, distal part .... Cochlidium serrulatum (Swartz) L. E. Bishop 9. Laminae sinuate, shallowly lobed, or more often pinnatifid to pinnate (rarely more di¬ vided), not differentiated into distinct sterile and fertile parts . 10 Novon 3: 478-489. 1993. Volume 3, Number 4 1993 Smith Terpsichore (Grammitidaceae) 479 1 0(9). Sori with conspicuous, whitish, spherical, waxlike paraphyses (most easily seen in young fronds); veins free . Ceradenia 10. Sori lacking evident waxlike, spherical pa¬ raphyses, or if paraphyses evident these brownish and not waxlike; veins often anas¬ tomosing to form regular or irregular ar- eoles . 11 1 1(10). Laminae sinuate, or lobed no more than % the distance to the midrib . Enterosora 1 1 . Laminae deeply pinnatifid or pinnatisect nearly to the midrib . Zygophlebia 12(8). Scales of rhizomes lacking setulae (often bearing papillate glands less than 0.1 mm at tip), strongly clathrate, often iridescent; rhizomes long-to short-creeping, strongly dorsiventral . Melpomene 12. Scales of rhizomes setulose or not, not clathrate, or if clathrate, then setulose; rhi¬ zomes generally short-creeping to suberect, radial or weakly dorsiventral, rarely strong¬ ly dorsiventral . 13 13(12). Main vein (costa) in the segments (or pin¬ nae) simple (lacking branches) or with a single acroscopic veinlet; sori only one per segment or pinna; laminae narrowly linear, usually less than 1 cm wide; rhizomes ra¬ dially symmetric . 14 13. Main vein (costa) in the segments (or pin¬ nae) with more than 1 branch; sori usually more than one per segment or pinna; lam¬ inae usually lanceolate to ovate, usually more than 1 cm wide; rhizomes radially to dorsiventrally symmetric . 15 14(13). Rhizome scales golden to orangish or light brownish, not clathrate; castaneous or red- brown simple (unforked) setae more than 1 mm long generally present on midribs and sometimes on lamina . Micropolypodium 14. Rhizome scales dark brown to blackish, clathrate; castaneous or red-brown setae generally lacking on midribs and lamina, if present then less than 0.5 mm long and often unequally forked . Lellingeria 1 5( 1 3). Scales clathrate; sori often slightly to deep¬ ly immersed in the lamina; stipes and ra- chises with hyaline to light reddish setulae 0. 1 0.3 mm, lacking dark setae; rhizomes ascending, radially symmetric . . . Lellingeria 15. Scales not clathrate; sori superficial; stipes and rachises generally with dark setae more than 0.5 mm; rhizomes short-creeping to ascending, dorsiventral to radial . Terpsichore Terpsichore A. R. Smith, gen. nov. TYPE: Po¬ lypodium asplenifolium L., Sp. PI. 2: 1084. 1753. = Terpsichore asplenifolia (L.) A. R. Smith. Figure 1. Plantae plerumque epiphyticae; rhizomata infirme dor- siventralia vel radialia; squamae rhizomatis non clathratae, plerumque brunneolae, atropurpureae, vel nigrescentes, concolores, margine plerumque setulosae; folia plerumque pinnatisecta, cum hydathodis adaxialiter, interdum exsu- datis calcareis efferentibus; laminae interdum fungis ateris clavatis praeditae; petioli et rhachides setosi, setis casta- neis vel atropurpureis, interdum albidis; venae librae non furcatae; sori rotundi superficales, sine glandibus. Plants epiphytic, rarely saxicolous or terrestrial; rhizomes short-creeping to ascending, generally weakly dorsiventral to radial, the scales nonclath- rate, brown to blackish, infrequently orangish, con- colorous, dull to usually shining, glabrous or with variously colored (hyaline to castaneous) setae; phyl- lopodia present or absent; petioles nearly absent to equaling the lamina, setose (especially proxiirtally) and sometimes also puberulent, the setae 0.5-3 mm, usually reddish or castaneous, numerous, spreading, the hairs 0. 1-0.2 mm, branched or unbranched, pale reddish, sometimes glandular; laminae pinnat¬ isect to 1 -pinnate, rarely 1 -pinnate-pinnatifid, fork¬ ing in a few spp., monomorphous, usually setose (at least along the rachis); hydathodes present, some¬ times producing calcareous secretions (whitish lime- dots); veins simple, pinnate from the costae, Iree, hidden or easily visible; sori round, not sunken into the lamina, without paraphyses; sporangial capsules glabrous or setose; spores globose-tetrahedral, ob¬ late, or reniform, with a trilete or monolete laesura. x = 37. Distribution. Terpsichore comprises about 50 neotropical species, with one, T. cultrata , extending into Africa (Schelpe, 1969), Madagascar (Tardieu- Blot, 1960), and the Mascarenes. It is uncertain whether African material called T. cultrata is really conspecific with New World ma¬ terial of the same species, but if not, plants from the two areas are certainly closely allied. It is highly interesting, and suggestive of antiquity for the Gram¬ mitidaceae, that most of the genera of neotropical Grammitidaceae, including Terpsichore, Ceradenia, Enterosora, Lellingeria, Melpomene, Zygophle¬ bia, Grammitis sensu stricto, and Cochlidium, are also represented by one or a few species in Africa and islands of the Indian Ocean (Smith, 1993); these same genera, with the exception of Lellingeria and Grammitis, are absent from southeast Asia and the Pacific. Characters defining Terpsichore include the pres¬ ence of rather conspicuous hydathodes on the ad- axial surface of the laminae; castaneous or atro- purpureous to whitish setae along the stipe, rachis, costae, and sometimes lamina; setae usually dense at the stipe bases, absence of glandular paraphyses in the sori; nonclathrate, usually castaneous or atro- purpureous (occasionally blackish or orangish), con- colorous rhizome scales that are usually setulose along the margins, or at least at the tip, and some¬ times on the surfaces; rhizomes generally weakly dorsiventral to radial in symmetry; and free, un- 480 Novon Figure 1. Characteristic species of Terpsichore. — A. T. alsophilicola. Group 1, habit (pendent, indeterminate epiphyte) and details to show rhizome scale, abaxial surface of segment, indument, and sporangium (Knapp & Kress 4315 , UC). — B. T. alsopteris. Group 2, habit (erect-arching) and details to show rhizome scale, fungal fruiting body from rachis, and calcareous secretions from hydathodes on adaxial surface ( van tier Werff & Palacios 10589, UC). — C. T. longa. Group 3, habit (pendent indeterminate epiphyte) and details to show sporangia, paired and clustered setae on pinna margin (van der Werff et al. 9488, UC). — D. T. sub tills. Group 4, habit (arching-pendent) and details to show rhizome scale, abaxial surface of segments, fungal fruiting body in sorus, and calcareous secretion from hydathodes on adaxial surface (van der Werff & Palacios 9147, UC). — E. T. longiselosa. Group 5, habit (± erect) and details to show rhizome scale, abaxial surface of segments, and branched laminar hairs (Herrera 3670, UC). Volume 3, Number 4 1993 Smith Terpsichore (Grammitidaceae) 481 branched veins from the costae of the segments or pinnae. The most similar species to Terpsichore currently are classified in Ctenopteris, a diverse and probably polyphyletic genus typified by the Old World species C. venulosa (Blume) Kunze. A synonym of Cten¬ opteris is Cryptosorus [synonym: Grammitis sect. Cryptosorus (Fee) R. M. Tryon & A. F. Tryon], type Cryptosorus blumei Fee [= Ctenopteris ob- liquata (Blume) Copeland], also Old World. The type species of both Ctenopteris and Cryptosorus, as well as many allied Old World species (see Parris, 1990: 82), lack hydathodes, have subclathrate rhi¬ zome scales, and have sori sunken in crypts. In these three characteristics, Ctenopteris differs from all species here included in Terpsichore. Some other Old World species included in Ctenopteris do have hydathodes, nonclathrate rhizome scales, and non- sunken sori, characteristics that might suggest a closer affinity to Terpsichore. The affinities of the “nontypical, ” Old World Ctenopteris species need further study, but it seems clear that Terpsichore is not congeneric with any genus of Grammitidaceae based on an Old World type. Certain species of Terpsichore, especially those in the T. taxifolia group (Group 2, below), also resemble Pecluma (Polypodiaceae) because of their pectinate laminae. Pecluma, however, has short phyllopodia, yellow, bilateral, monolete spores, and often comose rhizome scales; species of Pecluma also lack stout castaneous setae and black clavate fungi. It seems likely that the resemblance is the result of convergent evolution and is not an indi¬ cation of any close affinity between the two genera. In his revision of American Ctenopteris , Copeland (1956) treated the species of Terpsichore in seven of his nine species groups, exemplified bv the fol¬ lowing seven species: Ctenopteris senilis, C. cap- illaris, C. elastica, C. pendula, C. suspensa, C. sericeolanata, and C. meridensis. Copeland’s groups bear little or no resemblance to the genera as Bishop and I have circumscribed them, and in fact Cope¬ land's groups of species are mostly highly artificial assemblages based on size and dissection of the fronds, characters that are completely unreliable in defining natural groups of species in Grammitidaceae. Species of Terpischore can be rather easily di¬ vided into the following five groups, which could conveniently be regarded as natural sections of the genus (Fig. 1). Group 1. Terpsichore asplenifolia Group (12 spp.). — Fronds arching, blades determinate; rhi¬ zomes weakly dorsiventral to radial; rhizome scales orange to brown, setulose on margins and often surface; sporangia setose (except 7. atroviridis ); blades lacking black clavate fungal fruiting bodies; stipes distinct, the pinnae not reduced to base of frond; laminae with solitary setae; hydathodes lack¬ ing calcareous secretions (Fig. 1A). Group 2. Terpsichore taxifolia Group (13 spp.). — Fronds arching to erect, blades determinate; rhi¬ zomes generally dorsiventral; rhizome scales casta¬ neous, setose at tip and/ or margins; sporangia gla¬ brous; blades with black clavate fungal fruiting bodies (except T. paulistana); stipes distinct, the pinnae not reduced to base of frond; laminae with solitary setae; hydathodes often producing calcareous se¬ cretions (Fig. IB). Group 3. Terpsicore lanigera Group (16 spp.). — Fronds pendent, blades indeterminate; rhizomes gen¬ erally radially symmetric; rhizome scales hairy on margins and often surface, or sometimes absent (71 alfarii ); sporangia setose (except T. delicatula, T. jamesonioides, and 7. spathulata); blades lacking black clavate fungal fruiting bodies; stipes usually less than 1 cm long, the pinnae reduced nearly to base of frond; laminae with tendency for paired or clustered setae (except T. delicatula, T. jameson¬ ioides); hydathodes producing calcareous secretions in a few spp. (Fig. 1C). Group 4. Terpsichore subtilis Group (2 spp.). — Fronds arching or directed downward, sometimes pendent, blades indeterminate; rhizomes weakly dor¬ siventral to radial; rhizome scales blackish, with a single apical seta or marginally setulose; sporangia glabrous; blades with black clavate fungal fruiting bodies; stipes less than 1 cm long, the pinnae reduced nearly to base of frond; laminae with solitary setae; hydathodes producing calcareous secretions (Fig. ID). Group 5. Terpsichore achilleifolia Group (3 spp.). — Fronds arching, blades determinate; rhi¬ zomes generally weakly dorsiventral; rhizome scales orangish to orange-brown, entire to glandular to sparingly setose on margin; sporangia glabrous; blades lacking black clavate fungal fruiting bodies; stipes distinct, the pinnae not reduced to base of frond; laminae with solitary setae; hydathodes lacking cal¬ careous secretions (Fig. IE). The relationships among these groups are un¬ certain. The pendent, nearly estipitate fronds, ra¬ dially symmetrical rhizomes, clustered setae, and usually setose sporangia of the T. lanigera group are probably derived features. In frond form, the T. asplenifolia group seems to be ancestral, possibly having given rise to T. taxifolia and allies, all but one of which bear black clavate fungi. Species in both the asplenifolia and taxifolia groups have more or less weakly dorsiventral rhizomes, thought by Bishop (1989) to be an ancestral condition in the family. In its orangish, entire or sparingly setose rhizome scales, the T. achilleifolia group is aberrant 482 Novon Table 1. Spores of Terpsichore, T. lanigera group. Vouchers are all in UC. Less common conditions are in parentheses. Bo = Bolivia; Br = Brazil; C = Colombia; CR = Costa Rica; E = Ecuador; G = Guatemala; M = Mexico; Pa = Panama; Pe = Peru; V = Venezuela, e = ellipsoid; mo = monolete; o = oblate; t = tetrahedral; tr = trilete. Species Shape Laesura Number of cells Voucher Locality T. alfarii o tr 1 Fiedler 159 CR T. cultrata e mo 2 Steyermark 48601 G e mo 2(1) Silverstone-Sopkin 3614 C t-o tr 1 (2) Steyermark & Wurdack 1177 V e ino 1 (2) Steyermark 100715 V t-o tr 1 & 2 Krai 7197211 V t-o tr 1 & 2 Kummrow 2153 Br t-o tr 1 (2) Hutchison & Wright 5582 Pe e & o mo & tr 1 & 2 Fay & Fay 2184 B T. delicatula o tr 1 (2) Smith 471 M T. hetermorpha t-o tr 1 & 2 Ceron 1207 E t-o tr 1 0llgaard 34281 E T. jamesonioides o tr 1 & 2 Bishop 2520 Pe T. lanigera t-o tr 1 Herzog 2180 Bo t-o tr 1 Wiggins 10768 E t-o tr 1 Bishop 862 CR e (o) mo (tr) 1 & 2 Mickel 3025 CR e mo 1 & 2 Buchtien 5251 Bo t-o tr 1 Sodiro s.n. E T. laxa e (o) mo (tr) 1 Bishop 2509 Pe T. tonga e mo 1 (2) Gomez 21085 CR e mo 1 (2) Chacon 1518 CR e mo 1 (2) Palacios 5655 E e mo 1 & 2 van der Werff 9488 E T. mollissima e mo 1 Steyermark 49625 G e mo 1 & 2 Skutch 5204 CR e mo 1 & 2 Maguire 53790 V T. senilis e mo 2 Breedlove 68357 M e (t-o) mo (tr) 1 & 2 Bishop 861 CR e mo 1 & 2 Gomez 22285 Pa e mo 2(1) Smith et al. 1430 V e mo 1 or 2 Steyermark & Liesner 118283 V T. spathulata t-o tr 1 Pur pus s.n. M T. subflabelliformis e mo 1 & 2 van der Werff 9148 E e mo 1 (2) Bishop 2515 Pe e mo 1 (2) Herzog 1985 Bo T. turrialbae t-o tr 1 Smith 2073 CR t-o tr 1 Bishop 859 CR T. xanthotrichia e mo 2 Beitel 85083 V and may show a relationship to Micropolypodium or to some Old World species currently included in Ctenopteris (but not the type). The pinnate-pinnat- ifid fronds of three species belonging here are un¬ usual in Grammitidaceae but are found in several species of Lellingeria (e.g., L. suprasculpta, L. melanotrichia). This character state (pinnate-pin- natifid lamina) has undoubtedly arisen several times among neotropical grammitids and also in the Old World genus Ctenopteris (e.g., C. taxodioides (Ba¬ ker) Copeland, C. bipinnatijida (Baker) Copeland, and allies). A peculiar feature in nearly all species of the Terpsichore taxifolia and T. subtilis groups is the presence of small (ca. 0.5- 1.0 mm long), black, clavate fungi (presumably Acrospermum maxonii Farlow) on the abaxial laminae, particularly along the vascular tissue (rachis, costae, and veins) and in the sorus (Mickel, 1973). Not all fronds in a given collection will show this feature, but it is of such regular occurrence on at least some fronds of a gathering as to be diagnostic for the two groups. Most likely, the presence of this fungus on species in the two groups indicates a monophyletic origin Volume 3, Number 4 1993 Smith Terpsichore (Grammitidaceae) 483 for this suite of about 15 species (Groups 2 and 4, below). The only other fern outside these groups that is a known host for this particular fungus is the grammitid Melpomene anfractuosa (Kunze ex Klotzsch) A. R. Smith & R. C. Moran. Chromosome counts are known for a few species of Terpsichore, namely, T. asplenifolia {n = 37, 2n = 74; Walker, 1966), T. semihirsuta (2 n = ca. 148; Walker, 1966), from Jamaica, and T. lanigera (n = 37; Evans, 1963, reported as Ctenopteris cultrata, a misidentifieation, UC 58-234-1), from Costa Rica. From these fewr reports, it would appear that the base number for the genus is x = 37, the same number found in several other grammitid gen¬ era, including Ceradenia, Micropolypodium, En- terosora, and Melpomene. Tryon & Lugardon (1991: 363) provided an SEM photograph of a spore ol T. asplenifolia that shows a spore that is nearly globose, with an obscure trilete aperture and finely papillate surface. Spores of other genera of Grammitidaceae shown in this work show similar features, but are sometimes more tetrahe¬ dral-globose and have more prominent apertures and papillae or surface globules. However, insufficient material has been studied to indicate the taxonomic significance, if any, of these variations in spore mor¬ phology in the family. Wagner (1985) reported that spores of Gram- mitis lanigera, G. senilis, and several related gram- mitids had monolete laesurae. 1 confirm the presence of monolete spores in the first two species and in other species belonging to the Terpsichore lanigera group (Croup 3, see below). Ellipsoid, bilateral, monolete spores appear to be consistently present in T. longa, 7. mollissima, T. senilis, and T. xanthotrichia; they are also some¬ times present in T. cultrata, 7. lanigera, and /. laxa (Table 1). The other species for which spore material has been available in Group 3 have globose- tetrahedral, trilete spores (Table 1), as do species examined in Groups 1, 2, and 4. The significance of ellipsoid, monolete spores in the genus is not known, but this character state is undoubtedly de¬ rived and does not indicate affinity with Pecluma or any other extant genera of Polypodiaceae, which nearly always have bilateral, monolete spores. A peculiar feature of the spores of Grammitida¬ ceae is that they often undergo precocious division within the spore wall before being released from the sporangium. Bicellular or multicellular spores are a feature of several other groups of ferns that produce chlorophyllous spores (Tryon & Lugardon, 1991). In Terpsichore, it appears that spores often undergo division before the spore wall is ruptured and before release from the sporangia ( Fable 1; Stokey & At¬ kinson, 1958). New Species Terpsichore pirrensis A. R. Smith, sp. nov. TYPE: Panama. Darien: Cerro Pirre, 2,500- 4,500 ft., Duke & Elias 13812 (holotype, UC; isotypes, MO, NY not seen, SCZ not seen). Figure 2A. Ex affinitate T. taxifoliae (L.) A. R. Smith et T. ze- ledonianae (Lellinger) A. R. Smith, stipitis setis brevior- ihus hyalinis vel pallide rubellis 0.2-0. 3 mm, rhachide glabra abaxialiter, et setis inter sporangia plerumque car- entibus distinguenda; insuper differt a /'. taxifoliae setis margine segmentorum carentibus. Rhizome short-creeping to suberect; scales cas- taneous, concolorous, shining, 1.5-2. 5 x 0.2-0. 3 mm, entire, often with a single minute hyaline patent or oblique seta at tip; fronds mostly 15-30 cm; stipes dark brown, up to 8 cm x 0.5-1 mm, with hyaline to light reddish setulae 0.2-0. 3 mm; laminae thin-chartaceous, pinnatisect, narrowly elliptic with proximal pinnae (up to ca. 12 pairs) gradually re¬ duced to small lobes 1 -3 mm long; rachises glabrous abaxially or with very sparse setae to 1 .5 mm, also with black clavate fungi; segments mostly 30-55 pairs per frond, spreading to often ascending up to ca. 70° from rachis, linear-lanceolate, 1.5-2. 5 cm x 2-3.5 mm, acutish at tip, lacking marginal and laminar setae; veins simple, blackish and easily vis¬ ible abaxially, up to 15 pairs per segment; hyda- tliodes lacking calcareous secretions; sori up to 1 5 pairs per segment, lacking setae from receptacle or setae sparse, less than 0.3 mm; sporangia glabrous. This particular group, also including /. taxifolia (L.) A. R. Smith and T. alsopteris (C. V. Morton) A. R. Smith in Mesoamerica, is in need of revision, especially in South America where there are addi¬ tional undescribed species. Terpsichore pirrensis is known only from Cerro Pirre. Paratypes. PANAMA. Darien: same locality as type, Duke & Elias 13754 (MO, NY not seen, SCZ not seen, UC); Serrania de Pirre, along steep ridge from Altos de Nique to Cerro Pirre, ca. 8 km N of Alturas de Nique, ca. 8 km W of Cana Gold Mine, 1,430-1,480 m, Croat 37833 (MO); Cerro Pirre, valley between Pirre and next most southerly peak, Folsom 4447 (MO). Terpsichore spathulata A. R. Smith, sp. nov. TYPE: Mexico. Mexico: Sierra de las Cruces, 1 1,000 ft., Pringle 4145b (holotype, UC; iso¬ type, US). Figure 2B. A T. heteromorpha (Hooker & Greville) A. R. Smith rhizomatis paleis parvioribus aurantiacis vel brunneolis, sporangiis sine setis, et laminis gracilioribus plerumque non furcatis aut si furcatis nunc divisionibus ca. 30° div- ergentibus differt. 484 Novon Figure 2. New species of Terpsichore. — A. T. pirrensis A. R. Smith, habit and details showing stipe base with uniformly short hairs, abaxial surface of segments, and fungal fruiting body from rachis ( Duke & Elias 13842, UC). — B. T. spathulata A. R. Smith, habit and details showing rhizome scale, branched hair along rachis, stellate seta on rachis, and abaxial surface of segments, Pringle 4145b (UC). Plants epipetric in moist, shaded rock crevices and on cliffs, under rock overhangs, and in shallow caves and grottoes; rhizome very short-creeping to suberect; scales orangish to brownish, concolorous, dull, 0.3-1 x 0.2-0. 5 mm, somewhat obscured by dense, hyaline setae on margins and surfaces; fronds numerous, densely crowded, pendent, often more than 50 per plant, (12-)25-60(-90) cm, apices indeterminate; stipes brown, 20 150 x 0.2-0. 4 mm, with numerous, mostly unbranched, tawny se¬ tae 1-2 mm; laminae 10-85 x 0.4-1.2(-1.6) cm, 1 -pinnate, or occasionally rachises 1 -2-bifurcate with forks ca. 30° apart, pinnae very gradually reduced and more widely separated proximally, contiguous or often overlapping distally, without black clavate fungi, thin-chartaceous; rachises brown to blackish, with numerous dark, often paired or fasciculate setae 1 2 mm; pinnae often more than 50 pairs per frond, alternate, 2— 5(— 7) x 2-4 mm, ascending, sessile or proximal ones stipitate to 2 mm, irregularly oval, obovate, rhombic, or slightly lobed, larger ones often inequilateral (excised basiscopically), base cuneate, apex rounded or truncate, margin setose, setae pale reddish, not paired, paired, and/or sessile-stellate; veins obscure except for the darkened costa (basal half), 1-3 pairs per pinna; laminar surface between the veins glabrous or with pale setae on both sur¬ faces, often also with 1-3-branched hairs 0.1 -0.2 Volume 3, Number 4 1993 Smith Terpsichore (Gramm itidaceae) 485 mm; hydathodes without lime dots; sori usually 2- 6 per pinna, lacking setae from receptacle; sporangia glabrous. Specimens of Terpsichore spathulata from Mex¬ ico and Central America have previously been de¬ termined as Grammitis heteromorpha (Hooker & Greville) C. V. Morton. The latter species is re¬ stricted to the Andes from Colombia to Bolivia and differs by the larger (1-1.5 mm), black rhizome scales, thicker lamina and rachises, and presence of sporangial setae. In addition, Andean specimens have the lamina markedly and regularly bifurcate, with the blade forks diverging at 60-90°. Smith (1981) and Stolze (1981) have previously discussed some of the variation in Grammitis heteromorpha sensu lato. Paratvpes. MEXICO. Chiapas: Munic. Union Juarez, SE side of summit of Volcan Tacana, 12,500 ft., Breed¬ love 24297 , 24303, 24304 (DS); Volcan Tacana, 2,800 m, Matuda 2890 m (K, MEXU, NY, US). Distrito Federal: Desierto de los Leones, ca. 9,600 ft., Her¬ nandez X. s.n. (US), Lyonnet 232 (US). Jalisco: Nevado de Colima, above Jazmin, above El Isote, 2,600 m, McVaugh 10043 ( US), 3,200 m, McVaugh 10073 (V S), ca. 4,000 m, McVaugh 13820 (US). Mexico: Sierra de las Cruces, Pringle 4145 (US); Ixtaccihuatl, above San Rafael, ca. 4,040 m, Beaman 2817 (UC, US); Ixtacci¬ huatl, La Joya, 3,990 m, Beaman 3514 (US); Ixtacci¬ huatl, 12,000-13,000 ft., Pur pus s.n. (UC), 172 (US), 1596 (UC, US), 3,500 m, Matuda 27574 (US), 4,100 m, Rzedowski 25410 (US); La Cienega, cerca del Llano Grande, Ixtaccihuatl, Rzedowski 29309 (US); arriba de El Salto, Valle de Ayoloco, Ixtaccihuatl, Rzedowski 23465 (LJS); Volcan Ixtaccihuatl, Moore et al. 4526 (UC, US); SE de Ixtaccihuatl, 3,600 m, Matuda 26113, 26114 (US); Popocatapetl, 3,400 m, Sanchez M. 423 (US); between Km 76 and 77 on Amecameca-Popocatepetl road, ca. 3,250 m, Beaman 2060 (UC); Cerro Cabezas, 3,500 m, Lyonnet 1479, Sanchez S. 364 (US); Xitle Chico, Rzedowski 938 (US). Veracruz: Orizaba, 1 1,000 ft., Plantae Mexicanae Liebmann 2508 (US); Munic. Calcahualco, 5 km SW of Jacal, 19°05'N, 97°15'W, 3,450 m, Nee & Diggs 24843 (UC). GUATEMALA. Sacatepequez: Volcan de Agua, 3,400-3,752 m, Pit- tier 24 (US). San Marcos: upper slopes of Volcan Ta¬ cana, 4,100-4,400 m, Steyermark 36134 (US). New Combinations Terpsichore achilleifolia (Kaulfuss) A. H. Smith, comb. nov. Basionym: Polypodium achillei- folium Kaulfuss, Enum. Fil. 116. 1824. Cten- opteris achilleifolium (Kaulfuss) J. Smith. Grammitis achilleifolia (Kaulfuss) R. M. Tryon & A. F. Tryon. Distribution. Southern Brazil. Group 5. A species from Madagascar and the Mascarenes, Ctenopteris torulosa (Baker) Tardieu, is very similar to Terpsichore achilleifolia in blade dissection hut apparently lacks hydathodes or has only faint hy¬ dathodes (Barbara Parris, in litt.). Terpsichore alfarii (Donnell Smith) A. R. Smith, comb. nov. Basionym: Polypodium alfarii Donnell Smith, Bot. Gaz. 33: 262. 1902. Cten- opteris alfarii (Donnell Smith) Copeland. Grammitis alfarii (Donnell Smith) C. V. Mor¬ ton. Polypodium oligosorum Mettenius ex Kuhn, non Klotzsch. Distribution . Costa Rica, Venezuela, Colombia, Ecuador. Group 3. Terpsichore alsophilicola (H. Christ) A. R. Smith, comb. nov. Basionym: Polypodium al¬ sophilicola H. Christ, Bull. Soc. Bot. Geneve, II, 1: 219. 1909, as P. alsophilicolum. Gten- opteris alsophilicola (H. Christ) Copeland. Grammitis alsophilicola (H. Christ) F. Sey¬ mour. Distribution. Costa Rica, Panama, Co¬ lombia?, Ecuador. Group 1. Terpsichore alsopteris (C. V. Morton) A. R. Smith, comb. nov. Basionym: Grammitis al¬ sopteris C. V. Morton, Contr. U.S. Natl. Herb. 38: 112, t. 4. 1967. Distribution. Costa Rica, Panama, Colombia, Ecuador, Peru. Croup 2. Terpsichore amphidasyon (Kunze ex Mettenius) A. R. Smith, comb. nov. Basionym: Polypo¬ dium amphidasyon Kunze ex Mettenius, Abh. Senckenberg. Naturf. Ges. 2: 49. 1857. G ram- mitis amphidasyon (Kunze ex Mettenius) Duek & Lellinger. Xiphopteris amphidasyon (Kunze ex Mettenius) Alston. Distribution . Venezuela. Group 2. Terpsichore angustipes (Copeland) A. R. Smith, comb. nov. Basionym: Gtenopteris angustipes Copeland, Philipp. J. Sci. 84: 404. 1956. Grammitis angustipes (Copeland) Lellinger. Distribution. Colombia. Group ? Terpsichore asplenifolia (L.) A. R. Smith, comb, nov. Basionym: Polypodium asplenifolium L., Sp. PI. 2: 1084. 1753. Gtenopteris aspleni¬ folia (L.) Copeland. Grammitis asplenifolia (L.) Proctor. Distribution. Southern M exico to Panama, Colombia and Venezuela to Bolivia, Antilles. Group 1 . Terpsichore athyrioides (Hooker) A. B. Smith, comb. nov. Basionym: Polypodium athy¬ rioides Hooker, Sp. Fil. 4: 224, t. 27 > B. 1862. Ctenopteris athyrioides (Hooker) Copeland. Grammitis athyrioides (Hooker) C. \ . Morton. Distribution. Peru, Bolivia. Group 2. Polypodium yungense Rosenstock, Repert. Spec. Nov. Regni Veg. 5: 236. 1908. Xiphopteris yungensis (Rosenstock) Crabbe. 486 Novon Terpsichore atroviridis (Copeland) A. R. Smith, comb. nov. Basionym: Ctenopteris atroviridis Copeland, Philipp. J. Sci. 84: 461. 1956. Grammitis atroviridis (Copeland) F. Seymour. Distribution. Nicaragua, Costa Rica, Panama. Group 1 . Terpsichore attenuatissima (Copeland) A. R. Smith, comb. nov. Basionym: Ctenopteris at¬ tenuatissima Copeland, Philipp. J. Sci. 84: 456, t. 1 1. 1956. Grammitis attenuatissima (Cope¬ land) C. V. Morton. Distribution. Ecuador. Group 2. Terpisichore ehrysleri (Copeland) A. R. Smith, comb. nov. Basionym: Ctenopteris ehrysleri Copeland, Philipp. J. Sci. 84: 448. 1956. Grammitis ehrysleri (Copeland) Proctor. Dis¬ tribution. Costa Rica, Panama, Colombia and Venezuela to Bolivia, Brazil, Jamaica, Hispan¬ iola. Group 1 . Terpsiehore eoncinna (A. R. Smith) A. R. Smith, comb. nov. Basionym: Grammitis eoncinna A. R. Smith, Ann. Missouri Bot. Gard. 76: 338. 1989. Polypodia m coneinnum Mettenius ex Kuhn, Linnaea 36: 132. 1869, not Willdenow, Sp. PI., ed. 4. 5: 201. 1810 (= Thelypteris eoncinna (Willd.) Ching). Distribution. Vene¬ zuela. Group 3. Terpsichore cretata (Maxon) A. R. Smith, comb, nov. Basionym: Polypodium cretatum Maxon, Amer. Fern J. 5: 51. 1915. Ctenopteris cre¬ tata (Maxon) Copeland. Grammitis cretata (Maxon) Proctor. Distribution. Cuba, Hispan¬ iola, Jamaica. Group 4. Terpsichore cultrata (Bory ex Willdenow) A. R. Smith, comb. nov. Basionym: Polypodium eul- tratum Bory ex Willdenow, Sp. PI., ed. 4. 5: 187. 1810. Ctenopteris cultrata (Bory ex Willdenow) Copeland. Grammitis cultrata (Bory ex Willdenow) Proctor. Xiphopteris cultrata (Bory ex Willdenow) Schelpe. Distribution. Southern Mexico to Panama, Colombia and Venezuela to Bolivia, Brazil, Antilles; Africa, Madagascar, Mauritius, Reunion, and the Sey¬ chelles Islands. Group 3. Polypodium, elasticum Bory ex Willdenow, Sp. PI., ed. 4. 5: 183. 1810. Ctenopteris elastica (Bory ex Willdenow) Copeland. Xiphopteris elastica (Bory ex Willdenow) Alston. Polypodium reclinatum Brackenridge, Expl. Exp. 16: 11. 1854. Ctenopteris reclinata (Brackenridge) Copeland. Polypodium ciliare Fee, Crypt. Vase. Bresil 1: 94, t. 27, f. 2. 1869. Ctenopteris ciliaris (Fee) Copeland. Polypodium ovalescens Fee, Crypt. Vase. Bresil 1: 94 t. 27, f. 3. 1869. Terpsichore david-smithii (Stolze) A. R. Smith, comh. nov. Basionym: Grammitis david-smithii Stolze, Fieldiana, Bot., n.s. 32: 109. 1993. Distribution. Peru, Bolivia. Group 2. Terpsichore delicatula (M. Martens & Galeotti) A. R. Smith, comb. nov. Basionym: Polypo¬ dium delicatulum M. Martens & Galeotti, Nouv. Mem. Acad. Roy. Sci. Bruxelles 15(5): 35, pi. 7, f. 1. 1842. Ctenopteris delicatula (M. Martens & Galeotti) J. Smith. Grammitis delicatula (M. Martens & Galeotti) F. Sey¬ mour. Distribution. Southern Mexico, Guate¬ mala. Group 3. Terpsichore dependens (Baker) A. R. Smith, comh. nov. Basionym: Polypodium dependens Baker, Syn. Fil. 335. 1867. Grammitis de¬ pendens (Baker) C. V. Morton. Distribution. Colombia, Ecuador, Peru, Bolivia. Group 3. Terpsichore dolorensis (Hieronymus) A. R. Smith, comh. nov. Basionym: Polypodium do- lorense Hieronymus, Bot. Jahrb. Syst. 34: 512. 1905. Grammitis dolorensis (Hieronymus) Lellinger. Distribution. Colombia. Group 1. Terpsichore eggersii (Baker ex Hooker) A. R. Smith, comb. nov. Basionym: Polypodium eg- gersii Baker ex Hooker, Icon. PI. t. 1671. 1886. Grammitis eggersii (Baker ex Hooker) Proctor. Distribution. Lesser Antilles. Group 1. Terpsichore exornans (Maxon) A. R. Smith, comb. nov. Basionym: Polypodium exornans Maxon, Amer. Fern J. 18: 47. 1928. Cten¬ opteris exornans (Maxon) Copeland. Gram¬ mitis exornans (Maxon) Proctor. Distribution. Jamaica. Group 1. Terpsichore gradata (Baker) A. R. Smith, comh. nov. Basionym: Polypodium gradatum Baker, FI. Bras. 1(2): 513. 1870. Ctenopteris gra¬ data (Baker) Copeland. Grammitis gradata (Baker) R. M. Tryon & A. F. I'ryon. Distri¬ bution. Southern Brazil. Group 5. Polypodium schwackei H. Christ in Schwacke, PI. Nov. Mineiras 2: 20. 1900. Ctenopteris schwackei (H. Christ) Copeland. Terpsichore hanekeana (Proctor) A. R. Smith, comh. nov. Basionym: Grammitis hanekeana Proctor, Mem. New York Bot. Gard. 53: 341. 1989. Distribution. Puerto Rico. Group 1. Terpsichore heteromorpha (Hooker & Greville) A. R. Smith, comb. nov. Basionym: Polypo¬ dium heteromorphum Hooker & Greville, Icon. Fil. 1: pi. 108. 1829. Ctenopteris heteromor¬ pha (Hooker & Greville) Copeland. Grammitis Volume 3, Number 4 1993 Smith Terpsichore (Grammitidaceae) 487 heteromorpha (Hooker & Greville) C. V. Mor¬ ton. Xiphopteris heteromorpha (Hooker & Greville) Crabbe. Distribution. Colombia, Ec¬ uador, Peru. Group 3. Terpsichore immixta (Stolze) A. R. Smith, comb, nov. Basionym: Grammitis immixta Stolze, Fieldiana, Bot., n.s. 32: 115. 1993. Distri¬ bution. Peru. Group 3. Terpsichore jamesonioides (Fee) A. R. Smith, comb. nov. Basionym: Polypodium jameson¬ ioides Fee, Mem. Soc. Sci. Hist. Nat. Stras¬ bourg 5 [Mem. Foug. 7]: 59, t. 21, I. 4. 1857. Ctenopteris jamesonioides (Fee) Copeland. Grammitis jamesonioides (Fee) C. V. Morton. Distribution. Costa Rica, Panama, Venezuela and Colombia to Peru. Group 3. Ctenopteris nudipes Copeland, Philipp. J. Sci. 84: 405, t. 4. 1956. Terpsichore jenmanii (Underwood ex Maxon) A. R. Smith, comb. nov. Basionym: Polypo¬ dium jenmanii Underwood ex Maxon, Contr. U.S. Natl. Herb. 16: 62. 1912. Ctenopteris jenmanii (Underwood ex Maxon) Copeland. Grammitis jenmanii (Underwood ex Maxon) Proctor. Distribution. Jamaica and Hispaniola. Group 1 . Terpsichore kegeliana (Kunze) A. R. Smith, comb. nov. Basionym: Polypodium kegelian- um Kunze, Linnaea 21: 210. 1848. C tenop- teris kegeliana (Kunze) K. U. Kramer. Gram¬ mitis kegeliana (Kunze) Lellinger. Distribution. Surinam, French Guiana. Group ? Terpsichore lanigera (Desvaux) A. R. Smith, comb. nov. Basionym: Polypodium lanigerum Desvaux, Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Naturk. 5: 316. 1811. Ctenopteris lanigera (Desvaux) Cope¬ land. Grammitis lanigera (Desvaux) G. V. Morton. Xiphopteris lanigera (Desvaux) Crabbe. Distribution. Costa Rica, Panama, Co¬ lombia to Bolivia, Hispaniola, Lesser Antilles. Group 3. Terpsichore laxa (C. Presl) A. R. Smith, comb, nov. Basionym: Polypodium laxum G. Presl. Reliq. Haenk. 1: 23, t. 4, f. 1. 1825. Gram¬ mitis laxa (C. Presl) C. V. Morton. Distribu¬ tion. Ecuador, Peru, Bolivia. Group 3. Ctenopteris contacta Copeland, Philipp. J. Sci. 84: 447. 1956. Terpsichore lehmanniana (Hieronymus) A. R. Smith, comb. nov. Basionym: Polypodium leh- mannianum Hieronymus, Bot. Jahrb. Syst. 34: 513. 1904. Ctenopteris lehmanniana (Hi¬ eronymus) Copeland. Distribution. Guatemala to Panama, Colombia, Ecuador. Group 1 . Terpsichore leucosticta (J. Smith) A. R. Smith, comb. nov. Basionym: Ctenopteris leucosticta .1. Smith, Hist. Fil. 185. 1875, a norn. nov. for Polypodium leucosticton Fee, not Kunze ex Klotzsch (1847), Gen. Fil. 240. 1852. Poly¬ podium longiusculum C. Christensen, norn. nov. for P. leucosticton Fee. Ctenopteris lon- giuscula(C. Christensen) Copeland. Grammitis leucosticta (J. Smith) C. V. Morton. Distri¬ bution. Venezuela, Ecuador, Peru. Group 2. Terpsichore liogieri (Proctor) A. R. Smith, comb, nov. Basionym: Grammitis liogieri Proctor, Mem. New York Bot. Card. 53: 341. 1989. Distribution. Puerto Rico. Group 1. Terpsichore longa (C. Christensen) A. R. Smith, comb. nov. Basionym: Polypodium longum G. Christensen, Index. Fil. 541. 1906, a norn. nov. for P. alternifolium Hooker, Sp. Fil. 4: 222, t. 277A, 1862, not Willdenow, Sp. PL, ed. 4. 5: 168. 1810 (= Microsorum alterni¬ folium (Willdenow) Copel). Ctenopteris longa (G. Christensen) Copeland. Xiphopteris longa (C. Christensen) Alston. Distribution. Costa Rica, Ecuador. Group 3. Terpsichore longisetosa (Hooker) A. R. Smith, comb. nov. Basionym: Polypodium longise- tosum Hooker, Sp. Fil. 4: 225. 1864. Distri¬ bution. Costa Rica, Panama, Venezuela, Ec¬ uador to Bolivia. Group 5. Polypodium myriophyllum Mettenius ex Hooker & Ba¬ ker, Syn. Fil. 338. 1868. Ctenopteris myriophylla (Mettenius ex Hooker & Baker) Copeland. Gram¬ mitis myriophylla (Mettenius ex Hooker & Baker) C. V. Morton. Xiphopteris myriophylla (Mettenius ex Hooker & Baker) Crabbe. Polypodium piligerum Hooker (Hooker’s Icon. PI. 4: t. 321. 1841), type from Ecuador, appears to be this species and an earlier name, but the rhizome was described as with scales “nigro-fuscen- tibus, nitidis.” Specimens of T. longisetosa seen from Mesoamerica, as well as from Venezuela and Ecuador, have orange-brown, rather dull sc ales. Morton (1967) did not account for Polypodium piligerum in his treatment of Grammitidaceae for Ecuador. Terpsichore mollissima (Fee) A. R. Smith, comb, nov. Basionym: Polypodium mollissimum Fee, Hist. Foug. Antill. [Mem. Foug. 1 1]: 47, t. 12, f. 2. 1866. Ctenopteris mollissima (Fee) Cope¬ land. Grammitis mollissima (Fee) Proctor. 488 Novon Distribution. Southern Mexico to Panama, Venezuela, Ecuador, Antilles. Group 3. Terpsichore paulistana (Brade & Rosenstock) A. R. Smith, comb. nov. Basionym: Polypo¬ dium paulistanum tirade & Rosenstock, Arch. Inst. Biol. Veg. Rio Janeiro 2: 3, t. 1(4), 4. 1935. Distribution. Southern Brazil. Croup 2. Terpsichore pichinchae (Sodiro) A. R. Smith, comb. nov. Basionym: Polypodium pichinchae Sodiro, Crypt. Vase. Quit. 329. 1893. Cten- opteris pichinchae (Sodiro) Copeland. Gram- mitis pichinchae (Sodiro) C. V. Morton. Dis¬ tribution. Ecuador. Group 2. Terpsichore pichinchense (Hieronymus) A. R. Smith, comb. nov. Basionym: Polypodium pi¬ chinchense Hieronymus, Bot. Jahrb. Syst. 34: 506. 1904. Polypod ium ecuadorense C. Christensen, nom. superfl. Grammitis pi¬ chinchense (Hieronymus) C. V. Morton. Dis¬ tribution. Ecuador, Peru. Group 2. Terpsichore semihirsuta (Klotzsch) A. R. Smith, comb. nov. Basionym: Polypodium semihir- sutum Klotzsch, Linnaea 20: 379. 1847. Cten- opteris semihirsuta (Klotzsch) Copeland. Grammitis semihirsuta (Klotzsch) Proctor. Distribution. Southern Mexico to Panama, Ja¬ maica, Hispaniola, Venezuela and Colombia to Bolivia, Brazil. Croup 2. Polypodium gratum Fee, Crypt. Vase. Bresil 1: 242, t. 76, f. 2. 1869. Terpsichore senilis (Fee) A. R. Smith, comb. nov. Basionym: Polypodium senile Fee, Mem. Soc. Sci. Hist. Nat. Strasbourg 5 [Mem. Foug. 7]: 60, t. 25, 1. 1. 1857. Ctenopteris senilis (Fee) Copeland. Grammitis senilis (Fee) C. V. Mor¬ ton. Distribution. Southern Mexico to Panama, Colombia, Venezuela, Ecuador, Peru, and Bo¬ livia. Group 3. Polypodium subflabelliforme Rosenstock, Repert. Spec. Nov. Regni Veg. 7: 306. 1909. Ctenopteris subfa- belliformis (Rosenstock) Copeland. Grammitis subflabelliformis (Rosenstock) C. V. Morton. Terpsichore staheliana (Posthumus) A. R. Smith, comb. nov. Basionym: Polypodium stahelian- um Posthumus, Recueil Trav. Bot. Neerl. 23: 401. 1928. Ctenopteris staheliana (Posthu¬ mus) K. U. Kramer. Grammitis staheliana (Posthumus) Lellinger. Distribution. Nicara¬ gua, Costa Rica, Panama, Surinam, Venezuela, Colombia? Group 1. Terpsichore subtilis (Kunze ex Klotzsch) A. R. Smith, comb. nov. Basionym: Polypodium sub¬ tile Kunze ex Klotzsch, Linnaea 20: 375. 1847. Ctenopteris subtilis (Kunze ex Klotzsch) J. Smith. Grammitis subtilis (Kunze ex Klotzsch) C. V. Morton. Distribution. Southern Mexico to Panama, Venezuela, Colombia, Ecuador. Group 4. Terpsichore taxifolia (L.) A. R. Smith, comb, nov. Basionym: Polypodium taxifolium L.,Sp. PI. 2: 1086. 1753. Ctenopteris taxifolia (L.) Copeland. Grammitis taxifolia (L.) Proctor. Distribution. Costa Rica, Panama, Antilles, Venezuela, Ecuador, Surinam, Brazil. Croup 2. Polypodium Vherminieri Fee var. costaricense Rosen¬ stock, Repert. Spec. Nov. Regni Veg. 22: 17. 1925. Terpsichore turrialbae (H. Christ) A. R. Smith, comb. nov. Basionym: Polypodium turrialbae H. Christ, Bull. Soc. Roy. Bot. Belgique 35: Mem. 226. 1896. Ctenopteris turrialbae (H. Christ) Copeland. Grammitis turrialbae (H. Christ) F. Seymour. Distribution. Costa Rica, Panama. Group 3. Terpsichore variabilis (Mettenius ex Kuhn) A. R. Smith, comb. nov. Basionym: Polypodium variabile Mettenius ex Kuhn, Linnaea 36: 1 33. 1869. Grammitis variabilis (Mettenius ex Kuhn) C. V. Morton. Distribution. Colombia, Ecuador, Peru. Group 3. Terpsichore xanthotrichia (Klotzsch) A. R. Smith, comb. nov. Basionym: Polypodium xanthotrichium Klotzsch, Linnaea 20: 376. 1847. Grammitis xanthotrichia (Klotzsch) Duek & Lellinger. Distribution. Venezuela. Croup 3. Terpsichore zeledoniana (Lellinger) A. R. Smith, comb. nov. Basionym: Grammitis zeledoniana Lellinger, Proc. Biol. Soc. Wash. 98: 383. 1985, nom. nov. for Polypodium taxifolium L. var . fragillimum H. Christ, Bull. Herb. Bois- sier, ser. 2. 4: 1 103. 1904. Distribution. Costa Rica, Panama. Group 2. Literature Cited Bishop, L. E. 1988. Ceradenia, a new genus of Gram- mitidaceae. Amer. Fern J. 78: 1-5. - . 1989. Zygophlebia, a new genus of Gram- mitidaceae. Amer. Fern J. 79: 103-118. - & A. R. Smith. 1992. Revision of the fern genus Enterosora (Grammitidaceae) in the New- World. Syst. Bot. 17: 345 362. Copeland, E. B. 1 956 [“1955”]. Ctenopteris in Amer¬ ica. Philipp. J. Sci. 84: 381-473. Evans, A. M. 1963. New chromosome observations in the Polypodiaceae and Grammitidaceae. Caryologia 16: 671-677. Hooker, J. D. 1886. Hooker’s Icon. PI. 17: pi. 1673. Lellinger, D. B. 1989. The ferns and fern-allies of Costa Volume 3, Number 4 1993 Smith Terpsichore (Grammitidaceae) 489 Rica, Panama, and the Choco (Part 1: Psilotaceae through Dicksoniaceae). Pteridologia 2A: 1-364. Mickel, J. T. 1973. Fungi on ferns. New and Views 9: 4. - & J. M. Beitel. 1988. Pteridophyte flora of Oaxaca, Mexico. Mem. New York Bot. Card. 46: 1-568. Morton, C. V. 1967. The genus Grammitis in Ecuador. Contr. U.S. Natl. Herb. 38: 85-123. Parris, B. S. 1990. Noteworthy species of Grammiti¬ daceae from South-East Asia. Hooker’s Icon. PI. 40: 1-129. Proctor, G. R. 1977. Pteridophyta. Pp. 1-414 in R. A. Howard (editor), Flora of the Lesser Antilles, vol. 2. Arnold Arboretum, Jamaica Plain, Massachusetts. - . 1985. Ferns of Jamaica. British Museum (Nat¬ ural History), London. - . 1989. Ferns of Puerto Rico and the Virgin Islands. Mem. New York Bot. Gard. 53: 1-389. Schelpe, E. A. C. L. E. 1969. Reviews of tropical African Pteridophyta 1. Contr. Bolus Herb. 1: 1- 132. Smith, A. R. 1981. Pteridophytes. Pp. 1-370 in D. E. Breedlove (editor). Flora of Chiapas, Part 2. Cal¬ ifornia Academy of Sciences, San Francisco. - . 1992. A review of the fern genus Micropo- lypodium (Grammitidaceae). Novon 2: 419-425. - . 1993. Phytogeographic principles and their use in understanding fern relationships. J. Biogeog¬ raphy (in press). - & R. C. Moran. 1992. Melpomene , a new genus of Grammitidaceae (Pteridophyta). Novon 2: 426-432. - , - & L. E. Bishop. 1991. Lellingeria, a new genus of Grammitidaceae. Amer. Fern J. 81: 76 88. Stokey, A. G. & L. R. Atkinson. 1958. The gameto- phyte of the Grammitidaceae. Phytomorphology 8: 391-403. Stolze, R. G. 1981. Ferns and fern allies of Guatemala. Part II. Polypodiaceae. Fieldiana, Bot., n.s. 6: 1 522. Tardieu-Blot, M. L. 1960. Polypodiacees (sensu lato). Blechnacees-Polypodiacees. In: H. Humbert, Flore de Madagascar et des Comores (Plantes Vasculaires). Fam. 5. 2: 1-122, figs. 1-26. Paris. Tryon, A. F. & B. Lugardon. 1991. Spores of the Pteridophyta. Springer-Verlag, New York. Wagner, F. S. 1985. Bilateral spores in New World grammitid ferns. Amer. Fern J. 75: 6-11. Walker, T. G. 1966. A cytotaxonomic survey of the pteridophytes of Jamaica. Trans. Roy. Soc. Edin¬ burgh 66: 169-237. Notes on Elaeagia myriantha , Comb. Nov. (Rubiaceae) Charlotte M. Taylor and Harry E. Hammel Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. Abstract. Observations of recent fruiting collec¬ tions of Simira myriantha (Standley) Steyerrnark, a tree found in wet mid-elevation forests from south¬ ern Costa Rica to Peru, support the following trans¬ fer of this species to Elaeagia. Sichingia myriantha Standley was originally de¬ scribed from flowering material from northern Co¬ lombia, hut Standley himself later erected the mono- typic genus Holtonia to accommodate this species, as Holtonia myriantha (Standley) Standley. Even though classification within the Rubiaceae has been strongly dependent on knowledge of mature fruits, which were lacking for this taxon, Standley (1938) placed Holtonia in the Rondeletieae along with Sich¬ ingia Willdenow. Steyerrnark (1972) determined that Sichingia is a synonym of Simira Auhlet and published new combinations in Simira for all Sichingia species, including Sichingia myriantha. In that work he considered the placement of this species in Elaeagia, hut rejected this conclusion based on its corolla aestivation. He stated only that the aestivation dif¬ fered, hut did not describe its condition. Steyerrnark (1974) later presented a detailed description of this species, although the mature fruits were still un¬ known and his description of them was sketchy. Recent botanical exploration, particularly in Co¬ lombia, has made more material of this species avail¬ able, including mature fruiting specimens. The fruits are small (2-3 mm long), loculicidal woody capsules that dehisce only partially, apparently allowing a “salt-shaker'' dispersal of the numerous small an¬ gled seeds. The fruits and seeds are decidedly not those of Simira, which has larger (> 1 cm diam.), completely dehiscent capsules with broad, flat, sa¬ maralike seeds. Rather, they are characteristic of Elaeagia Weddell and Warszewiczia Klotzsch, sympatric genera of neotropical trees. These genera are distinguished in part by their corolla aestivation, convolute in Elaeagia and imbricate in Warszew¬ iczia. Aestivation of the species in question is nearly valvate to slightly convolute, more like that of Elae¬ agia, rather than imbricate as originally implied by Standley when he described this species in Sichin¬ gia. “ Holtonia ” also shares with Elaeagia species rounded to truncate stipules (vs. acute in Warszew¬ iczia), resinous terminal buds (vs. nonresinous in Warszewiczia), glomerulate cymes arranged in py¬ ramidal paniculate inflorescences, and white, short tubular-funnelform corollas with exserted stamens that are barbate near their insertion (similar in War¬ szewiczia). Thus, because of its fruits “ Holtonia myriantha" is not a Simira hut rather an Elaeagia, and a slight variation in corolla aestivation does not warrant maintaining Holtonia as a separate genus. We present below an expanded morphological description and geographic range for this species, and a selected list of specimens, in addition to types, on which these are based. Representative specimens from Venezuela were cited by Steyerrnark (1974). This species is distinguished within Elaeagia by its persistent, relatively short rounded to truncate stip¬ ules. It appears to be most closely related to E. cuhensis Britton, which shares membranaceous, rel¬ atively short stipules that persist on several distal nodes. However, the stipules of E. cuhensis are ultimately completely deciduous and shorter, and the corolla is campanulate. Elaeagia Weddell, Hist. Nat. Quinquinas 24. 1849. TYPE: Elaeagia utilis W eddell. Holtonia Standley, Trop. Woods 30: 37. 1932, syn. nov. TYPE: Holtonia myriantha (Standley) Standley, based on Sichingia myriantha Standley. Elaeagia myriantha (Standley) C. M. Taylor & Hammel, comb. nov. Basionym: Sichingia my¬ riantha Standley, Publ. Field Columbian Mus., Bot. Ser. 7(1): 27. 1930. Holtonia myriantha (Standley) Standley, Trop. Woods 30: 37. 1932. Simira myriantha (Standley) Steyerrnark, Mem. New York Bot. Card. 23: 306. 1972. TYPE: Colombia. Magdalena: Las Nubes road, region of Santa Marta, 1,200 m, 3 Dec. 1898, //. H. Smith 1810 (holotype, F; isotypes, MO, NY not seen, US not seen). Deppea panamensis Dwyer, Ann. Missouri Bot. Gard. 67: 145. 1980, syn. nov. TYPE: Panama. Panama: 5 10 km NE of Altos de Pacora, Mori & Kallunki 4965 (holotype, MO). Trees to 20 m tall, buds resinous; bark gray, rough; stems pilosulous to glabrescent. Leaves op- Novon 3: 490-491. 1993. Volume 3, Number 4 1993 Taylor & Hammel Elaeagia myriantha 491 posite, elliptic, 8-22 cm long, 2.5-10 cm wide, acute at base, acute to shortly acuminate at apex, chartaceous to stiffly so, glabrous above, below gla¬ brous to puberulent or pilosulous, frequently barbate with tufted domatia in vein axils; secondary veins 8-12 pairs; petioles 7-25 mm long; stipules per¬ sistent, inter- and intrapetiolar, resinous when young, glabrous to pilosulous, 2-5 mm long, subtruncate to lobed for ca. % of their length, not splitting to base, lobes rounded to obtuse. Inflorescences ter¬ minal, paniculate, pyramidal, peduncles 0 (and in¬ florescences tripartite) to 1-7 cm long, panicles 6- 1 1 cm long, 7-14 cm wide, bracts triangular, those subtending branches 13 mm long and those sub¬ tending flowers ca. 0.5 mm long, branches and hracts glabrous to pilosulous; flowers sessile or with pedicels to 1 mm long in glomerules of 2-8, homostylous, proterogynous; ovary turbinate, 1-1.5 mm long; calyx limb ca. 0.5 mm long, dentate for ca. % of its length, lobes 5, acute to obtuse; corolla shortly tubular-funnelform, white to creamy yellow, gla¬ brous externally, internally densely white-villous, tube 3.5-4 mm long, lobes 5, ca. 0.5 mm long, acute to obtuse, aestivation valvate to slightly convolute; stamens 5, inserted near apex of corolla tube, the anthers ca. 1.5 mm long, exserted by ca. 1 mm; stigmas 2, 1-1.5 mm long, linear, recurved, shortly exserted. Fruit capsular, ellipsoid to turbinate, woody, 2-3 mm long, 1.5-2. 5 mm wide, loculicidal, basip- etally dehiscent, opening for ca. '/2 of its length; seeds angled, pale brown, 0.2-0. 5 mm diarn. Distribution and habitat. Wet forests at (400 )1 ,600-2, 200(-2, 350) m, southern Costa Rica to Andean Venezuela and northern Peru, most fre¬ quently collected in northern and central Colombia. Phenology. Collected in flower January, March, April, and September to November, in fruit March, April, June, July, and October to December. Representative specimens studied. COLOMBIA. An- tioquia: autopista Medillin Bogota, sector rio Samana- rio Claro, San Luis, Hernandez et al. 522 (COL). Cauca: Popayan, Timbio en Hatoviejo, Perez & Cuatrecasas 6107 (COL). Magdalena: Campano, Sierra Nevada de Santa Marta, Minca, 1 1°08’N, 74°01'W, Gentry & Cuadros 64762 (MO). Quindio: municipio Salento, ver- eda rio Arriba, Hacienda El Cairo, Arbelaez et al. 2563 (COL, HLJQ). Valle: hoya del rio Cali, rio Pichinde, en Los Carpatos, Cuatrecasas 21625 (CUVC, VALLE). COSTA RICA. Puntarenas: Reserva Biologica Carara, sitio Bijagual, Zuniga 407 (CR). San Jose: Zona Pro- tectora Cerro Turrubares, Hammel et al. 18967 (CR); El General, Skutch 2387 (MO). ECUADOR. INapo: car- retera Hollin-Loreto, Km 40-50, Hurtado 777 (MO). Pastaza: Hacienda San Antonio de Baron von Humboldt, 2 km al norte de Mera, 1°27'S, 78°06'W, Neill et al. 6069 (MO). PERU. Cusco: Paucartambo, Atalaya a Chontachaca, alrededor de la carretera en la ruta hacia Shintuya, Nunez 8074 (MO). Acknowledgments. This work was stimulated by preparation for the Flora Costaricensis and the Manual of the Plants of Costa Rica, and was sup¬ ported in part by National Science Foundation grant BSR-9006449. We thank the Asociacion Colom- biana de Herbarios for making available Colombian collections of this species, and David Lorence for valuable comments. Literature Cited Standley, P. C. 1938. Flora of Costa Rica (Rubiaceae). Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1264- 1380. Steyermark, J. A. 1972. Simira. In: B. Maguire & Collaborators, The botany of the Guiana Highlands. Part. IX. Mem. New York Bot. Card. 23: 299-307. - . 1974. Rubiaceae. In: T. Lasser (editor), Flora de Venezuela 9: 1-2070. Ministerio de Agricultura y Cria, Direccion de Recursos Naturales Renovables, Instituto Botanico, Caracas. A New Species of Faramea (Rubiaceae) from Amazonian Peru Charlotte M. Taylor Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. ABSTRACT. A new species of Rubiaceae from Am¬ azonian Peru, Faramea vasquezii C. M. Taylor, described and illustrated here, appears to be related to F. capillipes and F. quadricostata. Faramea Aublet is a neotropical genus of perhaps 100 species distributed from southern Mexico and the Antilles to Paraguay and Bolivia. It is considered closely allied to Coussarea Aublet, and these two genera comprise the tribe Coussareae. Faramea is distinguished within the Rubiaceae by its one-locular ovary with separate basal ovules and fruits usually with only one developed seed. Additionally, Faramea can usually be recognized by its stipules that ter¬ minate interpetiolarly in a triangular, usually aristate segment, branches with the leaves usually held in a distichous attitude (and the internodes twisted ca. 90° to accommodate this arrangement), white or frequently bright blue corollas usually with a well- developed tube, subglobose to frequently strongly oblate leathery fruits, and seeds with a chartaceous testa. The aristae of the stipules frequently are elon¬ gated and crossed at the stem apex. The leaves of a few species have pronounced brochidodromus ve¬ nation similar to that of many Melastomataceae, with well-developed submarginal veins and second¬ ary veins spreading at right angles from the midrib; however, the venation of Faramea vasquezii C. M. Taylor is more typical of Faramea. During a review of the Rubiaceae known from Peru (Taylor & Pool, 1993), the following new species was discovered. Faramea vasquezii C. M. Taylor, sp. nov. TYPE: Peru. Loreto: Maynas, Alpahuayo, Estacion II AP, 20 Oct. 1984, R. Vdsquez & G. Criollo 5785 (holotype, MO; isotypes, AMAZ, USM). Figure 1. Faramea capillipedi Mueller Argoviensis affinis, sed ab ea internodiis costatis complanatis, foliis minoribus nerviis secundariis paucioribus munitis, lobulis calycinis longioribus ac lobulis corollinis brevioribus differt. Glabrous shrubs or small trees to 4 m tall (flow¬ ering at 0.3 m tall); stems strongly flattened and costate, the distalmost internodes 3-6 mm wide below the leaves. Leaves opposite; blades elliptic to rarely somewhat lanceolate, 13-23 cm long, 3-9.5 cm wide, at apex acuminate with tip 1 .2-2 cm long, at base cuneate to usually acute or sometimes at¬ tenuate, subcoriaceous, glabrous; secondary veins 10-14 pairs, looping to interconnnect, with l(-2) intersecondary veins usually well developed, without domatia, with the costa, secondary, and intersecon¬ dary veins prominulous above and below, the lesser venation reticulate and visible to slightly raised; pet¬ ioles stout, 3-6 mm long; stipules with sheath 0.3- 1 mm long, abruptly contracted to an awn 3-15 mm long. Inflorescences axillary and occasionally also terminal, 4.5-5 cm long, 4-7 cm wide, fasci¬ cled, ebracteate, peduncles 2-4 per axil, 12-28 mm long, cymes once or usually twice divided into 2-3 branches or pedicels, pedicels 6-18 mm long, filiform, flexuous; calyx limb with truncate tube 0.3- 0.5 mm long, lobes 4, narrowly triangular, 0.6- 1.5 mm long, acute; corollas funnelform, white to yellow, glabrous, tube 2-3 mm long, lobes 2.5-3 mm long, triangular to lanceolate. Fruiting pedicels markedly thickened distally; fruits subglobose, 6-7 mm long, 7-9 mm wide, smooth, becoming red to purple and then hlack. Distribution and habitat. Amazonian Peru, in upland or seasonally flooded forest on sandy soils at 100-400 m, most frequently collected in primary forest. Phenology. Collected in flower February, April, August, and October to December, in fruit January to May, July, August, and October to November. This species is distinguished by its usually axillary inflorescences with relatively long flexuous pedicels, strongly flattened and costate internodes, and co¬ rollas with the tube shorter than or equal to the lobes. It is similar to Faramea capillipes Mueller Argoviensis in general aspect and inflorescence structure, and these species are probably closely related. Faramea capillipes differs from F. vas¬ quezii in its slightly costate but generally rounded internodes, papyraceous to membranaceous leaf blades 5-12 cm long with 7-8 secondary veins, calyx limb with lobes 0.1 -0.2 mm long, and corollas with lobes 5-6 mm long. This new species is also similar to F. quadricostata Bremekamp (emend. Steyermark, 1967), which shares a similar general Novon 3: 492-493. 1993. Volume 3, Number 4 1993 Taylor Faramea vasquezii 493 Figure 1. Faramea vasquezii C. M. Taylor. — A. Habit, drawn from Vasquez 5710. — B. Fruit with pedicel, drawn from Vasquez 3974. Both to same scale. aspect and flattened costate internodes, but differs in its leaf blades 6 12 cm long, terminal inflores¬ cences with peduncles 0-5 mm long, truncate calyx limb 1.3- 1.5 mm long, and corollas with tubes 20 23 mm long and lobes 9.5-14 mm long. This species is named in honor of Rudolfo Vasquez of the Missouri Botanical Garden, the accomplished Peruvian botanist who collected most of the material studied. Additional collections examined. PERU. Loreto: Lo¬ reto, Nauta, rio Maranon, 4°29'W, 73°33'W, H. Vasquez & N. Jaramillo 3445 (AMAZ, MO); Maynas, Cahuide, rio Itaya, Vasquez & Jaramillo 5770(AMAZ, MO, USM); Maynas, Amazonas, ExplorNapo and Explorama Camps, rio Sucusari, ca. 3°1 5-30'S, 72°55'W, Gentry et al. 27695 (MO), 31594 (MO), 74316 (MO), Pipoly 14176 (AMAZ, MO), Vasquez & Jaramillo 13111 (AMAZ, MO); Maynas, Iquitos, rio Nanay, area of Mishana and Callicebus Biological Reserve, ca. 4°50'S, 73°30'\Y, Da¬ vidson 5232 (MO), Foster 4354 (MO), Gentry et al. 31610(MO ), Pipoly et al. 14915 (AMAZ, MO), Solomon 3572 (MO), Vasquez et al. 617 (AMAZ, MO, USM), 14196 (AMAZ, MO, USM); Maynas, Iquitos, near Puerto Almendras, ca. 3°45-50'S, 73°21-25'W, Croat 19036 (MO), Diaz <£ Jaramillo 275 (MO), Revilla 2309 (MO), Ruiz 1298 (MO), Vasquez & Jaramillo 159 (AMAZ, MO, USM), 3974 (AMAZ, MO), 6246 (AMAZ, MO), 6516 (AMAZ, MO), 8658 (AMAZ, MO), 10192 (AMAZ, MO), l asquez & Soto 13732 (AMAZ, MO); Maynas, Pebas, rio Ampiyacu, Revilla 970 (MO); Ucayali, Sa- puena, Jenaro Herrera CDJH-IIAP, 4°55'S, 73°45’\^ , Vasquez et al. 12013 (MO). Pasco: Oxapampa, Palcazu Valley, Iscozach, 10°12'S, 75°15'W, Foster 7974 (MO). Acknowledgments. I thank R. Vasquez and J. Pipoly for helpful comments and discussion. Literature Cited Steyermark, J. A. 1967. Faramea. In: B. Maguire & Collaborators, Flora of the Guayana Highlands. Mem. New York Bot. Card. 17: 371-396. Taylor, C. M. & A. Pool. 1993. Rubiaceae. In: L. Brako & J. Zarucchi, Catalogue of the Flowering Plants and Gymnosperms of Peru. Monogr. Syst. Bot. Missouri Bot. Gard. 45. Ancistrocladus korupensis (Ancistrocladaceae): A New Species of Liana from Cameroon Duncan W. Thomas Department of International Research and Development, Oregon State University, Corvallis, Oregon 97331, U.S.A. Roy E. Gereau Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A. ABSTRACT. Ancistrocladus korupensis, a species here described Irom the Southwest Province of Cam¬ eroon and adjacent Nigeria, is distinguished within the genus by sessile leaves, short petals, and long¬ winged fruits. The new species was investigated after michellamine B, a unique naphthyl isoquinoline al¬ kaloid isolated from a sterile Ancistrocladus leaf collection, showed in vitro activity against HIV-1 and HIV-2. Subsequent collections showed that the liana was undescribed and apparently narrowly en¬ demic to the Korup area. Ancistrocladus Wallich (norm conserv.) is a ge¬ nus of 19 known species: 9 in tropical Asia (Dyer, 1874; Gagnepain, 1909; Craib, 1925; Steenis, 1948; Ramamoorthy, 1976; Harriman, 1987) and 10 in tropical Africa (Pellegrin, 1951; Hutchinson & Dal- ziel, 1954; Leonard, 1982, 1986). The resemblance of the winged fruits of Ancistrocladus spp. to those of the Dipterocarpaceae Blume has led some authors (e.g., Oliver, 1868; Dyer, 1874) to include the genus in the latter family. However, Cilg (1925) noted that the resemblance is strictly superficial and con¬ sidered the Ancistrocladaceae Walpers (nom. con¬ serv.) to be a monogeneric family in the Parietales. The results of anatomical (Metcalfe, 1952; Schmid, 1964; Gottwald & Parameswaran, 1968) and pal- ynological (Erdtman, 1958) investigations support an affinity between Ancistrocladaceae and the en¬ demic African family Dioncophyllaceae Airy Shaw; Cronquist (1981, 1988) treated the two as closely allied families of the Violates. The Ancistrocladaceae are distinguished within the Violales (Cronquist, 1981) and among all of the dicotyledonous families of central Africa (Robyns, 1958) by the following diagnosis: plants lianas, not succulent, climbing by hooked apices of sympodial branches; leaves alternate (but often crowded in rosettes on flowering shoots), simple, entire, not sheathing at base, the blades symmetric; stipules present, very small and caducous; flowers in ra¬ cemes, spikes, or dichotomously branched panicles; sepals 5, accrescent and winglike in fruit; petals distinct or slightly connate at base; extrastaminal corona lacking; stamens 5 or 1 0— 15, erect in bud, the filaments slightly connate at base; ovary half¬ inferior, unilocular; ovule solitary, basilateral; fruit dry, indehiscent; endosperm present, ruminate. An¬ cistrocladus can sometimes be confused with the unrelated genus Hugonia L. (Hugoniaceae or Lin- aceae), but species of the latter genus can be veg- etatively distinguished by the usually pale stems, leaves usually toothed, less crowded, and with nu¬ merous more prominent secondary veins, and hooks borne on normal branches. Of the ten species of Ancistrocladus previously described from tropical Africa, six occur in tbe ev¬ ergreen forests near the Atlantic coast from Gabon to Sierra Leone (Pellegrin, 1951; Hutchinson & Dalziel, 1954), three are apparently restricted to central and northern Za'ire (Leonard, 1982), and one is endemic to the Buda Forest of coastal Kenya (Leonard, 1986). The new species herein described is apparently endemic to the immediate vicinity of the Korup National Park in the Southwest Province of Cameroon, including the adjacent part of Cross River State of southeastern Nigeria. Ancistrocladus korupensis D. W. Thomas & Gereau, sp. nov. TYPE: Cameroon. Southwest Prov.: Ndian Division, 0.6 km E of confluence of Mededibe (Moliba) and Ndian (Mana) Rivers, 90 m, 5°02'N, 8°53'E, 4 Mar. 1993, Gereau, Thomas, F. Namata & K. Jato 51 HO [infruc- tescences and fallen fruits] (holotype, MO; is¬ otypes, K, P, SCA, YA). Figure 1. Plantae omnino glabrae. Caules adulti parce ramosi; ramis principalibus elongatis, rectis subcurvatisve; ramis lateralibus extra-axillaribus, uncos gerentibus. Folia adulta sessilia, ex forrnis duabus constantia: folia ramorum prin- cipalium remote disposita, ex ellipticis oblonga, basi cu- neata, apice rotundata, venis secundariis paucijugis; folia Novon 3: 494-498. 1993. 5 mm Volume 3, Number 4 1993 Thomas & Gereau Ancistrocladus korupensis 495 Figure 1. Ancistrocladus korupensis D. W. Thomas & Gereau. — A. Lateral branch with rosette leaves and inflorescence. — B. Flower. — C, D. Fruit. Based on: A, B, Jato s.n. (Nov. 1992); C, D, Gereau et al. 5177. 496 Novon ramorum lateraliurn in rosulas apicales crebra, oblanceo- lata, (8-)l 4.5-33. 1 x (2.9-)4.6-8.5 cm, basi ex anguste cuneatis ad longe cuneata, apice ex acutis rotundata, rnargine prope basim valde revoluta, sursum nonnihil re- voluta, venis secundariis paribus 8-18 dispositis, brochi- dodromis; foliorum omnium costa infra prominente, in herbario supra parum impressa, venis secundariis utrinque prominulis, venis tertiariis tenue reticulatis. Inflorescentia ex panicula dichotoma laxa inter rosulas foliorum portata constans; pedunculo extra-axillari, (4-)5.5-9.8 cm longo; pedicellis 6. 5-8. 5 mm longis, sub calyce ca. 2 mm ar- ticulatis. Flores sepalis ex suborbicularibus oblongis, man- ifeste dimorphis, majoribus tribus ca. 5x4 mm, mino- ribus duobus ca. 2.5 x 3.5 mm, basi truncatis, apice rotundatis, sub anthesi ad ovarii basim breve decurren- tibus; petalis suborbicularibus quam sepalis brevioribus, prope basim convexis incrassatisque; staininibus 10 in verticillos duos dispositis, filamentis validis, internis ca. 1 mm longis, externis perbrevibus, antheris ca. 1 mm longis; stylis 3, distinctis, ca. 2 mm longis, distaliter incrassatis. Fructus hebetate brunneus; sepalis tenuibus, inaequalibus, majoribus spathulatis 3. 4-4. 8 x 1. 3-2.1 cm, minoribus ex anguste oblongis spathulatis 1.1 -2. 8 x 0.5- 1.1 cm, alarum marginibus 5 ad basim fructus decurrentibus cos¬ tas prominentes 5 formantibus; fructus corpu turbinato, receptaculo supra alas inflato tholiformique; styli basi per- sistente accrescente conica; pericarpio crasso sicco cori- aceo; semine diametro ca. 8 mm. Plants glabrous throughout. Saplings unbranched or very sparingly branched, to 4 m high, lacking hooked lateral branches; sapling leaves long-lived, in dense terminal rosettes, 38.8 75. 1 x 1 1.0 13.8 cm (L/W = 3. 5-5. 4), the secondary veins in 25- 32 pairs; saplings becoming lianas by branching from lateral bud some distance below leaf rosette, this “main branch" long and scandent, bearing hook- producing lateral branches. Adult stems climbing to 20 m, to 10.4 cm maximum diameter, sparingly branched, not normally rooting when in contact with ground, terete, with outer bark purplish brown, inner bark red and fibrous; main branches elongate, straight or somewhat curving, not twining; lateral branches extra-axillary, to ca. 50 cm long, bearing 1 -several hooks in a single plane. Adult leaves sessile, of 2 kinds: leaves of main branches widely spaced, elliptic to oblong (ca. 12x5 cm in type), cuneate at base, rounded at apex, with as few as 7 pairs of secondary veins; leaves of lateral branches crowded in rosettes produced near apex of current year’s growth and persisting up to three years, oblanceolate, (8—) 14.5— 33.1 x (2.9-)4.6 8.5 cm (L/W = 2.9 4.2), nar¬ rowly cuneate to long-attenuate at base, acute to rounded at apex, the margin strongly revolute near base, somewhat revolute distally, the secondary veins in 8-18 pairs, brochidodromous, forming an intra¬ marginal vein 2-4 mm from margin; all leaves dry¬ ing dull dark green above, yellow-green beneath, the midrib extending to apex, terminating in a gland, prominent beneath, slightly impressed above in her¬ barium specimens, the secondary veins prominulous on both surfaces, shortly decurrent on midrib, the tertiary veins finely reticulate, rather obscure. In¬ florescence a lax, dichotomously branched panicle, borne among leaf rosettes, sometimes bearing hooks; peduncle extra-axillary, (4-)5.5-9.8 cm; pedicels 6. 5-8. 5 mm long, articulated ca. 2 mm below calyx. Flowers with sepals suborbicular to oblong, distinctly dimorphic, the larger three ca. 5 x 4 mm, the smaller two ca. 2.5 x 3.5 mm, pale yellow-green with fine brownish veins, truncate at base, rounded at apex, shortly decurrent on base of ovary at an- thesis; petals suborbicular, shorter than sepals, pale yellow, entire, convex and thickened toward base; stamens 10 in 2 whorls, the filaments stout, those of inner stamens ca. 1 mm long, those of outer stamens very short, the anthers ca. 1 mm long with thecae separated by well-developed connective; styles 3, distinct, ca. 2 mm long, distally thickened, each terminating in a broad flattened stigma. Fruit dull brown; sepals thin, dry, brittle, unequal, the larger spathulate, 3. 4-4. 8 x 1. 3-2.1 cm, the smaller narrowly oblong to spathulate, 1.1-2. 8 x 0. 5-1.1 cm, with both margins of smaller wings and one margin of one larger wing decurrent on base of fruit, forming 5 prominent ribs; fruit body turbinate, broadest at level of wings with diameter of 1.0- 1.2 cm, the receptacle inflated and domelike above wings; scars of fallen petals and stamens forming a pale annulus 1-2 mm above wings; style base persistent, enlarged, conical; pericarp thick, dry, coriaceous; seed ca. 8 mm diam., with cerebriform-ruminate endosperm evident through thin testa; germination epigeal in moist litter layer, the cotyledons remaining inside fruit. The following preliminary key serves to distin¬ guish Ancistrocladus korupensis from the other species of Ancistrocladus in Africa: la. Adult leaves pseudopetiolate . A. abbreviatus Airy Shaw, A. barteri Scott-Elliot, A. congolensis J. Leonard, A. ealaensis J. Leonard, A. letestui Pellegrin, A. uncinatus Hutchinson & Dalziel lb. Adult leaves sessile. 2a. Petals equaling or longer than sepals . A. guineensis Oliver, A. pachyrrhachis Airy Shaw, A. robertsoniorum J. Leonard, A. sp. nov. ined. [Cheek & Ndumbe 3915, Thomas 901 6] 2b. Petals shorter than sepals. 3a. Longer sepals 3.4 4.8 cm long in fruit . A. korupensis 3b. Longer sepals 0.8- 1 .2 cm long in fruit . 4. likoko J. Leonard Additional material examined. CAMEROON. South¬ west Prov.: Ndian Div., ca. 10 km NW of Mundemba on Ekundu Kundu Rd., in disturbed primary forest on sandy clay soil, 90 m, 5°01'N, 8°52'E, 28 Feb. 1993, Gereau et at. 5172 [sterile] (K, MO, P, SCA, YA); Korup Volume 3, Number 4 1993 Thomas & Gereau Ancistrodadus korupensis 497 National Park, Science Camp 1, 50 m, 5°01'N, 8°48'E, 1 Mar. 1993, Gereau et al. 5175 [fallen fruits and infruetescences] (K, MO, P, SCA, YA); 1.2 km E of Science Camp 1 on trail to Ndian (Maria) River, 80-100 m, 5°02'N, 8°49'E, 1 Mar. 1993, Gereau et al. 5177 [fallen fr & infr] (K, MO, P, SCA, YA); type locality, 4 M ar. 1993, Gereau et al. 5179 [sterile sprouts from fallen stem] (K, MO, P, SCA, YA); type locality, 4 Mar. 1993, Gereau et al. 5181 [seedlings] (K, MO, P, SCA, YA); near S bank of Six Cup Garri Creek less than 1 km E of confluence with Ndian (Mana) River, 90 m, 5°03'N, 8°53'E, 7 Mar. 1993, Gereau et al. 5200 [sterile sapling] (K, MO, P, SCA, YA); Korup National Park, ca. 2 km N of Ikassa Last Bush, 60 m, 4°57'N, 8°50'E, 9 Mar. 1993, Gereau et al. 5203 [fallen fr] (K, MO, P, SCA, YA); ca. 10 km NW of Mundemba on Ekundu Kundu Rd„ 90 m, 5°0rN, 8°52'E, 10 Mar. 1993, Gereau & Thomas 5204 [flowers and buds from sapling flowering out of season] (K, MO, SCA, YA); forest ca. 6 km NW of Mundemba, 70 m, 5°01'N, 8°53'E, Nov. 1992, Jato s.n. [fls] (MO); Korup National Park, primary rainforest, 60 m, 5°02'N, 8°50'E, 28 Mar. 1987, Thomas 6 889 [sterile] (MO, YA), Mar. 1991, Thomas 8505 [sterile] (MO); forest ca. 6 km NW' of Mundemba, 60-70 m, 5°01'N, 8°53'E, Apr. 1992, 77 wmas 9020 [old infr] (K, MO, P, SCA, YA). NIGERIA. Cross River State: Oban, Talbot 1726 [sterile] (BM). Physical Environment The area in which Ancistrodadus korupensis is known to grow supports closed-canopy evergreen forest with areas of secondary growth following hu¬ man disturbance (Gartlan et al., 1986). The area lies at 50-100 in above sea level and is flat to gently undulating with numerous small creeks in shallow valleys. There are occasional outcrops and boulders of the hard, acidic metamorphic rocks of the African basement complex. The autochthonous soils are highly acidic (pH ca. 3. 9-4. 5), leached, and infer- tile, with a high to very high sand content (60- 91%) and the clay fraction greatly depleted near the surface. The rainfall pattern is pseudo-equato¬ rial, with only a single wet and dry season rather than the two wet and two dry seasons found further south. Rainfall at the nearby Ndian oil palm plan¬ tation averaged 5,460 mm annually in the period 1963-1983. The single long wet season peaks in July-September; November-March is the dry sea¬ son, with January the driest month. There is little annual variation in mean temperatures. Vegetation Letouzey (1985) classified the vegetation of the area as Atlantic-Biafran evergreen forests, rich in Caesalpiniaceae. This forest type is widespread at low elevations near the coast in Cameroon; however, the forest on low-lying sandy clay soils in which Ancistrodadus korupensis occurs has a unique and distinctive vegetation. Ancistrodadus korupensis occurs in forests of well-defined physical structure with a high degree of local endemism. Common or locally dominant trees of the upper canopy include Afzelia bipin- densis Harms, Berlinia bracteosa Bentham, Di- delotia africana Baillon, Krythrophloeum ivorense A. Chevalier, Julbernardia seretii (DeWildeman) Troupin, Microberlinia bisulcata A. Chevalier, Te- traberlinia bifoliolata (Harms) Hauman (Caesal¬ piniaceae), Lecomptedoxa klaineana (Pierre ex En- gler) Dubard (Sapotaceae), and Lophira alata Banks ex Gaertner (Ochnaceae). The lower canopy is dom¬ inated by Oubanguia alata Baker f. (Scytopetala- ceae), apparently common only in the Korup area; other common smaller trees include Dichostemma glaucescens Pierre, Klaineanthus gaboniae Pierre ex Prain, Mareyopsis longifolia (Pax) Pax & K. Hoffmann (Euphorbiaceae), Diogoa zenkeri (En- gler) Exell & Mendon$a, Strombosia pustulata Ol¬ iver (Olacaceae), Diospyros gabunensis Giirke (Ebenaceae), Hymenostegia afzelii (Oliver) Harms (Caesalpiniaceae), and Tabernaemontana brach- yantha Stapf (Apocynaceae). The genus Cola Schott & Endlicher (Sterculiaceae) is well represented in the understory, which is dominated by C. semecar- pophylla K. Schumann, a monocaulous tree with rosettes of very large leaves; C. cauli flora Masters, C. lateritia K. Schumann, C. megalophylla Brenan & Keay, and C. rostrata K. Schumann are also common. Deinbollia unijuga D. W. Thomas (Sap- indaceae) appears to be limited to the same area as A. korupensis, and additional narrow endemics to be described from Korup collections include new' species of Corymborkis Thouars (Orchidaceae), Uvariopsis Engler (Annonaceae), and I epris Com- merson ex A. Jussieu (Rutaceae). The area is rich in species of lianas. Ancistro¬ dadus guineensis occurs in the same area as the new species, while A. abbreviates is common in nearby seasonally flooded forests. Ancistrodadus letestui occurs on nearby bills, while A. uncinatus is known only from Cross River State, Nigeria, ad¬ jacent to the Korup area. Other climbers include many species in the genus Strychnos L. (Logania- ceae) and in the families Annonaceae, Apocynaceae, Connaraceae, Fabaceae, Icacinaceae, and Menis- perinaceae. Collecting History and Phytochemistry Ancistrodadus korupensis was apparently first collected near Oban in the Cross River State of southeastern Nigeria (Talbot 1726, BM); this sterile specimen was identified only as Ancistrodadus sp. The second collection, Thomas 6889 (MO, YA), was collected in 1987 in the Korup National Park, 498 Novon probably about 50 km from Talbot’s locality; this collection, also sterile, was tentatively identified as the widespread species A. abbreviatus. The Thomas collection was a voucher for a 0.5-kg sample of dried stems and leaves, collected under Contract No. N01-CM-67923 between the Missouri Botanical Garden and the Developmental Therapeutics Pro¬ gram of the United States’ National Cancer Institute (N.C.I.). Extracts of this sample tested for in vitro biological activity were found to inhibit the destruc¬ tion of cultured human lymphocytes by both HIV-1 and HIV-2 (Manfredi et alM 1991). Before it was realized that a new species was involved, the new alkaloids that inhibit HIV had been isolated and named michellamine A and B. These are apparently dimers of other naphthyl is¬ oquinoline alkaloids (Bringmann, 1986) previously identified from the genus (references in Leonard, 1982, 1986). At this point, the N.C.I. requested more material for further testing; samples of A. abbreviatus from Gabon, however, showed no in¬ hibitory activity against HIV. When the original population from which the biologically active sam¬ ples had been taken was relocated in Korup, it became clear that the plants belonged to a new species, apparently narrowly endemic to the Korup area. With the discovery of flowers and fruits, we have been able to complete the description of this species; the initial impetus for this task came from the plant’s pharmaceutical potential. Acknowledgments. We thank Wendy Madar and Mary Bucy for the illustration. Special thanks are due to the Jato family: Johnson Jato, who provided hospitality in Yaounde and transportation to Korup and obtained all necessary permits; and Emmanuel Jato, who tirelessly pursued the liana and finally collected its flowers and fruits. Staff of the WWF/ Cameroon Government’s Korup Project provided extensive assistance and field expertise. In partic¬ ular, we thank J. S. Gartlan (WWF Cameroon rep¬ resentative), Paul Symonds (Korup Project Man¬ ager), and Ferdinand Namata (Park Guide). Finally, we thank Gordon W. Cragg of the N.C.I. for his enthusiastic support and encouragement during sev¬ en challenging years of collaboration. This work was supported in part by N.C.I. Contracts Nos. N01- CM-67923 and N01-CM-17547. Literature Cited Bringmann, G. 1986. The naphthyl isoquinoline alka¬ loids. Alkaloids 29: 141-184. Craib, W. G. 1925. Contributions to the flora of Siam. Additamentum XV. Bull. Misc. Inform. 1925: 7- 23. Cronquist, A. 1981. An Integrated System of Classifi¬ cation of Flowering Plants. Columbia Univ. Press, New York. - . 1988. The Evolution and Classification of Flowering Plants, 2nd ed. New York Bot. Card., Bronx, New York. Dyer, W. T. T. 1874. Dipterocarpaceae. In: J. D. Hooker (editor). The Flora of British India 1: 294- 317. I. Reeve, London. Erdtman, G. 1958. A note on the pollen morphology in the Ancistrocladaceae and Dioncophyllaceae. Ver- off. Geobot. Inst. Riibel Zurich 33: 47-49. Gagnepain, F. 1909. Plantes nouvelles dTndo-Chine. Notul. Syst. (Paris) 1: 114-119. Gartlan, J. S., D. M. Newbery, D. W. Thomas & P. G. Waterman. 1986. The influence of topography and soil phosphorous on the vegetation of Korup Forest Reserve, Cameroon. Vegetatio 65: 131-148. Gilg, E. 1925. Ancistrocladaceae. Nat. Pflanzenfam., 2nd ed., 21: 589-592. Gottwald, H. & N. Parameswaran. 1968. Das sekun- dare xylem und die systematische stellung der An¬ cistrocladaceae und Dioncophyllaceae. Bot. Jahrb. Syst. 88: 49-69. Harriman, N. A. 1987. Ancistrocladaceae. In: M. D. Dassanyake & F. R. Fosberg (editors), A Revised Handbook to the Flora of Ceylon 6: 1-2. Amerind Publishing, New Delhi. Hutchinson, J. & J. M. Dalziel. 1954. Ancistroclada¬ ceae. In: J. Hutchinson & J. M. Dalziel, Flora of West Tropical Africa, 2nd ed. (revised by R. W. J. Keay) 1: 233-234. Crown Agents Overseas Governments and Administrations, London. Leonard, J. 1982. Ancistrocladaceae. In: P. Bamps (editor), Flore d'Afrique Centrale. Jardin botanique national de Belgique, Meise. - . 1986. Ancistrocladaceae. In: R. M. Polhill (editor), Flora of Tropical East Africa. A. A. Balkema, Rotterdam. Letouzey, R. 1985. Notice de la carte phytogeogra- phique du Cameroun au 1 : 500,000 : 4) Domaine de la foret dense humide toujours verte. Institut de la Carte Internationale de la Vegetation, Toulouse. Manfredi, K. P., J. W. Blunt, J. H. Cardellina, J. B. McMahon, L. L. Pannell, G. M. Cragg & M. R. Boyd. 1991. Novel alkaloids from the tropical plant Ancistrocladus abbreviatus inhibit cell killing by HIV-1 and HIV-2. J. Med. Chem. 34: 3402-3405. Metcalfe, C. R. 1952. The anatomical structure of the Dioncophyllaceae in relation to the taxonomic affin¬ ities of the family. Kew Bull. 1951: 351-368. Oliver, D. 1868. Dipterocarpaceae. In: D. Oliver, Flora of Tropical Africa 1: 172-175. L. Reeve, London. Pellegrin, F. 1951. Deux plantes congolaises a affinites asiatiques. Bull. Soc. Bot. France 98: 17-18. Ramamoorthy, T. P. 1976. Ancistrocladaceae. Pp. 171 - 172 in C. J. Saldanha & D. H. Nicolson (editors), Flora of Hassan District, Karnatka, India. Amerind Publishing, New Delhi. Robyns, W. 1958. Flore du Congo Beige et du Ruanda- Urundi, Spermatophytes, Tableau analytique des families. Institut National pour l’Etude Agronomique du Congo Beige, Brussels. Schmid, R. 1964. Die systematische stellung der Dion- cophyllaceen. Bot. Jahrb. Syst. 83: 1-56. Steenis, C. G. G. J. van. 1948. Ancistrocladaceae. In: C. G. G. J. van Steenis (editor), Flora Malesiana 4: 8-10. Noordhoff-Kolff N. V., Djakarta. Una Nueva Haploclathra (Clusiaceae) de la Amazonia Peruana Rodolfo Vasquez Proyecto Flora de las Reservas Biologicas de Iquitos, Apartado 280 Iquitos, Peru RESUMEN. El genero Haploclathra, hasta la fecha era desconocido de Peru; pero recientes colecciones en la Region Loreto, han reportado que se encuentra representado por dos especies: una es Haploclathra paniculata var. paniculata, conocida de Brasil; y la otra es Haploclathra cordata, nombre que se propone porque las hojas con base subcordada y el tamano de las inflorescencias y de los frutos indican que se trata de una nueva especie. ABSTRACT. Heretofore, the genus Haploclathra was not known from Peru, but recent collections from the Loreto Region have indicated that in fact two species are represented. One, Haploclathra paniculata var. paniculata, is known from Brazil, and the other, Haploclathra cordata, is a new species described and illustrated herein. El genero Haploclathra es conocido de la Ama¬ zonia de Brasil y de Peru: Haploclathra cordata, por ahora reportada de Peru, Loreto, Requena, Je- naro Herrera; Haploclathra grandijlora, conocida solo del tipo, Brasil, Amazonas, Taracua en el rio Llaupes; Haploclathra leiantha, en Brasil, Ama¬ zonas, rio Curicuriary afluente del alto rio Negro; M anaos, rioTaruma; Haploclathra paniculata var. paniculata, reportada de Brasil, Amazonas, alto y hajo rio Negro, Manaos, y de Peru, Loreto, Iquitos; Haploclathra paniculata var. verticillata, de Bra¬ sil, Amazonas, alto rio Negro. Clave Para las Especies de Haploclathra (Adaptada de LLeras, 1972) 1. Botones <10 mm de largo, petalos glabros 2 1 . Botones > 12 mm de largo, petalos pubes- centes o puberulentos en la porcion expuesta . 3 2(1). Peciolos 2-3 mm, base foliar subcordada; an- teras 2 mm de largo; capsula 1 6 x 11 mm . Haploclathra cordata 2. Peciolos 10-35 mm de largo, base foliar agu- da a obtusa; anteras 3-5 mm de largo; capsula 20-30 x 8-13 mm .... Haploclathra leiantha 3(1). Hojas 30 cm de largo; petalos > 30 mm de largo . Haploclathra grandijlora 3. Hojas < 28 cm de largo; petalos < 20 mm de largo . 4 4(3). Hojas opuestas o subopuestas . . Haploclathra paniculata var. paniculata 4. Hojas verticiladas . . Haploclathra paniculata var. verticillata Especies de Haploclathra de la Amazonia de Peru Haploclathra paniculata var. paniculata Haploclathra paniculata (Martius) Bentham, J. Linn. Soc. Bot. 5: 58. 1861. Especimenes examinadas. PERU. Loreto: Provincia de Maynas, Distrito de Iquitos, Puerto Alinendras, bosque primario, suelos con arena blanca y drenaje deficiente, Vasquez 9637, 966 0, 10158 (AMAZ, MO, USM). Haploclathra cordata, sp. nov. TIPO: Peru. Lo¬ reto: Provincia de Requena, Distrito de Jenaro Herrera, Jenaro Herrera, bosque primario (sue¬ los arenosos, con humedad permanente) “va- 01131,” 12 Nov. 1987, Vasquez 9996 (holotipo, AMAZ; isotipos, F, G, MO, USM). Figura 1. Arbor 24 m alta, ramulis inferne glabris, superne pu- berulis. Folia decussata, petiolus 2-3 mm longo, coriacea, oblonga vel elliptica, revoluta, apice rotundata, basi sub- cordata, supra glabra, infra glabra, in nervo puberula, 3- 5( 8) cm longa, 2-3(-5) cm lata. Inflorescentia terminalis paniculata ferrugineo-puberula, floribus minutis; sepala 2 mm longa, petala alba, 7 mm longa. Capsula ovata vel oblique ovata, glabra, 1.6 cm longa. Arbol 24 m de alto. Latex marron claro. Ramitas teretes y glabras, pero cerca de la inflorescencia tetragono-comprimidas, ferrugineo-puberulentas. Hojas decusadas; peciolo puberulento 2-3 mm de largo; limbo coriaceo con el borde revoluto, oblongo a eliptico, 3 — 5( — 8) x 2-3(-5) cm, apice redondeado a obtuso, base subcordada, haz glabro, enves tam- bien glabro excepto el nervio medio, que es pube¬ rulento, 10-14 pares de nervios secundarios, emer- gentes en el enves, venacion terciaria subparalela conspicua. Inflorescencia terminal, panicula ferru- gineo-puberulenta, 9 cm de largo; pedicelos 2 mm de largo; bracteas lanceoladas 5 mm de largo, 2 mm en la base, puberulas por fuera y glabras por dentro, caducas; bracteolas opuestas, lineares, pu¬ berulas por fuera y glabras por dentro, 3 mm de largo, 1 mm en la base; boton floral ovoide de 5 mm de largo. Sepalos 5 deltiformes, puberulos por Novon 3: 499-501. 1993. 500 Novon 5 mm Figura 1. Haploclathra cordata Vasquez. — A. Habito (Vasquez 9996, AMAZ, F, G, MO, USM). — B. Hoja de arbol juvenil. — C. Bractea. — D. Bracteola. — E. Sepalo. — F. Petalo. — G. Estambre. — H. Gineceo. —I. Fruto. fuera y glabros por dentro, 2 mm de largo, 1.5 mm en la base; petalos 5, libres, blancos, glabros en ambos lados, 7x3 mm. Estambres numerosos, con filamentos 2-4 mm de largo; anteras lineares, 2 mm de largo, subcordadas en la base; tecas onduladas, con dehiscencia longitudinal. Ovario glabro, rugoso, 1.5 mm de alto por 2 mm diam., tricarpelar, tri- locular, con placentacion axial, 1 o 2 ovulos por loculo; estilo 2.5 mm de largo con la seccion trans¬ versal fuertemente triangular. Capsula ovoide u ovoide-oblicua, con la superficie glabra y ligera- mente rugosa, 1.6 cm de largo, 1.1 cm de diam.; caliz persistente; generalmente con una semilla por fruto; exocarpo no separable del endocarpo. Ecologia. Bosque primario no inundable, por aguas de los rios; pero con humedad casi permanente por el drenaje deficiente del suelo, el cual es esencial- Volume 3, Number 4 1993 Vasquez Haploclathra 501 mente constituido por arena blanca; la infertilidad hace que esta especie se comporte como gregaria formando un bosque llamado “varillal.” Florece en noviernbre; fructifica en marzo. Nornbres vernaculares. Localmente se le conoce con los nombres siguientes: “Balatillo” y “Boa caspi.” Usos. Es la madera mas usada para la construc¬ tion de viviendas en la localidad de Jenaro Herrera. Esta especie tiene sepalos puberulos, petalos y ovario glabros, con dimensiones mas o menos como Haploclathra leiantha ; pero difiere de esta por tener inflorescencias y capsulas mas pequenas; y tambien porque las hojas tienen peciolo corto y limbo con la base subcordada, que es un caracter inusual en el genero. Pardtipos. PERU. Loreto: Provincia de Requena, Distrito de Jenaro Herrera, Jenaro Herrera, Vasquez 7664 (AMAZ, MO, USM); Valcarsel s.n. (AMAZ, G, USM). Literatura Citada Lleras, E. 1972. Review of the Genus Haploclathra (Bonnetiaceae). Mem. New York Bot. Card. 22(4): 129 136. A New Species of Ceratozamia (Zamiaceae) from Queretaro and Hidalgo, Mexico Andrew P. Vovides Institute) de Ecologia, A.C., Apartado Postal 63, 91000 Xalapa, Veracruz, Mexico Mario Vazquez Torres Instituto de Investigaciones Biologicas, Apartado Postal 294, 91000 Xalapa, Veracruz, Mexico Hart Schutzman Department ol Environmental Horticulture, University of Florida, Fifield Hall, Gainesville, Florida 32611-0670, U.S.A. Carlos G. Iglesias Instituto de Ecologia, A.C., Apartado Postal 63, 91000 Xalapa, Veracruz, Mexico Abstract. A new species, Ceratozamia sabatoi, is described and illustrated. Its closest affinities are with C. kuesteriana from Tamaulipas, but it differs from that species in both cone and leaf morphology. The chromosome number of Ceratozamia sabatoi is 2 n = 16, and the karyotype is similar to other species of the genus. During botanical explorations and ecological im¬ pact studies in the area that will be affected by the Zimapan hydro-electric dam project, a small-trunked Ceratozamia was found in pine-oak forest by Ri¬ cardo Zirahuen, a biologist collaborating with the aforementioned project in the state of Queretaro. This taxon also appears in an adjacent population in mixed oak forest on the Hidalgo side of the border at a similar elevation. Following further expeditions to both localities to procure live, vegetative, and fertile material to es¬ tablish at the Jardin Botanico Clavijero (Botanic Garden of the Instituto de Ecologia) for observation, it was concluded that this species is new to science. It is compared to Ceratozamia kuesteriana Regel, with which it is closely related, but differs in habit, leaf, female cone characteristics, and the light green color ol emergent leaves. Ceratozamia sabatoi Vovides, Vazquez Torres, Schutzman & Iglesias, sp. nov. TYPE: Mexico. Queretaro: 15 Apr. 1991, A. P. Vovides 1205 female (holotype, XAL). Figure 1 . Truncus globosus ad cylindricum, hypogaeus vel semi- hypogaeus, humilis ad 25 cm altus; cataphylla lanata, triangularia 4.55 cm longa, basi 2. 5-3. 5 cm lata; folia pauca, usque 6 pinnate, glabra; petiolus subteres vel cy- lindricus, 25-45 cm longus, parte infima dilatatus, validis spinis armatus; rachis semiteres in dimidio inferiore paucis spinis armata, supra fere inermis vel inermis, in cuspidem 5-11 mm longern excurrens; foliola subopposita vel al- terna, 15-35 juga, remota linearis, 12-25 longa, 1.2- 2.4 cm lata, basi cuneata, apicem symetrice, cuspidata, margine integerrima, 9-14 (x = 12)nervis; strobilus mas- culinus lineari-cylindricus 12-20 cm longus, 2-2.4 cm latus; pedunculus tomentosus 6-14 cm longus, 0.8-1 cm latus; strobilus femeninus cylindricus 8 10 cm longus, 4- 6 cm latus; pedunculus tomentosus 4-8 cm longus, 4- 5.5 cm latus; semina 1.5 longa, 1.3 cm lata; 2n = 16. Small palmlike plants; trunk partly subterranean, globose, readily producing offshoots, becoming cy¬ lindrical with age up to 25 cm long, 17.5 cm diam. protected with persistent leaf bases, dark brown in color. Leaves 2-6, pinnate, spirally arranged form¬ ing an open crown, up to 80 cm long, 52 cm wide; petiole and rachis ascending to horizontal, armed with short stout prickles, 0.08-0.4 cm long (x = 0.15, n = 40), petiole tomentose at base. Leaflets 12-136 (x = 69, n = 33 leaves) lanceolate, nar¬ rowly obovate to subulate, glabrous, margin entire, subrevolute, apex pungent, base attenuate, dark to light green on adaxial surface, lighter green on ab- axial, 9-29 cm long (x = 17, n = 47), 0.7-2. 4 cm wide (x = 1.2, n = 47); articulation zone 0.2 0.5 cm wide (x = 0.35, n = 43), venation ± visible on adaxial surface, more prominent on abaxial, number of veins 9-14 (x = 12, n = 20), distance between veins 0.75-1.4 mm. Microstrobili green when im¬ mature becoming light to dark brown at dehiscence, 6.5-23 cm long (x = 15.8), 1.9-3 cm diam. (x = 2.3) ; peduncle tomentose, 1.5-11 cm long (x = 6.3) , 0.5-1 cm diam. (x = 0.7, all measurements Novon 3: 502-506. 1993. Volume 3, Number 4 1993 Vovides et al. Ceratozamia sabatoi 503 Figure 1. a-k, Ceratozamia sabatoi Vovides, Vazquez Torres, Schutzman & Iglesias. — a. Habit of plant. — b. Leaf cataphyll. — c. Base of petiole. — d. Detail of leaflets and rachis. — e. Male cone cataphyll. — f. Male cone at dehiscence. — g. Microsporophyll. — h. Female cone cataphyll. — i. Mature female cone. — j. Megasporophyll. — k. Seeds. 504 Novon II II II II ii ii .. || || |l II II n ii || Figure 2. Diploid idiogram of Ceratozamia sabatoi (2 n = 16), bar = 2 /itm. n = 6); microsporophylls numerous, spirally ar¬ ranged forming apparently vertical rows, cuneiform, bicornate at distal end, fertile portion covering %- % of abaxial surface excluding horns and stalk, 0.9 1.4 cm long (x = 1.1), 0.4-0. 7 cm wide (x = 0.6, n = 6); microsporangia numerous in sori of 2 to 3, dehiscence by longitudinal slit, 0.9- 1.3 mm diam. (x = 1.1, n = 12). Megastrobili cylindrical to barrel¬ shaped slightly tapering toward apex, light blue- green when immature, turning green to light blue- brown at maturity, 6.0-12 cm long (x = 9.5), 3.4- 5.6 cm diam. (x = 4.8); peduncle tomentose, 2.0- 10 cm long (x = 5.4), 0.6- 1.3 cm diam. (x = 0.8); megasporophylls numerous, spirally arranged form¬ ing apparently vertical rows, cuneiform-peltate, dis¬ tal ends hexagonal, thickened, bicornate, with red¬ dish tomentum near horns, 1.7-2. 6 cm long (x = 2.1), 1.1 -2. 8 cm wide (x = 1.7, all measurements n = 7). Seeds ovate variably angulate, sarcotesta creamy-white when immature becoming blue-green to light brown when mature, sclerotesta light beige, smooth, 8-10 visible ridges radiating from micro- pyle, 1.3- 1.9 cm long (x = 1.5), 1.1 -1.4 cm diam. (x = 1.3, n = 6). Chromosome number 2 n = 16. We assign the specific epithet in honor of the late Sergio Sabato, distinguished professor at the University of Naples, Italy, for his outstanding and prolific fieldwork and experimental biology on neo¬ tropical Zamiaceae. Paratypes. MEXICO. Queretaro: 15 Apr. 1991, A. P. Vovides , K. Norstog & P. Fawcett 1200 , 1201, 1202, 1204, all 9, 1190, 1191, 1192, 1194 all 6 (XAL). Hi¬ dalgo: 12 Feb. 1991, A. P. Vovides, M. V azquez T. <£ C. lglesias 1156, 1157, 1158, 1159, 1160 all 9, 1161, 1162, 1163, 1164, 1165, 1 166, 1168, 1169, 1170, 1171, 1172, 1173, 1175, 1176, 1177, 1178, 1179, 1180, 1181, 1182, 1183, 1184, 1185, 1186 all <5 (XAL). The following key permits the separation of Cer¬ atozamia sabatoi from C. kuesteriana and C. rni- crostrobila. Diagnostic Key la. Median leaflet width greater than 2.8 cm, per¬ sistent leaf bases light brown, tightly appressed to trunk . C. microstrobila lb. Median leaflet width less than 2.8 cm, persistent leaf bases dark brown, not appressed to trunk 2a. Petiole and rachis coppery in adult leaves; leaflets falcate to subfalcate, linear-lanceo¬ late; veins 6-9; megastrobili reddish-brown to dark green, longer than 13cm . . C. kuesteriana 2b. Petiole and rachis light to dark green in adult leaves; leaflets not falcate to subfal¬ cate, not linear-lanceolate; veins 9 14; me¬ gastrobili blue-green to blue-brown, less than 13cm long . C. sabatoi Chromosomal Studies The chromosome number and karyotype were determined from three established specimens held at the Jardin Botanico Fco. J. Clavijero under the accession numbers: 91-028, 91-040, 91-041 and vouchers deposited at (XAL). The root tip mitosis technique was used described by Vovides (1983) and chromosome classification based on centromere position that of Levan et al. (1964) modified by Schlarbaum and Tsuchiya (1984). The diploid idi¬ ogram (Fig. 2) was constructed by taking the av¬ erage arm lengths of the best three metaphase cells examined (Fig. 3). Arm lengths, total chromosome length, chromosome index (short arm divided by long arm), and symmetry index (length of longest pair divided by length of shortest pair) were com¬ puted using the average arm lengths from the three metaphase cells (Table 1). The karyotype shows 1 1 median region (m) chromosomes, 1 submedian (sm) chromosome, 1 subterminal region (st) chromosome, and 2 terminal point (T) chromosomes. Satellite number and position vary with cells observed and a maximum of 5 satellites were recorded, but not considered in the calculations. Habitat The vegetation at the Queretaro locality is mainly pine-oak forest dominated by Pirius teocote Schle- chtendal & Chamberlain, Quercus crassifolia Hum¬ boldt & Bonpland, Q. macrophylla Martius & Gal- eotti, Q. germana Chamberlain & Schlechtendal, and Q. xalapensis Humboldt & Bonpland on reddish clay (laterite) soils. The vegetation of the Hidalgo locality, a mixed oak forest, appears much richer due to higher rainfall. The dominant tree species are; Arbutus xalapensis Humboldt, Bonpland & Kunth, Buddleja cordata Humboldt, Bonpland & Kunth, Carya ovata (Miller) K. Koch, Litsea glau- cescens Humboldt, Bonpland & Kunth, Meliosma alba (Schlechtendal) Walpers, Persea sp., Primus serotina Ehrenberg, Quercus germana Chamber- lain & Schlechtendal, Q. sartorii Liebmann, Q. polymorpha Schlechtendal & Chamberlain, and Q. Volume 3, Number 4 1993 Vovides et al. Ceratozamia sabatoi 505 r • •■‘v ' Figure 3. Chromosomes of Ceratozamia sabatoi at mi¬ totic metaphase, bar = 2 xalapensis Humboldt & Bonpland. The soil is a dark humus-rich clay on limestone. Discussion Precise information on localities has been pur¬ posely omitted in order to discourage indiscriminate commercial collecting, which could lead to the ex¬ tinction of this endangered species. Even though some vegetative characteristics, es¬ pecially the leaves, show on occasion morphological similarities between Ceratozamia kuesteriana and C. sabatoi , the consistent differences in cones, emer¬ gent leaves, seeds, and habitat lead us to consider the two taxa as separate species. Geographically speaking, several species are found in the areas between C. kuesteriana and C. sabatoi, including C. zaragozae Medellin, C. hidae Landry & Wilson, C. microstrobila Vovides & Rees, and C. mexicana var. robusta (Miquel) Dyer, species whose popula¬ tions are well delimited. Ceratozamia sabatoi occurs amongst low shrubs in oak forests and mixed oak- pine forests on volcanic soils in relatively dry hab¬ itats; C. kuesteriana , however, is found in much wetter cloud-forests on karst topography. The chromosome count and karyotype are con¬ sistent with that reported for the genus (2 n =16) by Marchant (1968), Vovides (1983; 1985) and Moretti (1990). The karyotype is nearly typical for the genus Ceratozamia (12m + 2sm + 2T), which appears to be stable. The msm and st chromosomes fall into these categories only by 0.03 and 0.01 gm respectively and, considering the considerable length of cycad chromosomes, these discrepancies are TABLE 1. Karyotypic data at metaphase in root-tip mitoses of Ceratozamia sabatoi (mean of three rneta- phase cells), m = median, msm = median-submedian, sm = submedian, st = subterminal, T = terminal. Chro¬ mo¬ some pair Arm length (Mm) Long Short (L) (S) Total Index (S/L) r value Centro¬ mere position i 6.67 6.62 13.29 0.99 1.01 III 6.92 6.10 13.02 0.88 1.13 m 2 6.78 6.15 12.93 0.91 1 . 1 0 m 6.67 6.15 12.82 0.92 1.08 m 3 6.61 5.96 12.57 0.90 1.11 m 5.67 5.30 10.97 0.93 1.07 m 4 5.53 5.41 10.94 0.98 1.02 m 5.56 5.15 10.71 0.93 1.08 m 5 5.43 4.78 10.21 0.88 1.14 m 5.54 4.18 9.72 0.75 1.33 msm 6 5.14 4.24 9.38 0.82 1.21 in 4.92 4.15 9.07 0.84 1.19 m 7 5.14 1.96 7.10 0.38 2.62 sm 4.99 1.66 6.65 0.33 3.01 st 8 6.64 0.00 6.64 0.00 0.00 T 6.71 0.00 6.71 0.00 0.00 T Symmetry index = 0.51. Average chromosome in¬ dex = 0.72. Total chromosome length = 162.73. probably due to differential contraction at meta¬ phase. The number and position of satellites varies between the species (Vovides, 1983). Two satellites have been reported by Vovides (1985) in C. kues¬ teriana and C. sabatoi has five. Acknowledgments. We thank Ricardo Zirahuen, Christina Jocabeth P., and colleagues of the Com- ision Federal de Electricidad (CFE) of the Zimapan Hydroelectric Project for permission, assistance, and use of helicopter in the field during the early ex¬ ploration of the habitat. We thank CONACyT for partial funding of this investigation through project No. P020CCOR-904 1 33. We also thank Edmundo Saavedra for the excellent botanical illustration of this species. Literature Cited Levan, A., K. Fredga & A. A. Sandberg. 1964. No¬ menclature for centromeric position on chromo¬ somes. Hereditas 52: 201-220. Marchant, C. J. 1968. Chromosome patterns and nu¬ clear phenomena in the cycad families Stangeriaceae and Zamiaceae. Chroinosoma (Berl.) 24: 100-134. Moretti, A. 1990. Karyotypic data on north and central American Zamiaceae (Cycadales) and their phylo¬ genetic implications. Amer. J. Bot. 77: 1016-1029. Schlarbaum, S. E. & T. Tsuchiya. 1984. The chro¬ mosomes of Cunninghamia konishi, C. lanceolata. 506 Novon and Taiwania cryptomerioides (Taxodiaceae). PI. Syst. Evol. 145: 169-181. Vovides, A. P. 1 983. Systematic studies on the Mexican Zamiaceae I. Chromosome numbers and karyotypes. Amer. J. Bot. 70: 1002-1006. - . 1985. Systematic studies on Mexican Zami¬ aceae II. Additional notes on Ceratozamia kuester- iana from Tamaulipas, Mexico. Brittonia 37: 226- 231. Volume 3, Number 1, pp. 1-92 of NOVON was published on 19 March 1993. Volume 3, Number 2, pp. 93-219 of NOVON was published on 1 July 1993. Volume 3, Number 3, pp. 221-310 of NOVON was published on 30 September 1993. Volume 3, Number 4, pp. 311-506 of NOVON was published on 29 December 1993.