Volume 5 Number 1 1995 GARDEN LIBRARY. Notes on the Asclepiadaceae of China Michael G. Gilbert Flora of China Project, Missouri Botanical Garden, c/o Department of Botany, Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom W. D. Stevens Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. Li Ping-tao College of Forestry, South China Agricultural University, Wushan, Guangzhou 510642, Guangdong Province, China Abstract. This paper is a precursor to the account of the Asclepiadaceae in the Flora of China. A new genus and species, Sichuania alterniloba, is de¬ scribed. Twenty-four new species are described in Biondia (B. crassipes, B. laxa, B. parviurnula, B. revoluta, and B. tsiukowensis ), Ceropegia (C. sinoerecta), Cynanchum (C. bicampanulatum, C. brevicoronatum, C. duclouxii, C. kingdonwardii, C. longipedunculatum, C. megalanthum, C. ping- shanicum, C. rock 'd, and C. sinoracemosum ), Hoya (H. commutata and II. mekongensis ), Marsdenia ( M. brachyloba, M. tend, and M. yuei), and Ty- lophora (T. forrestd, T. rockii, T. tuberculata, and T. uncinata). Jasminanthes is resurrected. Eleven new combinations are proposed in Ceropegia (C. exigua), Cynanchum ( C. boudieri subsp. cauda- tum), Heterostemma (//. menghaiense), Hoya (H. chinghungensis), Jasminanthes ( J. chunii, J. mu- cronata, J. pilosa, and J. saxatilis), and Tylophora (T. costantiniana, T. oligophylla, and 7. tsiangii). The new name Cynanchum triangulare is proposed to replace C. deltoideum Hooker, not Hance. New synonymy is proposed in Cynanchum, Lygisma, Marsdenia, and Tylophora. The status of Merril- lanthus is discussed. The following new taxa and new combinations are required for the forthcoming account of the Ascle¬ piadaceae for the Flora of China. Some new syn¬ onymy is also presented. The taxa are arranged alphabetically. Biondia Schlechter. Biondia, a genus of 13 species endemic to China, is characterized by narrow leaves with minute pale dots on the adaxial surfaces, small flowers mostly with well-developed corolla tubes, and reduced, often annular, coronas. Such leaf dots are also seen in some species of Tylophora, but they have restricted occurrence in genera related to Biondia. The most important exception is Cynanchum thesioides K. Schumann, which also has narrow leaves. This spe¬ cies is somewhat isolated in Cynanchum and has been placed in its own genus, Rhodostegiella C. Y. Wu & D. Z. Li. There is perhaps a case for regarding it as a Biondia with a more normally developed corona than usual. Most species of Biondia are too poorly repre¬ sented in herbaria to establish patterns of variation, and it is difficult to have much confidence in species delimitation. A number of collections cannot be placed within the known range of variation of existing taxa. These are herein described as distinct species. It is likely that more information on variation within spe- Novon 5: 1-16.1995. 2 Novon cies will require a reassessment of the number of species recognized. Biondia crassipes M. G. Gilbert & P. T. Li, sp. nov. TYPE: China. Sichuan: Yiajiang Xian, 2700 m, 10 Sep. 1977, B. Z. Guo & W. Y. Wang 23624 (holotype, HNWP 88274). Species nova Biondiae tsiukowensi M. G. Gilbert & P. T. Li similis sed ab ea foliis angustatis nervatione laterali non prominenti, rachidibus inflorescentiarum plus minusve pedicellos aequantibus, rachillis cymularum elongatis plus minusve incrassatis differt. Twining herbs. Stems puberulent along 2 sides and at nodes. Petioles 2-4 mm, adaxially densely puberulent; leaf blades linear-oblong, to 5 x 0.3 cm, paler abaxially, base cuneate, apex acute; midrib raised adaxially, lateral veins obscure, glabrous. In¬ florescences 2—3.5 cm, mostly with two cymules separated by a rachis ca. as long as pedicels; rachillas of cymules slightly elongated and ± fleshy, very sparsely puberulent. Pedicels 6-10 mm. Sepals el¬ liptic, ca. 1.5 x 0.6 mm, sparsely puberulent. Co¬ rollas apparently green, bell-shaped, 2.5 mm, ex¬ terior glabrous, interior minutely hairy, hairs short, stiff, white; tube ca. 1.5 mm; lobes triangular, rev¬ olute, twisted. Coronas reduced to inconspicuous collar concealing bases of anthers. Pollinia oblong- reniform. Follicles solitary, narrowly fusiform, ca. 5.5 x 0.4 mm. The narrow leaves and small flowers of Biondia crassipes suggest a relationship to B. longipes P. T. Li and B. revoluta. The corolla form is most similar to those of B. microcentra (Tsiang) P. T. Li and B. tsiukowensis, while the lax inflorescence is similar to that in B. laxa. Biondia crassipes differs from all other species of Biondia by the slightly elongated and distinctly thickened rachillas of the cymules; all other species have contracted umbel- liform cymules. Biondia longipes differs further from B. crassipes by the glabrous corollas, while B. revoluta differs by the longer and more distinctly tubular corollas. Biondia laxa M. G. Gilbert & P. T. Li, sp. nov. TYPE: China. Yunnan: Ouei cha (Yunpe), 13 May 1920, Ten 404 (holotype, E). Species nova Biondiae insigni Tsiang similis sed ab ea foliorum nervis lateralibus utroque costae latere 4-6, omnibus similibus, rachidi inflorescentiae longa cymulas plerumque 3 gerente differt. Twining herbs. Stems densely pubescent along 2 sides. Petioles to 7 mm; leaf blades lanceolate, 4.5- 6 x 1.2-2 cm, glabrous, base rounded, apex acute; lateral veins 4-6 pairs, at ca. 45° to midrib, all similar, adaxially slightly raised and minutely pu¬ berulent. Cymes extra-axillary; peduncles to 2.5 cm, glabrous; cymules umbel-like, up to 3 along zigzag rachis to 2 cm. Pedicels 1-1.3 cm, very slender. Sepals ovate, ca. 1 x 0.5 mm, glabrous or sparsely pilose on margins. Corolla tubes bowl-shaped, ca. 1 mm, much shorter than lobes; lobes triangular-acu¬ minate, ca. 2.5 mm, revolute, apex twisted, very minutely puberulent. Coronas ± annular with fleshy, square to ± rounded lobes to base of anthers. Gy- nostegium slightly exserted from corolla tube; pol¬ linia cylindric, as long as corpusculum. Fruits not seen. Biondia laxa is immediately separable from all other members of the genus by the inflorescence, which has a distinct rachis to 2 cm long and up to three separate umbel-like cymules. Older leaves have thick, pale veins very similar to those of the closely related B. insignis. However, B. insignis has a distinctive leaf venation with two major lateral veins at an acute angle to the midrib, as well as up to eight pairs of minor veins at a wider angle. Biondia insignis also differs by the ringlike corona. Paratypes. CHINA. Xizang (Tibet): Rong To Val¬ ley, Layul, 1800 m, 20 Apr. 1933, Kingdon-Ward 10352 (BM). Yunnan: Lao Kouy Chan, near My le, 1906, P. Ngueou in Ducloux 4171 (P); Shweli-Salwin Divide at 25°30'N, 2700 m, Aug. 1917, Forrest 15718 (BM, E). Biondia parviurnula M. G. Gilbert & P. T. Li, sp. nov. TYPE: China. Anhui: Jinzhai, Guan Cai Gou, Bai Ma Zhai, 800 m, 4 Mar. 1984, K. Yao 8956 (holotype. A; isotype, K). Species nova Biondiae hemsleyanae (Warburg) Tsiang similis sed ab ea nervatione foliorum prominentiore, in- florescentiis minimis, corollis minoribus urceolatis differt. Twining herbs. Stems densely minutely puberu¬ lent along 2 sides. Petioles to 7 mm, adaxially pu¬ berulent; leaf blades lanceolate, to 6.5 x 2.2 cm, base rounded, apex acute-apiculate, glabrous except for adaxially densely puberulent midrib; lateral veins 3 or 4 pairs, slightly raised on both sides; upper leaves relatively narrower, ± elliptic to elliptic-ob¬ long. Inflorescences umbel-like, ca. 4-flowered; pe¬ duncles ca. 2 mm, puberulent. Pedicels 1.5-2 mm, glabrous. Sepals lanceolate, ca. 1.5 x 0.7 mm, apex acute, pubescent, somewhat glabrescent. Corollas urn-shaped, 3-3.5 mm, interior pale purple, exterior glabrous; tube ca. 2.5 x 2 mm, throat ca. 1.2 mm wide, interior sparsely hairy; lobes ovate, ca. 1.2 mm, apparently not revolute. Corona a collar cov¬ ering bases of anthers, thin when dried. Stigma head conical. Fruits not seen. Volume 5, Number 1 1995 Gilbert et al. Asclepiadaceae from China 3 The type of Biondia parviurnula was identified as B. hemsleyana, but B. parviurnula is readily distinguished by having leaves with more prominent venation, very small inflorescences (less than 1 cm long), and smaller, more distinctly urn-shaped co¬ rollas. The corolla of B. parviurnula is most similar to those of B. microcentra and B. yunnanensis (H. Leveille) Tsiang but is glabrous inside. The type material also differs from those species by having internodes with two lines of pubescence. Biondia revoluta M. G. Gilbert & P. T. Li, sp. nov. TYPE: China. NW Yunnan/E Xizang (Ti¬ bet): Tung-chu-ling, 3000 m, May 1913, King- don-Ward 310 (holotype, E). Species nova Biondiae pilosae P. T. Li similis sed ab ea internodiis bilateraliter pubescentibus, foliis manifeste revolutis nervatione laterali prominentiore, pedicellis lon- gioribus differt; a speciebus aliis lobis corollarum tubum aequantibus differt. Twining herbs. Stems hairy along 2 sides. Petioles to 3.5 mm, puberulent; leaf blades linear-oblong, to 2.8-6 x 0.4-0.55 cm, base ± rounded, margins strongly revolute, apex subacute, apiculate, glabrous except for adaxial midrib, lateral veins not raised, underside distinctly paler, wrinkled when dried, probably fleshy when fresh. Peduncles 1-1.5 cm, hairy along 1 side; cymules umbel-like, 2-4-flow- ered, solitary or 2 separated by rachis to 5 mm, puberulent. Pedicels 7-9 mm, glabrous. Sepals ovate, 0.7-1 x 0.5-0.7 mm, sparsely puberulent, apex acute. Corollas bell-shaped, inside minutely puber¬ ulent, 2.5-3.5 mm, lobes ovate, ca. 1.5 x 1.5 mm, apex rounded, slightly revolute. Coronas annular, obscurely 5-toothed. Fruits not seen. Biondia revoluta is most closely related to B. pilosa, which differs by having internodes with only one line of hairs, leaves with acuminate apices and more conspicuous lateral veins, and shorter pedun¬ cles. The collection from Gansu ( Wang 19120) has smaller leaves and flowers than the type collection, and the lateral leaf veins are distinctly darker than the lamina and are not raised. In all other respects, it is a good match and is included here as a paratype despite the disjunct distribution. Paratype. CHINA. Gansu: Wen Xian, 1500 m, 30 Apr. 1966, Z. B. Wang 19120 (HNWP). Biondia tsiukowensis M. G. Gilbert & P. T. Li, sp. nov. TYPE: China. Yunnan: Salween-Me- kong Divide, Tsiu kow, 2400 m, May 1911, Kingdon-Ward 110 (holotype, E). Species nova Biondiae microcentrae similis sed ab ea internodiis bilateraliter pubescentibus, pedunculis longior- ibus, lobis corollinis ca. % partes longitudinis tubi ae¬ quantibus differt. Twining herbs. Stems densely pubescent along 2 sides. Petioles ca. 6 mm, adaxially puberulent; leaf blades lanceolate, ca. 6x1.2 cm, glabrous except for veins, base rounded, apex ± acute, slightly apic¬ ulate; lateral veins 5-7 pairs, adaxially slightly raised and minutely puberulent, at an acute angle to midrib. Cymes umbel-like; peduncles 2-18 mm, almost gla¬ brous. Pedicels to 11 mm. Sepals ovate, ca. 1.4 x 0.7 mm, glabrous. Corolla tube bell-shaped, ca. 2.3 mm long, longer than lobes; lobes triangular-acu¬ minate, 2x1.4 mm, revolute toward apex, mi¬ nutely puberulent inside. Corona lobes ± square, reaching base of anthers. Gynostegium ca. 1.4 mm high; pollinia cylindrical, as long as corpusculum. Fruits not seen. Biondia tsiukowensis is distinguished from other species of Biondia by having longer peduncles and corolla lobes only slightly shorter than the tube. Other species of Biondia have corolla lobes either less than half as long as or distinctly longer than the corolla tube. It also differs by its more or less square corona lobes that reach the bases of anthers. Biondia microcentra is in some ways most similar because it also has the lateral veins of the leaves raised adaxially and a hairy interior to the corolla, but it differs markedly by having much shorter (to 4 mm) peduncles and pedicels. Ceropegia L. Ceropegia , which includes about 170 species mostly in Africa, is represented in China by 17 species. Ceropegia exigua (Huber) M. G. Gilbert & P. T. Li, comb, et stat. nov. Basionym: Ceropegia longifolia Wallich subsp. exigua Huber, Mem. Soc. Brot. 12: 42. 1957. TYPE: China. Si¬ chuan: Tung Valley, Wilson 4112 (holotype, BM; isotypes, K, P). Ceropegia exigua is perhaps the most distinctive member of the C. longifolia Wallich complex. It can be recognized instantly by the very short corolla lobes, which are abruptly contracted from the prom¬ inently spreading top of the corolla tube to form a short narrow beak, even when the flower is fully open. The corolla lobes in the other members of this group are much longer (usually at least three-fourths as long as the corolla tube, sometimes longer), and the upper part of the tube does not spread and 4 Novon passes much more gradually into the lobes, which are not abruptly contracted into a beak. Further¬ more, in C. exigua the corolla lobes are only slightly darker than the rest of the corolla and are quite coarsely pilose, while in the rest of the complex, as in many other species in the genus, the corolla lobes are deeply colored and finely and densely pubescent, contrasting greatly with the rest of the corolla. Ceropegia sinoerecta M. G. Gilbert & P. T. Li, sp. nov. TYPE: China. Yunnan: au pied du Hee Chan men (Ho Kin), 2000 m, 30 May 1887, Delavay 2625 (holotype, P). Species nova Ceropegiae mairei (H. Leveille) Huber similis sed ab ea corollae angustioris basi quam ostio latiore lobis linearibus fere ad basim atratis manifeste differt; a C. wallichii White partibus omnibus gracilioribus differt. Erect herbs. Rootstock a cluster of fusiform roots. Stems erect, to 20 cm, uniformly finely puberulent. Petioles 4-6 mm, narrowly winged; leaf blades el¬ liptic, 2-5 x 0.6-1.6 cm, base cuneate, apex acute, lowermost leaves sometimes cuspidate, abaxially pale and sparsely hairy on veins only, adaxially densely puberulent. Peduncles 4-17 mm. Cymules umbel¬ like, 2-4-flowered, sometimes producing a second cymule separated by a short rachis. Pedicels 5-17 mm. Sepals linear-lanceolate, ca. 3 x 0.6 mm, glabrous. Corollas 3.6-4.3 cm, glabrous except for lobes, tube dark for basal half, pale for apical half, basal swelling ovoid, 4-6 mm wide, tube 1.3-1.6 mm wide, increasing gradually to 3.5-5.5 mm at base of lobes; lobes darkly colored almost to base, minutely puberulent, 14-15 mm, linear, slightly curved inward at base, otherwise parallel except for sharply incurved tip. Outer corona lobes each with 2 linear teeth ca. 1.5 mm, ciliate; inner corona lobes linear, ca. 2.5 mm, erect. Fruits not seen. Ceropegia sinoerecta is perhaps most closely re¬ lated to C. mairei, from which it differs by the more slender flowers widest at the basal inflation and by the differentiated corolla tips extending almost to the base of the lobes instead of less than halfway down. Ceropegia wallichii White (Nepal and west¬ ern Himalayas), which also has erect stems and similarly shaped corollas, is a robust plant with un¬ usually thick stems and much larger leaves and flowers, whereas C. sinoerecta is delicate in all fea¬ tures. The holotype has simple umbel-like inflorescences and parallel corolla lobes, while the paratype has inflorescences with pairs of umbels separated by a distinct rachis and corolla lobes bowing slightly out¬ ward. More collections are needed to properly assess the significance of these differences. There are sev¬ eral other collections by Delavay numbered as 2625, but these differ in details of date and locality, and all belong to C. mairei. Paratype. CHINA. Yunnan: Kou ty pres Pin tchouay, Apr. 1907, Jean Py in Ducloux 5305 (P). Cynanchum L. With 57 species, Cynanchum is the largest and most difficult genus of Chinese asclepiads. It is being treated here in the wider sense, i.e., including Vin- cetoxicum N. M. Wolf, Cyathella Decaisne, and Rhodostegiella. When the group as a whole is con¬ sidered, it is currently not possible to define adequate discontinuities among the potential segregate gen¬ era. Cynanchum acutum L. subsp. sibiricum (Willd- enow) F. Rechinger, FI. Iranica, Asclepiada- ceae 9. 1970. Cynanchum sibiricum Willd- enow, Ges. Naturf. Freunde Berlin Neue Schrif- ten 2: 124. 1799. TYPE: B-WILLD 5233 (microfiche). Cynanchum cathayense Tsiang & Zhang, Acta Phytotax. Sin. 12: 110. 1974. Syn. nov. Cyathella cathay- ensis (Tsiang & Zhang) C. Y. Wu & D. Z. Li, Acta Phytotax. Sin. 28: 465. 1990. TYPE: China. Gansu: Chiu-Chuan Hsien, 1350 m, 2 Dec. 1956, R. C. Ching 30 (PE). Cynanchum cathayense was not compared with C. acutum in the protologue. Numerous collections from Xinjiang match the description of C. cathay¬ ense and are not separable from C. acutum subsp. sibiricum. There seems to be no alternative but to include C. cathayense in that taxon. Cynanchum acutum L. subsp. acutum is wide¬ spread in the Mediterranean region from Spain east to Turkey, where there is an area of overlap with C. acutum subsp. sibiricum. It differs by the ovate leaves with convex rather than concave sides, but individuals with hastate leaves are found from throughout the range, and it does not seem possible to treat the two as species. Cynanchum ascyrifolium (Franchet & Savatier) Matsumura, Ind. PI. Jap. 2, 2: 509. 1912. Vincetoxicum ascyrifolium Franchet & Sa¬ vatier, Enum. PI. Jap. 2: 441. 1879. TYPE: “prob. ins. Nippon, cum V. multinerve mixtum, ex Tanaka accepit Dr. Savatier” (holotype, P). Cynanchum ascyrifolium has been used widely recently in both Japan and China for a very dis¬ tinctive erect plant with large membranous acumi¬ nate leaves and small inflorescences. Unfortunately, Volume 5, Number 1 1995 Gilbert et al. Asclepiadaceae from China 5 the type of C. ascyrifolium is a plant with oblong- elliptic leaves with a thick texture and rounded- apiculate tips. It is clearly different from any other species known to us. The correct name for most material commonly included under C. ascyrifolium is C. acuminatifolium Hemsley. Cynanchum auriculatum Royle ex Wight, Contr. Bot. India 58. 1834. SYNTYPES: India. Ka- moun, Wallich 8228 (“asclep. 137”) (K- WALL); Kimour, Royle s.n. (LIV not seen). Cynanchum saccatum W. T. Wang in Tsiang & P. T. Li, Acta Phytotax. Sin. 12: 91. 1974. Syn. nov. TYPE: China. Xizang (Tibet): Chang-Tu, Chi-na- tung, Tsa-wa-rung, 3000 m, Aug. 1935, C. W. Wang 65215 (holotype, IBSC). Cynanchum saccatum was described without the examination of authentic material of C. auricula- turn, from which it is indistinguishable. Specimens of C. boudieri were often misidentified as C. auri- culatum, but C. boudieri has strongly reflexed co¬ rollas, whereas C. auriculatum has rotate corolla lobes. Cynanchum bicampanulatum M. G. Gilbert & P. T. Li, sp. nov. TYPE: China. Sichuan: Kangting (Tachienlu), 2700 m, 11 July 1934, Harry Smith 10443 (holotype, UPS; isotypes, MO, S). Species Cynancho vincetoxico sensu lato affinis sed ab eo corollis coronisque campanulatis, lobulis coronae ad antheras corollasque adnatis facile distinguenda. Herbs. Roots many, little branched, fleshy. Stems forming fairly dense cluster, little branched, erect or ascending, to 30 cm, minutely puberulent mainly along 2 sides. Petioles 3-9 mm; leaf blades oblong- ovate, 5.5-7 x 4-4.5 cm, base shallowly cordate, truncate or rounded, apex acute to slightly acumi¬ nate; basal veins 5-7, lateral veins 4 or 5 pairs, glabrous or minutely puberulent on some. Inflores¬ cences ± umbel-like, cymules several flowered, sep¬ arated by a very short rachis; peduncles (5-)8-18 mm, puberulent; bracts linear, 0.9—2 mm. Pedicels 5—8 mm (to 12 mm in fruit), puberulent. Sepals oblong, ca. 1.5 x 0.5 mm, apex rounded, almost glabrous. Corollas bell-shaped, ca. 4 mm; glabrous except for a few stiff hairs within tube, purple (fide Guo & Wang 21046); tube ca. 2 x 3 mm; lobes oblong-ovate, ca. 1.7 x 1.5 mm, apex rounded, usually revolute. Corona tube ± as high as anthers, membranous with indexed, rounded, fleshy lobes, adnate to corolla tube and to anthers opposite lobes. Anthers square, slightly gibbous; anther appendage ovate, indexed over stigma head; pollinia ovoid, ca. 0.2 mm. Stigma head dark, rounded. Follicles paired or solitary, spreading, glabrous, beaked-fusiform, at least 3.5 x 1.5 cm. Distribution and ecology. China, Gansu and Si¬ chuan Provinces. In open fields (“prato aprico”); 2570-2700 m. At first glance Cynanchum bicampanulatum looks very similar to some of the forms of C. forrestii Schlechter, a member of the C. vincetoxicum (L.) Persoon complex. However, in C. bicampanulatum the corolla and corona are campanulate, in contrast to the shallow-tubed, rotate corolla and bowl-shaped corona of members of that complex. Cynanchum stenophyllum Hemsley resembles C. bicampanu¬ latum in having the corona adnate to both the an¬ thers and corolla tube but differs in having narrow leaves and almost no indumentum. Paratypes. CHINA. Gansu: Erh-lau-shan ad Min- chow, 2570 m, 30 June 1930 (fl), Hummel 3894 (S); Minchow, 2410 m, 8 Sep. 1930 (fr), Hummel 5097 (S). Sichuan: Daofu Xian, 5 July 1977 (fl), Guo & Wang 20861 & 21046 (HNWP). Cynanchum boudieri H. Leveille & Vaniot, Bull. Soc. Bot. France 51: CXLIV. 1904. TYPE: China. Sichuan: Kouy-Tcheou, Pin-fa, ruisseau du Tu-chang, 5 Oct. 1902, Cavalerie 620 (holotype, E). Cynanchum amphibolum C. K. Schneider in Sargent, PI. Wils. 3: 346. 1916. Syn. nov. TYPE: China. Hubei Province: Patung Hsien, Aug. 1907, Wilson 2247 (holotype, A; isotype, K). Cynanchum boudieri subsp. caudatum (Miquel) P. T. Li, M. G. Gilbert & W. D. Stevens, comb, et stat. nov. Basionym: Endotropis caudata Miquel, Ann. Mus. Bot. Lugduno-Batavum 2: 128. 1866. TYPE: Japan, Keiske (?) (not seen). The Chinese material of Cynanchum boudieri, which has often been misidentified as C. auricula¬ tum, has elongated inflorescences, whereas the Jap¬ anese collections have umbelliform inflorescences. Plants of the two countries are otherwise very sim¬ ilar, and they are best treated as subspecies. Cynanchum brevicoronatum M. G. Gilbert & P. T. Li, sp. nov. TYPE: China. Hubei (Hupeh): Ichang, A. Henry 6418 (holotype, BM; iso¬ types, K, P). Species Cynancho uiallichii simiiis sed ab eo calyce reflexo corona brevissima cupuliformi membranacea in- tegra appendicibus carente facile distinguenda. 6 Novon Twining herbs. Steins puberulent along 1 line. Petioles to 3 cm; leaf blades abaxially slightly glau¬ cous, ovate or ovate-triangular, 6-7.5 x 3-4.2 cm, base cordate, apex acute to slightly acuminate, basal lobes ± incurved, adaxially very sparsely and mi¬ nutely hairy, abaxially puberulent on veins, some¬ times pale gray-green; basal veins 3(-5), lateral veins 2 or 3 pairs; axils often with reduced stipule¬ like shoots. Inflorescences simple, shortly and dense¬ ly raceme-like, to 1.5 cm; peduncles 1-5 mm, pu¬ berulent; cymules 2-flowered. Pedicels to 5.5 mm, pubescent. Sepals lanceolate, ca. 1.5 x 0.7 mm, ciliate, minutely puberulent outside, strongly re¬ flexed at anthesis. Corollas ± erect, lobes oblong- ovate, ca. 3.5 x 1.7 mm, slightly fleshy, glabrous. Coronas a shallow ± membranous cup ca. half as high as gynostegium, internal appendages absent. Gynostegium ca. 2.5 mm high; anther appendages ovate, acute. Follicles often solitary, lanceolate in outline, ca. 6 x 1.5 cm. Distribution. China in the provinces of Hubei and Sichuan. There is no information on ecology. The corona in Cynanchum brevicoronatum is so short that at one point it was suspected to be a growth abnormality, but the flowers seem uniform and fully functional. Initially C. brevicoronatum was confused with C. decipiens, which also has a short corona, but closer examination shows that the co¬ rona is much shorter, membranous rather than fleshy, and lacks the inner appendages seen in C. decipiens. Another distinctive feature of C. brevicoronatum is the reflexed sepals. It is probably most closely related to species such as C. wallichii Wight and C. cal- lialatum Hamilton ex Wight. Similar very short coronas are seen in Biondia, which can easily be distinguished by the much narrower leaves with pale dots on the adaxial surface and the distinctly fleshy corona. Paratype. CHINA, t. Sichuan: Tchen-keou-tin, Farges 92 bis (P). Cynanchum duclouxii M. G. Gilbert & P. T. Li, sp. nov. TYPE: China. Yunnan: Yunnansen, 3 Aug. 1904, Ducloux 176 (holotype, E). Species Cynancho kintungensi similis sed ab eo corolla parva lobis erectis, corona brevioribus differt, ab speciebus aliis appendicibus coronae parvis vel reductis plus minusve minute longitudinaliter bicarinatis pilis caulium unifariis distinguenda. Twining herbs. Stems puberulent along 1 line. Petioles 2.5—3.2 cm; leaf blades triangular-ovate, 4.5-8 x 1.7-5.1 cm, adaxially uniformly minutely puberulent, abaxially slightly paler and glabrous ex¬ cept for puberulent veins, base deeply cordate, basal lobes slightly enlarged, sometimes overlapping, apex acute to slightly acuminate, basal veins 3-5, pedate, lateral veins 2 or 3 pairs. Inflorescences raceme¬ like, sometimes terminal on short axillary branches; peduncles 0.7-5.5 cm; rachis to 1.5 cm; cymules 2-flowered, in a lax spiral. Pedicels to 6 mm, pu¬ berulent. Sepals ovate, ca. 1 x 0.8 mm, sparsely puberulent, apex acute. Corollas white, ± erect, deeply divided, lobes 2.5-3 x 1-1.4 mm, glabrous outside, minutely puberulent inside. Coronas mem¬ branous, prominently 5-lobed, tube ca. as high as middle of anthers, shorter than lobes; lobes longer than anther appendages, often inflexed, with 2 prom¬ inent adaxial gibbosities at base, rarely with a small adaxial appendage. Anther appendages ovate, acute, prominent. Follicles solitary, lanceolate in outline, ca. 6.5 x 0.9 cm, base tapered, apex slightly acu¬ minate. Distribution. China, Yunnan and Sichuan Prov¬ inces. Bushland in valleys; no indication of altitude. Cynanchum duclouxii is related to C. wallichii Wight, C. sinoracemosum M. G. Gilbert & P. T. Li, and C. kintungense Tsiang. From these, C. du¬ clouxii is separated by its smaller flowers with erect corolla lobes and its coronas, which are shorter than the anthers. In C. wallichii the corolla lobes are reflexed, whereas in C. kintungense they are spread¬ ing, and in both species the corona tube is longer than that of C. duclouxii. The characters separating C. duclouxii from C. sinoracemosum are listed un¬ der the latter. Paratypes. CHINA. Sichuan: Tianquan Xian, Erlong Mountain, 14 Aug. 1953, X. Y. Jiang 35246 (HNWP). Yunnan: Yunnansen, E. E. Maire 1141 (E). Cynanchum kingdonwardii M. G. Gilbert & P. T. Li, sp. nov. TYPE: China. Yunnan: Yung¬ ning, 29 June 1922, Kingdon-Ward 5266 (ho¬ lotype, E). Species Cynancho callialato similis sed ab eo laminis foliaribus linearilanceolatis basibus cuneatis manifeste dis¬ tinguenda. 1 wining herbs. Internodes with poorly defined line of pubescence. Petioles to 11 mm; leaf blades linear- lanceolate, to at least 11 x 1.2 cm, adaxially thinly puberulent, abaxially glabrous, base cuneate, apex acute; lateral veins ca. 10 pairs. Inflorescences um¬ bel-like; peduncles to 3 mm, pubescent along 1 side. Pedicels to 7 mm, puberulent. Sepals lanceolate, ca. 2 x 0.8 mm, thinly hairy. Corollas very pale green, deeply divided, glabrous; lobes oblong-lanceolate, ca. 5 x 2.4 mm, blunt. Coronas white, cup-shaped, ca. 2.5 mm high, membranous, adnate to anthers, mar- Volume 5, Number 1 1995 Gilbert et al. Asclepiadaceae from China 7 gin with 5 low, acute teeth, smooth adaxially. Gy- nostegium slightly higher than corona; anther ap¬ pendages narrowly ovate. Fruits not seen. Distribution and ecology. China, Yunnan Prov¬ ince. On scrub-clad limestone slopes, in shade, at 3000-3300 m. Cynanchum kingdonwardii resembles C. calli- alatum and C. wallichii in having cup-shaped cor¬ onas with reduced or no internal appendages. It differs from those and all related species by the linear-lanceolate leaf blades with cuneate bases, in¬ stead of more or less ovate blades with deeply cor¬ date or sometimes rounded bases. Cynanchum longipedunculatum M. G. Gilbert & P. T. Li, sp. nov. TYPE: “Western China,” 12,000 ft. [3600 m], June 1904, Wilson 4103 (holotype, BM; isotype, P). Species Cynancho decipienti affinis sed ab eo inflo- rescentiis subcapitatis pedunculis longioribus corona long- iore cupuliformi non carnosa diifert. Twining herbs. Stems puberulent along 1 side. Petioles ca. 2.5 cm, slender; leaf blades lanceolate, ca. 5.6 X 2.3 cm, adaxially uniformly puberulent, abaxially slightly glaucous, hairy on veins only, base deeply cordate with incurved, often overlapping, basal lobes, apex long-acuminate; basal veins usually 7, lateral veins 3 or 4 pairs. Inflorescences long pedunculate, umbel-like, very congested; peduncles 6-9 cm, longer than subtending leaf, densely hairy on 1 side; cymules usually all aggregated, sometimes an isolated flower below main inflorescence. Pedicels 3.5-6 mm, puberulent on 1 side. Sepals oblong- lanceolate, ca. 2 x 0.9 mm, puberulent, sometimes only on margins. Corollas white (fide Wilson), ex¬ terior glabrous, interior densely pilose, tube ca. 0.5 mm; lobes ca. 5.5 x 1.7 mm, ± erect. Coronas cup-shaped, longer than stigma head, not very fleshy, tube longer than triangular marginal lobes, interior with 5 internal appendages opposite to and nearly as long as marginal lobes. Anther appendages ovate, nearly acute, scarious except for narrow midrib, higher than corona. Stigma head rounded. Fruits not seen. Distribution. Known only from the type collec¬ tion without precise locality or notes on ecology apart from the unusually high altitude, 3600 m; most likely collected in Sichuan Province. Cynanchum longipedunculatum differs from the related C. decipiens C. K. Schneider by having subcapitate inflorescences up to 2 cm in diameter and coronas longer than the gynostegium. Cynan¬ chum decipiens has umbel-like inflorescences 3-15 cm long and corona shorter than the gynostegium. Cynanchum lysimachioides Tsiang & P. T. Li, Acta Phytotax. Sin. 12: 89. 1974. TYPE: Chi¬ na. Yunnan: between Likiang, Youngnung & Yungpei, en route to Muli, Rock 5317 (holo¬ type, NY; isotype, E). Cynanchum likiangense W. T. Wang in Tsiang & P. T. Li, Acta Phytotax. Sin. 12: 89. 1974. Syn. nov. TYPE: China. Yunnan: Li-Kiang, 6 Aug. 1939, Y. C. Chao 21216 (holotype, KUN). Comparison of the types of Cynanchum lysi¬ machioides and C. likiangense shows that the sup¬ posed difference in habit, the one erect and the other twining, is not real because the habit is basi¬ cally erect but with a distinct tendency toward twin¬ ing. Other supposed differences in density and length of indumentum and flower size are well within the range found in many other species and seem unlikely to be of real significance. Cynanchum megalanthum M. G. Gilbert & P. T. Li, sp. nov. TYPE: China. Yunnan: 20 mi. S of Yung-ning, 2700 m, 27 May 1921, King- don-Ward 4103 (holotype, E). Species Cynancho corymboso similis sed ab eo corolla magna parce pubescenti, coronae lobulis marginalibus del- tatis tubo semper longioribus distinguenda. Robust twining herbs. Stems puberulent along 1 (or 2) lines. Petioles ca. 2.5 cm; leaf blades trian¬ gular-ovate, ca. 8.5 x 4.8 cm, thinly papery, abax¬ ially gray-green, minutely puberulent along veins, adaxially minutely puberulent, base deeply cordate, basal lobes rounded, not incurved, apex acute to acuminate; basal veins 5, lateral veins ca. 3 pairs. Inflorescences umbel-like or sometimes raceme-like with many-flowered cymules; peduncles robust, ca. 5 cm, thinly puberulent. Pedicels 10-14 mm, pu¬ berulent along 1 side. Sepals lanceolate, ca. 5 x 1.4 mm, sparsely and minutely puberulent. Corollas cream or pale greenish pink, deeply divided, rotate; lobes linear-lanceolate, ca. 10 x 2.2 mm, very sparsely hairy. Coronas cup-shaped, ca. 1 mm high, with 5 triangular marginal lobes to 3 mm high and small internal appendages. Anther appendages oval, incurved over stigma head. Fruits and seeds not seen. Distribution and ecology. China (Yunnan Prov¬ ince), Myanmar. Along thicket margins; 2700 m. Cynanchum megalanthum is superficially similar to C. corymbosum but with the corolla much larger and more sparsely hairy and with the corona tube 8 Novon shorter than the well-defined triangular marginal lobes. The internal corona appendages are always small and sometimes more or less obsolete, sug¬ gesting a relationship also to C. callialatum and C. duclouxii, which show comparable variation. Paratypes. CHINA. Yunnan: To ngay pres Lou Pou prefecture de Cong tibouray, June 1906, S. Ten in Du- cloux 4143 (P). MYANMAR. Forrest 10327 (E). Cynanchum pingshanicum M. G. Gilbert & P. T. Li, sp. nov. TYPE: China. P’ing Chan, ca. 500 km S of Beijing, mountains SW of Nan Yeli, Mei-huei-t’ouo, 5 June 1923, L. Chanet & J. H. Serre A178 (holotype, P). Species Cynancho acuminatifolio Hemsley affinis sed ab eo paginis adaxialibus foliorum uniformiter puberulis, lobis corollae angustioribus, inflorescentiis longioribus dif- fert. Erect herbs. Stems 30-60 cm high, ± tending to twine at tip in larger plants; internodes at first pubescent, soon glabrescent. Lower and uppermost leaves reduced, other leaves larger; petioles 1-2 cm; leaf blades ovate to elliptic, 7.5-15 x 3.7-8.5 cm, adaxially uniformly puberulent, abaxially pu- berulent along veins, base cuneate to ± truncate and decurrent along petiole; veins 5-8 pairs. Inflo¬ rescences at upper nodes, up to 4 per node, up¬ permost usually longer than subtending leaf, irreg¬ ularly forked to ± umbel-like, many flowered; pe¬ duncles 1-6.5 cm, glabrescent. Pedicels 6-9 mm, glabrous or sparsely puberulent. Sepals triangular- lanceolate, ca. 2 x 0.6-0.8 mm, glabrous except for ciliate margin. Corollas probably white, ca. 12 mm diam., glabrous; lobes oblong-triangular, ca. 6 x 2 mm, apex ± obtuse. Coronas fleshy, 5-lobed to near base, slightly shorter than anthers; lobes bluntly triangular, ± free from gynostegium, without internal appendages. Fruits not seen. Cynanchum pingshanicum , which is closely re¬ lated to C. acuminatifolium Hemsley and C. ja- ponicum Morren & Decaisne, is known only from five collections all made by Chanet and Serre be¬ tween 1923 and 1926. It differs from both of those species by the uniformly puberulent adaxial leaf surface, from C. acuminatifolium by the width of corolla lobes (2 vs. 3 mm wide), and from C. ja- ponicum by the shorter inflorescences. Paratypes. CHINA. P’ing Chan, Nan ye li, 1 June 1926, L. Chanet & J. H. Serre A724 (P); P’ing Chan, Pai lin t’ong, 4 June 1926, L. Chanet & J. H. Serre A722 (P); no data, L. Chanet & J. H. Serre 15 (P); no data, L. Chanet & J. H. Serre 1223 (P). Cynanchum rockii M. G. Gilbert & P. T. Li, sp. nov. TYPE: China. Sichuan: Muli, Mutirong, Muti Konka, snow range E of the Yalung, 3000 m, May-June 1932, Rock 23718 (holotype, K; isotypes, BM, E, HUH). Species Cynancho vincetoxico sensu lato affinis sed ab eo corolla atrorubenti lobis angustatis erectis glabris tubo interne lanato manifeste distinguenda. Erect herbs. Stems 2, to 30 cm, branched, pu¬ bescent. Petioles 5-11 mm; leaf blades ovate, ca. 5x3 cm, adaxially sparsely pubescent, abaxially densely puberulent along veins, base rounded to broadly cuneate, apex acuminate; lateral veins 3 or 4 pairs. Inflorescences up to 7-flowered, slender, often forked with umbel-like clusters of flowers, to 3.6 cm; peduncles ca. 1.8 cm. Pedicels ca. 6 mm. Sepals lanceolate, ca. 2.2 x 0.5 mm. Corollas dark red, tube ca. 2 mm, lobes erect, ca. 4 x 1.5 mm, triangular-acuminate, apex blunt, glabrous except the throat with woolly hairs. Corona lobes almost free, ovate, higher than gynostegium, apex rounded, adnate to anther, without internal appendage. Fruits not seen. Distribution and ecology. China, Sichuan. Al¬ pine meadow; 3000 m. The type of Cynanchum rockii was cited by Tsiang (1939) as C. muliense Tsiang. However, the type of C. muliense is clearly a member of the complex including C. vincetoxicum, C. inamoenum (Maximowicz) Loesener, and C. forrestii Schlechter. Cynanchum rockii is easily distinguished by the dark red, narrowly lobed corollas, which are glabrous except for the prominently woolly throat. It also grows at a higher altitude than members of the C. vincetoxicum complex. Cynanchum sinoracemosum M. G. Gilbert & P. T. Li, sp. nov. TYPE: China. Yunnan: Kou ly Region of Pin tchouan, 1910, J. Py in F. Ducloux 7653 (holotype, P). Species nova Cynancho corymboso, C. wallichii , et C. kintungensi similis sed ab eis internodiis uniformiter puberulis, foliis angustioribus, lobis corollarum erectis gla¬ bris differt; etiam C. duclouxii similis sed ab eo floribus majoribus foliis angustioribus basibus non dilatatis differt. Twining herbs. Stems puberulent all round when young, later along 1 or 2 sides only. Petioles 1.4- 3.5 cm, slender, puberulent along adaxial groove; leaf blades lanceolate, 4.3-6(-8) x 1.5-2.5(-4.5) cm, adaxially sparsely pubescent, abaxially paler and more sparsely hairy, sometimes ± glabrous except for veins, base deeply cordate, basal lobes incurved and sometimes overlapping, apex long acute to Volume 5, Number 1 1995 Gilbert et al. Asclepiadaceae from China 9 slightly acuminate; basal veins 5-7, lateral veins 3 or 4 pairs; minute stipule-like axillary leaves present. Inflorescences raceme-like, 2-4 cm, cymules 2- flowered, in regular spiral; peduncles 0.4-2.5 cm, puberulent. Pedicels 3-7 mm. Sepals ca. 1.3 x 0.7 mm, puberulent. Corollas white, erect, glabrous; lobes oblong-lanceolate, 3.3-4.5 x 1.3-1.5 mm. Coronas white, deeply cup-shaped, 2-3 mm high, concealing gynostegium, plicate, marginal lobes 5, ± bluntly triangular, to 0.4 mm, alternating with well-defined, ± appendage-like plicae. Anther appendages lan¬ ceolate, erect, white. Fruits not seen. Distribution. China, Sichuan and Yunnan Prov¬ inces. Growing in hedges, at least at 1900 m. Cynanchum sinoracemosum resembles C. du- clouxii in having erect glabrous corollas and reduced corona appendages. Cynanchum duclouxii has dis¬ tinctly smaller flowers with corona appendages re¬ duced to paired gibbosities, whereas C. sinorace¬ mosum has prominent longitudinal plicae and broad¬ er leaves often with enlarged basal lobes. Other species with similar coronas and inflorescences (e.g., C. callialatum, C. corymbosum Wight, C. kintun- gense, and C. wallichii ) differ by their spreading to reflexed, mostly hairy, corollas and broader leaves. In all these closely related species the internodes have the hairs restricted to one or two narrow lines, but in C. sinoracemosum the internodes are uni¬ formly hairy when young and do not develop the well-defined lines of hairs when glabrescent. Paratypes. CHINA. Sichuan: Kouy Tcheou, June 1914, J. Esquirol 5037 (P); Kouy Tcheou, near Gan- pin, 29 Aug. 1897, E. Bodinier & L. Martin 1958 (P). Yunnan: Yo lin chan, near Tong miy tcheou, 1911, F. Ducloux (P); near Ou se tchong, 1 Aug. 1904, J. Ten in F. Ducloux 2904 (P); Yunnan Fu, 1900 m, 30 July 1916, Schoch 255 (US). Cynanchum triangulare M. G. Gilbert, W. D. Stevens & P. T. Li, nom. nov. Replaced name: Cynanchum deltoideum J. D. Hooker, FI. Brit. India 4: 24. 1883, not Cynanchum deltoideum Hance, Ann. Sci. Nat., ser. 5. 5: 228. 1866. Vincetoxicum deltoideum [“deltodeum”] O. Kuntze, Rev. Gen. 2: 424. 1891.1 YPE: India. Khasia Mts.: Kalapanee and Moflong, 5000— 6000 ft.. Hooker & Thomson s.n. (holotype, K; isotype, P). western India. This leaves C. otophyllum as a Chi¬ nese endemic. Heterostemma Wight & Arnott. Heterostemma has about 30 species distributed from India east through the tropics to northern Australia and Samoa. It is represented in China by nine species, of which four are endemic. Heterostemma menghaiense (H. Zhu & H. Wang) M. G. Gilbert & P. T. Li, comb, et slat, nov. Basionym: Heterostemma villosum Cos- tantin var. menghaiense H. Zhu & H. Wang, Acta Bot. Yunnan. 16: 27, fig. 2. 1994. TYPE: China. Yunnan: Xishuangbanna, Menghai, Mengsong, 1000 m, 14 May 1989, Zhu H. & Wang H. 2443 (holotype, HITBC not seen). Heterostemma menghaiense was first collected in 1938 by T. T. Yii (Yu 16307), and it is strange that it remained undescribed for so many years. It has possibly been confused with the Indochinese H. villosum Costantin, which has a much less distinctive indumentum. The uniformly densely long pilose in¬ dumentum is not matched by any other member of Heterostemma. Several species have lines of similar long hairs along the internodes, but the hairs on the leaves and flowers of these other species are always much finer and sparser than in H. menghaiense and are often appressed. The large flowers of H. menghaiense are also distinctive, matched best by those of H. grandijlorum Costantin. Second collection. CHINA. Yunnan: Shunning, T. T. Yii 16307 (A, E). Hoya R. Brown. Hoy a is a genus of perhaps 100 species distrib¬ uted primarily in southeastern Asia into Oceania. Seventeen of the 32 Chinese species are endemic. Hoya chinghungensis (Tsiang & P. T. Li) M. G. Gilbert, P. T. Li & W. D. Stevens, comb. nov. Basionym: Dischidia chinghungensis Tsiang & P. T. Li, Acta Phytotax. Sin. 12: 130. 1974. TYPE: China. Yunnan: Che-li Hsien, Meng- soong, Dah-Meng-Lung, 1900 m, Sep. 1936, C. W. Wang 78311 (holotype, IBSC; isotype, A). Cynanchum triangulare has usually been dis¬ missed (e.g., Tsiang & Li, 1977) as a synonym of C. otophyllum C. K. Schneider, but it differs from that by the much better developed corona tube and appears to be a good species restricted to north¬ The collections cited in the protologue of Dis¬ chidia chinghungensis are all in fruit, and flowers were not known to the original authors. Flowering collections are now available, and it is obvious that this is a species of Hoya closely allied to H. Ian- 10 Novon ceolata D. Don. Material of H. chinghungensis collected in Thailand (at Tee-Lao-Su waterfall, Mae Sod District, Tak Province, near the border with Myanmar), which was not seen by us, along with a good color photo on the cover, were discussed by Thorut (1993) as “ Hoya bella subsp. nov.” How¬ ever, Thorut failed to make the connection with D. chinghungensis, even though he cited all the Chi¬ nese collections in the protologue of that species. There may well be a case for treating this and H. bella W. J. Hooker as subspecies of H. lanceolata, but this should only be done after a detailed study of the complex as a whole, which is otherwise entirely extra-Chinese. Material seen. CHINA. Yunnan: Jinghong Xian, 1700 m, 5 June 1992, Tsi Zhanhuo 92-362 (MO). MYAN¬ MAR. Northern Triangle, Kingdon-Ward 21152 (E). Hoya commutata M. G. Gilbert & P. T. Li, sp. nov. TYPE: China. Guangxi: Shap Man Taai Shan, near Iu Shan village, SE of Shang-sze, near border with Guangdong, W. T. Tsang 22375 (holotype, P, mixed with H. fusca). Species nova a congeneris ramis fioriferis dense pu- bescentibus foliis caducis, floribus purpureis, lobulis co- ronae acutis gracilibus sinus corollae excedentibus distin- guenda. Habit not known, possibly a hanging epiphyte. Stems pubescent. Leaves not known. Inflorescences pubescent, ca. 30-flowered; peduncles ca. 2 cm. Pedicels to 4 cm, slender, flexuous in dried speci¬ mens. Sepals triangular, ca. 2.5 x 0.9-1.3 mm, densely pubescent. Corollas rotate with revolute margins, “purple,” ca. 1.5 cm diam., densely pa¬ pillate-pubescent, limb ca. 0.8 cm wide. Corona lobes acute, stellately spreading, extending beyond sinus between corolla lobes; inner margins acute, ± meeting in center. Fruits not seen. The only piece of Hoya commutata seen from China is a bare twig with a single inflorescence included with a collection of H. fusca. Other sheets of Tsang 22375 have only young fruiting material of H. fusca. The two taxa can easily be separated by the fact that H. commutata is more or less uniformly pubescent, while H. fusca is glabrous. The paratype is very similar to the holotype, again a bare stem with leaves from another plant, this time belonging to Alstonia (Apocynaceae). It differs from the holotype only by the sparser indumentum. The inflorescence and the long corona lobes, which extend beyond the sinuses between corolla lobes, are very similar indeed, though there is no note on the color. It would seem that Hoya commutata flowers on rather specialized hanging branches with cadu¬ cous leaves, most unusual in a genus normally noted for its long-lived leaves. Paratype. MYANMAR. Northern Triangle, Kingdon- Ward 21276 (BM). Hoya mekongensis M. G. Gilbert & P. T. Li, sp. nov. TYPE: China. Xizang/Yunnan; Tsekou (Haut Mekong), 10 Oct. 1895, Biet in Soulie 1598 (holotype, P; isotypes, P 2 sheets). Species nova Hoyae lii Burton similis sed ab ea ramulis petiolisque non setosis inflorescentiis lateralibus differt. Lianas climbing by adventitious roots. Stems mi¬ nutely puberulent, glabrescent. Petioles 1.2-1.6 cm, minutely puberulent; leaf blades oblong-oblanceo- late, 10-13 x 4-5 cm, minutely hairy, base round¬ ed, apex acuminate to caudate-acuminate; lateral veins ca. 7, poorly defined. Pseudumbels extra-ax¬ illary, apparently flat topped, at least 8-flowered; peduncles (0.3—)1 —2.5 cm, pubescent. Pedicels 2- 2.7 cm, sparsely hairy on 1 side. Sepals ovate, ca. 1.5 x 1.5 mm, sparsely hairy. Corollas white, sweet scented, 1.5-1.7 cm diam., reflexed, lobes ca. 6 x 5 mm, minutely papillate with short-ciliate margins. Corona lobes ± ovoid, erect, ca. 2 times as high as gynostegium, dorsal grooves very narrow, inner tooth incumbent on gynostegium, just meeting in center. Anther margins short; appendages inconspicuous. Fruits not seen. Hoya mekongensis appears to be most closely related to H. lii Burton, which has a similar reflexed corolla and erect, rounded corona lobes. It differs by having minutely puberulent stems and petioles and extra-axillary inflorescences. In contrast, H. lii has setose stems and petioles and terminal inflores¬ cences. Jasminanthes Blume. We believe that the genus Stephanotis is best treated as a Malagasy endemic. Therefore, the fol¬ lowing new combinations are needed for the Flora of China. Jasminanthes chunii (Tsiang) W. D. Stevens & P. T. Li, comb. nov. Basionym: Stephanotis chunii Tsiang, Sunyatsenia 3: 165. 1936. TYPE: China. Guangdong: Tung-Wu Shan, 6 May 1928, W. T. Chun 6417 (holotype, IBSC, originally in SYS). Jasminanthes mucronata (Blanco) W. D. Ste¬ vens & P. T. Li, comb. nov. Basionym: Apo- cynum mucronatum Blanco, FI. Filip. 852. 1837. TYPE: from seed sent from China (ho¬ lotype, not preserved; see Merrill, 1918). Volume 5, Number 1 1995 Gilbert et al. Asclepiadaceae from China 11 Jasminanthes pilosa (Kerr) W. D. Stevens & P. T. Li, comb. nov. Basionym: Stephanotis pi¬ losa Kerr, Bull. Misc. Inform. 1938: 453. 1938. TYPE: Thailand. Loi, Dan Sai, Kao Keo Kang, 1300 m, Kerr 5769 (holotype, K). Jasminanthes saxatilis (Tsiang & P. T. Li) W. D. Stevens & P. T. Li, comb. nov. Basionym: Stephanotis saxatilis Tsiang & P. T. Li, Acta Phytotax. Sin. 12: 118. 1974. TYPE: China. Yunnan: Foo-Ning, 1100 m, 3 May 1940, C. W. Wang 89139 (holotype, KUN). Lygisma J. D. Hooker. A genus of 3(-6) species known from Malaysia, Myanmar, Thailand, Vietnam, and China. One spe¬ cies in China. Lygisma inflexum (Costantin) Kerr, Bull. Misc. Inform. Kew 1939: 457. 1939. Basionym: Pi- lostigma inflexum Costantin in Lecomte, FI. Indo-Chine 4: 73. 1912. TYPE: Vietnam. Cochinchine, Thorel s.n. (holotype, P). Tylophora hainanensis Tsiang & P. T. Li, Acta Phytotax. Sin. 10: 35. 1965. Syn. nov. TYPE: China. Hainan: Po-Ting, 120 m, 1 Oct. 1957, H. D. Zhang 948 (holotype, HC-HSNU). Lygisma is easily recognized by the small flowers with very characteristic inflexed corolla lobes. The correct generic position of the Chinese material had been questioned since it was first collected. It was placed in Tylophora, but with reservation. The type of T. hainanensis is a very good match with the type of Lygisma inflexum. Marsdenia R. Brown. Marsdenia includes about 100 species distributed in America, Asia, and tropical Africa. Of the 25 native species in China, 14 are endemic. Marsdenia braehyloba M. G. Gilbert & P. T. Li, sp. nov. TYPE: China. Yunnan: Feng Chen Lin Mountain, A. Henry 11196 (holotype, K). Species nova a speciebus aliis inflorescentiis umbelli- formibus, corolla purpurea, lobulis coronarum brevibus lateraliter compressis basibus profunde sulcatis distin- guenda. Lianas, glabrous except for sepal and corolla mar¬ gins. Nodes with distinct interpetiolar line. Petioles 2.5-4.5 cm; leaf blades elliptic, 10-14.5 x 5-7.5 cm, abaxially very pale, adaxially dark with paler zone along midrib, base cuneate, apex acuminate; lateral veins 5 or 6 pairs, raised on both sides. Inflorescences umbel-like, up to 20-flowered; pe¬ duncles to 3 cm. Pedicels 1-1.7 cm. Sepals ± circular, ca. 1.5 x 1.5 mm, margin ciliolate. Co¬ rollas bell-shaped with spreading lobes, purple; tube ca. 2 mm, broader than long; lobes ca. 4x2 mm, apex rounded. Corona lobes ca. as high as anthers, laterally compressed with almost hooked tip, basal half deeply grooved. Anther appendages oblong, rather fleshy toward base, half as high as stigma head. Stigma head ovoid with acuminate, divided beak, exserted from corolla tube. Follicles solitary, lanceolate in outline, ca. 16x4 cm, base narrowed into a stalk 1 cm long. Seeds ovate, ca. 17 x 7 mm; coma off-white, to 3 cm. Because of its somewhat reduced corona lobes, material of Marsdenia braehyloba has usually been filed under Gongronema. Each corona lobe is closely appressed to the anther in the form of an inverted “V” with the apex laterally compressed and some¬ what recurved and the lower part deeply grooved, with the two legs of the “V” slightly spreading and almost winglike. This contrasts very strongly with the short, spreading, dorsiventrally compressed scales inserted at the bases of the anthers that characterize Gongronema. The conspicuously exserted stigma head of M. braehyloba is also similar to that of many other species of Marsdenia in the narrow sense. Paratype. CHINA. Yunnan: Feng Chen Lin Moun¬ tain, A. Henry 11073 (K). Marsdenia tenii M. G. Gilbert & P. T. Li, sp. nov. TYPE: China. Yunnan: Siu tchoang, Sim¬ eon Ten 395 (holotype, E). Species nova Marsdeniae hainanensi Tsiang similis sed ab ea lamina foliari basi rotundata vel truncata, in¬ florescentiis longis simplicibus cymulos umbelliformes ger- entibus differt. Lianas, yellow-brown tomentose except for flow¬ ers. Petioles to 4 cm; leaf blades oblong-ovate, to 12.5 x 7.5 cm, sparsely hairy, base rounded to truncate, apex acute; lateral veins ca. 5 pairs. In¬ florescences with several umbel-like cymules along an unbranched rachis; peduncles to 3 cm; rachis to at least 2 cm. Pedicels ca. 5 mm. Sepals elliptic, ca. 3x2 cm. Corollas white, rotate, ca. 6 mm, glabrous except for lobes and retrorsely pilose throat; lobes ca. 3.5 x 2.5 mm, densely appressed-tomen- tose in center. Corona lobes to base of anthers, almost flat. Stigma head bifid-conical, equaling an¬ ther appendages. Fruits not seen. 12 Novon Marsdenia tenii is similar to M. hainanensis, M. tomentosa Morren & Decaisne, and M. sinensis Hemsley, with which it shares ovate tomentose leaves. It is easily separated from these by the inflorescence, which has a long, unbranched rachis. The other species have much-branched inflorescences that are often wider than long. Marsdenia tinctoria R. Brown, Mem. Wern. Soc. 1: 28. 1810. TYPE: Indonesia. Sumatra: “Tar- am akkar Mars. Sumat. 78” (holotype, BM). Marsdenia glubifera Tsiang, Sunyatsenia 3: 199. 1936. Syn. nov. TYPE: China. Kwangtung: Yung Yuen, S. K. Lau 24527 (isotype, P). Marsdenia tinctoria var. brevis Costantin in Lecomte, FI. Indo-Chine 4: 94. 1912. Syn. nov. SYNTYPES: Laos, Phon-thane, Spire s.n. (P); Pon-pissay, Thorel s.n. (P). At first glance, the flowers of Marsdenia globi- fera are very distinct, and it is difficult to compare them with those of any other species. However, the stems and leaves are so similar to those of the common and widespread M. tinctoria that the two plants have to be closely related. A closer exami¬ nation revealed that the flowers of M. globifera are diseased, and their lower part is an undifferentiated solid tissue. This is hinted at by the drawing of a longitudinal section of a flower in the protologue. Therefore, we have no hesitation in including this diseased material in M. tinctoria. Marsdenia yuei M. G. Gilbert & P. T. Li, sp. nov. TYPE: China. Yunnan: Mienning, Taheching, 2300 m, T. T. Yil 17801 (holotype, E). Species nova Marsdeniae koi Tsiang similis, sed ab ea pedunculis brevioribus, corollis glabris parvilobis, lobulis coronae tubo corollae longioribus differt. Lianas, glabrous except for flowers. Stems stout. Petioles 4 cm; leaf blades ovate, ca. 9.5 x 5.8 cm, base shallowly cordate, apex bluntly cuspidate to shortly acuminate, lateral veins 4 or 5 pairs, flat¬ tened. Inflorescences umbel-like, up to 9-flowered; peduncles ca. 15 mm. Pedicels to 9 mm. Sepals rounded, ca. 3 x 2.5 mm, finely appressed puber- ulent. Corollas white, ± campanulate, glabrous ex¬ cept for sparsely ciliate margins; tube ca. 4.5-5 x 4 mm; lobes oblong-obovate, ca. 5.5 x 2.5-3.2 mm, apex rounded. Gynostegium ca. 6 mm high. Corona lobes narrowly triangular, as long as anther appendages, apex rounded, exserted from corolla tube. Anther appendages ovate, membranous mar¬ gin relatively narrow. Stigma head hemispherical, conspicuously exserted from anther appendages and corolla tube. Fruits not seen. We take pleasure in dedicating Marsdenia yuei to the collector, one of the most highly regarded of all Chinese plant taxonomists. The type collection was cited as M. medogensis P. T. Li in the proto¬ logue of that species, which has much larger flowers with corolla lobes ca. 9x9 mm, discoid stigma head only slightly exserted from the anther ap¬ pendages, and oblong leaf blades more than four times as long as broad with rounded base and acute apex. Vegetatively M. yuei seems more closely re¬ lated to M. koi Merrill than to M. medogensis. Marsdenia koi has much longer pedicels, larger flowers, and corona lobes shorter than the anthers and corolla tube. Merrillanthus Chun & Tsiang. The original account of this monotypic genus described the pollinia as pendulous, which would make it a member of the Asclepiadeae. The general morphology, especially the form of the corona lobes, suggested that this might not be correct, but most material available had flowers too poorly preserved to check the pollinia. An isoparatype sheet, Lau 3550 (P), has flowers well enough preserved to examine the corona in more detail. This proved to be extremely Tylophora-like in all respects, includ¬ ing the horizontal pollinia. Tylophora augustiniana (Hemsley) Craib is particularly similar, differing only by the much smaller flowers. Merrillanthus can be easily separated from known species of Tylophora by the large fruit with a thick fibrous mesocarp, in contrast to the very thin, often almost papery follicle walls of all species of Tylophora familiar to us. Thus Merrillanthus is kept, at least until Tylophora as a whole is revised. However, the fruits of I. au¬ gustiniana have not been described, and Marsden¬ ia, even in the narrow sense adopted for the Flora of China, shows a comparable range of fruit types. A collection from Cambodia, M. A. Martin 310 ( P), is almost certainly M. hainanensis, which would mean that Merrillanthus is no longer endemic to China. Sichuania alterniloba M. G. Gilbert & P. T. Li, gen. et sp. nov. TYPE: China. Sichuan: S Wushan, A. Henry 7262 (holotype, E). Genus et species nova Holostemmate R. Brown similis sed ab eo inflorescencentiis racemiformibus, floribus mi- noribus, lobulis coronae liberis et antheris alternantibus, antheris pallidis lucentibus marginibus longis dorsaliter sulcatis appendicibus prominentibus scariis manifeste dif¬ fert. Robust twining herbs. Stems minutely puberulent along 1 side (sometimes along 2 at base of internode). Volume 5, Number 1 1995 Gilbert et al. Asclepiadaceae from China 13 Petioles 2—3 cm; leaf blades ovate-triangular, 5-8 x 3.3-7 cm, abaxially pale, adaxially much darker and apparently glossy, base cordate, thick, glabrous throughout; lateral veins 4-6 pairs. Inflorescences raceme-like, with few-flowered cymes arranged in lax spiral along rachis; peduncles 1.5-5.5 cm, some¬ times sharply reflexed; rachis up to 2 cm; bracteoles few, ca. 1 mm. Pedicels up to 1 cm, puberulent along 1 side. Sepals lanceolate, ca. 2.7 x 1.3 mm, apex acute, almost glabrous. Corollas very shallowly bowl-shaped, almost rotate, white, glabrous; lobes ovate, ca. 5.5 x 3.5 mm. Corona lobes separate, alternating with anthers, ovate to ± semicircular, short and inconspicuous. Gynostegium ca. 2.5 mm high; anthers ca. 2 mm, uniformly pale and glossy with long margins and dorsal groove; anther ap¬ pendages ovate, scarious except for narrow midrib. Retinaculum ca. 0.5 mm; translator arms short and rigid; pollinia cylindrical-ovoid, slightly compressed, ca. 0.6 mm. Stigma head apiculate, concealed by anther appendages. Fruits not seen. Distribution. Endemic to China, Sichuan Prov¬ ince (South Wushan and Tianquan Xian). The generic position of Sichuania alterniloba is so questionable that it seems best to place it in a genus of its own. The original collection had been named as Holostemma, a view supported by the presence of an interpetiolar line, fairly large, hard- textured anthers more or less decurrent at the base to the corolla, pendent pollinia, and fairly thin-tex- tured corona. However, the distinctive corona with separate, spreading, ovate scalelike lobes alternating with the anthers (i.e., opposite the corolla lobes) is quite different from the annular corona of Holos¬ temma and from that of any other genus known to us. Other genera such as Gongronema have free, scalelike corona lobes inserted at the base of the gynostegium. In all taxa with similar scalelike lobes known to us, these are opposite the anthers. Rou- linia parviflora Decaisne (Cynanchum contrape- talum E. Sundell; see Sundell, 1981) is described as having corona lobes opposite the corolla lobes, but these are erect and grooved. Cynanchum con- trapetalum is apparently in all other ways typical of Cynanchum subg. Mellichampia and thus is un¬ related to Sichuania. The anther form of Sichuania is also different from that of Holostemma in that the anther tips are more or less retuse with very sharply delineated scarious appendages, not acu¬ minate with poorly differentiated inconspicuous ap¬ pendages. Paratype. CHINA. Sichuan: Tianquan Xian, 1000 m, 11 Sep. 1963, West of Sichuan Expedition Group 3334 (HNWP). Tylophora R. Brown. Tylophora is in need of an overall revision. Most African material has never been named. The dis¬ tinction from allied genera such as Belostemma Wallich ex Wight, Merrillanthus, and Pentastelma Tsiang & P. T. Li needs closer investigation. Some of the species from tropical Asia look very different from typical members of the genus and equally deserve investigation. The separation between Cynanchum and Tylo¬ phora on floral morphology is sometimes difficult. It is often hard to interpret the orientation of the pollinia in the smaller-flowered asclepiads with only slightly elongated pollinia, especially in dried ma¬ terial. In most cases the characteristic erect corona lobes clearly inserted on the anthers of most species of Tylophora is diagnostic. However, there are some species, such as T. glabra Costantin, otherwise typ¬ ical of the genus, in which the corona lobes are almost free from the anthers, spreading and laterally joined at the base to form a very Cynanchum -like structure. Several non-floral characters often enable the confident recognition of virtually all species of Tylophora, at least in mainland Asia. These include often elongated, zigzag, inflorescence rachises, wide¬ ly spaced cymules frequently elongated with age, very narrow bracts, threadlike pedicels, and often paired, widely divaricate fruits. None of these char¬ acters is of itself diagnostic, but in combination they form a distinctive facies that leaves little doubt in the recognition of Tylophora. Tylophora costantiniana (Tsiang) M. G. Gilbert, W. D. Stevens & P. T. Li, comb. nov. Bas- ionym: Cynanchum costantinianum Tsiang, Sunyatsenia 4: 119. 1939, new name for Cy¬ nanchum hirsutum Costantin, in Lecomte, FI. Gen. Indo-Chine 4: 66, fig. 10 (12-13). 1912, not C. hirsutum Vahl, Eclog. Amer. 2: 24. 1798, nor Tylophora hirsuta (Wallich) Wight, Contr. Bot. India 49. 1834. TYPE: Vietnam. Tonkin, Mekong, Thorel s.n. (holotype, P). The type material of Cynanchum hirsutum Cos¬ tantin is mostly in fruit. However, one stem has an inflorescence with buds developed sufficiently to con¬ firm that this is a Tylophora closely related to 7! tsiangii. Tylophora forrestii M. G. Gilbert & P. T. Li, sp. nov. TYPE: China. Yunnan: hills NW of Tiugyueh(?) (25°30'N, 98°25'E), 2100 m. May 1931, Forrest 29601 (holotype, E). 14 Novon Species Tylophorae ovatae (Lindley) Hooker ex Steu- del similis sed ab ea internodiis glabrescentibus, foliis mem- branaceis, inflorescentiis longioribus cymulas umbelli- formes gerentibus, floribus majoribus differt; a T. brownii foliis membranaceis basi cuneatis vel rotundatis differt. Lianas. Stems uniformly pubescent when young but glabrescent and soon glabrous or pubescent along 1 side only. Petioles to 2 cm; leaf blades ovate- elliptic, to 9.4 x 4.2 cm, membranous, base broadly cuneate to ± rounded, apex acuminate; lateral veins 4 or 5 pairs, prominently pilose when young, gla¬ brescent. Cymes to 8 cm; peduncles to 4.5 cm, pubescent, cymules ± umbel-like, ca. 6-flowered; bracts linear, ca. 1.5 mm. Pedicels to 15 mm. Sepals linear-lanceolate, ca. 3 x 0.4 mm, apex acute, margins prominently ciliate. Corollas yellow-green, tube ca. 0.7 mm, lobes ca. 6 x 1.6 m wide, glabrous. Corona lobes narrowly ovoid, appressed to gynos- tegium and reaching base of anthers, ± acute. Gy- nostegium ca. 2 mm high; anther appendages ovate, erect. Corpusculum longer than ellipsoid pollinia. Fruits not seen. The indumentum suggests an affinity to Tylo- phora ovata, but T. forrestii is a more vigorous plant with softer leaves, larger flowers, more strictly umbel-like cymules, and readily glabrescent stems. Tylophora glabra Costantin in Lecomte, FI. Indo- Chine 4: 109. 1912. TYPES: Vietnam. Tonkin vers Ninh-Binh, Bon s.n. (syntype, P); vers Ouonbi, au nord de Guang-yen, Balansa 609 (syntypes, P, K); Sept Pagodes, Mouret s.n. (syntype, P). Tylophora longipedicellata Tsiang & P. T. Li, Act. Phytotax. Sin. 12: 136. 1974. Syn. nov. TYPE: China. Hainan: Lung-Kiang, 15 Sep. 1933, C. H. Tsoong 545 (holotype, IBSC). Tylophora renchangii Tsiang, Sunyatsenia 3: 232. 1936. Syn. nov. TYPE: China. Guangxi (“Kwangsi”): Tien- chen, N of Sup-man-ta Shan, 900 m, 13 Oct. 1928, R. C. Ching 7821 (holotype, IBSC; isotype, A). Tsiang (1936) first drew attention to the unusual corona of this species in his protologue for Tylo¬ phora renchangii. The corona lobes are almost Heterostemma- like and distinct from any other Ty¬ lophora seen from mainland Asia. Typical Tylo¬ phora corona lobes are closely appressed to the stamens, dorsally gibbous, and have an acuminate tip. In T. glabra the corona lobes are more or less circular, inserted at the base of the anthers, and spread at an angle of about 45° to them, while the acuminate tip is often strongly reflexed. Superficially these are so different that it is difficult to accept this species in Tylophora, but all other features of T. glabra are typical of that genus. The status of Tylophora longipedicellata de¬ serves more investigation. Most material from Hai¬ nan has the broad spreading corona lobes with a blunt inflexed tip characteristic of T. glabra. Other collections have short erect corona lobes with an erect acuminate tip, as illustrated in the protologue of T. longipedicellata. In other features, the two species are nearly identical. Henry 8275 (K) has corona lobes somewhat intermediate between those of the two species. Therefore, we tentatively reduce T. longipedicellata to synonymy of T. glabra. Tylophora oligophylla (Tsiang) M. G. Gilbert, W. D. Stevens & P. T. Li, comb. nov. Bas- ionym: Absolmsia oligophylla Tsiang, Suny¬ atsenia 6: 121. 1941. TYPE: China. Yunnan: Che-li Hsien, Aug. 1936, C. If . Wang 75455 (holotype not seen; isotype, PE). The generic placement of Absolmsia oligophylla is uncertain. It differs profoundly from the type and only other species of Absolmsia, A. spartioides (Bentham) Kuntze, which is a very specialized rel¬ ative of Hoya, known only from rot holes in large trees in Borneo. Absolmsia spartioides has fleshy leaves when young but then depends on persistent, long, stiff, photosynthetic peduncles, which seem to take over the function of leaves. A sheet labeled in Tsiang’s handwriting as the holotype of A. oligo¬ phylla, deposited in the herbarium of the Fan Me¬ morial Institute of Biology and now housed in the Institute of Systematic Botany, Beijing (PE), consists of the lower part of a single stem and completely lacks any trace of inflorescences and flowers. A photograph of a different sheet showing the upper parts was reproduced in the protologue. A search for an isotype at the South China Institute of Botany was not successful. The material seen appears to come from a prostrate plant with no sign of the stem’s twining and with soil adhering to the under¬ sides of the leaves, very like Tylophora rotundifolia Hamilton ex Wight, which also has similar almost circular leaves. The illustration of the flowers ac¬ companying the protologue, particularly the small gynostegium, is also suggestive of a Tylophora. The spreading corona lobes would be unusual for Tylo¬ phora, but T. glabra Costantin has even larger spreading lobes that are at least superficially similar. The description of pollinia as pendulous, if correct, would rule out a placement in Tylophora, but, as discussed above, Merrillanthus was described as having pendulous pollinia whereas examination of material cited in the protologue showed the pollinia Volume 5, Number 1 1995 Gilbert et al. Asclepiadaceae from China 15 to be horizontal. The large, stiffly papery leaves and indumentum of short stiff hairs of 71 oligophylla are distinctive, and it should be easy to make a positive match with new material, should this become available. Tylophora rockii M. G. Gilbert & P. T. Li, sp. nov. TYPE: China. Sichuan: Muti Konka, 3300 m. Rock 23717 (holotype, E). Species nova Tylophorae yunnanensi Schlechter sim- ilis sed ab ea inflorescentiis sessilibus, floribus majoribus, capite stigmatico elevato differt. Erect herbs to 70 cm. Stems little branched, tending to twine at tips, uniformly puberulent. Pet¬ ioles ca. 15 mm; leaf blades oblong-oblanceolate, to 9.5 x 4 cm, sparsely puberulent, uppermost and lowermost reduced, base cuneate to ± rounded, apex rounded-apiculate, margins ± undulate, veins 4 or 5 pairs. Inflorescences extra-axillary from up¬ per nodes, to 8 cm, mostly ± sessile with long first internode, cymules shortly and densely raceme-like to umbel-like. Pedicels to 1(—2) cm. Sepals linear- triangular, ca. 3 mm. Corollas purplish, stiffly rotate, 8-10 mm diam.; lobes oblong, 5(-7) x 2.5-3 mm, apex rounded; interior densely puberulent. Corona lobes reaching base of anther appendage, ± acute. Gynostegium ca. 1.5 mm high; anther appendages broader than long. Stigma head broadly dome-shaped with light-colored center. Fruits not seen. Tylophora rockii is most closely related to T. yunnanensis, which has distinctly smaller flowers, sepals ca. 1.5 mm long, and corolla lobes 3-4 x 1.5-2 mm. Tylophora rockii has mainly sessile inflorescences with long first internodes, whereas T. yunnanensis has peduncles to 6 cm long. Tylophora rockii has a broadly dome-shaped stigma head in contrast to the depressed stigma head of T. yun¬ nanensis. Tylophora tsiangii (P. T. Li) M. G. Gilbert, W. D. Stevens & P. T. Li, comb. nov. Basionym: Cynanchum tsiangii P. T. Li, Bull. Bot. Lab. North-E. Forest. Inst. 3(1): 103. 1983. TYPE: China. Guizhou: Xing-yi, 1300 m, 7 Oct. 1960, Y. T. Chang et al. 6880 (holotype, SCBI). The hispid indumentum, brownish coloring of the dried leaves, and inflorescence architecture (a long zigzag rachis with elongated raceme-like cymules and linear bracts) of Tylophora tsiangii are all very similar to those of Tylophora ovata (Lindley) Hook¬ er ex Steudel. The short, elliptic corona lobes adnate to and clearly shorter than the anthers are typical for Tylophora. Though placed in different parts of the family, the distinction in floral morphology be¬ tween these two genera can be very subtle. Tylo¬ phora tsiangii is clearly related to the Indochinese T. costantiniana, which is very similar in habit and was also misplaced within Cynanchum. Tylophora tuberculata M. G. Gilbert & P. T. Li, sp. nov. TYPE: China. Yunnan: Kouy- Tcheou, Echang loing, 800 m. May 1913, Esquirol 4377A (holotype, P, mixed with Cy¬ nanchum corymbosum Wight). Species nova Tylophorae tenui Blume similis sed ab ea et speciebus similibus caulium indumento ex trichom- atibus erectis crassisque constante, foliis adaxaliter tub- erculatis, lobulis coronae basaliter connatis, dorsaliter di- latatis umbonem subpatentem truncatum vel leviter emar- ginatum formantibus manifeste distinguenda. Lianas. Stems slender, short pilose along 2 sides. Petioles 5-10 mm, adaxially hairy, glabrescent; leaf blades lanceolate, 2.5-4 x 1.1-1.2 cm, thick pa¬ pery, base rounded-cordate, apex acute, glabrous, adaxially with prominent scattered tubercles. Inflo¬ rescences to 4 cm, much branched; peduncles to 6 mm, intemodes to 14 mm. Pedicels 4-6 mm, gla¬ brous. Sepals lanceolate, ca. 0.7 mm, tip recurved, acute, glabrous. Corollas rotate to shallowly bowl¬ shaped, ca. 5 mm diam., greenish white, glabrous outside, densely short pilose inside; lobes ovate, ca. 2 x 1.5 mm. Corona lobes joined at base, reaching base of anther, with very prominent spreading, trun¬ cate or slightly emarginate, dorsal projection. Anther appendages short, rounded; pollinia ellipsoid, almost vertical. Stigma head rounded. Old follicles linear lanceolate in outline, ca. 6.5 x 0.5 cm. Tylophora tuberculata resembles 7. tenuis Blume in having small lanceolate leaves and delicate much- branched inflorescences. It differs from that and all similar species of Tylophora by the thick stiffly erect hairs forming two lines down the internodes of young stems and along the margins of the petioles, the well-defined small tubercles scattered adaxially on leaf blades, the inflorescence with the peduncle clearly shorter than the intemodes of the branched rachis, and the distinctive corona lobes, which are enlarged dorsally into a more or less spreading, fleshy, trun¬ cate to slightly emarginate projection and joined laterally at the base by a thin flange. The corona can be interpreted as forming a link between the corona lobes of typical species of Tylophora, in which the lobes are only slightly swollen dorsally, and the corona lobes of 71 glabra, in which the dorsal swelling seems to be grossly expanded into a flat disc. The only sheet seen also has a fruit of Cynanchum corymbosum Wight mounted on it. 16 Novon Tylophora uncinata M. G. Gilbert & P. T. Li, sp. nov. TYPE: China. Hainan: Po-ting, How 73159 (holotype, IBSC; isotypes. A, IBSC). Species nova Tylophorae ovatae (Lindley) Hooker ex Steudel affinis sed ab ea lamina foliari adaxialiter glabra abaxialiter trichomatibus apice uncinatis basi papillosis manifeste differt. Lianas. Stems glabrescent. Petioles ca. 1.5 cm; leaf blades ovate, 3.5-9 x 1.3-5.5 cm, stiffly pa¬ pery, adaxially glabrous, abaxially uniformly pu¬ bescent, hairs with papillate bases and strongly hooked tips, base cordate, sinus usually deep and narrow, apex ± acute, apiculate. Cyme extra-ax¬ illary; peduncles to 2 cm, rachis zigzag, sometimes ± absent, cymules densely raceme-like; bracts lin¬ ear-lanceolate, to 1.5 mm. Pedicels 6-8 mm. Sepals ca. 1.7 x 0.5-0.7 mm, oblong-lanceolate, apex acute, with few to many hooked hairs. Corolla pale green at apex, dull green at base, 3.5-5 mm; tube ca. 0.5 mm; lobes ovate to ovate-oblong, 2.5-3.5 x 1.2-1.3 mm. Corona lobes ovate, ca. 0.5 x 0.2 mm, pouched, apex obtuse, covering base of anthers. Anther appendage rounded; pollinia globose, hori¬ zontal. Stigma head ± flat, 5-angled, depressed in center. Follicles linear-lanceolate in outline, ca. 5 x 0.6 cm, apex acuminate. Flowering August-Sep- tember. Tylophora uncinata had been named mostly as T. ovata (Lindley) Hooker ex Steudel var. brownii (Hayata) Tsiang & P. T. Li, but closer examination has shown that it differs from all species of Tylo¬ phora known to us by the very distinctive hooked hairs on the underside of the leaves. Tylophora uncinata differs further from otherwise similar spe¬ cies such as T. ovata by the more papery, adaxially glabrous leaves. Paratypes. CHINA. Guangdong: Gao-zhou, 4 Apr. 1929, K Tsiang 2088B (IBSC). Guangxi: Shang-si, 2 Aug. 1951, W. C. Ko 901 (CANT); without locality, Nov. 1889, A. Henry 8105 (K, P). Acknowledgments. Thanks are due to Roy Ger- eau for correcting the Latin diagnoses, to Marshall R. Crosby, Ihsan Al-Shehbaz, Amy McPherson, and Diana Gunter for facilitating the production of this paper in many ways, and to the directors of the following institutions for help during visits and/or loan of material: E, HNWP, IBSC, K, P, PE, S, SYS, UPS, US. The Stanley Smith Horticultural Trust is thanked for the financial support of Gilbert. Literature Cited Merrill, E. D. 1918. Species Blancoanae. Publ. Dept. Agric. Nat. Resour. Manila 12: 1-423. Sundell, E. 1981. The New World species of Cynan- chum , subgen. Mellichampia (Asclepiadaceae). Evol. Monogr. 5: 1-63. Thorut, C. 1993. A very special Hoya. The Hoyan 15: 29-30. Tsiang, Y. 1936. Notes on the Asiatic Apocynales III. Sunyatsenia 3: 121-239. -. 1939. An enumeration of the Asclepiadaceae plants collected by Tsai and Wang from Yunnan in 1933-1936. Sunyatsenia 4: 95-130. - & P.-T. Li. 1977. Asclepiadaceae. In: FI. Reipubl. Popularis Sin. 63: 249-575. Nomenclatural Notes on the Boraginaceae of China Chu Ge-lin Institute of Botany, Northwest Normal University, Lanzhou, Gansu 730070, People’s Republic of China Rudolf Kamelin Herbarium, V. L. Komarov Botanical Institute, Russian Academy of Sciences, Prof. Popov Street 2, 197376 St. Petersburg, Russia Robert R. Mill Royal Botanic Garden, 20A Inverleith Row, Edinburgh EH3 5LR, Scotland, U.K. Michael G. Gilbert Flora of China Project, Missouri Botanical Garden, c/o Department of Botany, Natural History Museum, Cromwell Road, London SW7 5BD, U.K. ABSTRACT. Three new combinations, Tournefortia sibirica L. var. angustior (A. DC.) G. L. Chu & M. G. Gilbert, Lappula ferganensis (Popov) Ka¬ melin & G. L. Chu, and L. lasiocarpa (W. T. Wang) Kamelin & G. L. Chu, plus a new name, Trigonotis harrysmithii R. R. Mill, are validated. Two precursory papers on the Boraginaceae for the Flora of China have already been published (Kamelin, 1993; Riedl, 1994). The following ad¬ ditional nomenclatural adjustments are herein pro¬ posed to make the names available for the forth¬ coming Volume 16 of the Flora of China. Tournefortia sibirica L. var. angustior (A. DC.) G. L. Chu & M. G. Gilbert, comb. nov. Bas- ionym: Tournefortia arguzia Roemer & Schultes p angustior A. DC., Prodr. 9: 514. J845. SYNTYPES: China. Nei Mongol (“Chi¬ nese Mongolia”), Turczaninow s.n., Bunge s.n.; Kazakhstan. E shore of Caspian Sea, Karelin s.n. and Hohenacker s.n.; Russia. Astrakan, Bieberstein s.n.; Turkey (“Asia Minor”), Aucher-Eloy s.n.; Ukraine. S Podolia, Tur¬ czaninow s.n. (All syntypes in G, not seen.) The new combination is needed because the va¬ riety does not have a valid combination in Tour¬ nefortia sibirica. A lectotype should be designated from de Candolle’s original material at G. Such material has not been examined by us. Trigonotis harrysmithii R. R. Mill, nom. nov. Replaced name: Trigonotis smithii W. T. Wang, Bull. Bot. Res., Harbin 13(1): 3. 1993, not Trigonotis smithii Banerjee, Bull. Bot. Surv. India 8; 325. 1966 (1967). TYPE: China. Sichuan: Sungpan, 3200 m, 14 July 1922, Harry Smith 2775 (holotype, UPS; isotype, PE).' The earlier name Trigonotis smithii Banerjee, which applies to a species from Bhutan and the east Himalaya so far not recorded from China, was named after William Wright Smith, the collector of the type specimen and a former Regius Keeper of the Royal Botanic Garden, Edinburgh. The later hom¬ onym was dedicated to the Swedish botanist Harry Smith, an expert on the Gentianaceae who collected in China. There has been a reappraisal of the characters dividing Lepechiniella from Lappula. Lappula is readily distinguished from Lepechiniella in having styles exserted from the nutlets instead of concealed by them. No Chinese material belongs to Lepechi¬ niella as thus defined, which makes necessary the following generic transfers. Lappula ferganensis (Popov) Kamelin & G. L. Chu, comb. nov. Basionym: Lepechiniella fer¬ ganensis Popov, FI. USSR 19: 715. 1953. TYPE: [Tajikistan]. Alai Valley, Mount Kultz- cha, 26 June 1931, Lipschitz 248 (holotype, MW). Novon 5: 17-18. 1995. 18 Novon Lappula platyptera C. J. Wang, Bull. Bot. Res., Harbin 1(4): 91. 1981. Syn. nov. TYPE: China. Xinjiang: Wuqia Xian, 3300 m, 4 Aug. 1978, Xinjiang Ex- pediton 1911 (holotype, WUK). Lappula lasiocarpa (W. T. Wang) Kamelin & G. L. Chu, comb. nov. Basionym: Lepechi- niella lasiocarpa W. T. Wang, Bull. Bot. Res., Harbin 4(2): 7. 1984. TYPE: China. Xinjiang: Fuhai, 21 May 1959, Y. F. Wang s.n. (holo¬ type, PE). Lepechiniella balchaschensis M. Popov, Spisok. Rast. Gerb. FI. SSSR 12: 50. 1953. Syn. nov. Lappula betpakdalensis Nabiev, Opred. Rast. Sred. Azii 8: 138. 1986, not Lappula balchaschensis M. Popov ex N. Pavl., Zhurn. Bot. U.R.S.S. 30(4): 190. 1945. TYPE: Kazakhstan. Lake Balchasch, southern shore at Bertys Bay, 15 June 1934, M. Popov s.n. (ho¬ lotype, LE). Lepechiniella balchaschensis is the oldest name for this species, but the epithet is not available in Lappula because Popov had already described Lap¬ pula balchaschensis based on a different type. The transfer to Lappula was made in 1986 by Nabiev, who introduced the replacement epithet betpak¬ dalensis. However, Lepechiniella lasiocarpa W. T. Wang is the older name that should be used in Lappula. Literature Cited Kamelin, R. 1993. Nomenclatural adjustments in Chi¬ nese Boraginaceae. Novon 3: 263. Riedl, H. 1994. A new species and new combinations of Boraginaceae from China. Novon 4: 46-47. Onosma apiculatum (Boraginaceae), a New Species from China Harald Riedl Botanische Abteilung, Naturhistorisches Museum, Postfach 417, Burgring 7, A-1014 Wien, Austria ABSTRACT. Onosma apiculatum from Xinjiang, China, is described. Its relationship to the three species of series Tianschanica is discussed, and its distinguishing characters from these are given. Onosma apiculatum H. Riedl, sp. nov. TYPE: China. Xinjiang: Zhao Moa Hong Qi, 2nd wood¬ cutting grounds, 2100 m, 9 June 1965, Cheo Tai-yien et al. 650713 (holotype, SZ; isotype, SZ; photos, W). Herba perennis, foliis rosulae linearibus vel lineari- subspathulatis, basin versus petiolatim attenuatis, 4-13 cm longis, 4-8 mm latis; foliis caulinis inferioribus bas- alibus similibus basin versus paulo tantum attenuatis, su- perioribus prope basin latissimis manifeste brevioribus, omnibus viridibus, supra setis patentibus densis tuberculis albis vix elevatis e circulis 2 vel 3 cellularum compositis insidentibus et papillis nonnullis minutis, infra setis im¬ primis in nervo mediano densis, ceterum laxis pilisque brevibus numerosis densissimis vestitis, cymis singulis vel geminatis, calyce ca. 1.5 cm longo, laciniis linearibus, subobtusis, usque ad basin liberis, corolla 2-2.2 cm longa, compressa 8 mm fere lata, extra minutissime papillosa, intus glabra, antheris tota longitudine cohaerentibus apice sterili vix 1 mm longe exsertis; filamentis antheris paulo brevioribus, in parte tertia inferiore corollae insertis, stylo corollam 2.5-3 mm superante stigmate bilobo insignis. Herbs perennial, with a single flowering stem and sterile rosette of leaves arising from either a stout vertical rhizome or from its ± horizontal branches. Stem lateral, ascending, 20-35 cm high, un¬ branched, straw-colored when dry, densely covered by a mixture of white patent bristles ca. 2 mm long and very short patent hairs. Leaves of the sterile rosette 4-13 cm long, 4-8 mm wide, linear-spat- ulate to linear oblanceolate, tapering into a petiole¬ like base, revolute at margin, acute to subobtuse at apex, green, densely covered adaxially with patent to subappressed white bristles arising from a white, nearly flat tubercle composed of 2 or 3 circles of roundish to radially elongated, strongly convex cells and numerous minute papillae, abaxially covered with similar bristles mainly along the midrib and only sparsely on other parts, with dense short hairs as those on stem; lower stem leaves similar to rosette leaves in size and shape but only a little narrowed toward base; uppermost leaves much shorter, 2.5- 4 cm long, widest near base, acute, bristles and hairs as in rosette leaves. Cymes single or paired, involute, 4-6 cm long in flower; lower bracts similar to upper stem leaves, to 2 cm long, broad at base, decreasing in size upward. Lower pedicels to 5 mm long, upper ones to 2 mm. Calyx ca. 1.5 cm long in flower, divided to base into 5 linear, subobtuse lobes densely covered with fairly soft, appressed or subappressed bristles and short hairs, elongated soon after flow¬ ering. Corolla pale yellow, turning brown after flow¬ ering and when dry at least in apical part, club- shaped, 2-2.2 cm long, ca. 8 mm wide just below lobes, ca. 3 mm wide at base in the compressed state, very minutely papillate outside, glabrous inside and on nectary; lobes ca. 1.5 mm long, recurved, ca. 4.5 mm wide at base, abruptly tapering into an obtuse apex ca. 1.5 mm wide. Anthers coherent for their entire length, ca. 9.5 mm long, fertile part ca. 3.5 mm; sterile tips horizontally truncate at apex, exserted from corolla for less than 1 mm; filaments ca. 7 mm long, ca. 0.8 mm wide, tapering toward anther. Style surpassing corolla by 2.5-3 mm; stig¬ ma 2-lobed. Nutlets not seen. Onosma apiculatum belongs to series Tian¬ schanica Popov, the three other species of which are 0. gmelinii Ledebour, O. irritans Popov ex Pavlov, and O. baldshuanicum Lipsky. From O. gmelinii, the new species is distinguished by its green instead of grayish leaves, bracts longer than calyx, club-shaped corolla, and slightly longer an¬ thers; from 0 . irritans by its markedly wider leaves without a distinct petiole, presence of short hairs between the bristles, bracts longer than calyx, short¬ er calyx, and club-shaped corolla; and from O. bald¬ shuanicum by its narrower leaves without a distinct petiole, much denser cover of bristles and short hairs, bracts longer than calyx, and shorter anthers. Onosma apiculatum also differs from these three species by its shorter corolla and by the shape of its corolla lobes, which are very broad at base and suddenly contracted into a narrow tip, hence the specific epithet apiculatum. The type collection was misidentified as O. echioides L., but this species is native to Europe and adjacent parts of Turkey and differs by several characters, the most notable of which are the shorter calyx scarcely half as long as corolla and the asterotrichous indumentum. Acknowledgment. I thank Ihsan Al-Shehbaz for his help in the preparation of this manuscript and in obtaining the loan of specimens. New Combinations in Neotropical Grammitidaceae (Pteridophyta) Alan R. Smith University Herbarium, University of California, Berkeley, California 94720, U.S.A. Abstract. Continuing study on neotropical Gram¬ mitidaceae for various floras in press or in progress necessitates the following new combinations: Cera- denia oidiophora, C. phalacron, Lellingeria cili- olepis, L. dissimulans, L. hombersleyi, Melpomene gracilis, M. zempoaltepetlensis, Terpsichore bi- pinnata, T. elastica, and 7. Jlexuosa. Examination of additional types of several rare and geographically restricted species of Grammiti¬ daceae, as well as further study of problematic groups, has resulted in the need for several new combina¬ tions. This work is a continuation of observations previously published on the genera Ceradenia, Lel¬ lingeria, Melpomene, and Terpsichore (Bishop, 1988; Smith et ah, 1991; Smith & Moran, 1992; Smith, 1993). Ceradenia oidiophora (Mickel & Beitel) A. R. Smith, comb. nov. Basionym: Grammitis oi¬ diophora Mickel & Beitel, Mem. New York Bot. Gard. 46: 202. 1988. TYPE: Mexico. Oaxaca: Dto. Ixtlan, trail from San Pedro No- lasco N to the Llano Verde, Mickel 3822 (ho- lotype, NY; isotype, UC). Known only from four collections from Oaxaca (Mickel & Beitel, 1988). Ceradenia phalacron (Stolze) A. R. Smith, comb, nov. Basionym: Grammitis phalacron Stolze, Fieldiana, Bot. 32: 84. 1993. TYPE: Peru. Cuzco: Valley of Rio Urubamba, Bites A29 (holotype, US). Ceradenia phalacron is known only from the type. It is somewhat aberrant in Ceradenia because of its relatively simple or only shallowly lobed blades, lack of laminar setae, and prominulous veins, but it fits more comfortably there than in any other de¬ scribed genus. With other ceradenias, it agrees in the absence of hydathodes, castaneous rhizome scales with setulose margins, lack of spongy parenchyma, the prominulous venation, the lack of laminar or circumsoral setae, and the glandular paraphyses within the sorus. Lellingeria ciliolepis (C. Christensen) A. R. Smith, comb. nov. Basionym: Polypodium ciliolepis C. Christensen in Asplund, Ark. Bot. 20A: 21. 1926. Ctenopteris ciliolepis (C. Christensen) Copeland, Philipp. J. Sci. 84: 395. 1956 [1955]. Grammitis ciliolepis (C. Christensen) Lellinger, Amer. Fern J. 74: 58. 1984. TYPE: Bolivia. Sur Yungas, El Chaco valley, Asplund 1501 (isotype, US). Lellingeria dissimulans (Maxon) A. R. Smith, comb. nov. Basionym: Polypodium dissimu¬ lans Maxon, Contr. U.S. Natl. Herb. 10: 502. 1908. Ctenopteris dissimulans (Maxon) Cope¬ land, Philipp. J. Sci. 84: 415. 1956 [1955]. Grammitis dissimulans (Maxon) F. Seymour, Phytologia 31: 179. 1975. Melpomene dis¬ simulans (Maxon) A. R. Smith & R. C. Moran, Novon 2: 429. 1992. TYPE: Guatemala. Alta Verapaz: near Coban, von Tiirckheim (ed. Don¬ nell Smith 884) (holotype, US). On the basis of descriptions, this rare and poorly known species was mistakenly transferred to Mel¬ pomene (Smith & Moran, 1992), but examination of the type and several additional specimens from Guatemala (Johnson 678, NY, US; Steyermark 48781, US; Standley 71160, US) indicates that it belongs in Lellingeria (Smith et al., 1991). Lellin¬ geria dissimulans is similar to L. randallii (Maxon) A. R. Smith & R. C. Moran, from Panama and Jamaica, except that the rhizome scales are larger and lack marginal setae, and the blades completely lack hairs and setae on the lamina and rachis abax- ially; the sori are somewhat impressed. The species is so far known only from Guatemala. Lellingeria hombersleyi (Maxon) A. R. Smith, comb. nov. Basionym: Polypodium homber¬ sleyi Maxon, Amer. Fern J. 20: 1. 1930. Cten¬ opteris hombersleyi (Maxon) Copeland, Phi¬ lipp. J. Sci. 84: 430. 1956 [1955]. Grammitis hombersleyi (Maxon) Lellinger, Proc. Biol. Soc. Wash. 89: 714. 1977. TYPE: Trinidad. Hom- bersley 331 (holotype, US). This is known only from Trinidad and, like / dissimulans, is also related to L. randallii. Novon 5: 20-21. 1995. Volume 5, Number 1 1995 Smith Neotropical Grammitidaceae 21 Melpomene gracilis (Hooker) A. R. Smith, comb, nov. Basionym: Polypodium gracile Hooker, Bot. Misc. 2: 239. 1831. Grammitis gracilis (Hooker) Stolze, Fieldiana, Bot. 32: 96. 1993. TYPE: Peru. Pasco: Huayllay, Cruckshanks s.n. (holotype, K not seen; isotype, GH not seen). Melpomene zempoaltepetlensis (Mickel & Bei- tel) A. R. Smith, comb. nov. Basionym: Gram¬ mitis zempoaltepetlensis Mickel & Beitel, Mem. New York Bot. Card. 46: 205. 1988. TYPE: Mexico. Oaxaca: Dto. Mixe, NW slope of Cerro Zempoaltepetl, trail from Yacoche to Totontepec, Mickel 4638 (holotype, NY). I now believe this is sufficiently distinct from M. pilosissima to warrant species status. Melpomene zempoaltepetlensis differs in lacking setae at the margins of segments and appears to be confined to southern Mexico. Terpsichore bipinnata (Stolze) A. R. Smith, comb, nov. Basionym: Grammitis bipinnata Stolze, Fieldiana, Bot. 32: 92. 1993. TYPE: Peru. Cuzco: Prov. La Convencion, Loma Grande, Biles 2167 (holotype, US not seen; isotype, F not seen). Terpsichore elastica (Bory ex Willdenow) A. R. Smith, comb. nov. Basionym: Polypodium elasticum Bory ex Willdenow, Sp. PL, ed. 4. 5: 183. 1810. Ctenopteris elastica (Bory ex Willdenow) Copeland, Philipp. J. Sci. 84: 426. 1956 [1955]. TYPE: Bourbon [Reunion]. Bory s.n. (holotype, B-Herb. Willd. 19675 not seen; microfiche, UC). I have now seen sufficient material from Mada¬ gascar and the Seychelles to be convinced that this species is not synonymous with T. cultrata (Bory ex Willdenow) A. R. Smith, as previously thought (Smith, 1993). It differs by the narrower blades and by the absence of rhizome scales, thus agreeing with T. mollissima (Fee) A. R. Smith. Terpsichore elas¬ tica is the only known species in the genus from Africa and islands of the Indian Ocean. Like many of its neotropical congeners allied to 7. cultrata (Group 3 of Smith, 1993), the spores are ellipsoid, monolete, and binucleate when shed (van der Werff 12890 , UC, from Madagascar). Terpsichore flexuosa (Maxon) A. R. Smith, comb, nov. Basionym: Polypodium Jlexuosum Max¬ on, Contr. U.S. Natl. Herb. 17: 597, t. 42. 1916. Grammitis maxoniana Lellinger, Amer. Fern J. 74: 58. 1984 (nom. nov., not G. flex¬ uosa Kunth = Eriosorus flexuosus). Lellin- geria flexuosa (Maxon) A. R. Smith & R. C. Moran, Amer. Fern J. 81: 84. 1991. TYPE: Cuba. Oriente: Camp La Gloria, S of Sierra Moa, Shafer 8037 (holotype, US). On the basis of descriptions, this species was mistakenly transferred to Lellingeria by Smith and Moran (in Smith et al., 1991). Examination of the type and other specimens from Cuba shows that Terpsichore flexuosa is closely related to T. sta- heliana (Posthumus) A. R. Smith, also with setose sporangia, and not Lellingeria pendula (Swartz) A. R. Smith & R. C. Moran, as Maxon thought. Literature Cited Bishop, L. E. 1988. Ceradenia, a new genus of Gram- mitidaceae. Amer. Fern J. 78: 1-5. Mickel, J. T. & J. M. Beitel. 1988. Pteridophyte flora of Oaxaca, Mexico. Mem. New York Bot. Card. 46: 1-568. Smith, A. R. 1993. Terpsichore, a new genus of Gram¬ mitidaceae (Pteridophyta). Novon 3: 478-489. - & 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. Arracacia ravenii (Apiaceae), a New Species from Southern Mexico Lincoln Constance Herbarium and Department of Integrative Biology, University of California, Berkeley, California 94720, U.S.A. James Affolter The State Botanical Garden of Georgia, The University of Georgia, 2450 South Milledge Avenue, Athens, Georgia 30605-1624, U.S.A. Abstract. A new species of Arracacia (Apiaceae) is described from the high mountains of Chiapas and Oaxaca, Mexico. Although sharply distinct in many respects, it is contrasted with A. pringlei J. Coulter & Rose, the Oaxacan taxon it most closely resem¬ bles. Arracacia Bancroft is one of the largest and most important apioid genera of Umbelliferae in the New World, not least because some of its members served as important food plants for the Incas. The genus is central to a number of other Mesoamerican gen¬ era, such as Tauschia , Coaxana, Coulterophytum, Myrrhidendron, and Neonelsonia, all of which have been confused with it in one way or another. It continues to yield well-marked but previously un¬ described forms from throughout its extensive range. Attention is called to the following example to fa¬ cilitate its inclusion in Flora Mesoamericana. Arracacia ravenii Constance & Affolter, sp. nov. TYPE: Mexico. Chiapas: Municipio de Tene- japa, steep heavily wooded slope of Cerro Zon- tehuitz, with Quercus, Drimys & Magnolia on NE side of hill, paraje of Matsab, alt. 9200 ft. (2804 m), 25 Aug. 1966, D. E. Breedlove 15311 (holotype, DH; isotype, UC). Figure la-f. Species Arracaciae pringlei J. Coulter & Rose affinis sed habitu procumbenti, non glauco-caulibus, divisionibus foliorum confertis acutis non acuminatis, pedunculo so- litario terminali, involucello evidenti, floribus flavo-viren- tibus, fructibus late ovoideis differt. Slender, caulescent, branching, woody-based pe¬ rennial herbs forming trailing mats to 60 cm long, the foliage glabrous, the inflorescence scaberulous; leaves triangular-ovate, 5-15 cm diam., temate- bipinnate or 2-3-pinnately dissected, the ultimate divisions oblong-ovate to lanceolate, 5-15 mm long, 1-10 mm broad, crowded, sessile, acute, spinulose- serrate and pinnately incised toward base, the up¬ permost confluent, conspicuously reticulate, squa- mosely tufted on the rachis above, a well-defined nodal ring lacking; petiole 5-10 cm long, oblong¬ sheathing; cauline leaves numerous, like the basal; inflorescence of a solitary rather stout terminal pe¬ duncle 10-20 cm long; umbels 8-12 cm diam.; involucre lacking, or of inconspicuous filiform bracts; rays 12-18, mostly fertile, slender, 3.5-5 cm long, subequal, slightly webbed; umbellets 30-40-flow- ered, the 2-8 fertile pedicels slender, 5-10 mm long; involucel of 5-8 linear-lanceolate, scarious- margined, acuminate bractlets to 10 mm long, equal¬ ing flowers but shorter than fruit; flowers greenish yellow, the petals oval; stylopodium conical, the styles slender; carpophore apparently bipartite; fruit broadly ovoid, 4-5 mm long, 3-4 mm broad, gla¬ brous, tapering at apex, the ribs very prominent; vittae large, usually solitary in intervals, 2 on com¬ missure; seed channeled under intervals, the face sulcate; chromosome number unknown. Distribution. Known only from mountains of cen¬ tral Chiapas (Cerro Zontehuitz) and southern Oaxaca (Cerro Zempoaltepetl). Habitat. Steep slopes with Quercus, Drimys, Magnolia, Ilex, and Podocarpus, in wet montane or cloud forest, at 2400-3000 m. Arracacia ravenii keys readily to the group of woody-based species comprising A. fruticosa Rose, A. ebracteata (Rose) Mathias & Constance, A. ae- gopodioides (Kunth) J. Coulter & Rose, and A. pringlei J. Coulter & Rose, more particularly the last. It differs from A. pringlei in its sprawling habit, green rather than glaucous stems and rachises, crowded leaf divisions, solitary terminal peduncle rather than terminally clustered peduncles, evident involucel, yellowish green rather than purple flowers, and broadly ovoid rather than oblong-fusiform fruit. We are pleased to name the plant for Peter Raven, who, among so many other accomplish¬ ments, played a very significant role in initiating the Flora of Chiapas project. Volume 5, Number 1 1995 Constance & Affolter Arracacia ravenii 23 Figure 1. Arracacia ravenii Constance & Affolter. —a. Habit. —b. Basal leaf. —c. Fruiting umbellet. —d. Fruit, lateral view. —e. Mericarp transection. —f. Petal, (a, Breedlove & Raven 8105; b-f, Breedlove 15311.) Paratypes. MEXICO. Chiapas: Mun. of San Cristobal las Casas, SW slope of Cerro Zontehuitz, 19 Jan. 1965, Breedlove & Raven 8105 (DH, UC), 24 May 1965, Breedlove 10020 (DH, MICH, UC); Santa Cruz en San Felipe, 15 Nov. 1986, Ton & M. C. Mtz. de Lopez 9529 (MO); Mun of Tenejapa, slope at Paraje Matsab, 5 Jan. 1966, Ton 487 (DH, UC); slope in Paraje Pahal Ton, 20 Jan. 1966, Ton 668 { DH, UC), Cerro Sontehuits, Las Casas-Tenejapa, 28 June 1942, M. C. Carlson 2411 (MEXU); summit of Cerro Tzonohuitz, 92°35'W, 16°49'N, 26 Mar. 1986, Snow & Whittemore 212 (MO). Oaxaca: steep N slope of Cerro Zempoaltepetl, 35 km N of Ayutla, Mitla-Choapam, 17 Apr. 1988, Breedlove & Barthol¬ omew 66871 (CAS). Acknowledgment. The figure was prepared by Charlotte Mentges Hannan. Two New Species of Araceae for the Guianas Thomas B. Croat Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A. Abstract. Two species of Araceae from the Guianas are described. Philodendron billietiae, known from Guayana, French Guiana, and Brazil, is most closely related to P. acutatum. Xanthosoma granvillei, known only from French Guiana, is dis¬ tinguished mainly by its large, blackish-drying blades. In order to complete the Araceae treatment for the Flora of Saul and the Flora of the Guianas it has become necessary to describe a number of new species. Descriptions of other new species of Araceae will follow. Philodendron billietiae Croat, sp. nov. TYPE: French Guiana. National Hwy. 2, Cayenne- Regina, vie. km 93, 100-150 m, 4°18'N, 52°10'W, Croat 74321 (holotype, MO 4343643-5; isotypes, B, BBS, BR, BRG, CAS, CAY, CM, F, G, GH, INPA, K, M, MG, NY, P, US, VEN). Figures 1-4. Planta hemiepiphytica; internodia brevia, usque 2.5- 5 cm diam.; petiolus subteres, 25-57 cm longus, pallide aurantiacus-flavus vel viridi-flavus; lamina plus minusve anguste triangularis et sagittata, 30-85 cm longa, 8.1- 21 cm lata; nervis lateralibus primariis 3-5 utroque; pedunculus 9.4-27 cm longus; spatha 17.5-18.5 cm longa, extus viridis; lamina intus alba; tubus intus pur- purascens; spadix 16.5-18 cm longus; ovaria 6-11 lo- cularia cum ovulis plus quam 10 in quoque loculo; affinis P. acutato Schott sed internodis proportione brevioribus, petiolo in vivo fiavido vel ochraceo versus constanter viridi, lamina proportione angustiore basique lobis 35- 40° versus 60-75° divergentibus differt. Appressed-climbing hemiepiphyte, sometimes on rocks; stems about 1 m long, sometimes longer; internodes shorter than broad, 2.5-5 cm diam., light brown with irregular, weak, reticulate to mostly transverse fissures, turning yellow-brown; petiole scars to 4.5 cm diam.; cataphylls sharply D-shaped near apex, otherwise terete and unribbed or some¬ times bluntly 2-ribbed, green to pale orange (some¬ times brown, orange at base) or reddish, promptly caducous; roots moderately few, stout, ca. 5 mm diam., purplish brown, drying smooth, semiglossy, the epidermis often fissured; leaves usually clustered near the end of the stem, with erect spreading pet¬ ioles and pendent blades; petioles broadly spreading, 25-57 cm long, averaging 40 cm long, broadly Novon 5: 24-29. 1995. sulcate at base, obtusely flattened toward the apex, weakly angular near the apex, sheathed 4-8 cm at the base, firm, olive green to golden-yellow, pale yellow-orange or greenish yellow, when fresh, some¬ what asperous and rather densely purplish-lineate, drying light yellow-brown to dark brown, typically closely fissured; blades pendent, narrowly triangular to narrowly ovate-triangular to oblong-triangular, 30-85 cm long, 8.1-21 cm wide (averaging 52 x 15 cm), 2.7-4 times longer than wide, averaging 3.4 times longer than wide, broadest at the base or near the petiole attachment, sometimes weakly con¬ stricted above the petiole attachment, 0.85-1.4 times as long as the petioles (averaging 1.07 times longer than petioles), moderately coriaceous, dark green and semiglossy above, much paler and matte to weakly glossy below; the margins often weakly un¬ dulate, hyaline or reddish; posterior lobes directed at about 170° from the midrib (the two lobes sep¬ arated by an angle of 35-40°), sometimes directed up slightly at an angle to the midrib, 8-22 cm long, 3—10 cm wide, about twice as long as wide; sinus much longer than wide, acute at apex, usually barely naked along the margin, but sometimes weakly na¬ ked up to 2.5 cm (the leaf tissue merging almost imperceptibly with the posterior rib); midrib flat to very broadly raised and paler (sometimes orange- yellow) with purplish dots above, narrowly raised and convex, yellow-brown to yellow-red or yellow- green below; primary lateral veins 3-5, widely spaced and usually not prominent, weakly sunken to flat and yellow-green above, convex to weakly raised and sometimes reddish to purplish below, arising at 70-105° angle, mostly at nearly 90° angle, some¬ times prominently down-turned at the midrib, some¬ times splayed out at the midrib; minor veins nu¬ merous, moderately distinct when fresh, weakly raised above, flat and distinctly visible below, drying prom- inulous on both surfaces, with alternate veins fre¬ quently turned to join another minor vein just before the margin; cross veins inconspicuous; young leaves reddish. Inflorescences 1 3 per axil, erect; peduncle 9.4-27 cm long, ca. 1 cm diam., somewhat flat¬ tened, ribbed on one side, medium yellow-green, green-lineate to purplish-lineate; spathe 17.5-18.5 cm long, yellow-green to dark green outside, slightly paler toward apex, densely pale purplish-lineate on Volume 5, Number 1 1995 Croat Araceae from the Guianas 25 Figures 1-4. Philodendron billietiae Croat. —1. Habit, plant creeping over rocks in full sun; Croat 74230. — 2. Close-up of stem showing old intact cataphyll and leaf scars, roots, and petioles; Moonen s.n., cultivated in Cayenne. — 3. Inflorescence at anthesis showing male portion of spadix barely emergent; Moonen s.n. —4. Open inflorescence; Moonen s.n. 26 Novon tube, dark purplish-lineate on blade, the outer mar¬ gin of the tube slightly paler, the blade white to greenish white within, the margins weakly turned back at anthesis; tube sometimes reddish-spotted outside, mauve to maroon within, becoming dark maroon-purple within after anthesis with conspicu¬ ous, closely spaced resin canals restricted to the area of the constriction, blending in with the red of the tube and the white of the blade; spadix 16.5- 18 cm long; staminate portion white, to 9 cm long, to 11 mm diam. at base, weakly constricted to ca. 9 mm somewhat above the sterile portion, tapered to the narrowly rounded apex; pistillate portion pale green, to 7.2 cm long on front side, 5.3 cm long on back side; pistils ca. 3 mm long, pale yellow- green, turning pale yellow post-anthesis; ovaries 6- 11-locular; ovules more than 10 per locule; sterile male flowers broader than the pistillate portion, scal¬ loped and undercut at base, 2.2 cm diam., 10-11 mm long; fertile portion constricted 8.5 cm above base, creamy white, bluntly pointed at apex. Philodendron billietiae is known from Guyana, French Guiana, and Brazil (Para) in tropical rain¬ forest at 140 to 340 m elevation; certainly it is to be expected in Suriname as well. In Brazil it has been collected only near Belem, and only a live collection cultivated at the Moscow Main Botanical Garden has been seen. The species is a member of Philodendron subg. Philodendron (formerly Polyspermium) and is characterized by its long, mostly triangular leaf blades with a very narrow sinus, elongated posterior lobes, relatively few, widely spaced primary lateral veins, and especially by its greenish yellow or pale orange petioles, which are about as long as the blade. Philodendron billietiae was first collected by H. A. Gleason of the New York Botanical Garden in 1921 in Guyana, where it has not been collected subsequently. It is named in honor of Frieda Billiet of the Jardin Botanique National de Belgique, Meise, who studied the species at several localities in French Guiana, brought it into cultivation, and first brought it to my attention. The species is in cultivation in the botanical garden at Meise, as well as at the Missouri Botanical Garden and the Moscow Main Botanical Garden. Philodendron billietiae is most easily confused with and probably most closely related to P. acu- tatum Schott, a similar species with which it may occur. Philodendron acutatum differs in having proportionately longer intemodes, consistently me¬ dium green petioles, and proportionately wider, mostly narrowly ovate blades that often have the posterior lobes divergent from one another at an angle of 60° or more. In addition, P. acutatum is ecologically different from P. billietiae in French Guiana, occurring principally in the coastal swamp forests with Avicennia. Paratypes. FRENCH GUIANA. Pic Coudreau-Mont Bakra, Region des Emerillons, 670 m, 3°18’N, 52°57'W, de Granville & Cremers 11833 (CAY, MO, P, U, US); Savane Roche de Virginie-Approuague River basin, 140 m, 4°1 l'N, 52°9'W, Cremers & Petronelli 11683 (CAY, K, MO, P, U); Montagnes de Kaw, 4°33'N, 52°09'W, Cremers 12718 (CAY); Montagnes de Kaw, along road to Montagne Favard, vie. jet. to Fourgrassie, 200 m, 4°38'N, 52°17'W, Croat 74335 (CAY, K, MO, US); Route Nationale, RN 2, 4°35'N, 52°20'W, Cremers 12723 (CAY); km 95.5, Prevost 1739 (CAY); km 67, Crique Tibourou, along old route for RN 2, 5 m, 4°29'N, 52°19'W, Billiet & Jadin 5740 (B, BR, CAY, K, MO, NY); km 93, 100-150 m, 4°18'N, 52°10'W, Croat 74321 (B, BR, CAY, K, NY, MG, P, US, VEN); road to Montagnes de Kaw, km 34, 4°33'N, 52°09'W, 340 m, Billiet & Jadin 4600 (BR); La Compte River, vie. km 40 on RN 2, 4°N, 52°21'W, Billiet & Jadin 4714 (BR, CAY, MO); between La Compte & Cacao, S of Cayenne, 1981, Billiet 81-0879 (BR), Croat 76441 (MO); Saul, vie. Eaux Claires, Sentier Botanique near headwaters of St. Eloi River, 350 m, Croat 74230 (CAS, CAY, CM, M, MO, W). GUY¬ ANA. Rockstone, Gleason 846 (GH). In addition to the localities listed above, the spe¬ cies has been seen in French Guiana on the road to Petit Saut, km 20 (F. Billiet, pers. comm.). Xanthosoma granviliei Croat & Thompson, sp. nov. TYPE: French Guiana. Saul, vie. Eaux Claires, along road between Eaux Claires and Belizon, 230 m, 3°37'N, 53°12'W, Croat 74131 (holotype, MO-4342303; isotypes, B, CAY, CM, K, NY, P, US). Figures 5-8. Planta terrestris; caulibus 15-30(-50) cm longis; in- ternodia usque 6(—15) cm diam.; petiolus teres, 39- 75(-100) cm longus; lamina ovato-hastata, 19-70 cm longa, (16-)21-50 cm lata, 2.1-2.3 plo longior quam lata, in sicco subdenigrata; cum costa postica nuda 1.0- 5.5 cm; nervis primariis lateralibus 3-6 utroque; inflo- rescentiae 3-6 per axila; pedunculus 8-30 cm longus; spatha 11.5-20 cm longa; tubus 4.5-6.5 cm longus, extus viridus, intus atropurpureus; lamina 8-8.5 cm lon¬ ga, extus et intus albida; spadix usque 18 cm longus, parte pistillata plus minusve aurantiaca; ovaria 2-3 lo- cularia, loculi 2-3 ovulati; fructus aurantiacus. Terrestrial; stems rhizomatous, the older portion creeping over ground, the younger portion erect, 15—30(—50) cm long; internodes short, to 6(— 15) cm diam.; cataphylls to 25 cm long, persistent, turning dark brown, the uppermost persisting semi¬ intact; petioles 39-75(-100) cm long, terete but somewhat flattened near apex, succulent and juicy, tinged purplish at base, sheathed usually less than 1 4 its length (up to 20 cm when subtending an in¬ florescence); blades ovate-hastate, 19-70 cm long. Volume 5, Number 1 1995 Croat Araceae from the Guianas 27 Figures 5-8. Xanthosoma granvillei Croat & Thompson; Croat 74131. —5. Habit showing an inflorescence and an emerging new leaf. —6. Stem with old persistent leaf sheaths. —7. Leaf blade. —8. Close-up of inflorescence showing an unopened spathe. 28 Novon (16-)21-50 cm wide, 2.1-2.3 times longer than broad, prominently lobed at base, acuminate at apex, dark green and semiglossy above, moderately paler and weakly glossy below, drying somewhat black¬ ened, dark olive-green above, dark brownish green below; the lateral margins usually convex, rarely concave; posterior lobes 1.2-1.8 times longer than wide, subacute at apex, sometimes turned weakly outward, often held somewhat upward at an angle to the midrib; sinus hippocrepiform or sometimes obovate, 9-19 cm deep; posterior rib straight and extending outward at ca. 60° angle to the midrib, naked 1-5.5 cm along sinus; basal veins 6-9 pairs, the uppermost free, the remainder pinnately ar¬ ranged along the posterior rib; primary lateral veins 3-6 pairs, arising at 40-60° angle; minor veins weakly etched above, flat and darker than surface below. Inflorescences 3-6 per axil; peduncle green¬ ish, 8-30 cm long, longer than or shorter than the spathe; spathe 11.5-20 cm long, constricted at low¬ er !4, the tube 4.5-6.5 cm long, the blade white, sometimes lined with pink, the tube green outside, sometimes tinged purplish, dark purple within, some¬ times becoming deep red outside; spathe blade 8- 8.5 cm long, whitish on both surfaces; spadix to 18 cm long, several centimeters shorter than spathe, female portion bright orange to yellow-orange; ova¬ ries 2-3-locular; locules usually 2-3-ovular. In- fructescences with peduncles to 22 cm long, fruiting spathe to 7 cm long, fruiting spadix 3.5-4.5 cm long, 2-2.5 cm diam. Fruits orange; immature seeds 1.6-1.8 mm long, 1 mm diam., obtuse at apex, cordulate at base, ca. 10-ribbed longitudinally. Very common and conspicuous in wet season but surviving as an underground stem during the dry season. Xanthosoma granvillei is known only from French Guiana, mostly between 200 and 500 m, but rarely to nearly sea level; it is to be expected in adjacent Suriname. The species was first collected in 1966 by Oldeman at Cacao, south of Cayenne. It is named for Jean Jacques de Granville from ORSTOM, in Cayenne, who collected more well- prepared specimens of the species than any other collector. Xanthosoma granvillei is recognized by its short stems emerging usually only a short distance above the ground level, its ovate-cordate, blackish-drying blades with triangular posterior lobes, sunken minor veins, and a partly naked, usually hippocrepiform sinus. Additionally, the cluster of 3—6 inflorescences with peduncles about as long as the spadix, with orange female spadices, distinguish it from other species of Xanthosoma. Xanthosoma granvillei is most easily confused with another blackish-drying species that also ap¬ pears to be new to science. That species differs by having tuberous growth, smaller, more hastate leaf blades, and an inflorescence with slender peduncles that are 3-4 times longer than the spathe. It is represented by collections from the Montagnes de Kaw (Cremers 2719, 8702) and from the nearby region at Crique Armontabo, a tributary of the Oya- pock River ( Cremers 7055). It will be described later. Xanthosoma granvillei might also be confused with X. undipes K. Koch & Bouche, which differs in having a longer stem, foul-smelling sap, and blades that dry greenish rather than blackened. Superfi¬ cially, X. granvillei is very similar to X. holivar- anum Bunting from the El Dorado-Gran Sabana area of Venezuela. That species, described from sterile material, has similar stems and blades but differs in having obtusely sulcate petioles sheathed to about the middle (versus merely obtusely flattened and usually less than ‘4 its length for X. granvillei), and leaf blades that dry gray-green above and light yellow-green below (not at all blackened). In addi¬ tion, X. bolivaranum lacks any obvious lateral veins directed off the posterior rib toward the base, where¬ as X. granvillei has 2-3 prominent lateral veins branched off the posterior rib and directed toward the base. Paratypes. FRENCH GUIANA. Cacao, 60 km S of Cayenne, Oldeman B-450 (P); Montagne Cacao, SE of Cayenne, Cremers 13031 (CAY, MO), Feuillet 548 (CAY); Basin de la Comte River basin, 4°34'N, 52°27'W, Cre¬ mers 12728 (CAY); Region de Paul Isnard, Citron Petite piste Boeuf Mort, Feuillet 653 (CAY, K); Mont Atachi Bacca-Inini River region, 9 km SE of Gobaya Soula, 420 m, 3°33'N, 53°55'W, de Granville et al. 10649 (CAY, K); E of Gobaya Soula, 440 m, 3°33'N, 53°55'W, de Granville et al. 10482 (B, CAY, FTC, K, US); Montagnes de Kaw, road to Montagne Tresor, side trail to Placer Tresor, 4°35'N, 52°18’W, 150 m, Croat 74283 (CAY, MO); Montagnes de Kaw, Cascade de Fourgassie, 4°37'N, 53°18'W, Cremers & Feuillet 12968 (CAY, MO); Mon¬ tagnes de Kaw, road to Degrad Lalanne, 300 m, 4°34' N, 52‘TO’W, Hoff et al. 6308 (CAY); Station des No- uragues, Prevost 2503 (CAY); Saul, Mont Galbao, 550 m, 3°36'N, 53°17'W, de Granville et al. 8608 ( B, CAY, K, NY, P, U, US), 500 m, de Granville et al 8837 (CAY, K, US); Saiil, in village, jardin de Richard, Capus 2 (CAY); 10 km SSW of village, de. Granville 5420 (BR, CAY, K, P); 18 km S of village, de Granville 5531 (BR, CAY, K, P); Saiil, Pied de Mont Galboa, 200 m, 3°37'N, 53°16’W, de Granville et al. 9042 (CAY); Mont La Fumee, 3°37'N, 53°12'W, Mori & Pipoly 15575 ( NY); Saiil, Fumee, Prevost 1798 (MO); Saul, La Fumee Moun¬ tain trail, 250 m, Mori & Grade 18865 (NY); Mont Galboa, 10 km SW of Saiil, 650 m, de Granville 2401 (CAY, P); Saul, road to Eaux Claires, 250 m, Mori & Grade 21083 (NY); Saiil, vie. Eaux Claires, along road between Eaux Claires and Belizon, 220-240 m, 3°37'N, 53°12'W, Croat 74131 (B, CAY, CM, K, MO, NY, P, Volume 5, Number 1 1995 Croat Araceae from the Guianas 29 US), de Granville 4480 (K); Saul, road to Belizon, 0-2 km S of Eaux Claires, Mori et al. 22194 (MO, NY); Saiil, Grand Boeuf Mort trail, 300 m, Mori & Grade 18935 (CAY, NY); vie. Saiil, trail to Mont Galboa, de Granville 4486 (CAY, K); Rte. de Belizon, km 7.7, 4°20'N, 52°23'W, Billiet & Jadin 4424 (BR, CAY); Saiil, ENE of village, Pic Matecho, Cremers 6184 (CAY, K); Saiil Region, Saut Quaimicouare, Yaroupi River (trib¬ utary of the Oyapock), de Granville 345 (P); Bassin du Sinnamary, Crique Kourcibo, Saut Kawene, 12 m, 4°53'N, 53°3'W, Hoff et al. 6475 (CAY, MO); Chemin de fer Gare-Tigre-St. Elie, km 0-5, 20 m, 4°57'N, 53°8'W, Hoff et al. 6585 (CAY, MO). Paspalum niquelandiae (Poaceae: Paniceae), a New Species from the Serpentine Outcrops of Central Brazil Tarciso S. Filgueiras Reserva Ecologica do IBGE, Caixa Postal 08770, 70312-970 Brasflia, DF, Brazil ABSTRACT. Paspalum niquelandiae Filgueiras, a new species of Poaceae from the serpentine outcrops of Niquelandia, a municipality in the Brazilian state of Goias, is described, illustrated, and compared with Paspalum soboliferum Chase, P. humboldtianum Flugge, and P. inconstans Chase. The conserva- tional importance of a serpentine endemic grass flora in central Brazil is briefly discussed. Recent fieldwork carried out in the municipality of Niquelandia, in the Brazilian state of Goias, where serpentine soils have been reported (Brooks et al., 1990), resulted in the description of several new taxa in the Poaceae (Davidse & Filgueiras, 1993; Filgueiras et al., 1993; Filgueiras & Davidse, 1994). A new species of Paspalum L. from the same area is herein described, illustrated, and compared to similar species. Paspalum niquelandiae Filgueiras, sp. nov. TYPE: Brazil. Goias: Municipio de Niquelandia, ca. 14'T8'S-48°23'W, morro pedregoso, ca. 2 km Leste da localidade de Macedo, planta crescendo entre pedras, proximo a pequena floresta de galeria, ocasional na area, 19 maio 1993, T. S. Filgueiras & F. C. de A. Oliveira 2461 (holotype, IBGE; isotypes, F, G, ICN, K, MEXU, MG, MO, R, SI, SP, US). Figure 1. Paspalo sobolifero Chase spiculis ellipticis subtiliter pilosis, gluma superiore marginibus ciliatis simile, sed cul- mis non ramosis, ligula 0.2-0.3 mm longa, laminis 20- 45 cm longis et 8-22 mm latis, communi rhachidi 9-15 cm longa, gluma superiore ciliis densis, numerosis, co- roniformibus abunde differt. Coarse, caespitose perennial. Rhizomes well de¬ veloped, knotty, covered with cataphylls. Culms 120- 185 cm tall, erect, unbranched in the vegetative portion, with 5-8 elongated internodes; internodes hollow, stramineous, glabrous; nodes dark, glabrous. Prophyll not seen. Leaves mostly cauline; sheaths strongly striate, glabrescent to pilose in the lower half, becoming papillose-hispid upward, the margins free, both ciliate or one glabrous toward the base; Novon 5: 30-33. 1995. auricles absent; ligule a minute, brown, glabrous membrane, v-shaped, 0.2-0.3 mm long; cilia at the ligular area 5-8 mm long, colorless; blades 20-45 cm long, 8-22 mm wide, flat, lanceolate, glabrescent to papillose-hispid, acuminate apically, the midrib projecting abaxially, the margins serrate-denticulate to ciliate; uppermost blade reduced, minute. Inflo¬ rescence of unilateral, ascending racemes, divergent in maturity; peduncle well exserted, 12-22 cm long, glabrous; common rachis 9-15 cm long, glabrous, cylindrical at the base, becoming 3-4-angulate up¬ ward, the margins denticulate, the internode at the base of each raceme with a tuft of colorless hairs 2-2.5 mm long; racemes 8-30, 2.5-8.5 cm long; rachis 1-1.5 mm wide, zig-zag, flat abaxially, with a central ridge adaxially, arcuate in maturity, bear¬ ing spikelets to the tip, the margins ciliate or den¬ ticulate, or both. Pedicels of unequal length, 0.1- 0.3 mm long, scabrid, the apex discoid. Spikelets paired, each pair alternate on each side of the rachis, pedicelled, 2.3-3.3 mm long, 1-1.2 mm wide, broadly elliptic-lanceolate in outline, pale to purplish, dorsally compressed, abaxial, awnless, with 2 florets, disarticulating below the glume and falling as a unit; lower glume absent; upper glume slightly to strongly convex on the back, 2.3-3.3 mm long, 3-nerved, glabrous to finely appressed pubescent, the margins densely ciliate-pectinate; the cilia colorless, 0.3-0.7 mm long, with bulbous bases, the apex apiculate; lower floret sterile, consisting only of a lower lemma; lower lemma 2.2-2.9 mm long, narrower than the upper glume, 3-nerved, navicular, hyaline, with a depressed area at the base, the surface glabrous to short-pubescent, the apex glabrous to slightly ciliate; lower palea absent; upper floret 2.2-2.7 mm long, bisexual, slightly chartaceous, pale; upper lemma obscurely 5-nerved, navicular, glabrous, the surface finely papillose-striate, the apex acute; germination flap evident; upper palea 2.1-2.2 mm long, acu¬ minate, glabrous, slightly chartaceous, the surface finely papillose-striate; lodicuies 2, fleshy, condu- plicate, 0.1-0.2 mm long; stamens 3; anthers 1.2- 2 mm long, laterally exserted; ovary 0.5—0.8 mm long, glabrous; styles 2, separate, 1.5-1.8 mm long; stigmas 2, plumose, yellow or purplish. Caryopsis 1.8-1.9 mm long, 0.9-1 mm wide, light-colored. Volume 5, Number 1 1995 Filgueiras Paspalum niquelandiae 31 Figure 1. Paspalum niquelandiae Filgueiras, illustrated from the type collection (Filgueiras & Oliveira 2461 ). — A. Habit. — B. Ligular area of leaf. —C. Portion of a raceme with paired spikelets. — D. Dorsal view of spikelet. —E. Upper floret, lemma side. —F. Upper floret, palea side. 32 Novon Table 1. Main distinguishing characteristics of Paspalum niquelandiae , P. humboldtianum , and P. soboliferum. Character P. niquelandiae P. humboldtianum P. soboliferum Habit Erect Reptant Erect Height (cm) 120-185 40-120 ca. 60 Culm Unbranched Branched Branched Nodes Glabrous Pilose Glabrous Ligule (mm) 0.2-0.3 1.8-3.8 2-2.5 Blades (cm x mm) 20-45 x 8-22 1-8 x 6-17 5-15 x 6-9 Common rachis (cm) 9-15 1.5-6 4-6 Number of racemes 8-30 1-8 3-7 Raceme length (cm) 2.5-8.5 4-10 2-4.5 Rachis width (mm) 1-1.5 1-3 1-1.2 Spikelet size (mm) 2.3-3.3 3-4 2.8-2.9 Cilia on glume (mm) 0.3-0.7 1.9-2.1 0.4-0.5 with persistent stylar bases; embryo %-% as long as the caryopsis; hilum punctate, basal, Z l0 -'/ s as long as the caryopsis. Paspalum niquelandiae is morphologically most closely related to Paspalum soboliferum Chase be¬ cause of the shape of the spikelets and especially because of the cilia on the margins of the upper glume; it differs in the unbranched culms, smaller ligule, longer blades, longer common rachis, greater number of racemes, and size and density of the cilia on the margins of the upper glume. Paspalum so¬ boliferum is known only from a single plant (the holotype) from Ecuador (Hitchcock, 1927). The new species is more distantly related to Paspalum hum- boldtianum Flugge; the latter is readily distinguished by its sprawling habit, the pilose nodes, the size of the ligule, blades, and common rachis, and the num¬ ber of racemes. Paspalum humboldtianum occurs from Mexico to Argentina (Chase, 1929). Table 1 summarizes the main differences among P. nique¬ landiae , P. humboldtianum , and P. soboliferum. It is evident from Table 1 that these species form a trio of closely related species with some overlap¬ ping characteristics, but whose distinctiveness is also readily apparent. The spikelet shape and the ciliate lower lemma remotely link Paspalum niquelandiae to P. incon- stans Chase, a species from Bolivia, Ecuador, and Peru. The latter is of smaller stature (culms 50 to 60 cm high), has both terminal and axillary racemes, and possesses a lower number of racemes (1 to 8, but commonly 2 to 5). A further distinguishing char¬ acteristic is the presence of a lower glume in the lower spikelets of P. inconstans (occasionally lack¬ ing in some spikelets). In addition, the cilia on the lower lemma of P. inconstans are smaller and much less dense than in P. niquelandiae. The classification of Paspalum niquelandiae is not yet clear. It is obviously related to a group of species included by Chase in her unpublished mono¬ graph of the South American species of Paspalum [accessible from the Hitchcock & Chase Grass Li¬ brary of the Department of Botany, Smithsonian Institution, Washington, D.C., U.S.A] in subgenus Ceresia (Persoon) Reichenbach. This group of spe¬ cies (which includes Paspalum bicilium Mez, P- carinatum Kunth, P. polyphyllum Nees, among others) has a characteristic upper glume with con¬ spicuous marginal cilia. The inclusion of P. nique¬ landiae in subgenus Ceresia is not acceptable be¬ cause this would make the subgenus exceedingly heterogeneous. Although related to some species in subgenus Ceresia, P. niquelandiae is best left un¬ grouped until more convincing evidence of relation¬ ships becomes available. Paspalum niquelandiae is known from two pop¬ ulations in the municipality of Niquelandia in the Brazilian state of Goias. It is apparently restricted to the serpentine outcrops in the area. It grows in scattered clumps among rocks and boulders, near the edge of some gallery forests, but was never found inside the gallery forest itself. Although the clumps seen were rather robust and, therefore, easy to spot, large populations of the species were never located. This is the third species of Paspalum described from the locality of Macedo in Niquelandia. The other two are P. longiaristalum Davidse & f “• gueiras (Davidse & Filgueiras, 1993) and P■ biar- istatum Filgueiras & Davidse (Filgueiras & Davidse, 1994). Ophiochloa Filgueiras, Davidse & Zuloaga, a panicoid grass genus, has also been recently de¬ scribed from the same area (Filgueiras et al., 1993). The discovery of so many new taxa in such a small area (ca. 3 km 2 ) in central Brazil suggests that Ni¬ quelandia is a center of high plant diversity and Volume 5, Number 1 1995 Filgueiras Paspalum niquelandiae 33 endemism, primarily due to the presence of serpen¬ tine soils, with their high concentrations of nickel, chromium, cobalt, and other minerals (Brooks, 1987). The presence of a highly specialized flora in Ni- quelandia is of utmost concern because the area is the center of the largest nickel mining and smelting operation in Brazil (Brooks et al., 1990). There is not a single conservation unit in the area to grant legal protection to this unique flora. A detailed bo¬ tanical survey should be undertaken in the region to assess the diversity of the entire flora and its level of endemism. The conservational implications of the presence of such a peculiar flora in Niquelandia will be addressed in a forthcoming publication. The specific epithet refers to the town of Nique¬ landia, in whose municipality the new species was collected. Paratypes . BRAZIL. Goias: municipio de Niquelan¬ dia, ca. 1 km Leste da localidade de Macedo, morro pedregoso, 19 maio 1993, T. 5. Filgueiras & F. C. de A. Oliveira 2456 (IBGE, RB, UB), 4 ago. 1992, T. S. Filgueiras & R. D. Lopes 2427 (B, IBGE). Acknowledgments. I am grateful to CAPES (Bra¬ zilian Post-Graduation Federal Agency) and IBGE (Instituto Brasileiro de Geografia e Estatistica) for a post-doctoral fellowship that enabled me to spend 18 months at the Missouri Botanical Garden. I also thank the Missouri Botanical Garden staff for mak¬ ing their facilities available for my research. I es¬ pecially thank my friend Clarence J. Lovejoy, Jr., and his family for their friendship and support during my stay in St. Louis. Vladimiro S. Dudas, San Isidro, Argentina, provided the excellent illustration. Literature Cited Brooks, R. R. 1987. Serpentine and Its Vegetation. Dioscorides Press, Portland. Brooks, R. R., R. D. Reeves, A. J. M. Baker, J. A. Rizzo & H. D. Ferreira. 1990. The Brazilian serpentine expedition (Braspex). 1988. Natl. Geogr. Res. 6: 205-219. Chase, A. 1929. The North American species of Pas¬ palum. Contr. U.S. Natl. Herb. 28: 1-310. Davidse, G. & T. S. Filgueiras. 1993. Paspalum lon- giaristatum (Poaceae: Paniceae), a new serpentine endemic from Goias, Brazil, and the first awned spe¬ cies in the genus. Novon 3: 129-132. Filgueiras, T. S. & G. Davidse. 1994. Paspalum biar- istatum (Poaceae: Paniceae), a new serpentine en¬ demic from Goias, Brazil, and the second awned species in the genus. Novon 4: 18-22. -,- & F. O. Zuloaga. 1993. Ophioch- loa , a new endemic serpentine grass genus (Poaceae: Paniceae) from the Brazilian cerrado vegetation. No¬ von 3: 360-366. Hitchcock, A. S. 1927. The grasses of Ecuador, Peru, and Bolivia. Contr. U.S. Natl. Herb. 24: 291-556 [Paspalum, contributed by Agnes Chase, pp. 434- 455]. A New Combination in Peltostigma (Rutaceae) Roy E. Gereau Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. Abstract. Peltostigma parviflorum Q. Jimenez & Gereau is reduced to synonymy under P. gua- temalense (Standley & Steyermark) Gereau, comb, nov., based on Galipea guatemalensis Standley & Steyermark. Peltostigma Walpers (Rutaceae: Rutoideae) be¬ longs to subtribe Choisyinae of the tribe Zanthox- yleae, while Galipea Aublet (Rutaceae: Rutoideae) belongs to subtribe Cuspariinae of the tribe Cus- parieae. The two subtribes may be contrasted as follows (Engler, 1931): Choisyinae: flowers actinomorphic; corolla poly- petalous; stamens all antheriferous; seeds with endosperm. Cuspariinae: flowers usually zygomorphic; corolla usually gamopetalous; some stamens often reduced to staminodes; seeds with little or no endosperm. Peltostigma parviflorum Q. Jimenez & Gereau is the only species of the genus with unifoliolate leaves and small flowers (petals 8.2-10.1 mm long) (Jimenez & Gereau, 1991); its congeners (P. pte- leoides (Hooker) Walpers and P. eximium C. V. Morton) possess 3—5-foliolate leaves and large flow¬ ers (petals 15-25 mm long) (Morton, 1933; Stan¬ dley & Steyermark, 1946). Galipea guatemalensis Standley & Steyermark was named from a fruiting specimen lacking the floral characters necessary for correct generic (or even subtribal) placement in the Rutaceae. Examination of its holotype has shown that G. guatemalensis, an earlier name, is the same species as P. parviflorum, necessitating the follow¬ ing new combination. Peltostigma guatemalense (Standley & Stey¬ ermark) Gereau, comb. nov. Basionym: Gali¬ pea guatemalensis Standley & Steyermark, Publ. Field Mus. Nat. Hist., Bot. Ser. 23: 165. 1944. TYPE: Guatemala. Izabal: Rio Dulce, between Livingston and 6 mi. upriver, on N side, 1—25 m, 14 Apr. 1940 (fr), Steyermark 39448 (holotype, F). Peltostigma parviflorum Q. Jimenez & Gereau, Arm. Missouri Bot. Gard. 78: 527. 1991. Syn. nov. TYPE: Costa Rica. San Jose: 9°43T0"N, 84°23T0"W, Zona Protectora La Cangreja, Santa Rosa de Puriscal, 400-500 m, 4 sep. 1987 (fl), Q. Jimenez M. & L. J. Poveda A. 494 (holotype, CR; isotypes, K, MO). Jimenez & Gereau (1991) discussed geographical trends in the morphology of Peltostigma guate¬ malense (as P. parviflorum) in terms of a “northern race” consisting of the collections from Guanacaste and Puntarenas, Costa Rica, and a “southern race comprising the type and four collections from An- tioquia, Colombia. The “races” may be contrasted as follows: “Northern race”: petioles 0.2-2.1 cm long, not extremely variable, the longest in a given specimen exceeding the shortest by 0.5-1.5 cm; leaflet blade 3.2-16.0 x 1.2-7.3 cm, the apex obtuse or obtusely acuminate, the base acute to rounded; petals 8.2-9.3 x 8.0-8.5 mm. “Southern race”: petioles 0.3-5.6 cm long, extremely variable, the longest in a given specimen exceeding the short¬ est by 2.0-4.5 cm; leaflet blade 4.2-20.6 x 2.2- 10.2 cm, the apex acutely acuminate, the base usually attenuate; petals 9.8-10.1 x 8.5-9.7 mm. The type of Galipea guatemalensis has larger leaflets (to 19.5 x 8.7 cm) and longer petioles (to 2.6 cm) than is characteristic for the “northern race,” but is otherwise more similar in foliar char¬ acters to the northern than to the southern popu¬ lations. A previously unreported collection of this species from Peru has foliar characters entirely with¬ in the parameters of the “southern race,” but in¬ sufficiently mature flowers to assess the correlation between geography, floral measurements, and foliar characters. With few collections and little available data on flower size, it is preferable to continue to consider Peltostigma guatemalense a single species with some observable correlation between geography and morphological characters. Further investiga¬ tions based on more ample collections could reveal the desirability of recognizing the two informal “rac¬ es” as subspecies. Specimen examined. PERU. Madre de Dios: Manu, Parque Nacional Manu, Rio Manu, Pakitsa station, Tach- igali Trail, 4 km N of camp, 11°56'S, 71'16'W, 350 m, 11 Sep. 1989 (fl), R. B Foster & H. Beltran 11371 (F, MO). Novon 5: 34-35. 1995. Volume 5, Number 1 1995 Gereau Peltostigma 35 Acknowledgments. I thank J. A. Kallunki and A. Pool for bringing the type of Galipea guate- malensis to my attention; J. S. Miller and G. E. Schatz for useful discussions of geographic vari¬ ability; and C. M. Taylor for a careful reading of the manuscript. Literature Cited Engler, A. 1931. Rutaceae. Nat. Pflanzenfam. ed. 2, 19a: 187-359. Jimenez M., Q. & R. E. Gereau. 1991. Peltostigma parviflorum (Rutaceae), nueva especie de Costa Rica y Colombia. Ann. Missouri Bot. Gard. 78: 527-530. Morton, C. V. 1933. A new species of Peltostigma from Mexico. Proc. Biol. Soc. Wash. 46: 83-84. Standley, P. C. & J. A. Steyermark. 1946. Rutaceae. In: P. C. Standley & J. A. Steyermark, Flora of Guatemala, Fieldiana, Bot. 24(5): 389-425. Triniochloa talpensis (Poaceae: Pooideae), a New Species with Deciduous Leaves from Mexico Manuel Gonzalez-Ledesma, Maricela Gomez-Sanchez, Stephen D. Koch Centro de Botanica, Colegio de Postgraduados, 56230 Chapingo, Edo. de Mexico, Mexico Teresa Mejia-Saules Instituto de Ecologia, A.C., Apartado Postal 63, 91000 Xalapa, Veracruz, Mexico ABSTRACT. Triniochloa talpensis from Mexico is described and illustrated. Its morphology and leaf anatomy are compared with those of T. laxa Hitch¬ cock. RESUMEN. Se describe e ilustra Triniochloa tal¬ pensis de Mexico. Se comparan caracteristicas de morfologia y de anatomia foliar con aquellas de 7. laxa Hitchcock. Triniochloa is a small genus of grasses that was described by Hitchcock in 1913, the members of which grow in temperate forests from Mexico to Bolivia. It is distinguished from other genera of the Pooideae by its spikelets 1-flowered without a rach- illa extension, floret callus obtuse and bearded, and lemma 5- or 7-nerved with a bifid apex and genic¬ ulate awn inserted above the middle. This genus was included in the tribe Meliceae of the subfamily Pooi¬ deae by Reeder (1968) on the basis of its connate leaf sheaths, membranous ligules, truncate lodicules that lack vascular traces, and small chromosomes. He suggested that it is closely related to Schizachne. This placement is supported by Clayton & Renvoize (1986) and Watson & Dallwitz (1992). Triniochloa, as presently circumscribed, includes four species: T. laxa Hitchcock and T. micrantha (Scribner) Hitchcock, which are restricted to Mex¬ ico; T. stipoides (H.B.K.) Hitchcock, distributed from central Mexico to Colombia, Venezuela, and Bolivia; and T. andina Luces, known only from Venezuela. The new species from western Mexico increases the number to five. Mature blades of basal leaves from herbarium specimens were used for anatomical studies. A seg¬ ment about 10 mm long was removed from the middle of the blade, rehydrated by boiling in water, sectioned free-hand, and mounted in glycerin jelly. Epidermis preparations were made following Met¬ calfe’s (1960) method. Drawings were made using an X-100-1 Ken-A-Vision Microprojector. Novon 5: 36-39. 1995. Triniochloa talpensis M. Gonzalez-Ledesma & M. Gomez-Sanchez, sp. nov. TYPE: Mexico. Jalisco: Municipio de Talpa de Allende, alrede- dores de la mina Zimapan en la Sierra de Cuale, ca. 2150 m, 10 Feb. 1992, M. Gonzalez- Ledesma & ]. A. Perez de la Rosa 496 (ho- lotype, CHAPA; isotypes, IBUG, MEXU, MICH, MO, US, XAL). Figure 1. Plantae perennes, rhizomatosae. Culmi 60-110 cm alti. Vaginae foliorum pubescentes prope nodos. Ligulae 1.5- 2.5(-5) mm longae. Laminae foliorum planae 22- 27 cm longae et 3.0-5.0 mm latae inferioribus caducis. Panicula erecta, angusta, 16-34 cm longa, ex vagina exserta. Spiculae 11.0-16.8 mm longae. Glumae inae- quales, prima spiculis breviore, 7.5-13.6 mm longa; se- cunda quam lemma aequanti vel longiore, 11.0-16.5 mm longa. Lemma 11.0-14.5 mm longum, 7-nerve, arista dorsali geniculata, 13.0-23.0 mm longa. Antherae (4.8-) 5.5- 7.0 mm longae. Perennial herbs with rhizomes to 20 cm long, 2- 3 mm diam., rhizome sheaths strigose near the nodes. Culms 60-110 cm tall, 1.5-2.0 mm diam. near the base, robust, strongly decumbent, pubescent below the upper nodes. Leaves, at least the lower, ciliate near the collar. Leaf sheath margins connate, the upper sheaths shorter than the internodes and pu¬ bescent below. Ligules 1.5-2.5(-5) mm long, mem¬ branous. Blades 22 27 cm long, 3.0-5.0 mm wide, flat, deciduous at level of the collar or beneath, adaxial surface scabrous to pubescent, with promi¬ nent ribs, abaxial surface glabrous. Panicle 16—34 cm long, 1.0—2.0(—2.5) cm wide, erect, narrow, completely exserted, with 30-65(-90) spikelets; panicle axis glabrous below, scabrous on the angles toward the apex; lowest node 1-2-branched; branch¬ es appressed or ascending, scabrous at least toward the apex. Spikelets 11.0-16.8 mm long, 1-flowered. Glumes unequal, hyaline except near the nerves. First glume 7.5—13.6 mm long, shorter than the spikelet, 1-nerved. Second glume 11.0-16.5 mm long, as long as the spikelet, 3-nerved. Lemma 11.0— 38 Novon Figure 2. Leaf blade anatomy. A-D. Triniochloa talpensis (Gonzalez-Ledesma & Perez de la Rosa 496 , CHAPA). —A. Outline of half of the blade. —B. Detail of transverse section, including the midrib. —C. Abaxial epidermis as seen in surface view. —D. Prickle. E-G. Triniochloa laxa (T. Mejia-Saules 1820 , CHAPA). —E. Outline of the blade. —F. Detail of transverse section, including the midrib. — G. Abaxial epidermis as seen in surface view. Stippling indicates the costal region in both C and G. 14.5 mm long, with a basal stipe 0.5 mm long, rounded dorsally, glabrous or scabrous at the apex, 7-nerved, the central 3 converging at or above the middle to form the awn; apex bifid; awn 13.0-23.0 mm long, geniculate; callus with hairs up to 5 mm long. Palea shorter than lemma. Rachilla not pro¬ longed beyond floret. Stamens 3, anthers (4.8-)5.7- 7.0 mm long. Lodicules 2, rounded, connate, short, and turgid. Caryopsis not seen. Leaf Blade Anatomy (Fig. 2A-D) Transverse section. Outline of the blade flat to involute. Adaxial surface with squared, flat-topped ribs associated with first-order vascular bundles and rounded ribs associated with second-order bundles; furrows narrow, moderately deep. Abaxial surface smooth, midrib only slightly projecting. Outer vas¬ cular bundle sheaths complete or interrupted by abaxial or adaxial sclerenchyma, round in outline. Sclerenchyma girders present, the adaxial T-shaped. Chlorenchyma nonradiate, its cells of various sizes and shapes, continuous between adjacent vascular bundles. Colorless parenchyma cells absent. Bulli- form cells in fan-shaped, adaxial groups not pro¬ jecting above the adjacent epidermis. Abaxial epidermis. Long cells longer than wide with undulating anticlinal horizontal walls. Stomata absent. Intercostal short cells solitary, square or tall and narrow. Papillae absent. Prickles absent, or a few in the costal zone near the margin of the blade. Silica cells in costal zone with dumb-bell-shaped silica bodies, sometimes associated with cork cells. Costal short cells solitary or in pairs, seldom in short rows. Triniochloa talpensis is clearly a member of the genus Triniochloa in its anatomical features (Wat¬ son & Dallwitz, 1992) and morphology, but it is unique in the genus because of its deciduous leaves. The basal and lower culm leaves simply die, and if the blade becomes detached it does so at the apex of the sheath, as in most grasses. The upper culm leaves have a special way of separating; when they are mature and dry, the blade and the portion of the sheath above the point of closure separate in Volume 5, Number 1 1995 Gonzalez-Ledesma et al. Triniochloa talpensis 39 Table 1. Salient characters distinguishing Triniochloa talpensis and T. laxa. T. talpensis T. laxa 2.0-3.0 0.8-1.0 60-110 40-80 deciduous completely exserted Rhizome diameter (mm) Plant height (cm) Leaf blade Panicle Panicle length (cm) Spikelets per panicle Adaxial ribs of first-order bundles Adaxial furrows Abaxial projection of the midrib Adaxial sclerenchyma girders Bulliform cells Anticlinal horizontal walls of long cells Prickles (abaxial epidermis) 16.0-34.0 30-65(-90) squared narrow inconspicuous T-shaped fan-shaped groups not projecting above the epidermis undulating absent or a few in the costal zone near the margin of the blade persistent partially exserted 7.0-13.0 6-18 rounded wide and shallow conspicuous I-shaped rows projecting above the level of the epidermis straight abundant in all costal zones one piece from the remainder of the sheath (Fig. 1C). This is especially noticeable in pressed, dried material when the upper blades are moved in order to expose the ligule or when they are slightly pulled. Detachment is along a well-defined line, but there is no articulation nor evidence of an abscission layer. Detachment was not seen in fresh material when the plants were collected and pressed. It appears that an abscission layer develops when the plants are drying. This suggests that the phenomenon is possibly a response to drought conditions. Triniochloa talpensis grows in open pine-oak forests and is only known from the Sierra de Cuale in the municipio of Talpa (whence the specific ep¬ ithet), state of Jalisco, in the western part of the Eje Neovolcanico. It is closely related to T. laxa , a species that grows in forests of spruce and pine mixed with poplar, and is restricted to the northern Sierra Madre Occidental in the state of Chihuahua. Although both species have rhizomes and similar spikelets, Triniochloa talpensis is more robust than T. laxa, and there are various other characters that distinguish them (Table 1). In addition, important anatomical differences were found in the arrange¬ ment of the adaxial sclerenchyma, the protrusion of the bulliform cells, the shape of adaxial ribs, the abundance and distribution of the prickles, and other features (Fig. 2, Table 1). Deciduous leaves, narrow adaxial leaf furrows, and more abundant adaxial prickles (Fig. 2B) suggest that T. talpensis is adapt¬ ed to drier habitats than T. laxa. Acknowledgments . We thank Fernando Chiang Cabrera for reviewing the Latin description. We also thank the curators of COL, ENCB, MEXU, MO, and US for lending specimens under their care, and the anonymous reviewers for their valuable obser¬ vations. Literature Cited Clayton, W. D. & S. A. Renvoize. 1986. Genera Gra- minum. Kew Bull., Addit. Ser. 13: 1-389. Hitchcock, A. S. 1913. Mexican grasses in the United States National Herbarium. Contr. U.S. Natl. Herb. 17(3): 181-389. Metcalfe, C. R. 1960. Anatomy of Monocotyledons. I. Gramineae. Oxford Univ. Press, Oxford. Reeder, J. R. 1968. Systematic position of the genus Triniochloa (Gramineae). Amer. J. Bot. 55: 735. Watson, L. & M. J. Dallwitz. 1992. The Grass Genera of the World. C. A. B. International, Wallingford, U.K. Didymocarpus miniatus (Gesneriaceae), a New Species from Peninsular Malaysia Ruth Kiew Department of Biology, Universiti Pertanian Malaysia, 43400 UPM Serdang, Selangor, Malaysia Abstract. Didymocarpus miniatus (Gesneri¬ aceae), a new species in Didymocarpus sect. Het¬ eroboea, is described from Bukit Bauk in Treng- ganu, Peninsular Malaysia. Didymocarpus miniatus Kiew, sp. nov. TYPE: Peninsular Malaysia. Trengganu: Bukit Bauk, R. Kiew RK 3792 (holotype, UPM; isotypes, L, SING). Differt a Didymocarpo atrosanguineo Ridley foliis petiolatis, pedunculis brevioribus et floribus omnino min- iatis. Herb. Stem woody, up to 1 m tall and 0.7 cm thick. Indumentum of young stem, petioles, and lower surface of veins silky ferrugineous with uni- seriate 3-4-celled trichomes ca. 1.5 mm long. Leaves petiolate, spirally arranged on the upper 4-12 cm of stem, separated by internodes up to 2 cm long. Lamina oblanceolate, (15-)17-24 x (5.5-)6-10 cm, apex acuminate, acumen 1-2 cm long, strongly narrowed to base, drying chartaceous, margin cre- nate, indumentum of 2-celled uniseriate hairs ca. 0.5 mm long, dense and roughly velvety above, less dense beneath, in life dark green above and pale grayish green beneath. Midrib prominent above and beneath, secondary veins 20-21 pairs, plane above and prominent beneath, tertiary veins obscure above, prominent beneath, areoles near the midrib irregular in shape, those toward the margin rectangular. Pet¬ iole (1.5—)2(—3) cm long. Flowers solitary. Peduncle slender, 6-12 mm long. Bract pairs ligulate, 3 mm long, finely hairy. Calyx divided to base, lobes linear, 4 mm long, densely hairy. Corolla completely ver¬ milion with two yellow longitudinal nectar guides in the base of the throat, finely pubescent outside, hairs glandular uniseriate, 2-3-celled stalk up to 0.5 mm long, nectar guides with scattered minute ellipsoid vesicles, tube 2.5-3.7 cm long, narrow, 3-4 mm wide in the lower half, dilating distaliy to 10 mm wide, lobes 5, broadly rounded, upper two 4x7 mm, lower three 8x6 mm and projecting beyond the upper two. Stamens 2, filament slender, 10 mm long, anthers white, oblong, 2x1 m m, connivent, positioned within corolla tube. Ovary 5 x 0.5 mm, Novon 5: 40-41. 1995. style slender, 30 mm long, densely pubescent with glandular hairs, stigma discoid, peltate, 1 mm across. Nectary encircling base of ovary, 0.75 mm high, distaliy lobed for ca. one-third of its length. Capsule slender, 7.5-8 cm long, finely pubescent when young. Among the 85-odd Didymocarpus species in Peninsular Malaysia, only D. miniatus has com¬ pletely vermilion flowers. Only one other species, D. atrosanguineus, has flowers with red lobes, but its corolla tube is usually cream-colored. (One or two plants of this latter species in Sekayu Forest Reserve, Trengganu, produce flowers that are com¬ pletely red, but they are deep crimson not vermilion.) In lamina indumentum, D. miniatus most closely resembles the broad-leaved form of D. atrosangui¬ neus (Kiew, 1989), but it is readily distinguished from this form by its petiolate leaves. Didymocarpus miniatus belongs to section Het- eroboea, which includes species with large, trumpet¬ shaped, solitary flowers and large oblanceolate leaves (Ridley, 1923). However, it is atypical of this sec¬ tion, in which the species have leaves with a winged petiole, by its petiolate leaves. In addition, D. min¬ iatus is distinct in its short peduncles, 0.6-1.2 cm long, as opposed to peduncles 2.5-10 cm long in other species in this section. In common with other large tubular Didymo¬ carpus flowers (Kiew, 1992), the stigma in D. min¬ iatus is discoid and peltate, and the nectary is large and cylindrical (although it is the first species in section Heteroboea to be described with a lobed nectary). The glandular indumentum on the style resembles that of D. quinquevulnerus Ridley, which is also a member of section Heteroboea. Didymocarpus miniatus has an extremely local distribution, at the foot of Bukit Bauk, where it grows on slopes above small swampy areas or seasonal streams, and at about 20 km south on Bukit Cha- bang, 5 km inland of Kerteh (G. W. H. Davison, pers. comm.). Both of these hills belong to the same coastal sandstone formation. Paratypes. PENINSULAR MALAYSIA. Bukit Bauk, S. Anthony SA 602 (UPM), G. IP. H. Davison GD5 (UPM), K. M. Kochumen KEP 9491 (KEP). Volume 5, Number 1 1995 Kiew Didymocarpus miniatus 41 Acknowledgments. This work was supported by IRPA grant number 4-07-05-813. I thank Geoffrey Davison, who first drew my attention to this new species. Literature Cited Kiew, R. 1989. Didymocarpus (Gesneriaceae) on Gun- ung Tahan, Malaysia. Gard. Bull. Singapore 42: 47- 64. -. 1992. Five new species of Didymocarpus (Gesneriaceae) from Peninsular Malaysia. Gard. Bull. Singapore 44: 23-42. Ridley, H. N. 1923. Didymocarpus. Flora Malay Pen¬ insula. 2: 506-523. Reeve, London. A New Species of Campyloneurum (Polypodiaceae) from Northwestern Ecuador Blanca Leon 1 Department of Botany, Field Museum, Chicago, Illinois 60605, U.S.A. and Museo de Historia Natural, Av. Arenales 1256, Apartado 14-0434, Lima-14, Peru ABSTRACT. Campyloneurum oellgaardii is newly described from a humid western premontane Andean forest of Ecuador. It appears to belong to the sphen- odes group, which is characterized by undivided primary areoles and long petiolate leaves, and which includes C. coarctatum, C. inflatum, C. sphenodes, and C. sublucidum. During a study of the fern genus Campyloneurum C. Presl, a distinctive new species was recognized and is described below. Campyloneurum oellgaardii B. Leon, sp. nov. TYPE: Ecuador. Carchi: drainage of Cerro Go- londrinas mountains, 0°52'N, 78°07'W, 21 Dec. 1987, Hoover 2211 (holotype, MO; isotype, QCA). Figure 1. Species C. inflato proxima, a qua rhizomate longe repenti, 6 mm crasso, atrofusco, dense paleaceo, squamis adpressis, brunneolis, foliis ampliori lanceolatis, 147 cm longis, 22 cm latis differt. Epiphyte; stem long-creeping, black, not prui- nose, 6 mm wide. Stem scales lax, light brown in mass, ovate, 3-4 mm long, 2-2.5 mm wide, bases auriculate, apices obtuse, margins entire with scat¬ tered hairs, slightly clathrate, the cells oblong or broadly oblong, cell walls 6—9 pm wide, several central cells with dark brown walls, walls of marginal cells yellowish or brownish, cell lumina transparent. Phyllopodia 4-5 mm long, 7-10 mm wide, 1-15 cm distant. Leaves pendulous (?), 147 cm long, petiole 55 cm long, shiny dark stramineous; laminas lanceolate, 22 cm wide, herbaceous-chartaceous, base cuneate, apex long-acuminate, margins carti¬ laginous, sinuate, leaves with inconspicuous bicel- lular glandular hairs scattered abaxially; stomata 1 Present addresses: Department of Geography, Uni¬ versity of Maryland Baltimore County, Baltimore, Mary¬ land 21228, U.S.A. and Department of Botany, NHB- 166, Smithsonian Institution, Washington, D.C. 20560, U.S.A. polocytic or rarely copolocytic; costa prominent, slightly angular abaxially; primary veins prominent, 75° divergent from the costa, straight, lighter in color than the adjacent tissue, 7-9 mm distant; secondary veins slightly prominulous on both sides of the lamina, transverse secondary veins forming 19—21 primary areoles between the costa and mar¬ gin; primary areoles undivided, with 2(—3) excurrent free veinlets, the marginal ones sometimes divided with (0)— 1 veinlet. Sori subapical on the excurrent veinlet; paraphyses and spores not seen. Campyloneurum oellgaardii is known only from the type material, collected in northwestern Ecua¬ dor, at 1200 m elevation in a perhumid premontane forest. This locality borders on what Dodson & Gen¬ try (1991) considered an extension of the Choco pluvial forest. The specimen was found growing as an epiphyte several meters above the ground. Based on the curvature of the petiole (Fig. la), the leaves are probably pendently arched. Campyloneurum oellgaardii is characterized by well-spaced and extraordinarily large leaves, more than 1 m long, in addition to its undivided primary non-costal areoles. Because of its habit, leaf mor¬ phology, and pattern of venation this species fits within the sphenodes group of Leon (1992). Besides this new species, the sphenodes group consists of C. chrysopodum (Klotzsch) Fee, C. coarctatum (Kunze) Fee, C.falcoideum (Kuhn ex Hieronymus) M. Meyer ex Lellinger, C. inflatum M. Meyer ex Lellinger, C. sphenodes (Kunze ex Klotzsch) Fee, and C. sublucidum (Christ) Ching. Campyloneurum oellgaardii differs from the other species of the group in having a larger stem diameter (6 mm vs. 2-3 mm), obtuse stem scales, and longer (more than 100 cm vs. (17-)30-70(-85) cm) and broader (20 cm vs. (2-)3-6(-10) cm) leaves. Most species of Campyloneurum have leaves less than 1 m long. Leaves longer than this are otherwise found only in: (1) those species with entire leaves, short-creeping stems and phyllitidis-, brevifolium- venation ( phyllitidis -, latum -venation of Lellinger, 1988) such as C. abruptum (Lindman) B. Leon, 6. Novon 5: 42-44. 1995. Volume 5, Number 1 1995 Leon Campyloneurum oellgaardii 43 Figure 1. Campyloneurum oellgaardii B. Leon (from Hoover 2211). a. Stem scale. b. Cellular detail of stem scale. —c. Habit. —d. Pattern of venation. (Drawn by the author.) brevifolium (Link) Link, C. pascoense R. M. Tryon & A. F. Tryon, C. phyllitidis (L.) C. Presl, and C. tucumanense (Hieronymus) Ching; and in (2) those pinnate-leaved species with undivided primary ar- eoles, such as C. decurrens (Raddi) C. Presl and C. magnificum T. Moore. All these species are low epiphytes, epipetrics, or terrestrials, and they usu¬ ally grow in partially closed forests. In contrast, C. oellgaardii is a high-canopy epiphyte, according to the collector, and its gigantism may be an adaptation 44 Novon to a dense canopy and/or low risk of leaf damage due to breakage. This species is dedicated to Benjamin 011gaard in honor of his knowledge of the Ecuadorean flora, especially the pteridophytes. Acknowledgments. I thank David B. Lellinger and Kenneth R. Young for comments on the manu¬ script. Thanks are also due to Altec Tangerini and David Brunner for assistance. Literature Cited Dodson, C. & A. H. Gentry. 1991. Biological extinction in western Ecuador. Ann. Missouri Bot. Card. 78: 273-295. Lellinger, D. B. 1988. Some new species of Campy- loneurum and a provisional key to the genus. Amer. Fern J. 78: 14-35. Leon, B. 1992. A Taxonomic Revision of the Fern Genus Campyloneurum (Polypodiaceae). Ph.D. The¬ sis, University of Aarhus, Denmark. A New Species of Beilschmiedia (Lauraceae) from Guerrero, Mexico Francisco G. Lorea Hernandez Departamento de Biologla, Facultad de Ciencias, UNAM, Ciudad Universitaria, Circuito Exterior, 04510 Mexico, D.F., Mexico. Current address: Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. Abstract. Beilschmiedia angustielliptica is de¬ scribed and illustrated. Morphologically this species is easy to recognize because of its long, reticulate (but not foveolate) lanceolate leaves, its glabrous anthers, and its glabrate ovary. So far, this species is known only from the central mountains of Guer¬ rero, Mexico. Resumen. Se describe e ilustra Beilschmiedia an¬ gustielliptica. Entre las otras especies del genero esta nueva especie es facil de reconocer por sus hojas grandes, lanceoladas, reticuladas (pero no fo- veoladas), sus anteras glabras, y su ovario glabrado. Hasta ahora esta especie solo se conoce de las mon- tanas centrales de Guerrero, Mexico. Extensive collections carried out during the last ten years in Guerrero, Mexico, under the auspices of the “Flora del Estado de Guerrero” project, or¬ ganized by the Laboratorio de Plantas Vasculares (Facultad de Ciencias, UNAM), have resulted in more complete collections for several formerly poor¬ ly known families in that state. For example, about a dozen species of Lauraceae were previously re¬ ported from Guerrero (Standley, 1922; Allen, 1945). Today, the assembled material represents more than 35 species. Several collections belong to undescribed species in different genera. A new species of Beilsch¬ miedia is here described. The Latin American species of Beilschmiedia were last revised more than 50 years ago by Kos- termans (1938), who recognized 15 neotropical spe¬ cies in the genus. Currently, the increase in collec¬ tions suggests the merger of some names, but also the recognition of new species, so the actual number of taxa is around 10-20 species (van der Werff, 1991). In Mexico there exist about eight species of Beilschmiedia. For a time, only B. mexicana (Mez) Kostermans was recognized as occurring in the country (Kostermans, 1938; Allen, 1945). Several Mexican specimens of Beilschmiedia have been an¬ notated in herbaria as B. pendula (Swartz) W. B. Hemsley, implying the presence of this species in the region. But B. mexicana is suspected to be a synonym of B. pendula (Burger & van der Werff, 1990). Early in the second half of this century B. riparia F. Miranda was described from Chiapas (Miranda, 1953) as a distinct species with pilose and subcoriaceous leaves; this species is now known from the tropical semideciduous forests in Guerrero, Oa¬ xaca, and Chiapas. Besides B. pendula, B. riparia, and the newly described species, several collections from different places in Mexico indicate the presence of at least five other taxa, some of them conspecific with or related to species known in Mesoamerica, namely: B. anay (S. F. Blake) Kostermans, B. hon- durensis Kostermans, and B. ovalis (S. F. Blake) C. K. Allen. Until a thorough revision of the American species of Beilschmiedia is done, I present the following provisional key to separate the Mexican species. Key to the Mexican Species of Beilschmiedia la. Leaf undersurface glabrous, rarely some straight appressed hairs on main veins. 2a. Leaf undersurface finely reticulate (can- cellate), minor veins evenly thickened forming areolae 0.1-0.3 mm diam., no or almost no free veinlets. 3a. Leaves stiff coriaceous, rather thick, obtuse to acute . B. aff. mexicana 3b. Leaves chartaceous, acute to long acuminate. B. mexicana 2b. Leaf undersurface roughly reticulate, mi¬ nor veins unevenly thickened, then areolae usually 1.0 mm diam. or larger and ap¬ pearing with several free veinlets. 4a. Secondary veins in 14-18 pairs, leaves narrowly elliptic. . B. angustielliptica 4b. Secondary veins in 8-10 pairs, leaves elliptic. B. hondurensis lb. Leaf undersurface pubescent, if hairs only on main veins, these not straight appressed. Novon 5: 45-47. 1995. 46 Novon ^'7 ; B p U ^'"‘ l ? d i a angushelliptica Lorea-Hernandez. -A. Branchlet with inflorescence. -B. Branchlet Tiew/of a Za k i ? ' ‘ aSt d ‘ visi ° ns - “ D ' °P en flower (°™y removed). -E. Lateral and abaxial vtews of a thtrd whorl stamen. -F. Stammode. -G. Ovary. -H. Adaxial and lateral views of an external stamen. 5a. Hairs on undersurface straight and appressed. fi. aif. pendula 5b. Hairs on undersurface straight to curly, erect or loosely appressed. 6a. Hairs curly, erect to loosely appressed. B. ovalis 6b. Hairs straight and erect. 7a. Leaf blades widely elliptic to ob- lanceolate; widest leaves 11-14 cm. B. aff. anay 7b. Leaf blades elliptic, widest leaves 5-8.5 cm. B. riparia Volume 5, Number 1 1995 Lorea Hernandez Beilschmiedia angustielliptica 47 Beilschmiedia angustielliptica Lorea-Hernan- dez, sp. nov. TYPE: Mexico. Guerrero: Mpo. Atoyac de Alvarez, aprox. 2 km S to El Molote, on the trail to El Eden, 1580 m, 19 May 1993 (fl, fr), F. Lorea & L. Lozada 5540 (holotype, FCME; isotype, MO). Figure 1. Arbores parvae (7-8 m altae). Folia opposita vel su- bopposita, lamina (7-)ll-18(-24.5) x (1.5-)2.5-4.5(- 5.5) cm, anguste elliptica, aliquantum oblanceolata, basi apiceque acuta vel acuminata, utrinque glabra, margine interdum Ieviter undulata, nervis lateralibus subtus prom- inentibus, costa supra complanata, subtus valde promi- nenti ac in sicco manifeste rubiginosa. Inflorescentiae 2.5- 5.5(-9.5) x 1.5-3 cm, paniculatae, axillares in ramulos aphyllos breves crebiter positae, vel terminales; axibus puberulis vel tomentosis. Flores hermaphroditi, 2.2-2.7 mm longi, tepalis sex, ovatis vel late ovatis, erectis, abax- ialiter pro parte maxima puberulis, adaxialiter praecipue basi puberulis, margine glabris; staminibus novem, 1.1- 1.4 mm longis, antheris glabris, eis verticillorum primi et secundi complanatis, late ovatis, eis verticilli tertii pyr- amidalibus, filamentis omnino pilosis vel tomentosis; stam- inodiis fere sessilibus, apice triangularibus, basi truncatis vel subcordatis, ovario 0.7-0.85 mm longo, pyriformi, glabrato; hypanthio tomentoso. Fructus 3-3.5 x ca. 1.7 cm, ellipticus, anthracinus. Low trees (7-8 m), young branches compressed to flattened, yellow-golden puberulent to tomentose, bark reddish brown, glabrous and light brown with rather smooth bark when older, buds densely to¬ mentose. Leaves opposite, a few subopposite, peti¬ oles (0.5-)0.7-1.3(-1.8) cm, slightly canaliculate, puberulent, soon glabrous, blades (7—)11 — 18(—24.5) x (1.5-)2.5-4.5(-5.5) cm, narrowly elliptic, some oblanceolate, base and apex acute or acuminate, chartaceous, glabrous on both surfaces, sparsely pu- berulous when young, margin sometimes slightly undulate, veins on upper and lower surface slighty elevated, secondaries in 14-18 pairs, ascending, prominent on lower surface, midvein flattened on upper surface, very prominent and distinctly reddish brown when dry on lower surface. Inflorescences 2.5-5.5(-9.5) x 1.5-3 cm, axillary, close to each other on short aphyllous branchlets, or terminal, paniculate, pedunculate, peduncle 0.6-2.5(—4.5) cm, major axes 2.5-5.5 mm, puberulent to tomentose, flower pedicels (2-)2.6-3.6(-4.2) mm, puberulent, bibracteolate close to the base or not, bracteoles minute and soon falling. Flowers hermaphroditic, 2.2-2.7 mm long, tepals six, 1.5-1.7 x 1-1.3 mm, ovate to widely ovate, erect, outside puberulent for most of the central area, inside mainly on the base, margins glabrous, stamens nine, 1.1-1.4 mm long, anthers of first and second whorls 0.7-0.8 mm long, flattened, widely ovate, thecae introrse, anthers of third whorl 0.5-0.6 mm long, pyramidal, thecae extrorse-lateral, all filaments pilose to tomentose, anthers glabrous, connectives protruding beyond the anther cells, apparently glandular, glabrous, glands irregularly round, 0.4-0.5 mm, staminodes 0.4- 0.6 mm long, almost sessile, apex triangular, acute, base truncate or subcordate, adaxial face plane, abaxial face convex and tomentose in the middle, ovary 0.7-0.85 mm long, pyriform, glabrate, style 0.4-0.5 mm long, stigma apical, oblique, hypan- thium 0.5-0.8 mm, tomentose. Fruit 3-3.5 x 1.7 cm, when mature, elliptic, black, smooth, and shiny. Beilschmiedia angustielliptica is well distin¬ guished morphologically from other species in Amer¬ ica by its narrow, elliptic leaves conspicuously re¬ ticulate (but not foveolate), and glabrous anthers. The general appearance of leaf reticulation in B. hondurensis suggests some similarity with B. an¬ gustielliptica, but the former has more or less abruptly acuminate, short elliptic leaves and the ovary is glabrous, whereas in B. angustielliptica the leaves are long lanceolate and the ovary is sparsely pilose. So far, B. angustielliptica is known only from mesophyllous mountain forests in Guerrero, where the canopy is dominated by Quercus spp.. Magnolia schiedeana D. F. L. v. Schlechtendal, Pinus strobus L. var. chiapensis M. Martinez, Per- sea americana P. Miller, Licaria aff. triandra (Swartz) Kostermans, Alchornea latifolia Swartz, Cinnamomum sp., Prunus sp., and Pouteria sp. Paratypes. MEXICO. Guerrero: Mpo. Atoyac, El Molote, 1630 m, (fr), A. Nunez 1159 (FCME); Mpo. Tecpan, km 16 on the road from El Porvenir to San Antonio de Las Tejas, 1360 m, 7 m tree, (fl), G. Lozano 227 (FCME). Acknowledgments. This paper was carried out with support of a scholarship from DGAPA (Univ- ersidad Nacional Autonoma de Mexico), and partial support for fieldwork from the Missouri Botanical Garden. I thank Henk van der Werff for critical comments on the manuscript, and Roy Gereau for having called my attention to some recommenda¬ tions of the Code of Botanical Nomenclature, and for checking the Latin description. The illustration was drawn by John Myers. Literature Cited Allen, C. K. 1945. Studies in the Lauraceae. VI. Pre¬ liminary survey of the Mexican and Central American species. J. Arnold Arbor. 26: 280-434. Burger, W. C. & H. van der Werff. 1990. Lauraceae. In: W. Burger (editor). Flora Costaricencis. Fieldiana, Bot. n.s. 23: 1-138. Kostermans, A. J. G. H. 1938. Revision of the Lau¬ raceae V. Recueil Trav. Bot. Neerl. 35: 834-931. Miranda, F. 1953. Plantas nuevas o notables de la flora de Chiapas. Anales Inst. Biol. Univ. Nac. Mexico 24: 69-96. Standley, P. C. 1922. Lauraceae. In: Trees and shrubs of Mexico. Contr. U.S. Natl. Herb. 23(2): 286-298. Werff, H. van der. 1991. A key to the genera of Lauraceae in the New World. Ann. Missouri Bot. Gard. 78: 377-387. Passiflora sanctae-mariae in Series Luteae (Passifloraceae), a New Species and Series in Subgenus Decaloba John M. MacDougal Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. ABSTRACT. Passiflora sanctae-mariae, restricted to Chiapas, Mexico, and adjacent Guatemala, is new¬ ly described and assigned to a species group that includes P. lutea L. of the United States. The five species of this group lack extrafloral nectaries and have very reduced or no bracts. The group is rec¬ ognized as Passiflora series Luteae (Small) J. MacDougal, comb, et stat. nov. Passiflora sanctae-mariae is a new species of small vining passionflower, known from nine dried collections and one living collection. The living col¬ lection, field collected by the author, was grown and observed in the greenhouse for several years, and it furnished the type collection. The species is re¬ stricted to the volcanic mountains of northwestern Guatemala and adjacent Chiapas, Mexico. It is close¬ ly related to P. lutea L. of the United States, the Mexican and Guatemalan P. pavonis Masters, an as yet undescribed species in east-central Mexico (Passiflora dictamo sensu Killip, not DC.), and P filipes Bentham, which ranges from Texas to Ec¬ uador. These five species lack extrafloral nectaries and have very reduced or no bracts, and form a related species alliance here recognized as a series. Passiflora series Luteae (Small) J. MacDougal, comb, et stat. nov. Passiflora [no rank] Luteae Small, Man. S. E. FI. 895. 1933. TYPE: Pas¬ siflora lutea L. Small (1933), in his flora of the southeastern United States, subdivided the seven species of Pas¬ siflora listed there into four categories, one of which he named Luteae. It contained P. lutea and P. sexflora A. L. Jussieu. This infrageneric unit was published without rank, among the many such key¬ ing categories used throughout that work. Small’s name is valid under Article 35.2 of the Code and serves as the basionym for the name of the series. Killip, in his taxonomic monograph of 1938, placed P. lutea and P. sexflora in subgenus Plectostemma Masters but separated the species into their own separate series under section Decaloba (DC.) Mas¬ ters. He correctly associated P. pavonis and P. filipes with P. lutea. Killip made no reference to Novon 5: 48-51. 1995. Small’s work, even in the bibliography of the mono¬ graph, and appears to have independently used the name Luteae. Killip’s “series” are invalidly pub¬ lished, however, as they lack Latin diagnoses, and the name is now validated as a series here. The series is most easily recognized within subgenus De¬ caloba by the absence of extrafloral nectaries, ex¬ treme reduction or absence of floral bracts, and the small purple berry. Further characteristics include small flowers with the coronal filaments in two series, the outer series filiform, greenish white to greenish yellow or pale yellow, and sometimes dull purplish or dull violet at base. The ovaries are glabrous or with few short hairs. The Passiflora bilobata A. L. Jussieu complex of the West Indies appears to be the sister group to series Luteae based on floral morphology and lack of extrafloral nectaries. The delineation and lectotypification of subgenus Plectostemma and the earlier and generally over¬ looked subgenus Decaloba (DC.) Reichenbach is nearly the same (see discussion in MacDougal, 1994), so the series and following new species are here placed in subgenus Decaloba section Decaloba. Passiflora sanctae-mariae J. MacDougal, sp. nov. TYPE: Cultivated at Duke University, June 1982, from stems of MacDougal & Miley 602 collected Jan. 1980 in Guatemala, MacDougal & Miley 602GR (holotype, DUKE; isotypes, BM, CAS, CR, EAP, HUA, MEXU, MO, USCG, US). Figures 1, 2b. Passiflora scandens; petioli eglandulosi; folia eglan- dulosa bilobata vel trilobata, lobis lateralibus acutis, mar- ginibus integris, lobo centrali obtuso vel obsolete, vel trun- cata; bracteae nullae; coronae filamenta biseriata, fila- mentis exterioribus filiformis, 2.1-3.0 mm longis; oper¬ culum plicatum; ovarium glabrum; bacca atropurpurea; semina 10-13 sulcata. Slender perennial vine 2-5 m long, minutely pu- berulent throughout at the shoot tip with trichomes (0.05-)0.08-0.2(-0.3) mm long, except the adaxial surface of the lamina glabrous (or a few trichomes at the base of the primary veins), older growth sparsely puberulent to glabrescent. Stems terete, tardily glabrescent, drying subangulate and striate, 2.7cm Volume 5, Number 1 1995 MacDougal Passiflora sanctae-mariae 49 Figure 1. Passifiora sanctae-mariae J. MacDougal (from MacDougal & Miley 602GR, except as noted). A. Habit. —B. Stipule. —C, D. Seed. —E-G. Leaves at prereproductive nodes; F (Steyermark 37098), G (Steyermark 37215). —H, I. Leaves at flowering nodes; H (Breedlove 50768), I (Purpus 7292). 3mm 50 Novon with little secondary growth; phyllotaxy 2/5; nodes sometimes purplish (especially prophylls and junc¬ tion of stipules); posture of shoot tip cernuous. Stip¬ ules (1.1 —)2—3.5 x 0.3-0.5(-1.0) mm, linear-lan¬ ceolate to broadly lanceolate, attenuate to acumi¬ nate, falcate. Petioles 1.1-2.6 cm long, eglandular, minutely puberulent. Laminas at fertile nodes (2.5-) 4-7.3 X (4.7-)6-l 1.2 cm, entire, not variegated, the general outline depressed obovate, truncate-ob¬ late, semi-circular, or transversely truncate-elliptic, subtruncate to shallowly 2(-3)-lobed less than 0.15 the distance to the very shallowly cordate to broadly rounded base, the lateral lobes acute (rarely slightly obtuse or somewhat rounded), the central lobe obtuse to obsolete, lobes not mucronulate or the mucro 0.5 mm long or less, the angle between the primary lateral veins (62-)67-90(-11 2f , the ratio of central to lateral vein lengths 1.1 — 1,5(—1.7), the ratio of laminar width to length 1.2-2.0(-2.3); leaves of juvenile plants often variegated along lateral veins, narrowly depressed obovate to transversely trun¬ cate-elliptic in outline, 2(-3)-lobed to !4 the distance to the base, the angle between the lateral veins 60- 130°, the ratio of central to lateral vein lengths 1.3- 2.1, the ratio of laminar width to length 1.2-3.6; laminar nectaries absent. Tendrils straight during development at shoot apex. Prophylls of the vege¬ tative ramifying bud 2, broadly lanceolate to ovate, usually purplish, long acuminate or 3-toothed with central tooth caudate. Peduncles 1.1-2.6 cm long, geminate, uniflorous. Bracts absent. Hypanthium 4.5-5 mm diam., flowers ca. 1.5 cm diam., sub¬ erect, pale yellowish green with a dark center, with faint sweet odor; stipe 3-6 mm long (to 8 mm in fruit); sepals 7.5-10.5 x 2.8-4.2 mm, elliptic- oblong to ovate-oblong, rounded to subacute, ecor- niculate; petals 5-8 x 2.4—2.6 mm, oblong to ovate- oblong, rounded, whitish; filamentous corona in 2 series, the outer filaments ca. 37-40, 2.1-3.0 mm long, ca. 0.2-0.3 mm diam., filiform, curved- spreading, slightly thicker in the distal half but at¬ tenuate at the very apex, cream to whitish, basally light greenish and often with a small purplish mark, the inner filaments 1.5 mm long, clavate, erect, purplish with pale apex; operculum 1.5 mm long, membranous, plicate, dark purple; nectary not con¬ cave; limen (disk) ca. 2.0 mm diam., uniformly dark purple, the raised edge only ca. 0.2 mm high; sta- minal filaments connate 3.1-4.1 mm along andro- gynophore, unmarked except dark purple at junction with limen, the free portions ca. 3-3.5 mm long, unmarked; anthers 2.8-3.1 mm long, not marked with purple; ovary 1.5-1.9 x 1.2-1.5 mm, widely ellipsoid, glabrous, light green; styles ca. 3-3.5 mm long; stigmas ca. 0.8 mm diam., cream. Fruit 1.1- Figure 2. —A. Silhouette of flower of P. filipes (MacDougal & Miley 579GR ). — B. Silhouette of flower of P. sanctae-mariae (MacDougal & Miley 602GR ). 1.4 x 0.9-1.1 cm, widely ovoid-ellipsoid, purplish black, estipitate; seeds 3.0-3.3 mm long, 2.0-2.3 mm wide, 1.2-1.3 mm thick, obovate to widely obovate, nearly symmetric to slightly campylotro- pous, transversely sulcate with 10-13 sulci, the ridges somewhat verrucose, the chalazal beak broad¬ ly conical and slightly inclined toward the raphe. Germination epigeal. Passiflora sanctae-mariae is restricted to the western slopes of the Serrania Transistmica, or Cor¬ dillera Madre, of Chiapas, Mexico, and adjacent western Guatemala. It may be distinguished from the other members of series Luteae in Mesoamerica by the following key. la. Plant completely glabrous; peduncles (1.5—)3— 5(-6) cm long; petals 3-4.5(-5) mm long; seeds usually 3-4(-5)-sulcate; elevation 1100 m or less. P. filipes lb. Plant minutely puberulent to pubescent (except adaxial surface of lamina) at least on young growth and petioles; peduncles 1.1-2.6 cm long; petals 5-8 mm long; seeds usually 6-13-sul- cate; elevation 1300-2800 m. 2a. Stems pubescent, or sparsely so below; leaves 1.5-5.5(-7) cm wide; corona 3-5 mm long; operculum pale greenish; andro- gynophore 4.8-5.5 mm long; seeds 6-7- sulcate (rarely irregularly rugose); eleva¬ tion (1350-)l 900- 2800 m. P. pavonis 2b. Stems minutely or sparsely puberulent, or glabrescent below; leaves (4.7-)6-11.2 cm wide; corona 2-3 mm long; operculum dark purple; androgynophore 3.0-4.1 mm long; seeds 10-13-sulcate; elevation 1300-1500 m. P. sanctae-mariae Passiflora sanctae-mariae was collected in the field in flower and fruit, then cultivated as a single Volume 5, Number 1 1995 MacDougal Passiflora sanctae-mariae 51 individual 1980-1986 at Duke University, with the type specimens collected in June 1982. Observations made on living material allowed the detailed de¬ scriptions of the flowers provided above. The flowers opened in the greenhouse around midday and closed in the late afternoon (the other species in the series open in early to mid-morning and close in the early to late afternoon). No autogamy was observed in six years of cultivation, and fifteen self pollinations failed to set fruit, good evidence that the species is self¬ incompatible. Fruits and seeds are known only from my field collection; the number of seeds per fruit was 26, 29, 30, and 38 + 2 aborted, i.e., seeds 26-40 (N = 4). The number of seeds per fruit is significantly higher than its relatives in the series, which range from 2 to 12 (MacDougal, unpublished). A few of the seeds were germinated in the greenhouse to determine germination type, but the seedlings were not grown. The germination type is epigeal with elliptic foliaceous cotyledons, the norm for the ge¬ nus, but is worth noting because the close relative Passiflora lutea has hypogeal germination, ex¬ tremely rare in the genus (MacDougal, 1994). Ger¬ mination type for the other species in the section is unknown. As is the case in many of the passifloras in section Decaloba, the leaves of the juvenile plants of this species are often variegated (cf. Fig. IF), but this condition is not retained at reproduction. The leaves tend to be larger than the other species of the section and are characteristically apically truncate or sub¬ truncate, often giving the leaves a hemispherical or bat-wing shape. The common name ala murcielago is attached to some of the specimens, a name gen¬ erally associated with many of the bilobed passifloras in northern Mesoamerica. One collector noted its use as a diuretic, a use also shared by several other bilobed passifloras in this area. Passiflora sanctae-mariae is found only in pre- montane and lower montane wet forest. In Quezal- tenango it is associated with P. prolata Masters, P. helleri Peyritsch, P. biflora Lamarck, and P. mem- branacea Bentham. This species is dedicated to the Blessed Mother who is the namesake of the volcano where the clone of the type was collected, and who bore Jesus Christ whose sufferings are commemorated in the generic name. Paratypes. GUATEMALA. Quezaltenango: 100 m up dirt road at Km post 197 on Hwy. 9S ca. 8 km below tunnel near Santa Maria de Jesus, old finca Pirineos on slopes near Volcan Santa Maria, ca. 1340 m, 20 Jan. 1980 (fl, fr), MacDougal & Miley 602 (BM, DUKE, F, MEXU, MO); rich forested slopes along Quebrada San Geronimo, finca Pirineos, lower S-facing slopes of Volcan Santa Maria between Santa Maria de Jesus and Cala- huache, 1-2 Jan. 1940, Steyermark 33331 (F); top of ridge between finca Pirineos and finca Soledad, lower S- facing slopes of Volcan Santa Maria, 1300-1400 m, 5 Jan. 1940, Steyermark 33525 (F). San Marcos: half¬ way up slopes above finca El Porvenir on “Todos Santos Chiquitos,” lower S-facing slopes of Volcan Tajumulco, 1300-1500 m, 7 Mar. 1940, Steyermark 37093 (F), Steyermark 37215 (F). MEXICO. Chiapas: Mpio. de Angel Albino Corzo, above finca Tuxtepec, 1380 m, 21 Oct. 1980 (fl), Breedlove & Strother 46719 (CAS), 7 Apr. 1981 (fl), Breedlove 50768 (CASJ, 24 June 1981 (fl), Breedlove 51184 (CAS); finca Irlanda, June 1914, Purpus 7292 (UC). Acknowledgments. The type material was col¬ lected during fieldwork supported by NSF grant DEB-7912607 while I was a graduate student on an NSF Fellowship. I am grateful to Donald E. Stone and the other members of my graduate committee at Duke University, as well as the staff of the green¬ houses there, for their guidance and assistance. John Myers prepared the drawing. Literature Cited Killip, E. P. 1938. The American species of Passiflor- aceae. Publ. Field Mus. Nat. Hist., Bot. Ser. 19: 1 — 613. MacDougal, J. M. 1994. Revision of Passiflora sub¬ genus Decaloba section Pseudodysosmia (Passiflor- aceae). Syst. Bot. Monogr. 41: 1-147. Small, J. K. 1933. Manual of the Southeastern Flora. New York. [Published by the author.] New Combinations in Asian Krascheninnikovia Gueldenstaedt (Chenopodiaceae) Sergei L. Mosyakin N. G. Kholodny Institute of Botany, 2 Tereshchenkivska Str., Kiev, 252601 Ukraine ABSTRACT. New combinations in the genus Kras¬ cheninnikovia are proposed: K. arborescens (Los- ina-Losinskaja) Mosyakin, K. compacta (Losina- Losinskaja) Grubov var. longipilosa (Tsien Cho-po & Ma Cheng-gung) Mosyakin, and Krascheninni¬ kovia sect. Caudalae (Aellen) Mosyakin. Krascheninnikovia Gueldenstaedt (1772: 551) is the correct name for the genus previously com¬ monly known as Eurotia Adanson and Ceratoides Gagnebin. This nomenclatural problem has been dis¬ cussed by several authors over the years (Lozina- Lozinskaya, 1930; Reveal & Holmgren, 1972; Gru¬ bov, 1976; Guterman, 1975; Tomsovic, 1990; Tsvelev, 1993). Unfortunately, when establishing the genus Cer¬ atoides, Gagnebin (1755: 59) did not write a di¬ agnosis, but rather only mentioned a pre-Linnaean work by Tournefort (1703). In the latter work the genus Ceratoides included both an annual species (later named by Linnaeus as Ceratocarpus aren- arius L.), and a perennial one, which is conspecific with Krascheninnikovia ceratoides (L.) Guelden¬ staedt. At the time of its establishment (Gagnebin, 1755) the genus Ceratoides included Ceratocarpus arenarius, the type of Ceratocarpus L. (1753); thus Ceratoides is a synonym of Ceratocarpus. The next available name for the genus is Krascheninnikovia. [More detailed discussion is provided by Guterman (1975) and Tsvelev (1993).] In the Flora Reipub- licae Popularis Sinicae (Tsien Cho-po & Ma Cheng- gung, 1979) the name Ceratoides was accepted for this genus. Thus, the following new nomenclatural combinations are necessary for a forthcoming edition of the Flora of China. Krascheninnikovia arborescens (Losina-Los- inskaja) Mosyakin, comb. nov. Basionym: Eu¬ rotia arborescens Losina-Losinskaja, Izv. Akad. Nauk SSSR, Ser. 7, Otd. Fiz.-Mat. Nauk 1930, No. 9: 999. 1930. Ceratoides arborescens (Losina-Losinskaja) Tsien Cho-po & Ma Cheng- gung, Flora Reipublicae Popularis Sinicae 25(2): 27. 1979. Krascheninnikovia compacta (Losina-Losinska¬ ja) Grubov var. longipilosa (Tsien Cho-po & Ma Cheng-gung) Mosyakin, comb. nov. Bas¬ ionym: Ceratoides compacta (Losina-Losin¬ skaja) Tsien Cho-po & Ma Cheng-gung var. longipilosa Tsien Cho-po & Ma Cheng-gung, Flora Reipublicae Popularis Sinicae 25(2): 28. 1979. When first describing the new variety Ceratoides compacta (Losina-Losinskaja) Tsien Cho-po & Ma Cheng-gung var. longipilosa Tsieng Cho-po & Ma Cheng-gung, the authors did not cite the basionym for their new combination at the species level (see Kung Hsien-wu et al., 1978). Thus, the complete combination (species plus variety) was validated later in the Flora Reipublicae Popularis Sinicae (Tsien Cho-po & Ma Cheng-gung, 1979). Two sections, based mostly on the differences in the structure of trichomes, were established by Ael¬ len (1952) for the genus Eurotia. The correct names for these sections, as well as a new combination for one of them, are given below. Krascheninnikovia sect. Krascheninnikovia. TYPE: Krascheninnikovia ceratoides (L.) Gueldenstaedt (type of the genus). Eurotia sect. Ecaudatae Aellen, Verh. Naturf. Ges. Basel 63: 267. 1952. TYPE: not designated. Because in Aellen’s (1952) treatment the section Ecaudatae included the type of the genus, this name should be replaced by the appropriate autonym. Krascheninnikovia sect. Caudatae (Aellen) Mo¬ syakin, comb. nov. Basionym: Eurotia sect. Caudalae Aellen, Verh. Naturf. Ges. Basel 63: 267. 1952. TYPE: Eurotia compacta Losina- Losinskaja = Krascheninnikovia compacta (Losina-Losinskaja) Grubov. The native North American species previously commonly known as Eurotia lanala (Pursh) Mo- quin-Tandon was transferred to Krascheninnikovia by Meeuse & Smith (1974). If accepted as a distinct Novon 5: 52-53. 1995. Volume 5, Number 1 1995 Mosyakin Asian Krascheninnikovia 53 entity, another North American taxon described by Rydberg (1912) as Eurotia subspinosa is in need of the same transfer or a new infraspecific combi¬ nation. However, in view of the current work on the Flora of North America project, I believe this nomenclatural decision should be left to American botanists. Literature Cited Aellen, P. 1952. Ergebnisse einer botanisch-zoologisch- en Sammelreise durch den Iran. Botanische Ergeb¬ nisse II. Chenopodiaceae: Agriophyllum, Esfandi- aria, Eurotia. Verh. Naturf. Ges. Basel 63: 253- 272. Gagnebin, A. 1755. Observations faites sur le systeme des autheurs de botanique et sur l’Ophris minima C.B. Acta Helv. Phys. Math. 2: 56-75. Grubov, V. I. 1976. Proposal for conservation of the genus name Eurotia Adans. against Axyris Linn. Taxon 25: 362. Gueldenstaedt, A. I. 1772. Krascheninnikovia , novum plantarum genus. Novi. Comment. Acad. Sci. Imp. Petrop. 16: 548-560. Guterman, W. 1975. Notulae nomenclaturales. Phyton (Horn) 17 (1-2): 31-50. Kung Hsien-wu, Chu Ge-ling, Tsien Cho-po, Li An-jen & Ma Cheng-gung. 1978. The Chenopodiaceae in Chi¬ na. Acta Phytotax. Sin. 16(1): 117. Lozina-Lozinskaya (Losina-Losinskaja), A. S. 1930. Ma- terialy po izucheniyu roda Eurotia (Materials for the study of the genus Eurotia). Izv. Akad. Nauk SSSR, Ser. 7, Otd. Fiz.-Mat. Nauk 1930, No. 9: 977- 1007. Meeuse, A. D. J. & A. Smith. 1974. A new combination in Krascheninnikovia (Chenopod.). Taxon 20: 644. Reveal, J. L. & N. M. Holmgren. 1972. Ceratoides , an older generic name for Krascheninnikovia and Eurotia. Taxon 21: 209. Rydberg, P. A. 1912. Studies of the Rocky Mt. Flora — XXVII. Bull. Torrey Bot. Club 39: 310-328. Tomsovic, P. 1990. Patri druh Ceratoides latens (Eu¬ rotia ceratoides ) do ceskoslovenske kveteny? (Does Ceratoides latens (Eurotia ceratoides ) belong to the flora of Czechoslovakia?). Preslia (Praha) 62: 33- 39. Tournefort, J. P. 1703 (repr. 1719). Corollarium in- stitutionum rei herbariae. Lugdunii. Tsien Cho-po & Ma Cheng-gung. 1979. Ceratoides (Tourn.) Gagnebin. Pp. 24-28 in Flora Reipublicae Popularis Sinicae. Vol. 25(2). Beijing. Tsvelev, N. N. 1993. Zametki o marevykh (Cheno¬ podiaceae) Vostochnoy Evropy (Notes on Chenopo¬ diaceae of Eastern Europe). Ukrayins’k. Bot. Zhurn. 50(1): 78-85. A New Species of Oserya (Podostemaceae) from Jalisco, Mexico Alejandro Novelo R. Departamento de Botanica, Instituto de Biologia, Universidad Nacional Autonoma de Mexico, Mexico, D.F. 04510, Mexico C. Thomas Philbrick Department of Biological and Environmental Sciences, Western Connecticut State University, Danbury, Connecticut 06810, U.S.A. ABSTRACT. A new species of Podostemaceae, Os¬ erya longifolia, from Jalisco, Mexico, is described and illustrated. Leaf length is the most prominent feature that distinguishes this species. RESUMEN. Se describe e ilustra una especie nueva de la familia Podostemaceae, Oserya longifolia, del estado de Jalisco, Mexico. La longitud de la hoja es la caracteristica mas prominente que distingue a esta especie. Oserya Tulasne & W eddell is an American genus of six species, most recently monographed by van Royen (1954). Like all Podostemaceae, species of Oserya grow attached to rocks in swift-moving river currents. Five of the six species ( O. biceps Tulasne & Weddell, 0. flabellifera Tulasne & Weddell, 0. minima van Royen, O. perpusilla (Went) van Roy¬ en, O. sphaerocarpa Tulasne) occur in northern Brazil and northeastern South America, while 0. coulteriana Tulasne occurs in Mexico. Little is known of the geographic distributions of species of Oserya. In fact, four of the South Amer¬ ican species are known only from the type collec¬ tions. At the time of van Royen’s monograph, 0. coulteriana was known from few collections in the Mexican states of Jalisco and Michoacan. During our recent studies of Mexican Podostemaceae we have found this species to be relatively widespread in Colima, Nayarit, and Guerrero, in addition to Jalisco and Michoacan. A collection from southern Sinaloa is also now known (Novelo & Philbrick, unpublished). While conducting field studies of Mexican Podo¬ stemaceae, we made collections that did not cor¬ respond to any described species. These collections serve as the basis for the description of a new species, Oserya longifolia Novelo & Philbrick. Oserya longifolia Novelo & Philbrick, sp. nov. TYPE: Mexico. Estado de Jalisco: mumcipio de La Huerta, Rio Purificacion, 4 km al oeste de La Huerta, 19°30'N, 104°40'W, 300 m, 26 Mar. 1993, Novelo & Philbrick 1166 (ho- lotype, MEXU; isotypes, MO, NY, WCSU). Figure 1. Herbae aquaticae plerumque caulibus stoloniferis ap- planatis. Folia circinata usque ad 40 cm longa; petiolus laevis; lamina repetite 2-4(-5)-divisa, divisionibus ultunis 0.3-0.6 mm latis, applanatis, apice acutis. Flores her- maphroditi, zygomorphi, pedicellati, axillares, solitarii. Te- pala 3, filamentis alternata, uno in furca duorum stam- inum affixo. Stamina 2, andropodio portata; antherae el- lipticae, dorsifixae, per 2 rimas laterales longitudinaliter dehiscentes. Ovarium 2 carpellis, 2 stylis 0.7-1.2 mm longi. Fructus 2 valvis, quaque valva 3-costata. Aquatic herbs usually with stoloniferous and flat¬ tened stems 0.8-1.0 mm diam., strongly adhering to rocks. Leaves alternate, circinate, up to 40 cm long, petiole 2-17 cm long, cylindrical, smooth, 0.7-1.6 mm diam., with a broadened base; blade 2—4(—5) divided, the ultimate divisions 0.3-0.6 mm wide, flattened, apex acute. Flowers hermaphroditic, zygomorphic, pedicellate, borne singly, axillary, pro¬ tected by a spathella; spathella up to 7 mm long, thin clavate. Pedicels 3.5-8.0 mm long, elongating during anthesis, not expanded at capsule base, re- pals 3, alternate with the filaments, free, membra¬ nous, subulate, one of them attached in the fork between the two stamens; lateral tepals 0.9-1-2 mm long, medial tepal 0.7-0.9 mm long. Stamens 2, borne by an andropodium, andropodium 0.8-1-3 mm long, elongating during anthesis; filaments 1 -0" 1.8 mm long, subulate, flattened, elongating during anthesis, deciduous; anthers 0.4-0.6(—0.8) mm long, elliptic, dorsifixed, 2-celled, dehiscing longitudinally by 2 lateral slits. Ovary superior, 1.7—2.1 mm long, Volume 5, Number 1 1995 Novelo & Philbrick Oserya longifolia 55 Figure 1. Oserya longifolia Novelo & Philbrick. Drawings based on the holotype. —A. General habit of plant with several parts of the leaves interrupted (longest leaf 40.1 cm). — B. Top of the stem with several circulate new leaves. —C. Detail of a single leaf (26.7 cm long). —D. Several stages of flower development from between leaf bases, before (left) and after (right) the spathella is ruptured. — E. Lateral view of the mature flower showing orientation of the styles, anthers, and tepals at the time of dehiscence. —F. Abaxial view of the mature flower showing the andropodium and tepals. —G. Mature styles with receptive stigmas. —H. Cross section of ovary showing placenta and numerous ovules. —I. Abaxial view of a single anther. —J. Adaxial view of a single anther. —K. Cross section of anther. —L. Mature fruit. —M. Cross section of a capsule showing ribs and dehiscence points of the capsule valves. —N. Seed. 56 Novon ellipsoid, 2-locular; styles 2, 0.7-1.2 mm long, free, conical, ovules numerous, placenta axile. Fruit 1.7— 2.6 mm long and 0.8-1.2 mm diam., 2-locular capsule, suture margins thickened; valves 2, each 3-ribbed. Seeds 0.24-0.26 mm long, 0.16-0.20 mm wide, obovoid. Pollen 14.88 gm (SD 0.9, N = 30) diam., tricolpate. Oserya longifolia grows submerged in the swift currents of river rapids, attached directly to rocks of various sizes. No other species of Podostemaceae were observed growing with Oserya longifolia, al¬ though a large population of Eichhornia crassipes (Martius) Solms-Laubach occurs in areas of slow current in association with 0. longifolia. Thus far, Oserya longifolia has only been col¬ lected in the Purificacion River, in areas of full sun. The largest leaves (up to 40 cm long) were found on plants that are submerged in areas of very fast current. In areas of slow current, or when the plants are exposed as the water level drops, the terminal portions of the leaves fall away, leaving the short, coarse bases. Two species of Oserya are now known from Mex¬ ico: O. coulteriana and O. longifolia. These species are readily distinguished. Oserya longifolia is the larger, more robust of the two (Table 1). The longer leaf of O. longifolia (15-40 cm vs. 2-10 cm for O. coulteriana) is particularly notable. However, the tepal that occurs between the anthers at the apex of the andropodium is shorter in O. longifolia than in O. coulteriana (Table 1). Two qualitative features also distinguish the species: the petioles of O. longifolia are cylindrical in cross section, where¬ as those of O. coulteriana are markedly flattened, and distinct petiolar spines are common in O. coul¬ teriana whereas they are lacking in O. longifolia. Leaf length also distinguishes Oserya longifolia from the five South American species of the genus. Leaf length of all South American species ranges Table 1. Vegetative and floral characteristics that distinguish Oserya longifolia from O. coulteriana. Characteristic O. longifolia 0. coulteriana Petiole shape (cross cylindrical flattened section) Petiolar spines absent usually present Leaf length 15-40 cm 2-10 cm Width of ultimate leaf 0.3-0.6 mm 0.1-0.3 mm division Spathella length 5-7 mm 3-6 mm Pedicel length 3.5-8.0 mm 2.0-6.0 mm Length of medial tepal 0.7-0.9 mm 1-1.3 mm Andropodium length 0.8-1.3 mm 0.2-0.5 mm Anther length 0.4-0.6 0.75-1.10 mm Fruit length (-0.8) mm 1.7-2.6 mm 1.8-2 mm from 2 to 3 cm, while leaves of O. longifolia range from 15 to 40 cm. In addition, all five South Amer¬ ican species have a single stamen; in contrast, both O. longifolia and O. coulteriana possess two sta¬ mens (occasional plants of the latter species have a single stamen). Acknowledgments. We thank Fernando Chiang for providing the Latin description, Fernando Chiang, Ronald Aajkar, and Donald H. Les for helpful com¬ ments on the manuscript, and Albino Luna for draw¬ ing Figure 1. Support for this study was provided by The Intercambio Academico Office at the Na¬ tional University of Mexico, a grant to CTP by the National Science Foundation, and a grant to ANR by the Consejo Nacional de Ciencia y Tecnologia. Literature Cited Royen, P. van. 1954. The Podostemaceae of the New World. III. Acta Bot. Neerl. 3: 215-263. Impatiens kulamavuensis , a New Species of Balsaminaceae from India A. G. Pandurangan and V. J. Nair Botanical Survey of India, TNAU—Campus, Coimbatore—641 003, India. Present address of A. G. Pandurangan: Tropical Botanical Garden and Research Institute, Thiruvananthapuram- 695 562, India. Present address of V. J. Nair: The Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, U.K. ABSTRACT. Impatiens kulamavuensis , a distinct new species from India allied to I. parasitica Bed- dome, is described. Impatiens kulamavuensis A. G. Pandurangan & V. J. Nair, sp. nov. TYPE: India. Kerala: Idukki Dist., Meenmutty-Kulainavu, ± 700 m, 6 Oct. 1983, A. G. Pandurangan 82037 (ho- lotype, CAL; isotype, MH). Figure 1. Ab Impatiente parasitica Beddome caulibus stoloni- feris, foliis glandulosis, nervis lateralibus non manifestis; sepalis lateralibus et petalis pilosis; stigmatibus curvatis, sericeo-pubescentibus et seminibus pyriformibus differt. Epiphytic stoloniferous herbs; stems short, stout, fleshy, less than 1 cm long. Leaves alternate; petioles gland-dotted, 2-7 mm long, with 2 prominent stip- itate glands near the base of the blade; lamina ovate- lanceolate, 2-5.5 x 1.5-2.5 cm, membranous, gland-dotted above, glabrous below, deep green, the base cuneate, the apex acuminate, margins entire to obscurely crenulate-serrate, crenules 2-4 mm apart; lateral veins 4—6 pairs, indistinct. Inflores¬ cence a 1-3-flowered raceme; peduncles 2.5-5 cm long; pedicels 3.5-5 cm long. Flowers greenish yel¬ low. Lateral sepals linear, 8-10 x 2-3 mm, hairy outside, glabrous inside. Lower sepal saccate, com¬ pressed, 12-15 x 5-7 mm, glabrous; spurs strongly curved, 7-8 x 3-5 mm, tip obtuse. Dorsal petal erect, orbicular to ovate, 6-7 x 8-9 mm, hairy at base. Lateral petals united close to the base, 3-lobed, upper lobes smaller in size, 7-8 x 9-10 mm. An- droecium 3-3.5 mm long. Ovary ellipsoid, glabrous, 5-6 x 3-3.5 mm; ovules many, 1-seriate; style short; stigma curved, with an obtuse tip, silky pu¬ bescent. Capsules ellipsoid, 10-12 x 4-5 mm, gla¬ brous. Seeds pear-shaped, smooth. Flowering and fruiting October-December. Distribution. Endemic to Meenmutty-Kulamavu, Idukki Dist., Kerala, India. Table 1. Comparison of major differences between Impatiens parasitica Beddome and Impatiens kulama¬ vuensis A. G. Pandurangan & V. J. Nair. I. parasitica I. kulamavuensis Stems 10-12 cm long, not stoloniferous, mo- niliform Stems less than 1 cm long, stoloniferous, not moniliform Leaves glabrous on both surfaces or glabrescent above; lateral nerves prominent Lateral sepals glabrous outside, glandular in¬ side at base Leaves glanduliferous above, glabrous be¬ neath; lateral nerves indistinct Lateral sepals hairy out¬ side, glabrous inside Petals glanduliferous in¬ side at base Petals hairy inside at base Stigmas straight; stigmatic lobes 5-toothed, not pubescent Seeds rounded to ellipsoid Stigmas curved; lobes not 5-toothed, obtuse, silky pubescent Seeds pear-shaped Habitat and ecology. Epiphytic herbs, growing on moss-clad tree trunks at an altitude of 600-700 m. The host trees are Elaeocarpus tuberculatus Roxburgh and Pygium wightiana Blume. Impatiens kulamavuensis was collected in the Idukki hydroelectric project area, Kerala. It is allied to I. parasitica Beddome, but differs in the char¬ acters listed in Table 1. Paratype. INDIA. Kerala: Idukki dist., Meenmutty, ± 600 m, 30 Dec. 1983, A. G. Pandurangan 62585 (MH). Acknowledgments. We thank N. P. Balakrish- nan. Joint Director, and N. C. Nair, former Joint Director, Botanical Survey of India, for help and encouragement. Novon 5: 57-58. 1995. 58 Novon 7 mm 1 J iomrr\ 7 mm Figure 1. lmpatiens hulamavuensis A. G. Pandurangan & V. J. Nair. —A. Habit. —B. Flower. —C. Lateral sepals. D. Lip. E, G. Lateral petals, outer and inner views. —F. Dorsal petal. —H. Stamens. —I. Pistil. —J- Transection of ovary. —K. Fruit. —L. Seeds. Two New Taxa for the Flora of Thailand John A. N. Parnell School of Botany, Trinity College, Dublin 2, Ireland ABSTRACT. The new subspecies Mitrasacme ero- phila subsp. grandijlora (Loganiaceae) and Pedi- cularis lhailandica subsp. parvula (Scrophularia- ceae) are described. Mitrasacme is a widespread Australasian genus, represented in Thailand by three undercollected spe¬ cies. In order to make a name available for the forthcoming Flora of Thailand account, the follow¬ ing new subspecies is described. Mitrasacme erophila Leenhouts subsp. gran- diflora J. Parnell & 0. Griffin, subsp. nov. TYPE: Thailand. Chiang Mai, Omkoi, ca. 35 km from pine improvement centre to Omkoi, 29 Dec. 1978, C. Niyomdham, M. Suangtho & B. Sangkhachand 168 (holotype, L; iso¬ type, AAU). A subsp. erophila foliis glandulis minutis instructis, corolla grandiore, stamenibusque basim corollae versus affixis differt. Herb to 20 cm. Stem < 1 cm high, unbranched, minutely papillose. Leaves 2—4 pairs at the base of the plant, ovate, < 4 mm long, with a single distinct mid-vein; apex acute to blunt. Inflorescence ter¬ minal, lax, branched and umbellate, many-flowered; peduncles 1-3 arising from the stem with 1-5 min¬ ute, triangular bracteoles either sterile or subtending the inflorescence branches; pedicels to 4.2 cm, gla¬ brous. Flowers white. Calyx 1.5-2 mm long, cam- panulate; lobes 0.7-1 mm. Corolla 5-6.7 mm long, campanulate; lobes 1.2-1.7 mm, oblong, mucro- nate. Fruit 1.5-2 mm high; styles connate at the tips; becoming freer with maturity. Mitrasacme erophila subsp. grandijlora differs from subspecies erophila in its leaves, which bear minute peltate scales on their upper surface, its longer corolla (the corolla in subspecies erophila is 3-3.7 mm long) with larger, rounded corolla lobes (the corolla lobes in subspecies erophila are oblong and 1.2-1.7 mm long) and in the position of the stamens’ insertion in the corolla tube (the stamens in subspecies erophila are attached at or above the center). Subspecies grandijlora is known only from the type collection. Pedicularis in Thailand has been recently revised for the Flora of Thailand, where four species are recognized (Yamazaki, 1990). This account does not fully document the great variability seen in Thai material, and some herbarium material does not match that of any of the described species. Material collected by the first joint expedition of the Royal Botanic Gardens, Kew, Trinity College, Dublin, and Khon Kaen University, although conspecific with Pedicularis thailandica, is sufficiently distinct to require circumscription. Pedicularis thailandica Yamazaki subsp. par¬ vula J. Parnell, subsp. nov. TYPE: Thailand. Chiang Mai, Doi Inthanon National Park, forest trail at km 42 along summit road, hill evergreen forest, 7 Oct. 1990, P. Chantaranothai, J. Parnell, D. Simpson & R. Pooma 90/613 (holotype, TCD). A subsp. thailandica statura minore; caulibus vix lan- atis; foliis minutis una cum floribus ternis verticillatis et corolla alba differt. Herb to 17 cm. Stems erect, simple, subglabrous. Leaves 3-verticillate; petioles 2 mm, sparsely lanate; blades ovate, deeply pinnate, the pinnules crenate. Flowers solitary, 3-verticillate; pedicels 0.2-0.5 mm. Calyx tubular-campanulate, 5.5 x 4 mm, sparsely lanate, 5-toothed; teeth rounded, ca. 2 mm long. Corolla white, 18-20 mm long, tube straight, 12 x 2 mm, densely glandular-pilose especially on the lobes and hood; hood (galea) ca. 3.5 mm; straight near the mouth, abruptly curved near the tube, two¬ toothed; lower lip three-lobed, all lobes broadly ob¬ long. Stamens with glabrous filaments ca. 10 mm long; anthers deltoid, ca. 1.5 mm long. Mature cap¬ sule unknown. Subspecies parvula is known only from the type collection. Pedicularis thailandica subsp. parvula differs from subspecies thailandica in being only half its height, with the main stem almost glabrous (vs. spreadingly lanate); the leaves being much smaller and not duplicate-serrate; and the flowers being in whorls of three (vs. four) with a white (vs. yellow) corolla. Novon 5: 59-60. 1995. 60 Novon Acknowledgments. I thank P. Chantaranothai and R. Pooma, who were jointly responsible for arranging the field excursion to Doi Inthanon, D. A. Webb for helping with the Latin, and D. Mid¬ dleton for commenting on a draft of this paper. Literature Cited Yamazaki, T. 1990. Scrophulariaceae. In: T. Smitinand & K. Larsen (editors), Flora of Thailand, 5: 139- 238. New Species of Siparuna (Monimiaceae) I. Four New Species from Ecuador and Colombia Susanne S. Renner and Gerlinde Hausner Institute of Systematic Botany, University of Mainz, Bentzel-Weg 2, D-55099 Mainz, Germany Abstract. Siparuna croatii and S. palenquensis from central and Pacific Ecuador, respectively, S. gigantotepala from Pacific Colombia and Ecuador, and S. harlingii from the eastern Andean slopes of Colombia and Ecuador are described, illustrated, and discussed as to their relationships with morpholog¬ ically similar species. Comprising an estimated 150 species, Siparuna is the largest and least understood genus of the Monimiaceae. The genus was last revised by Perkins (1901, 1911), whose typological species concept overstates diversity in certain areas, such as plan- altan and southeastern Brazil. In Colombia, Ecuador, and Peru, however, increased collecting activity over the past 20 years has turned up numerous new species, and it now appears that most species of Siparuna are Andean in distribution. Material in the major relevant herbaria (acronyms following Holmgren et a]., 1990), such as AAU, BM, COL, F, G, GB, GH, GOET, K, M, MA, MJG, MO, NY, OXF, P, QCA, QCNE, S, SEL, TCD, UC, US, W, WU, and Z, currently amounts to over 2500 col¬ lections representing approximately 100 species from the Andes. Continued discovery of new taxa in upper Amazonia strongly suggests the need for additional collecting. As a result of ongoing work toward a monograph of Siparuna, we herein describe four new species in order to make their names available for the forthcoming treatment of the Monimiaceae for the Flora of Ecuador. Work on the Peruvian species is still in progress, rendering description of several apparently new species extending south be¬ yond Ecuador inadvisable. Flowers of Siparuna are small and unisexual, and plants are monoecious or dioecious (Perkins, 1901; Feil, 1992). The flowers have an obconical or ur- ceolate floral cup in which the free carpels or sta¬ mens are more or less completely enclosed (e.g.. Figs. 1A, and 4B and F) and usually minute (1-4 mm) tepals. A feature characteristic of the genus is that the floral apex forms a roof, called a velum, with a central pore through which the upper parts of the stamens or styles protrude (for details of floral ontogeny and a morphological interpretation of the velum, see Endress, 1980). In the male flowers, the velum is more or less raised at anthesis and the pore is relatively large (Fig. 1A); in the female flowers, the inner margin of the velum is always distinctly raised and the pore is narrower (Fig. 1C). The stamens are sessile and initiated centripetally, the innermost ones often remaining small or even staminodial. The anthers are two-locular and open around the perimeter of both pollen sacs except at the distal end (Figs. IB, G, N; 4G; compare also Endress & Hufford, 1989, figs. 97-100). Dehis¬ cence results in a flaplike structure that is hinged distally and bent upward. In fully anthetic male flowers, these flaps usually can be seen with the unaided eye. Anthesis typically lasts 9-16 days in male flowers and 16-24 days in females (Feil, 1992), and the fertilized flowers then expand. Measurements of flo¬ ral parts and descriptions of their shape are therefore of limited value. More diagnostically useful are the shape, color, and pubescence of the leaves and the number of stamens and styles. Siparuna croatii Renner & Hausner, sp. nov. TYPE: Ecuador. Cotopaxi: 63.4 km SE of Quevedo, 6 km NW of El Corazon, primary forest on steep slopes near a waterfall above Rio Angamarca, 1030 m, 4 Apr. 1983 (female, fl), T. B. Croat 55752 (holotype, QCNE; iso¬ types, AAU, MO, QCA). Figures 1A-D, 2. Species ramulis quadrangularis valde sulcatis et foliis late ellipticis (18-42 x 10-20 cm) a congeneribus div- ersa. Dioecious shrub or tree, 4-5 m tall, partly with a liana-like habit, the twigs quadrangular, deeply sulcate, and khaki-colored due to a dense indumen¬ tum of minute stellate hairs. Leaves opposite, the petioles 4-5 cm long; the lamina drying brownish to olive-green, chartaceous, elliptic to broadly ellip¬ tic, 18-42 x 10-20 cm, the base obtuse or sub¬ acute, the apex shortly apiculate, the lower surface with minute appressed stellate hairs, the upper sur- Novon 5: 61-70. 1995. 62 Novon Volume 5, Number 1 1995 Renner & Hausner Siparuna 63 face scantily stellate-pubescent to glabrescent and rather smooth, with 22-24 pairs of secondary veins, the secondary and tertiary veins distinctly promi- nulous and yellow below, the margin inconspicuously denticulate. Cymes of both sexes axillary in groups of 2-4, ample and multiflorous, 3-5 cm long, with minute yellowish khaki stellate hairs. Male flower at anthesis 1.9-2.1 x 1.8-2.1 mm, the floral cup urceolate to globose with minute stellate hairs, the floral roof moderately raised, glabrous and drying dark brown, the 4-5 tepals obtusely triangular and 0.4-0.6 mm long, when fresh greenish; stamens 6, fleshy and containing white globules (large oil cells), the 4 outer ones slightly exserted at anthesis. Female flowers at anthesis of the same size and shape as the males but the floral roof raised to a cylindrical bulge separated by a distinct groove from a second innermost tube sheathing the style bases; the styles 5-7, basally united. Fruit fleshy and red when fresh; mature fruit and seeds unknown. Distribution, habitat, and phenology. Endemic in central Ecuador; growing in primary montane forest on steep slopes and in secondary scrub at elevations of 1000-1400 m; collected flowering and with young fruits in April and May. Siparuna croatii differs from all other species in the genus in its khaki-colored, deeply sulcate, qua¬ drangular branchlets, olive-green leaves with sec¬ ondary and tertiary veins distinctly raised below, and, for the genus, ample, khaki-colored inflores¬ cences with small flowers. Paratype. ECUADOR. Manabi: trail from El Corazon to Facundo Velo, 1-3 km S of El Corazon, 1400 m, 17 May 1980 (male, fl), Harling & Andersson 19216( AAU, GB, MJG, QCA). Siparuna palenquensis Renner & Hausner, sp. nov. TYPE: Ecuador. Pichincha: Hcda. Cov- adonga on Rio Pilaton, 1000 m, 2 July 1955 (male, fl), E. Asplund 16764 (holotype, S; is¬ otypes, MJG, QCA). Figures 1K-N, 3. A Siparuna eggersii Hieronymus foliis obovatis (lb- 27 x 10-17 cm) dilfert. Dioecious tree, 4-10 m tall, young branchlets densely pubescent with minute stellate hairs, older branches dark brown and subglabrous, terete. Leaves opposite, the petioles 1.5-4.5 cm long; the lamina drying olive-green or brown, chartaceous, obovate, 16-27 x 10-17 cm, the base cordate, truncate, or rounded, the apex shortly apiculate, both surfaces rather rough and scantily to moderately stellate- pubescent, with 10-13(-16) pairs of secondary veins, veins to the third order impressed above, distinctly visible below, the margin minutely denticulate. Cymes of both sexes axillary in groups of 2-4 and multi¬ florous (less so in the females), 1.5-4 cm long, and almost glabrous. Male flower at anthesis ca. 5.5- 6.3 x 5-5.5 mm, the floral cup obconical with scant stellate hairs, the floral roof strongly raised, glabrous and drying dark brown or black, the 4-5 tepals nearly completely fused, fresh greenish yel¬ low; stamens 6, often with 4 outer and 2 central ones, fleshy and containing white globules (oil cells), the outer stamens distinctly exserted at anthesis. Female flowers at anthesis of the same size and shape as the males; the styles ca. 15-30, basally united. Fruit fleshy, ca. 1.3-2 x 1-2 cm, fresh immature light green with sparse blackish scales, mature yel¬ lowish or reddish orange and smelling strongly of lemon; the 15-30 seeds distinctly visible in dried fruits, verrucose, gray, and with a red aril when fresh. Distribution, habitat, and phenology. Restrict¬ ed to Pacific Ecuador and described by collectors as infrequent or rare; growing in disturbed premon- tane forests on steep slopes or on cliffs; sea level to 1200 m; collected flowering and fruiting mainly from April to September. The suggested relative, Siparuna eggersii Hi¬ eronymus, has smaller, obovate leaves (usually 12- 15 x 8-10 cm) with cuneate bases. The two species have previously been confused, as, for example, in the florula of the Rio Palenque Biological Station by Dodson & Gentry (1978), which describes and il¬ lustrates specimens of S. palenquensis as S. eg¬ gersii. Paratypes. ECUADOR. Esmeraldas: NE across Rio Blanco from Quininde, Little 6230 (F, K, US). Los Rios: Rio Palenque Biological Station, Dodson 5134 (AAU, F, MO, QCA, SEL, US), Dodson 5723 (MO, QCA, SEL, Figure 1. A-D. Siparuna croatii Renner & Hausner (A, B, Harling A Andersson 19216; C, D, Croat 55752). —A. Part of male inflorescence. —B. Stamen. —C. Female flowers. —D. Young fruit. E-I. Siparuna gigantotepala Renner & Hausner (E, Barfod et al. 48903; F, G, Cuatrecasas 15746; H, I, Rubio et al. 999). — E. Young male flower. —F. Mature male flower. —G. Stamens. — H. Female flower. —I. Young fruit. K-N. Siparuna palenquensis Renner & Hausner (K, 0llgaard 98054; L, Jaramillo 44; M, Dodson 5134; N, Asplund 16764). —K. Female flowers. —L. Mature fruit. —M. Male flower. —N. Stamens. The following parts share the same magnification: A, C, E, F, H, I, K, M; B, G, N; and D, L. 64 Novon Figure 2. Female specimens of Siparuna croatii Renner & Hausner (Croat 55752 ). —A. AAU isotype. — B- MO isotype. Volume 5, Number 1 1995 Renner & Hausner Siparuna 65 Figure 3. Female specimens of Siparuna palenquensis Renner & Hausner. —A. Croat 55702. B. 0llgaard 98054, AAU, paratype. 66 Novon US), Dodson 5930 (MO, NY, SEL, US), Gentry & Dod¬ son 17970 (MO, QCA). Pichincha: Pichincha, ca. 8 km SE of La Aurora, km 7 on Sto. Domingo-Quevedo rd., at bridge over Rio Baba, 0llgaard 98054 (AAU, QCA); Tinalandia, 9.6 km E of Santo Domingo de los Colorados, above Rio Toachi, Croat 55702 (AAU, MO); km 3 of Toachi-Las Pampas rd., Dodson & Gentry 13708 (F, MO, QCNE, SEL); old rd. Quito-Santo Do¬ mingo, 2-9 km NE of turn-off to old rd. of AUuriquin, Luteyn et al. 8736 (AAU, NY, QCA, QCNE); Aloag- Santo Domingo, Toachi, at the confluence between Rio Pilaton and Rio Toachi, Sparre 13829 (MJG, S). Co¬ topaxi: island in Rio San Pablo, near La Mana, Webster 22727 (UC). Siparuna harlingii Renner & Hausner, sp. nov. TYPE: Ecuador. Morona-Santiago: 7-8 km N of Gualaquiza on rd. to Indanza, 1500 m, 16 Apr. 1985 (male, fl), G. Harling & L. An- dersson 24175 (holotype, QCA; isotypes, AAU, GB). Figure 4. Species Siparunae asperae (Ruiz & Pavon) A. DC. proxima, cujus flores foliorumque texturam et colorem habet. Differt floribus fructibusque minus pubescentibus et pedicellis paullo incrassatis. Dioecious tree or shrub, sometimes scandent, 2- 6(-12) m tall, the young and older branchlets dense¬ ly pubescent with stellate hairs, subangular. Leaves opposite and those of a pair slightly unequal in size, the petioles (2-)2.5-4(-7) cm long; the lamina dry¬ ing dark green to dark brown, papery, brittle, and subbullate to rather smooth, elliptic to broadly el¬ liptic, sometimes narrowly elliptic, 18-35 x 9-22 cm, the base truncate to cordate, occasionally round¬ ed or acute, the apex acuminate, both surfaces densely stellate-pubescent, glabrescent on the upper surface, with (8—)12— 13(—15) pairs of secondary veins, these distinctly visible on the upper surface and slightly raised and yellowish brown pubescent on the lower surface, the tertiary venation dense and distinctly visible, the margin finely denticulate. Cymes of both sexes axillary in groups of 2-4, much- branched (less so in the females), 2-5 cm long, with minute stellate hairs. Male flower at anthesis ca. 4- 4.5 x 4-5 mm, the floral cup obconical with minute, pale yellowish stellate hairs, the floral roof distinctly raised, glabrous and drying black, the 4-5 tepals obtuse or triangular and 1-1.5 mm long, glabrous or occasionally with a few stellate hairs, when fresh greenish yellow to creamy white, turning red; sta¬ mens 6, often with 4 outer and 2 central ones, fleshy and containing whitish globules (oil cells), the outer stamens distinctly exserted at anthesis. Female flow¬ ers at anthesis of the same size and shape as the males but the floral roof centrally conspicuously raised; the styles 20-30, free. Fruit fleshy, 1.5 x 1.5 cm (when fresh to 2 cm diam.), when immature green with pink spots, when mature light red and with a strong lemon smell when crushed, glabrescent; the 20-30 seeds distinctly visible in dried fruits, verrucose, gray and with a red aril when fresh. Distribution, habitat, and phenology. On the eastern Andean slopes from Putumayo and Caqueta in Colombia to Napo, Pastaza, Morona-Santiago, and Zamora-Chinchipe in Ecuador (and expected in Loja); usually collected in disturbed terra firme forest from 300 to 2000 m altitude; in Ecuador flowering and fruiting year-round. Siparuna harlingii can only be confused with S. aspera from which it differs especially in the fruit and pedicels, which in fruit remain slender in S. harlingii and become fleshy and almost part of the fruit in S. aspera. The two also differ in pubescence, S. harlingii having generally much shorter hairs, and in the leaf venation, S. harlingii having (8-) 12— 13(— 15) lateral nerves, S. aspera (15-)16-20(- 26). Finally, the two may be distinguished by the number of styles (20-30 in S. harlingii vs. (10-) 12— 15(— 18) in S. aspera). As is the case with several species of Siparuna, the crushed leaves of this plant are rubbed on the body by the Quichua Indians in Ecuador to cure “mal aire,” a general term used for various diseases of the nervous system and stomach. The Quichua name of the species is “Malaire panga,” panga meaning leaf ( Neill & Palacios 6993). Paratypes. ECUADOR. Zamora-Chinchipe: horse- trail Guadalupe-San Jose de Yacuambi, along Rio Ya- cuambi, potreros and riverside woods, Harling & An- dersson 13937 (AAU, GB). Tungurahua: between Ban¬ os and Rio Verde, Acosta-SoKs 10267 (F); Rio Topo, Harling et al. 10073 (AAU, GB); 30 km on rd. from Mera towards Banos, Lawesson et al. 43293 (AAU, QCA); Rio Margaritas, Penland & Summers 141 (F, GH, NY). Napo: Tena, Asplund 8938 (QCA, S); Tena, rd. to Archidona, Harling 3660 (S); Mission Shandia, Jatun Yaku River, Barclay 4932 (COL); Puerto Napo, Benoist 4757 (P); Puerto Napo, path to Latas, Harling 3537 (S), Ceron & lguago 5518 (AAU, QCNE); 8 km down¬ river from Misahualli, Neill & Palacios 6993 (AAU, F, MO, QCNE); km 32-35 Puyo-Tena rd., Jorgensen et al. 61248 (AAU, QCA); Zatzayacu, Mexla 7097 (NY, UC, US). Pastaza: Mera, Asplund 18357 (S), Harling et al. 7807 (AAU, GB), 7840 (AAU, GB). Morona- Figure 4. Siparuna harlingii Renner & Hausner (A, Harling 928; B, Cuatrecasas 11167; C, Harling & Andersson 13937; D, Neill & Palacios 6993; E-G, Harling & Andersson 13917). —A. Habit. —B. Female flower. C. Volume 5, Number 1 1995 Renner & Hausner Siparuna 67 Young fruits. —D. Mature fruit. —E. Male inflorescence. —F. Male flower. - G. Stamen. The following parts share the same magnification: A, E; B, F; and C, D. 68 Novon Santiago: rd. Limon-Macas, 96 km NE of Limon, Boh- lin et al. 1477 (GB, QCA); near Mendez, Camp E-851 (NY, S), Harling 928 (S); Mendez-Limon rd., ca. 3.2 km S of turnoff to Mendez, Dorr & Valdespino 6345 (AAU, NY, QCA, QCNE); Rio Tutanangosa, rd. Sucua- Huarani, Holm-Nielsen 20508 (AAU). Zamora—Chin- chipe: rd. Zamora-Zumba, km 5-12, Harling & An- dersson 13917 (AAU, GB); 10 km S of Zamora on rd. along left shore of Rio Jamboe, Harling & Andersson 24014 (AAU, GB, QCA); rd. La Saquea-Yacuambi, 1 km N of Chapintza, Harling & Andersson 23893 (AAU, GB, QCA); Zamora, Harling 5930 (NY, S), Knight 713 (S); Cumbaratza, Jaramillo & Winnerskjold 5922 (NY, QCA). COLOMBIA. Putumayo: Rio Guamues, San An¬ tonio del Guamues, Cuatrecasas 11167 (F, US); Mocoa, Cuatrecasas 11278 (F, US), Schultes 2031 (F, K, US), Schultes & Cabrera 19064 (US); Rio Mocoa drainage at San Antonio, Ewan 16704 (US); Puerto Umbria, King 1764 (BM, F, K, MO, NY, S, US); near Rio Putumayo, Soejarto et al. 1277 (US). Caqueta: 62 km SE of Guadalupe, along rd. to Florencia, Davidse et al. 5633 (COL, NY). Siparuna gigantotepala Renner & Hausner, sp. nov. TYPE: Ecuador. Carchi: San Marcos val¬ ley, 600 m, 20 Nov. 1983 (male, fl), A. Barfod, L. P. Kvist & D. Nissen 48903 (holotype, QCA; isotype, AAU). Figures 1E—I, 5. Siparuna subscandens A. C. Smith tepalis longe acu- tatis (2-3 mm) conspicuis distinguenda. Dioecious shrub or treelet, 2-3(-5) m tall, bran- chlets densely covered with minute sessile stellate hairs, quadrangular and slightly sulcate. Leaves op¬ posite, the petioles 2.5-4.5 cm long; the lamina drying umber, chartaceous, obovate, 18.5-33 x 8.5-14 cm, the base cuneate to acute, the apex cuspidate with the tip (l-)1.5-2(-3) cm long, both surfaces densely covered with minute stellate hairs, with 9— 10(—13) pairs of secondary veins, veins to the third order slightly raised above and distinctly raised below, the margin doubly serrate or subentire. Cymes of both sexes axillary in pairs and multiflorous (less so in the females), 1.5-5(-9) cm long, densely covered with minute stellate hairs. Male flower at anthesis ca. 2.8—3.2 x 2.5-3 mm, the floral cup obconic with stellate sessile hairs, a few of these also on the inside of the tepals and the floral roof, the floral roof distinctly raised, the (4-)5-6 tepals ob¬ long and 2-3 mm long, when fresh greenish cream or white; stamens 4-5(-6), the outer ones somewhat exserted at anthesis. Female flowers at anthesis of the same size and shape as the males but the velum distinctly raised to a central tube surrounding the style bases; the styles 5-8, free. Fruit fleshy, ca. 1.3-2 x 1-2 cm, subglobose and crowned by the persistent tepals, when mature red or purple with whitish spots and strongly lemon-scented; the ca. 5-8 seeds verrucose, gray and with a red aril when fresh. Distribution, habitat, and phenology. Collected in very humid primary pluvial forests in Pacific Colombia and Ecuador at elevations of 20-1500 m; in Esmeraldas and Carchi collected flowering and fruiting from November to April. Siparuna gigantotepala differs from all other Andean species in the elongate, oblong tepals that persist in fruit. A few specimens of S. gigantotepala (e.g., Forero et al. 6740) possess small domatia at the leaf bases. In this they resemble the western Ecuadorean S. eggersii Hieronymus, the western Colombian S. subscandens A. C. Smith, and the Panamanian S. domatiata A. H. Gentry. All three differ from S. gigantotepala in lacking elongate tepals. Siparuna eggersii also has smaller, paler green-drying leaves that lack the pronounced drip tip of S. gigantotepala, S. subscandens has nar¬ rower leaves, and S. domatiata has much larger domatia (as far as known) than S. gigantotepala. There is one Amazonian species, S. macrotepala Perkins, which has similarly conspicuous tepals. It differs in pubescence (simple hairs in S. macrotepala vs. stellate hairs in S. gigantotepala), number of secondary veins in the leaves (5-6(-8) in the first vs. 9— 10(— 13) in the second), and number of styles (9-12 in the first vs. 5-8 in the second). Moreover, S. gigantotepala has free styles, while those of S. macrotepala are united into a tube. The Coaiquer (Awa) Indians inhale the penetrat¬ ing smell of the fruits and leaves of S. gigantotepala to clear the nasal passages (A. Barfod et al. 48903). The local names “Limon de monte” or “Diablo de monte limon” also refer to the strong smell of the fruits. Paratypes. ECUADOR. Esmeraldas: Eloy Alfaro, Reserva Cotacachi-Cayapas, Parroquia Luis Vargas Tor¬ res, Rio Santiago, Tirado et al. 804 (QCNE); trail to Rio Mataje Awa encampment from Rio Palavi, Hoover et al. 4007 (AAU, MO, QCA); Reserva Awa, 25 km NW of El Chical, Municipio Maldonado, Rubio et al. 999 (AAU, MO, QCNE), Rubio et al. 1036 (AAU, MJG, MO, QCNE); Reserva Awa, Mataje, Jorgensen et al. 65316 (AAU, QCA), Rubio & Quelal 1356 (AAU, MJG, MO, QCNE). Carchi: Reserva Forestal Awa, trail Gualpi bajo Iara- bita, Jorgensen et al. 65232 (AAU, QCA); Gaulpi Chico, Awa encampment, Hoover et al. 2602 (AAU, MO, QCA); Reserva Awa, Centro San Marcos, Mendez et al. 162 (QCNE); wet plateau above San Marcos de los Coaiqueres, 0llgaard et al. 57274 (AAU, QCA), 0llgaard et al. 57584 (AAU, QCA). COLOMBIA. Choco: Quibdo-Tu- tunendo rd., 14 km E of Quibdo, Gentry & Renteria 24139 (MO); San Jose de Palmar, hoya del Rio Torito, finca Los Guaduales, quebrada Santa Fe, Forero et al. 6740 (MO). Valle: Rio Naya, Puerto Merizaldo, Costa Pacifica, Cuatrecasas 13964 (F, US); Rio Yurumangui. Cuatrecasas 15746 (F, US); Rio Calima, ca. 10 km N Buenaventura, Gentry 35263 (COL, MO). Narino: Mun¬ icipio de Tumaco, Mora 4212 (COL). Volume 5, Number 1 1995 Renner & Hausner Siparuna 69 Figure 5. Female specimens of Siparuna gigantotepala Renner & Hausner. —A. 0llgaard et al. 57274. B. Barfod et al. 48903, AAU isotype. 70 Novon Acknowledgments. We thank A. Berg and A. Horn for help with the illustrations, D. Neill for information on the whereabouts of duplicates in Qui¬ to, and D. Lorence and P. M. Jorgensen for insightful reviews. Dan Nicolson went over the Latin diag¬ noses; his input is gratefully acknowledged. Literature Cited Dodson, C. H. & A. H. Gentry. 1978. Flora of the Rio Palenque Science Center. Selbyana 4: 1-628 [Mon- imiaceae, pp. 434-436]. Endress, P. K. 1980. Ontogeny, function and evolution of extreme floral construction in Monimiaceae. PI. Syst. Evol. 134: 79-120. - & L. D. Hufford. 1989. The diversity of sta¬ men structures and dehiscence patterns among Mag- noliidae. Bot. J. Linn. Soc. 100: 45-85. Feil, J. P. 1992. Reproductive ecology of dioecious Siparuna (Monimiaceae) in Ecuador—A case of gall midge pollination. Bot. J. Linn. Soc. 110: 171-203. Holmgren, P. K., N. H. Holmgren & L. C. Barnett. 1990. Index Herbariorum 1. The Herbaria of the World, Ed. 8. New York Botanical Garden, New York. Perkins, J. R. 1901. Beitrage zur Kenntnis der Mon¬ imiaceae. III. Monographic der Gattung Siparuna. Bot. Jahrb. 28: 660-705, t. 12-14. -. 1911. Monimiaceae. Pp. 1-67 in A. Engler (editor). Das Pflanzenreich IV, 101 (Nachtrage). W. Engelmann, Leipzig. Two Lesquerellas (Cruciferae) of South Central and Western Montana Reed C. Rollins Gray Herbarium of Harvard University, 22 Divinity Avenue, Cambridge, Massachusetts 02138, U.S.A. ABSTRACT. Two heretofore unknown species of Lesquerella are described. These occur in limited areas in the mountains of central and western Mon¬ tana. Both species, here named L. lesicii and L. pulchella, occur in patches of limestone-derived soils and rubble that are conspicuously barren of trees and shrubs and other types of plant cover, or on the borders of such areas where the limestone detritus meets soils and rubble of crystalline rocks. Also, they are often found in open areas in sparse stands of low-growing trees or shrubs. Recent samplings of populations of Lesquerella in Montana show that species diversity in this genus goes somewhat beyond that previously recognized in monographic treatments (Payson, 1922; Rollins & Shaw, 1973; Rollins, 1993). What has been known for the area is that species of Lesquerella are often abundant and are an important component of the flora of specialized habitats, especially where limestone or derivatives of limestone are involved. The nature of some of these habitats is well shown in Figures 1 and 2, where the highly reflective limestone-derived soil and rubble contrast with the more vegetated surrounding areas. These windswept whitish areas are the places where L. pulchella grows. Also known is that a high degree of com¬ plexity exists within some species, such as L. alpina (Nuttall ex Torrey & A. Gray) S. Watson, which is frequently encountered in the general region. But now we see that there are existing taxa not previ¬ ously recognized, and these have complicated re¬ lationships with species south of Montana in Idaho and Wyoming. The main purpose of this paper is to describe two new species of Lesquerella and ex¬ plain their relationships to previously known taxa. It is somewhat ironic that I had only recently (Rol¬ lins, 1993) completed a review of Lesquerella as it occurs in North America when these two species showed up too late to be included in my treatment. Lesquerella lesicii Rollins, sp. nov. TYPE: U.S.A. Montana: abundant in gravelly limestone-de¬ rived soil at edge of limber pine woodland on the ridge W of Layout Creek, % mi. S of Mys¬ tery Cave, Pryor Mountains, Carbon County, 7500 ft., T85, R28 E, S21, SW '/„ with Shosh - onea pulvinata, Astragalus aretoides, and A. miser, 20 June 1992, Peter Lesica 5707 and Rob DeVelica (holotype, GH). Herba perennis, caudicibus simplicibus, folia basibus erectis dense stellatis argenteis 0.5-1 cm longis, caulis erectis vel decumbentibus gracilibus 1-1.5 dm longis, petalis aureis spathulatis vel lingulatis 6-7 mm longis, pedicellis fructiferis recurvatis vel divaricatis 5-10 mm longis, siliquis globosis vel subglobosis 3-4 mm diametro pilis stellatis adspersus, loculis 3-5 ovulatis. Delicate perennial; caudex simple, sometimes elongated and with old leaf bases; basal leaves erect, usually fewer than 10, entire, 1.5-3 cm long, pet¬ ioles slender, abruptly expanded to blade, 1-2.5 cm long, blades broadly ovate to elliptical, 0.5-1 cm long, silvery from a dense cover of stellate trichomes; leaf trichomes small, ca. 0.02 mm diam., ray tips 15-25, each primary ray forked near its base; flow¬ ering stems very slender, mostly filiform, simple, erect to decumbent, 1-1.5 dm long; cauline leaves few, remote, ± spatulate, lower somewhat petiolate, upper cuneate at base; inflorescences lax, rarely nodding, usually with fewer than 10 flowers; sepals erect, densely pubescent, oblong, nonsaccate and without scarious margins, 3.5-4 mm long; petals yellow, often fading to light purple toward their tips, spatulate to nearly Ungulate, 6-7 mm long; stamens strongly tetradynymous; infructescences lax, greatly elongated; fruiting pedicels filiform, recurved in a single arch to widely spreading, 5-10 mm long; siliques globose or subglobose, spreading at right angles to rachis to pendent, 3-4 mm diam., ± densely pubescent on exterior, valves glabrous on the interior; styles ca. 1.5 mm long, glabrous or Novon 5: 71-75. 1995. 72 No von with a few trichomes near their bases; ovules 3-5 per locule; mature seeds not seen. Lesquerella lesicii is most closely related to L. fremontli Rollins & E. Shaw. Both species are ap¬ parently endemic to very limited areas and appear to be restricted to Madison limestone derivatives. Lesquerella lesicii occurs in the Pryor Mountains of Montana, which are northwest of the Big Horn Mountains and separated from that range by the canyon of the Big Horn River. Lesquerella fremontii is found at the southeastern end of the Wind River range in Wyoming. These areas are about 200 miles apart. The two species are singular in having small unbranched caudices (although definitely perennial), erect basal leaves, simple flowering stems, more or less recurved fruiting pedicels, and globose siliques. Both species have small, stellate, many-rayed tri¬ chomes and weak stems, which are particularly frag¬ ile in L. lesicii. Lesquerella lesicii differs from L. fremontii in having basal leaves with very thin petioles that abruptly expand to a broadly ovate to orbicular blade, whereas in L. fremontii the petioles are stouter and expand gradually to a spatulate or nearly linear blade. The flowering stems of L. fremontii are hor¬ izontal along the ground and the infructescences are secund. In L. lesicii these stems are erect and the infructescences have the usual racemose pattern. The fruiting pedicels are definitely recurved in a single arch and the siliques are more or less pendent in L. fremontii, but in L. lesicii the pedicels, al¬ though tending to be recurved, are more often just slightly arched and are widely spreading, bearing siliques that are horizontal. The pedicels are so finely filiform that they are extremely fragile, while those of L. fremontii are thicker and much less fragile. According to Peter Lesica (in litt.), Lesquerella lesicii is not the only endemic species that occurs in the Pryor Mountains. He points out that Penste- mon caryi Pennell and Erigeron allocotus Blake are endemic there and that the recently described monotypic Shoshonea pulvinata Evert & Con¬ stance is restricted to Madison limestone in the Pryors, and along the front of the Beartooth, Ab- saroka, and Owl Creek mountains in Wyoming and Montana. Paratypes. U.S.A. Montana: Carbon County, very common in gravelly limestone derived soil on a NE-facing slope at the head of Big Coulee, Pryor Mtns., 6400 ft., T9S, R28E, NE /„ with Cercocarpus ledifolius, Musi- neon vaginatus, and Agropyron spicatum, 19 June 1992, Peter Lesica 5700 and Rob DeVelice (GH, MONTU); locally abundant in stony limestone derived soil in a wind¬ swept Douglas Fir forest on a ridge ca. % mi. S of Mystery Cave, Pryor Mtns., 7500 ft., T8S, R28E, Sec. 21, SW /„ with Hymenoxys torreyana and Eritrichium howar- dii, 27 June 1991, Peter Lesica 5449 (GH, MONTU). Lesquerella pulchella Rollins, sp. nov. TYPE: U.S.A. Montana: Beaverhead County, common in barren calcareous soil in open spruce wood¬ land on S-facing slope along Trapper Creek just S of Hecha Mines, Pioneer Mtns., T3S R11W, Sec. 32, 8600 ft., with Potentilla fruticosa and Zigadenus elegans, 25 July 1991, Peter Lesica 5525 and Steve Cooper (holotype, GH; isotype, MONTU). Herba perennis, caudicibus simplicibus, folia radicalibus simplicibus argenteis petiolata 1-2 cm longis laminis late ovatis vel ellipticis, caulis tenuibus prostratis 2-7 cm longis, folia caulina petiolata vel ad basi cuneatis, sepalis oblongis 3-4(-5) mm longis dense pubescentibus, petalis aureis spathulatis vel lingulatis (6—)7—8(—9) mm longis, pedicellis fructiferis plerumque patuleis ± sigmoideis 5- 7 mm longis, siliquis orbicularibus vel ellipticis pubescen¬ tibus (3-)4-6(-7) mm longis, 3-4 mm latis, stylis 1-1.5 mm longis, loculis 3-5 ovulatis, seminibus oblongis crassis ± 2 mm longis. Perennial; caudex mostly simple; caudex leaves several to numerous, simple, silvery from a dense cover of appressed radiately branched trichomes, blades broadly ovate or elliptical to wider than long, obtuse, 1-2 cm long including the slender petioles; leaf trichomes in several layers, branches usually forked near their bases, primary branches 6-7; stems several to many, slender, simple or occasionally branched, mostly prostrate, 2-7 cm long; cauline leaves petiolate below to cuneate above, spatulate to narrowly elliptical or broadly ovate; inflorescences congested, flowering pedicels divaricately ascending to nearly erect, straight or nearly so; sepals narrowly oblong, 3-4(-5) mm long, nonsaccate, densely pu¬ bescent; petals yellow, spatulate to Ungulate, (6-)7- 8(-9) mm long; stamens erect, filaments slender, anthers oval; fruiting pedicels divaricately ascending to widely spreading, sigmoid to slightly so or nearly straight, slender, 5-7 mm long, not swollen toward the apex; siliques nearly orbicular to efliptical or broadly oblong, rounded above and below, strongly compressed contrary to the septum but with a def¬ inite ridge down the middle formed over the replum edges, densely pubescent but not silvery, (3-)4-6(- 7) mm long, 3-4 mm wide; septum entire or with a narrow perforation; styles 1-1.5 mm long, not expanded at summit; seeds plump, oblong, brownish, ca. 2 mm long, wingless, cotyledons accumbent. Lesquerella pulchella (Figs. 3, 4) is most closely related to L. paysonii Rollins. Both species have sfliques compressed contrary (perpendicular) to the plane of the septum. Their fohage and sfliques are Volume 5, Number 1 1995 Rollins Lesquerella from Montana 73 Figures 1 and 2. Typical habitats of Lesquerella pulchella Rollins. —1. Upper, East Pioneer range, viewed from subsummit of Black Lion Mountain. —2. Lower, north of Black Lion Mountain, south of Vipond Park. Photos by Peter Lesica. 74 Novon Figures 3 and 4. Lesquerella pulchella Rollins (ca. natural size). —3. Upper, flowering plant (Lesica 5528). 4. Lower, fruiting plants (Lesica 5525). Photos by Peter Lesica. covered with relatively small, radiately branched trichomes, and in each species the foliage is silvery from the multiple layers of trichomes present. A single layer of trichomes on the silique surfaces does not produce the same silvery effect. The silique compression is more severe in L. paysonii than in L. pulchella. The result is that the replum margin does not protrude and the replum itself is narrower in L. paysonii than in L. pulchella. In the latter, the replum margin does protrude to form a riblike ridge where the valves come together. Generally in Lesquerella, the presence or absence Volume 5, Number 1 1995 Rollins Lesquerella from Montana 75 of trichomes on the interior of the valves is not an infallible mark of distinction between species, but most of the specimens of Lesquerella pulchella I have examined do have trichomes on the valve in¬ terior. On the other hand, none of the specimens of L. paysonii I have seen have trichomes there. This is worth noting because it may turn out to be a significant difference once enough populations have been sampled to establish the surety of the feature difference. Some variation in the length of the styles is to be expected and does occur in both species. However, this variation ranges from 2 mm in length downward in L. pulchella , but from 2 mm upward to 4 mm in L. paysonii. There is a basic difference in the outline shape of the siliques of Lesquerella paysonii and L. pul¬ chella. In the latter, the siliques are nearly orbicular in outline or only slightly longer than broad, while in L. paysonii the siliques are definitely longer than broad, usually nearly twice as long as wide. The effect of this pattern, combined with the degree of compression on the shape and length of the replum, is seen in the differences of the replum shape in the two species. The replum of L. paysonii is longer, narrower, and tapers more gradually toward the apex than in that of L. pulchella, where the replum tends toward a more oblong shape. In L. pulchella the valve margins are tightly pressed together so that the replum margin does not show and there is a continuous cover of trichomes over the ridge formed between the valves. But in L. paysonii the valve margins are rarely sufficiently together to touch each other. Here the replum margins show a break be¬ tween the valves, and the trichome cover is not continuous from one valve to the other. Lesquerella pulchella ranges in elevation from around 6300 to about 9600 feet. As with other plant species with such a wide elevational occur¬ rence, there is considerable variation from popula¬ tion to population. Some of this variation appears to be correlated with elevation, some not. I note that the flowers on the high elevation variants are both larger and more numerous per plant than in the populations of lower elevations. This was called to my attention by Bonnie Heidel. Paratypes. U.S.A. Montana: Beaverhead County, common on gentle E-facing slope at the top of Keokirk Mt., near contact between crystalline and calcareous par¬ ent materials, T3S, R11W, Sec. 10, 9600 ft., 25 July 1991, Peter Lesica 5528 and Steve Cooper (GH, MON- TU); common in portion of upper S-facing slope on ridge point, S of Bridger Pass, T7S, R11W, Sec. 27, 7140 ft., 5 Aug. 1992, B. Heidel 938 (GH); common in barren calcareous soil on a moderate E-facing slope of the ridge N of Black Lion Mt., T2S, R11W, Sec. 29, 9200 ft., 24 July 1991, Peter Lesica 5522 and Steve Cooper (GH, MONTU); steep slopes of upper montane zone, Nemesis Mt., T14S, R2E, Sec. 31, 9050 ft., 27 June 1992, B. Heidel 722 (GH); near same location, 27 June 1992, B. Heidel 721 (GH); above Scudder Creek on ridges at the edge of Pioneer Mtns., T6S, R12W, Sec. 21, 6720 ft., 24 June 1992, B. Heidel 700 (GH); open scree of S-facing slope, ridge E of Nemesis Mt., T14S, R2E, Sec. 21, 9000 ft., 26 July 1992, Lisa Schassberger Roe 493 and Jonathan Stewart Roe (GH). Acknowledgments. I am much indebted to, and thank, Peter Lesica of the University of Montana, and Bonnie L. Heidel and Lisa Schassberger Roe of the Montana Natural Heritage Program, for pro¬ viding me with excellent study material. Peter Les¬ ica, in addition to excellent collections, also provided significant notes on the field populations of Les¬ querella lesicii and additionally the photographs of Figures 1-4, for which I am grateful. My sister, Aileen G. Roads Carter, has generously provided funds to offset the cost of the colored plates, for which I thank her sincerely. Literature Cited Payson, E. B. 1922. A monograph of the genus Les¬ querella. Ann. Missouri Bot Gard. 8: 103-236. Rollins, R. C. 1993. The Cruciferae of Continental North America. Stanford Univ. Press, Stanford, Cal¬ ifornia. -& E. A. Shaw. 1973. The Genus Lesquerella (Cruciferae) in North America. Harvard Univ. Press, Cambridge, Massachusetts. Merostachys multiramea (Poaceae: Bambusoideae: Bambuseae) and Similar Species from Brazil Tatiana Sendulsky Herbario, Instituto de Botanica, Caixa Postal 4005, 01061-970 Sao Paulo, Brazil ABSTRACT. The true identity of Merostachys mul¬ tiramea Hackel is discussed. As a result of a review of the group of the species similar to M. multira¬ mea, the following are described as new: Meros¬ tachys abadiana, M. caucaiana, M. filgueirasii, M. kleinii, M. magellanica, M. pilifera, M. scan- dens, and M. skvortzovii . The description of floral morphology is added to M. glauca McClure & L. B. Smith, which was originally described from sterile material. A key to the species is provided. All the species are fully illustrated. In the past the name Merostachys multiramea Hackel has been applied erroneously to a number of different species, both described and undescribed. Because true M. multiramea is a distinct and rather rare species, represented in herbaria by only a few collections, a considerable amount of misidentified material has been placed under the specific epithet multiramea and thus left unstudied. Among these are species that present no difficulties in identifi¬ cation: either they are clearly new or they have already been described. However, there is a group of about 10 species that are difficult to distinguish one from another, but which cannot be aggregated under one name as has so far been the case. As botanists working with herbarium collections of Brazilian bamboos have tended, over the years, to assign almost any specimen of Merostachys with medium-sized, 1- or occasionally 2-flowered spike- lets to M. multiramea Hackel, the misidentification has had fourfold results. A number of validly pub¬ lished species, including M. argyronema Lind man, M. claussenii Munro, and M. clausenii Munro var. mollior Doell, were for a long time disregarded in the belief that they were all M. multiramea. Two names, M. anomala Dutra and M. burchellii Mun¬ ro emend. Dutra (not M. burchelii Munro), crept into the group. Collections that would later be con¬ sidered as distinct though unpublished taxa (M. ma¬ gellanica , for instance) went unrecognized. And the identity of M. multiramea itself, the “umbrella” for so many other taxa, was largely obscured. The name Merostachys multiramea was published in 1909 by Hackel, and M. claussenii Munro var. Novon 5: 76-96. 1995. mollior by Doell in 1880 (see von Martius, 1840- 1906). The two species were described again, under new names, by Dutra in 1938: the former as M. anomala Dutra and the latter as M. burchellii Munro emend. Dutra. Dutra’s emendation was based on the assumption that unidentified collections of flowering material in his possession were a good match for the vegetative and sterile material on which Munro (1868) based his original description. This “graft” was pub¬ lished by him (Dutra, 1938) as a distinct species. The situation was further complicated by the in¬ version of the captions to Dutra’s figures: his figure 2 (Dutra, 1938: 149), purporting to show leaves and inflorescences of M. burchellii Munro emend. Dutra, is, in fact, of M. anomala (= M. multira¬ mea), while his figure 3 (Dutra, 1938: 149), which claims to be of M. anomala, is of M. burchellii Munro emend. Dutra (= M. claussenii Munro var. mollior Doell). Return of the captions to their rightful figures eliminates a misleading interpretation of inflorescence and spikelet morphology in the two taxa concerned, and permits their proper identification as synonyms of previously published species. In this context, M. multiramea also assumes its true identity, as a rare bamboo from the southern part of Brazil; Dutra s figure 2 makes it clear that it is not one of the group of rather common species with only one floret in the spikelet. Merostachys multiramea Hackel is char¬ acterized by a 2-flowered spikelet and by its dark brown (nearly black), conical caryopses. Both florets are fertile—the rudimentary apical floret and the upper and lower glumes, called “florets” by authors of the last century, obviously not included. Thus, far from being widely distributed, M. multiramea has been collected only infrequently, and mostly by Dutra, under the name M. anomala. Key to the Species of Merostachys Similar to M. MULTIRAMEA 1. Culm internodes densely pilose, hairs sericeous, translucent, ca. 1 mm long, in some areas sharp, globose prickle hairs also present. M. pilifera 1. Culm internodes glabrous, smooth or scabrous, with prickle hairs or mottled with dark spots and stripes. ^ Volume 5, Number 1 1995 Sendulsky Merostachys from Brazil 77 2( 1). Spikelets always 2-flowered or occasionally 2- flowered (both florets fertile), otherwise bearing different stages of development of the upper floret . 3 2. Spikelets always 1-flowered. 5 3(2). Spikelets always 2-flowered. 4 3. Spikelets occasionally 2-flowered (both florets fertile), otherwise bearing different stages of development of the upper floret; florets ca. 9 mm long, caryopsis ellipsoid, 6 mm long, 1.5 mm wide. M. caucaiana 4(3). Florets 7-9 mm long, 3 mm wide, laterally strongly compressed, imbricate; caryopsis con¬ ical, attenuate at the apex, dark brown (nearly black), 6-7 mm long, 3 mm wide. . M. multiramea Hackel 4. Florets ca. 10-12 mm long; caryopsis broadly ovoid, basally and apically rounded, dark yel¬ low, 6-7 mm long, 4-4.5 mm wide. . M. kleinii 5(2). Spikelets always with two parallel, longitudinal, white lines, formed by the keels of the palea. . M. magellanica 5. Spikelets without parallel, white lines. 6 6(5). Caryopsis ovoid . 7 6. Caryopsis ellipsoid, 7.5 mm long, 3 mm wide, grayish brown, culm leaf blades narrow, 4-5 cm long, 4 mm wide, branch leaf blades lan¬ ceolate, 4-9 cm long, 8-12 mm wide, L:W = 5-7.5. M. jilgueirasii 7(6). Branch leaf blades lanceolate, 3-11 cm long, 6-16 mm wide, L: W = 5-6.9 or 3.5-10 cm long, 8-14 mm wide, L: W = 4.4-7.1; branch complements with 40-200 branches. 8 7. Branch leaf blades broadly lanceolate, 4-12 cm long, 15-20 mm wide, L:W = 2.6-6 or narrowly linear-lanceolate, 5-13 cm long, 6- 12(—17) mm wide, L: W = 8.3-10.8; branch complements with 20-60 branches. 9 8(7). Branch complement with 150-200 branches; branch leaf blades lanceolate, 3-11 cm long, 6-16 mm wide, L: W = 5-6.8; culm leaf blades lanceolate, 6-10 cm long, 10 mm wide; inflo¬ rescences pectinate; caryopsis 5 mm long, ca. 3 mm wide, apically rounded. M. skvortzovii 8. Branch complement with 40-60(-100) branches; branch leaf blades lanceolate, 3.5- 10 cm long, 8-14 mm wide, L: W = 4.4-7.1, culm leaf blades lanceolate, 8-12 cm long, 4- 5 mm wide; inflorescence distichously arranged, caryopsis 5.5-6.5 mm long, ca. 3 mm wide, slightly attenuate apically. M. scandens 9(7). Branch leaf blades broadly lanceolate, 4-12 cm long, 1.5-2 cm wide, L: W = 2.6-6; lemma finely puberulent . M. abadiana 9. Branch leaf blades narrowly linear-lanceolate, 5-13 cm long, 6-12(-17) mm wide, L: W = 8.3-10.8, lemma pilose, the hairs ca. 0.5 mm long, silver-grayish. . M. glauca McClure & L. B. Smith Merostachys abadiana Sendulsky, sp. nov. TYPE: Brazil. Estado de Sao Paulo: Municipio Itatinga, Abadia, 26 sep. 1978 (fl), Campos Neto s.n. (holotype, SP 154498). Figure 1. Culmi ca. 2 m longi. Fragmenta ad extremitates cul- morum internodium 2-5 mm diametro, laevia. Nodi prom- inentes, crista supranodali angusta, tubiformi. Folia cul- morum ignota. Rami in complemento 20-30. Foliorum ramorum laminae late lanceolatae, 4-12 cm longae, 1.5- 2 cm latae. Inflorescentia speciformis, pectinata, 2-4 cm longa. Spiculae 11-13 mm longae, late ellipticae, 1-flos- culatae; gluma infera 3 mm longa; gluma superior 8.5 mm longa, 11-13-nervia, acuta; lemma 9 mm longum, 15- 16-nervium, late lanceolatum, puberulum; palea fu- siformis, 9 mm longa, 10-nervia, anguste sulcata. Cary¬ opsis ovoidea, 4.5 mm longa, 3.4 mm lata, cinereo-lutea. Woody bamboo ca. 2 m high, according to col¬ lector’s note. Fragments of internodes from tip of the culm fistulose, 2-5 mm diam., yellowish, smooth. Nodes salient, supranodal ridge narrow, tubiform. Culm leaves not seen. Branch complement with 20- 30 branches, these 8-40 cm long, 1 mm thick, glabrous, 1 -2-noded before the leaves. Branch leaves of the flowering branches 4-5 per complement: sheaths tight, finely puberulent, the overlapping margin glabrous; adaxial surface dark-spotted; oral setae 2-3 mm long, distally crisped; pseudopetiole grayish yellow, 2 mm long, finely pubescent; inner ligule a thick membrane, pubescent on the back; outer ligule a slightly pubescent rim; blades 4-12 cm long, 1.5-2 cm wide, L:W = 2.6-6, broadly lanceolate, glabrous adaxially except for a scabrous base, with rows of fine prickle hairs and 2-3 mar¬ ginal nerves toward the apex with fine prickle hairs, slightly pubescent abaxially, more densely so at the base, the hairs whitish, margins glabrous or finely scabrous toward the apex. Inflorescences terminal on leafy branches, spikelike, pectinate, 2-4 cm long, with 4-15 spikelets. Rachis densely pubescent. Ped¬ icels pubescent, short. Spikelets solitary, 1 -flowered, bisexual, broadly ellipsoid, somewhat inflated, 11- 13 mm long, 3-4 mm wide. Glumes 2, unequal. Lower glume 1-nerved, carinate, attenuate toward the apex, finely puberulent, 3 mm long, 1 mm wide at the base. Upper glume 8.5 mm long, 3.5 mm wide, broadly lanceolate, acute, awnless, finely pu¬ berulent, 11-13-nerved, adaxially shiny, dark-spot¬ ted. Lemma 9 mm long, 6 mm wide, very broadly lanceolate, deeply concave, enclosing a palea, 15- 16- nerved, finely puberulent abaxially, the adaxial surface dark brown, shiny, densely spotted, the mar¬ gins densely ciliate, the hairs dark, reddish. Palea fusiform, 9 mm long, 5 mm wide, 2-keeled, narrowly sulcate abaxially, 10-nerved, the nerves densely anastomosing, the veinlets uneven; the keels finely ciliate; abaxial surface densely puberulent at the apex and along the sulcus, glabrous in the lower part; adaxial surface shiny, dark. Rachilla extension bristle-like, with a small rudiment on the tip. Car¬ yopsis 4.5 mm long, 3.4 mm wide, ovoid, rostrate. 78 Novon Figure 1. Merostachys abadiana Sendulsky. —a. Branch complements from the upper part of the culm- b. Apical part of the inflorescence. c. Spikelet. —d. Caryopsis, lateral view. —e. Caryopsis, embryo view. —f. Lower glume. g. Upper glume. h. Lemma. i. Palea. —j. Caryopsis, hilum view. —k. Upper portion of the branch leaf sheath and base of the blade (abaxial surface). Based on Campos Neto s.n. (SP 154498). grayish yellow, the embryo and the hilum scarcely manifest externally. Phenology. Known only from the type collection. Distribution. Known only from the type locality, in the State of Sao Paulo, Brazil. Merostachys abadiana was collected on the grounds of a Benedictine abbey in Sao Paulo in 1978 and sent to the Institute de Botanica, Sao Paulo, for identification. No further information about the plant was obtained, except that some clumps of Volume 5, Number 1 1995 Sendulsky Merostachys from Brazil 79 Figure 2. Merostachys caucaiana Sendulsky. —a. Flowering branch complement. —b. Spikelet, lower floret with mature caryopsis, upper floret not fully developed. —c. Spikelets with two fertile florets. —d, e. Spikelets in different stages of development (upper florets). —f. Lodicules. —g. Caryopsis, embryo view. —h. Inflorescence. —i. Caryopsis, lateral view. —j. Caryopsis, hilum view. —k. Upper portion of the branch leaf sheath and base of the blade (abaxial surface). —1. Lower glume. —m. Upper glume. —n. Lemma. —o. Palea. —p. Palea and rachilla extension with underdeveloped upper floret. Based on O. Handro & Menezes s.n. (SP 262720). the bamboos are still growing in that area. It is details of the lemma but differs in the glabrous somewhat similar to M. pilifera (described later in internodes and in the wider branched leaf blades, this paper) in the morphology of the spikelet and which are hairy on the abaxial surface. 80 Novon Merostachys caucaiana Sendulsky, sp. nov. TYPE: Brazil. Estado de Sao Paulo: Municipio de Cotia, Caucaia, 1978 (fl), 0. Handro & Menezes s.n. (holotype, SP 262720). Figure 2. Habitus ignotus. Fragments ad extremitates culmorum internodium fistulosa, laevia, opaca, maculis atro-viridibus rotundatis. Nodi pilis albis, retrorsis ad basin. Folia cul- morum ignotae. Rami in complemento 40-50. Foliorum ramorum laminae lanceolatae, 2-7 cm longae, 5-12 mm latae. Inflorescentia spiciformis, pectinata, 2-3 cm longa, ramum foliorum terminans. Spiculae 1 aut rare 2-flos- culatae, 9-10 mm longae, fusiformes, lateraliter com- pressae; flosculus inferior bisexualis, 9-10 mm longus vel rudimentalis. Gluma inferior 2 mm longa; gluma superior 7 mm longa, 9-11 -nervia; lemma 7 mm longum, 9- nervium; palea 7 mm longa, late sulcata. Caryopsis ellip- tica, 6 mm longa, pallide brunnea. Habit unknown. Fragments of the culm inter- nodes from the tip of the culm delicate, with thin pendent or clambering tips. Intemodes fistulose, ca. 1 cm diam., glabrous, smooth, but not lustrous, opaque, finely mottled with very small, dark green, rounded spots (not stripes) and with a glaucous- whitish touch toward the upper part. Upper culms sometimes rugose with small, sharp, retrorse prickle hairs. Culm walls 1 mm thick. Nodes salient with white retrorse hairs below. Culm leaves not seen. Branch complement (upper portion of plant) with 40—50 branches, these 10—15 cm long, 1 mm thick, glabrous, 1-2-noded before the leaves. Branch leaves 3-4 per complement: sheaths tight, glabrous, adax- ially dark-spotted, the overlapping margin finely cil- iate or glabrous, the intercostal zone finely trans- versally rugose; oral setae 13-15 cm long, dimin¬ ishing in size downward, along the margins of the sheath; pseudopetiole dark, rugose, with a few fine prickle hairs at the base adaxially; inner ligule 0.5 mm long; outer ligule a salient rim; blade 2-7 cm long, 5-12 mm wide, L:W = 4-5.8, lanceolate, with an acuminate apex and slightly asymmetrical at the base, glabrous on both surfaces except for the base abaxially, the hairs whitish, hyaline, ap- pressed, on one side of the midnerve; adaxial surface with 2-3 marginal nerves on one side with rows of fine prickle hairs; margins finely scabrous. Inflores¬ cences spikelike, pectinate, 2-3 cm long, half in¬ serted in ultimate sheath. Rachis and pedicels pilose. Spikelets 1- or rarely 2-flowered. Glumes 2, unequal. Lower glume 2 mm long, 1 mm wide, acuminate, carinate, 1-nerved, puberulent, ciliate at the mar¬ gins. Upper glume 7 mm long, 3 mm wide, broadly lanceolate, 9-11 -nerved, finely puberulent, ciliate at the margins, the nerves anastomosing. Lower floret bisexual, 9-10 mm long, 2-3 mm wide, fu¬ siform, laterally compressed. Lemma 7 mm long, 4 mm wide, broadly lanceolate, finely puberulent, 9- nerved, the nerves anastomosing, the margins cili¬ ate. Palea 7 mm long, glabrous, 2-keeled, broadly sulcate between the keels, the sulcus finely pilose, the keels ciliate. Rachilla extension flat, ca. 0.5 mm wide, puberulent and ciliate laterally, bearing one of several different stages of development of the upper floret, these varying in size and shape, from a small rudiment to a fully formed fertile floret, identical to the lower one. Rachilla extension of the upper floret bristle-like, with a small rudiment at the tip. Lodicules 3, 2 mm long, finely ciliate apically. Ovary not seen. Caryopsis 6 mm long, 1.5 mm wide, ellipsoid, rostrate, light brown, shiny, the embryo and the hilum scarcely manifest externally. Phenology. The only available flowering speci¬ men of this species is the holotype. Distribution. Known only from the type locality, in the State of Sao Paulo, Brazil. Merostachys caucaiana was found growing in a secondary forest in the Municipality of Cotia, ca. 40 km from Sao Paulo. Merostachys filgueirasii Sendulsky, sp. nov. TYPE: Brazil. Distrito Federal: Catetinho, 15°57'S, 47 <> 59 , W, 7 Jan. 1982 (fl), Filgueiras & Pereira 953 (holotype, IBGE; isotypes, CCN, MG, SP, UEC). Figure 3. Culmi 2-10 m longi, 1-1.5 cm diametro; internodia fistulosa, scabra. Nodi glabri; crista supranodali tubiformi. Foliorum culmorum laminae angustae, 4-5 cm longae et 4 mm latae. Rami in complemento 20-50. Foliorum ramorum laminae lanceolatae, 4-9 cm longa, 8-12 mm latae. Inflorescentia spiciformis, pectinata, 3.5-4.5 cm longa, ramorum foliarum terminans. Pedicelli pilosi, base dilata in projecturam nudam, globosam. Spiculae 13.5- 14 mm longae, fusiformes, 1-flosculatae. Gluma infera 2 mm longa; gluma supera 8-10 mm longa, 7 - 9 -nervia, aristulata; lemma 8-9 mm longum, 21-23-nervium, sub- tiliter puberulum, ad marginem strigosum, pilis fuscis; palea 11 mm longa, 12-nervia, anguste sulcata. Caryopsis elliptica, 7.5 mm longa, cinereo-brunnea. Caespitose, woody bamboo, forming dense clumps. Rhizomes pachymorph, sympodial, densely covered by yellow scales. Culms 2-10 m long, 1-1.5 cm diam., flexuous at the tips, according to Filgueiras (1988). Intemodes fistulose, scabrous, green when young, yellowish when mature, finely mottled with small, dark green stripes. Culm walls of the middle intemodes 1-1.5 mm thick. Nodes glabrous, salient, supranodal ridge tubiform, 1-2 mm long, especially evident in young culms. Culm leaves deciduous: sheaths 12-15 cm long, 5-6 cm wide, rounded at the summit, fringed at the apex with dense tufts of Volume 5, Number 1 1995 Sendulsky Merostachys from Brazil 81 Figure 3. Merostachys filgueirasii Sendulsky. —a. Flowering branches. —b. Base of the branch complement. — c. Lower glume. —d. Upper glume. —e. Lemma. —f. Palea. —g. Spikelet. —h. Apical part of the inflorescence. —i. Caryopsis, embryo view. —j. Caryopsis, hilum view. —k. Rhizome. —1. Upper portion of the branch leaf sheath and base of the blade (abaxial surface). —m. Culm leaf. Based on Filgueiras <£ Pereira 953 except for k and m, which are based on Filgueiras & Brochado 2009. hairs; hairs 12 mm long; abaxial surface glabrous, densely nerved, dark; adaxial surface somewhat shiny, dark, striate, densely nerved; margins gla¬ brous or finely ciliate; inner ligule thick, finely ciliate; outer ligule a short, inconspicuous rim; blades re¬ flexed, deciduous, narrow, 4-5 cm long, 4 mm wide. Branch complement with 20-50 branches, these 30-45 cm long, 1 mm thick, glabrous, smooth, yellowish, 5-6-noded before the leaves, the nodes with fine, whitish pubescence below. Branch leaves 4-7 per complement: sheaths tight, glabrous, the overlapping margin finely ciliate toward base; oral 82 Novon setae fine, 5-7 mm long, minutely scabrous, crisped distally; inner ligule solid, finely ciliate, ca. 0.5 mm long; outer ligule a short inconspicuous rim; pseu¬ dopetiole flat, grayish yellow, slightly puberulent, 3 mm long; blades 4-9 cm long, 8-12 mm wide, L: W = 5-7.5, lanceolate, light green, finely pilose on abaxial surface, more densely pilose at the base, the hairs whitish; adaxial surface glabrous, 2-3 marginal nerves on one side with rows of fine prickle hairs; margins and the apex finely scabrous. Inflorescence terminal on leafy branches, spikelike, pectinate, se- cund, 3.5-4.5 cm long, with 16-20 spikelets. Ra- chis densely pilose. Pedicel pilose, at the base swollen into a lateral, globose, tuberous, naked projection. Spikelets 13.5-14 mm long, 3 mm wide, solitary, 1-flowered, bisexual, attenuate, fusiform. Glumes 2, unequal. Lower glume 1-nerved, carinate, mucro- nate, finely puberulent, 2 mm long, ca. 1 mm wide at the base. Upper glume 8-10 mm long, 3 mm wide, 7-9-nerved, broadly lanceolate, finely puber¬ ulent, the apex scabrous, aristulate, the awn ca. 1 mm long; adaxial surface shiny, dark-spotted. Lem¬ ma 8-9 mm long, 6 mm wide, broadly lanceolate, 21-23-nerved, the nerves anastomosing in the up¬ per part; abaxial surface puberulent, apically densely strigose near the margin, the hairs thick, short, dark; adaxial surface dark-spotted. Palea 11 mm long, ca. 5 mm wide, glabrous, 2-keeled, narrowly sulcate abaxially, 12-nerved, the nerves obliquely anasto¬ mosing; keels finely ciliate; adaxial surface shiny. Rachilla extension bristle-like, 7 mm long, with a minute rudiment. Lodicules not seen. Ovary elon¬ gate, style single, the stigmas 2, plumose, the sta¬ mens 3, the anthers 6 mm long, dark. Caryopsis 7.5 mm long, ca. 3 mm wide, ellipsoid, rostrate, grayish brown, finely striate, the embryo and the hilum scarcely manifest externally. Phenology. According to Filgueiras (1988 and pers. comm.) two populations of Merostachys fil- gueirasii bloomed and died gregariously in 1980- 1982, at the Reserva Ecologica do IBGE (Brazilian Institute of Geography and Statistics), and no re- growth was observed. A few months later, hundreds of seedlings could be seen in the area; these were collected by Filgueiras (Filgueiras 1021, IBGE). This collection contains seedlings in different stages of growth. The smallest specimen is 10 cm long, with 5 leaves and the caryopsis still attached. The seedlings of M. filgueirasii grew fast and occupied the bare ground left by the parent clump very quick¬ ly. By 1992, the two populations had completely re-established themselves at approximately the same site as the previous ones. No previous flowering is recorded. Distribution. Brazil. Distrito Federal: grounds of the Reserva Ecologica do IBGE, gallery forest in cerrado vegetation, 850-1340 m, and grounds of Catetinho. However, the population of the type spec¬ imen at Catetinho was completely wiped out because the site was cleared for tourist interests. The swollen lateral projections at the base of the pedicels are very peculiar; these are also found in M. plurijlora Munro ex Camus and in M. skvort- zovii (described later in this paper). In the former they are rather prominent, whereas in the latter they are small and are located on the inner side of the pedicel. This species was cited by Tarciso S. Filgueiras (1988) as Merostachys multiramea and is named for him, a friend and colleague, for his contributions to the knowledge of the Brazilian grasses. Paratypes. BRAZIL. Distrito Federal: Reserva Ecologica do IBGE, 15°57'S, 47“52'W, proximo a cerca W da Recor, 26 Sep. 1980 (fl), Heringer et al. 5812 (IBGE, K, US), 18 Jan. 1982 (fl), Filgueiras 955 (IBGE), 8 Jun. 1982 (seedlings), Filgueiras 1021 (IBGE); Reserva Ecologica do IBGE, mata do ribeirao Taquara, 20 Nov. 1990 (st), Filgueiras & Brochado 2009 (IBGE, SP). Merostachys glauca McClure & L. B. Smith. TYPE: Brazil. Estado de Santa Catarina: Pal- hoga, Piloes, mata, 200 m, 24 feb. 1956 (st), Reitz <6 Klein 2737 (holotype, US). Figure 4. Habit unknown. According to McClure & L. B. Smith (1967) only fragments of the midculm are known. Internodes ca. 30 cm long, 1.2-2.5 diam., glabrous and smooth, greenish yellow, whitish glaucous upward, the thinner internodes greenish, smooth, somewhat shiny, finely mottled with very small, dark green rounded spots. Culm walls 1-3 mm thick. Nodes salient, with retrorse white hairs below; supranodal ridge tubiform. Culm leaves de¬ ciduous: sheaths oblong, 29 cm long, 11 cm wide, striate, rather rough, with a trace of wax and a few scattered white hairs, the blades not seen. Branch complement with 30-60 branches, these 1 - 2 -noded before the leaves, glabrous. Branch leaves 5—7 per complement: sheaths tight, striate; inner ligule dark, 0.5 mm long, ciliate at the margin; outer ligule a salient rim; pseudopetiole 3-4 mm long, dark, gla¬ brous; oral setae 5 mm long, deciduous; blades nar¬ rowly linear-lanceolate, attenuate at the apex, 5- 13 cm long, 6-12(-17) mm wide, L:W = 8.3- 10.8; adaxial surface glabrous, with 1-2 marginal nerves on one side with rows of fine, sharp, sparse prickle hairs; abaxial surface glabrous except for densely pubescent area at the base beside the mid- Volume 5, Number 1 1995 Sendulsky Merostachys from Brazil 83 Figure 4. Merostachys glauca McClure & L. B. Smith. —a. Flowering branch complement. —b. Part of the inflorescence. —c. Spikelet, showing lower glume and lemma. —d. Spikelet, showing upper glume and lemma. —e. Lower glume. —f. Upper glume. —g. Rachilla segment. —h. Lemma, adaxial surface. —i. Lemma, abaxial surface. —j. Palea with rachilla and rudiment. —k. Upper part of the branch leaf sheath and base of the blade (abaxial surface). Based on Klein & Sousa Sob. 10480. nerve; margins sparsely scabrous. The epidermis of the sheath and the blade presents extremely small, rounded, flat papillae and some small scattered, pointed or unpointed prickle hairs. Inflorescences terminal on leafy branches, spikelike, pectinate, se- cund, 1.5-3 cm long, with 8-10 spikelets. Rachis and pedicels pilose. Spikelets 1-flowered, solitary, fusiform, 10 mm long, 2.5 mm wide. Glumes 2, unequal. Lower glume 2.5 mm long, 1 mm wide, 1 -nerved, finely puberulent, ciliate at the upper mar¬ gin. Upper glume lanceolate, apiculate, finely pu¬ berulent, 6.5-7 mm long, 3 mm wide, 10-nerved, 84 Novon some nerves anastomosing; adaxial surface shiny, brownish, with a few dark spots; margins ciliate. Lemma broadly lanceolate, 7.5-8 mm long, 4 mm wide, 9-nerved, adaxial surface shiny; abaxial sur¬ face densely pilose, the hairs ca. 0.5 mm long, silver- grayish. Palea 8 mm long, ca. 2 mm wide (folded), finely puberulent, 8-nerved, 2-keeled, broadly sili¬ cate abaxially, 8-nerved, the keels ciliate, the sulcus puberulent; adaxial surface shiny, light brown, in¬ conspicuously anastomosing. Rachilla extension about half the length of the palea, finely ciliate. Caryopsis not seen. Phenology. According to Smith et al. (1981) flowering collections are known from 1971-1973. Distribution. Brazil, State of Santa Catarina. Merostachys glauca was originally described from vegetative material only (McClure & L. B. Smith, 1967). Later, the same species, but in flowering stage, was described again (Smith et al., 1981), although without a Latin diagnosis for the floral part. Merostachys glauca is distinguished by an unusu¬ ally densely hirsute lemma and by the long, narrow branched leaf blades. The smooth intemodes, mot¬ tled with dark green rounded spots, the glaucous- whitish touch upward, the dark-spotted adaxial sur¬ face of the branch leaf sheath, and the size of the 1-flowered spikelet suggest that this species resem¬ bles M. caucaiana. Additional specimens examined. BRAZIL. Santa Catarina: Paulo Lopes, Bom Retiro, mata, 400 m, 13 Dec. 1972 (fl), Klein & Sousa Sob. 104HO f\Y); Joinville, Estrada Dona Francisca, mata, 15 Jan. 1974 (st), Klein & Klein 10996 (NY). Merostachys kleinii Sendulsky, sp. nov. TYPE: Brazil. Santa Catarina: Papanduva, E.R.F., 1 km ao sul da entrada para Papanduva, 800 m, 16 Jan. 1974 (fr), Klein & Klein 11008 (ho- lotype, RB; isotype, US). Figure 5. Habitus ignotus. Fragmenta culmorum internodiorum 2-3.5 cm diametro, fistulosa, scabra, maculis et striis atro-viridibus. Nodi pilis densis appressis, retrorsis, albis ad basin. Folia culmorum ignota. Rami in complemento 50-120. Foliorum ramorum laminae 3-9 cm longae, 7- 15 mm latae, ianceolatae. Inflorescentia spiciformis, se- cunda, 3-6 cm longa. Spiculae 2-flosculata, usque 16 mm longa, congestae, dense aggregatae, 2-seriatae; gluma inferior 4 mm longa; gluma superior 12-14 mm longa, 15-nervia, apiculata; flosculi bisexuales, late fusiformes, 10-12 mm longi; lemma 10 mm longum, 16-17-nervium; palea fusiformis, 11-12 mm longa, 12-14-nervia, late sulcata. Rachillae projectura plana, ca. 1 mm lata; pro- jectura flosculi superioris setiformis, rudimento ornata. Caryopsis late ovoidea, 6-7 mm longa, atro-lutea aut brunnea. Habit unknown. Fragments of the culm inter¬ nodes 2-3.5 cm diam., fistulose, scabrous, gray- greenish, with dark green spots and stripes. Culm walls of the middle intemodes 2-3.5 mm diam. Nodes salient, prenodal zone with dense, appressed, retrorse, whitish silky hairs below. Culm leaves not seen. Branch complement dense, with 50-120 branches, these 15-40 cm long, 1-1.5 mm thick, glabrous, smooth, grayish green, 2-3-noded before the leaves, nodes dark. Branch leaves 4-5 per com¬ plement: sheaths tight, glabrous, the margins gla¬ brous; oral setae fine, finely scabrous, deciduous; inner ligule short, dark, finely ciliate; outer ligule a salient rim; pseudopetiole dark, 2.5 mm long, 1 mm wide; blades 3-9 cm long, 7-15 mm wide, L: W = 4.2-6, lanceolate, apically attenuate, adaxially somewhat rough, 2-3 marginal nerves on one side with rows of fine prickle hairs, abaxially glabrous except for the densely pubescent base, the hairs long, villous, whitish, beside the midnerve. Inflores¬ cences terminal on leafy branches, spikelike, secund, congested, 3-6 cm long, 2 cm wide, with 28-32 densely agglomerated spikelets in double rows. Ra- chis pilose. Pedicels short, pilose. Spikelet always 2- flowered, 16 mm long, florets bisexual, broadly fu¬ siform, 10-12 mm long, 4 mm wide (with mature fruit), glabrous. Glumes 2, very unequal. Lower glume ca. 4 mm long, 1.5 mm wide, 1-nerved, puberulent, ciliate at the margins. Upper glume 12- 14 mm long, 5 mm wide, broadly lanceolate, apic- ulate, finely puberulent, 15-nerved, some nerves anastomosing; adaxial surface shiny, brownish, densely dark-spotted. Lemma 10 mm long, 6 mm wide, broadly lanceolate, puberulent, 16-17-nerved, some nerves anastomosing; adaxial surface shiny, brownish, densely dark-spotted. Palea fusiform, 11" 12 mm long, ca. 6 mm wide, glabrous, 12-14- nerved, broadly sulcate abaxially, the keels ciliate, some nerves anastomosing; adaxial surface shiny, golden. Rachilla extension between the florets flat, ca. 1 mm wide, finely puberulent, hairs whitish, bearing a fertile upper floret identical to the lower one. Rachilla extension of the upper floret bristle¬ like, with a small rudiment at the tip. Caryopsis broadly ovoid, basally and apically rounded, rostrate, dark yellow, 6-7 mm long, 4-4.5 mm wide, the embryo and the hilum scarcely manifest externally- Phenology. Flowering collections are known only from 1974. Distribution. Southern Brazil. Merostachys kleinii seems to be a robust plant, with thick culms. This species can be found in several herbaria, but complete vegetative collections are still Volume 5, Number 1 1995 Sendulsky Merostachys from Brazil 85 Figure 5. Merostachys kleinii Sendulsky. —a. Flowering branch complement. —b. Caryopsis, hilum view. —c. Caryopsis, embryo view. —d. Spikelet with two fertile florets and a rudiment. —e. Palea. —f. Lemma. —g. Upper glume. —h. Lower glume. —i. Upper portion of the branch leaf sheath and base of the blade (abaxial surface). Based on Klein & Klein 11008. not available. The species is distinct in its 2-flowered spikelet, the stout appearance of its branch com¬ plements, inflorescences and spikelets, and in having the largest caryopsis in the multiramea group. For many years this species was conceived as being a 2-flowered form of “multiramea. ” The true M. mul¬ tiramea Hackel already is a 2-flowered species. Mer¬ ostachys kleinii is named in honor of the late Rob¬ erto M. Klein, a distinguished botanist from Itajai, Santa Catarina, Brazil. 86 Novon Paratype. BRAZIL. Santa Catarina: 2 km a leste de Bocaina do Sid, 900 m, 22 Jan. 1974 (fr), Klein & Klein 11062 (SP). Merostachys magellanica Sendulsky, sp. nov. TYPE: Brazil. State of Sao Paulo: Municipio Anhembi, Fazenda Barreiro Rico, 22°40'S, 48°09'W, 510 m, 11 Jan. 1988 (fr), Reis de Magalhaes s.n. (holotype, SP 248339; iso¬ types, BLA, BR, IBGE, ICN, K, MO, NY, P, R, RB, UB, UEC, US, W, WIS). Figure 6. Culmi ca. 6-8 m alti, 12-14 mm diametro, fistulosi; internodia 60-80 cm longa; infera laevia, luteo-cinerea, striis maculisque viridi-fuscis minutis picta; supra cinereo- viridia, aculeis retrorsis munita; nodi prominentes, supra subterque nodum pilis lanatis densis brevibus vestita. Folia culmorum laminae reflexae, ca. 5-12 cm longae, 12-15 mm latae. Rami verticillati, in complemento 12-130. Folia ramorum laminae 8-17 cm longae, 1.5-2 cm latae, lanceolatae. Inflorescentia spiciformis, pectinata, imbri- cata, 6-8 cm longa, ramum foliorum terminans. Spiculae 11-14 mm longae, fusiformes ubi juvenes, ad maturita- tem ovoidae, apicibus attenuatis, duabus lineis albis par- allelis, longitudinalibus, carenis palearum formatis. Car- yopsis oblonga, ovoidea, asymmetries, lageniformis, ros- trata, 6-7 mm longa, castanea. Rhizomes sympodial, pachymorph. Culms woody, ca. 6-8 m tall, 12-14 mm diam., fistulose, erect; intemodes 60-80 cm long; lower intemodes grayish yellow, with minute dark green stripes and spots, smooth and shiny above the nodes; upper intemodes greenish gray, with small, retrorse prickle hairs; culm walls 2 mm thick; nodes salient, with a band of short, dense, woolly hairs above and below. Culm leaves deciduous: sheath 25-30 cm long, 5-7 cm wide, rounded and asymmetrical at the summit, fringed with fine, wavy hairs, these reddish brown, but white at the base, 5—6 mm long, diminishing in size downward along the margins of the sheath; abaxially olive-green, many-nerved, the nerves fine, light-colored, smooth, shiny; blade reflexed, 5-12 cm long, 12-15 mm wide, attenuating gradually to a fine point. Branch complement with 12-130 sub¬ equal branches, these 15-65 cm long, 1-2 mm thick, 2-3-noded before the leaves, yellowish, nearly glabrous and shiny, with green stripes, sometimes with minute retrorse prickle hairs, the base with a few densely strigose, deciduous scales, these 3-20 mm long; first intemode above the scales densely and retrorsely strigose. Foliage leaves 4-7 per com¬ plement. Sheath tight, glabrous, striate, the lower sheath aphyllous or with a rudimentary leaf and a few oral setae, the uppermost a little loose at the summit. Oral setae erect or crisped distally, 3-10 mm long, dark yellow or red-brown, slightly dimin¬ ishing in size downward along the upper margin of the sheath, then giving way to a fine row of short, silver cilia. Inner ligule a thick, dark yellow struc¬ ture, ca. 0.5 mm long, finely ciliate on the margin; outer ligule a salient, fight-colored rim. Pseudope¬ tiole dark, dorsiventrally compressed, adaxially sca- berulous and waxy, glabrous on abaxial surface. Blades linear-lanceolate, 8-17 cm long, 1.5-2 cm wide, L:W = 5.3-8.5 (the lowermost and the up¬ permost smaller, 2.5-6 cm long, 3-4 mm wide), the base slightly asymmetrical, rounded to cuneate, gradually attenuate toward a scabrous apex, sub- glabrous on both surfaces, the adaxial surface some¬ times finely hirsute near the base and also bearing 2-8 rows of hyaline, antrorse-strigose prickle hairs along the nerves near one margin; the nerves in¬ conspicuous on the adaxial surface, weakly salient on the abaxial surface. Inflorescences terminal on leafy branches, spikelike, falcate, congested, second, 6-8 cm long, with distinctive row of upper glumes when young or in early flowering stage, when mature and fruiting, 12-18 cm long, with loose, open, broadly spreading spikelets; rachis finely strigose, hairs white, sometimes with a rudimentary leaf emerging at the top of the oblique scar at the base of the inflorescence and subtending the lowermost spikelet. Spikelets 11-14 mm long, 1.8-2(-4) mm wide, 1 -flowered, slightly curved, attenuate, narrow¬ ly fusiform when immature, ovoid when mature, swollen at the base, attenuate at the apex, grayish olive-green, except for two parallel, longitudinal, white, evident lines formed by the keels of the palea. Pedicels short, densely hispid. Glumes 2, unequal. Lower glume 3-3.5 mm long, 1 mm wide, trian¬ gular, 1-nerved, slightly curved, shortly strigose at the apex, ca. '/ t the spikelet length, the mid-nerve spotted adaxially, the spots dark, purplish. Upper glume 5.5-7.5 mm long, 3 mm wide, lanceolate, apiculate, 9-nerved, shortly strigose, % the spikelet length, the adaxial surface shiny, with large, dark spots; when young, glumes forming a clear fine along the row of densely imbricated spikelets. Rachilla segment between the upper glume and the lemma 2.5 mm long, cup-shaped, 1.2 mm diam., finely strigose, the hairs white. Lemma 10-11.5 mm long, 5-5.5 mm wide, oblong, lanceolate, 14 - 15 -nerved, finely strigose abaxially, shiny, dark-spotted adaxi¬ ally, ciliate at the margins, embracing the palea (at maturity only at the base). Palea a little longer than the lemma, 10.5-12.5 mm long, 4 mm wide, curved, 8-12-nerved, with oblique transverse veinlets, the dorsal keels white, minutely ciliate, the adaxial sur¬ face shiny, dark-spotted. Prolongation of the rachilla bristle-like, nearly as long as the palea, pubescent near the minute rudiment. Lodicules 3, 3.3 mm long, finely ciliate at the apex, faintly 2 - 3 -nerved. Volume 5, Number 1 1995 Sendulsky Merostachys from Brazil 87 Figure 6. Merostachys magellanica Sendulsky. —a. Lodicules. —b. Lower glume. —c. Upper glume. —d. Detail of the abaxial surface of the upper glume. —e. Lemma. —f. Palea. —g. Spikelet, lemma side view. —h. Spikelet, palea side view. —i. Spikelet, lateral view. —j. Bisexual flower. —k. Palea. —1. Mature inflorescence, fruiting stage. —m. Base of the spikelet. —n. Oral setae. —o. Caryopsis hilum view. —p. Caryopsis, embryo view. —q. Caryopsis, lateral view. —r. Apex of a blade. —s. Base of a blade, adaxial surface. —t. Branch complement of the upper and slender part of a culm, with young inflorescences. —u. Mature spikelet, with parallel, white keels on the palea. All illustrations based on J. G. Kuhlmann s.n. (US 1255439) except I, o, p, q, and u, which are based on Reis de Magalhaes s.n. (SP 248339). Stamens 3, the anthers 9 mm long. Ovary oblong, subcylindrical; style single; stigmas 2, long-fringed when mature. Caryopsis oblong, 6-7 mm long, (la- genoid) curved or asymmetrical in the dorsiventral plane, sulcate, apically long rostrate; pericarp gla¬ brous, of uniform thickness, light brown; embryo small, basal; hilum linear, but the embryotegium weakly manifested externally. 88 Novon Phenology. Merostachys magellanica has been collected in its fertile state in 1891, 1921-1922, 1956, and 1988-1990. These collections suggest a flowering cycle of 30-34 years. Distribution. Brazil, in the States of Rio de Ja¬ neiro and Sao Paulo, from sea level to ca. 700 m. The first known collection of Merostachys ma¬ gellanica was made in Peruibe, Sao Paulo, in 1891, followed by the collections in 1921 and 1922 in Rio de Janeiro. Mature caryopses were collected in 1956 by Jose Carlos Reis de Magalhaes on his ranch, “Barreiro Rico,” in the State of Sao Paulo (this is, in fact, a well-kept private forest reserve). Werner Bockermann, a zoologist at the Sao Paulo Zoological Garden, has studied these caryopses as a food supply for birds in the reserve. When Merostachys magellanica flowered again in 1988 and set seed in quantity, a large collection was made by Reis de Magalhaes and sent to the Institute of Botany in Sao Paulo, together with ac¬ curate data and observations concerning the habitat: soil analysis, rainfall, climate, behavior of the bam¬ boos, etc. According to Reis de Magalhaes, the poor¬ est soil was exactly in the area where the species was collected. The adjacent forest area, 500 m away, with much better soil, has few or no bamboos. Thus, this new species shows a distinct preference for poor soil. Merostachys magellanica is characterized by the spikelets, which have two white, longitudinal lines formed by the keels of the palea, and by the general aspect of the congested inflorescences when young, which present upper glumes in a distinctive row. When mature, the inflorescences are twice as long as wide, with loose, open, broadly spreading spikelets and have light brown, dorsiventrally asymmetrical caryopses. Merostachys magellanica is named for Jose Car¬ los Reis de Magalhaes, collector of the type collec¬ tion, who in his respect for and love of nature knows few equals. Paratypes. BRAZIL. Rio de Janeiro: Botafogo, Mundo Novo, 11 Nov. 1921 (early fl), J. G. Kuhlmann s.n. (RB 79198, US 1447816k Corcovado, Morro do Mundo Novo, Oct. 1922 (fl), J. G. Kuhlmann s.n. (US 1255439); Reserva Florestal da Fabrica Alian^a, Sep. 1922 (fl-abnormal), J. G. Kuhlmann s.n. (RB 3852, US 1447817). Sao Paulo: Peruibe, 1891 (fl), Loefgren & Edwall s.n. (US 1021511); Municipio Itatins, Iguape, Esta^ao Ecologica Jureia, 19 Nov. 1987 (fl), Catharino 1334 (SP); Municipio Anhembi, Fazenda Barreiro Rico, 21 fev. 1956 (fr), Reis de Magalhaes s.n. (SP 248383); Cananeia, Out. 1956 (fr), Pereira s.n. (SP 169270); Parque Estadual da Ilha do Cardoso, 8 Dec. 1987 (fl), Kirizauia 1984 (SP); Ilha do Cardoso, estrada para tur- bina, 3 Dec. 1990 (fl), Barros 2024 (SP); Ilha do Cardoso, Cambriu, 23 Aug. 1988 (late fr), Rossi et al. 486 (SP). Merostachys multiramea Hackel. In Feddes Re- pert. Nov. Spec. Regni Veg. 7: 326. 1909. TYPE: Brazil. Rio Grande do Sul: Municipio Rio Pardo, Fazenda Itacolomi, elev. 70 m, Apr. 1906 (fl), Jurgens s.n. (label G-308) (holotype, W). Figure 7. Merostachys anomala Dutra, Revista Sudamer. Bot. 5: 151. 1938. Fig. 2, p. 149, under M. burchellii Munro emend. Dutra (figures misplaced). Syn. nov. TYPE: Brazil. Rio Grande do Sul: Sao Leopoldo, Fazenda da Pedreira, 6 Oct. 1906 (fl), Dutra 518 (holotype, BLA; isotype, US). Culms woody, fistulose, terete, yellowish gray, glabrous, smooth, but densely pilose near the nodes, 12 m long, according to C. Jurgens s.n. (label G- 308); pendulous above, 6-9 m high, 3-4 cm diam. below, according to Dutra (1938); nodes prominent, glabrous, densely pilose above, pilose and also with a row of white, dense, brushlike, retrorse hairs below. Culm leaves of an incompletely developed shoot (see discussion): sheaths 20-25 cm long, 5-6 cm wide, symmetrical, papery, opaque, rugose adaxially, fringed with dense, wavy, reddish hairs at the sum¬ mit, diminishing in size downward along the upper margin; inner ligule a thick membrane, 1-2 mm long, opaque; outer ligule a salient rim; blades lan¬ ceolate, asymmetrical, ca. 7 cm long, 1 cm wide, gradually attenuate to a very fine point. Branch complement with 30-70 unequal leafy branches, these 5-50 cm long, 1-1.2 mm thick, glabrous, smooth, grayish green, 2-3-noded before the leaves; nodes prominent, dark, the base densely covered with small, deciduous, apiculate, glabrous scales. Foliage leaves 3-8 per complement, the leaves of the flowering branches smaller than those of the sterile ones. Sheaths tight, glabrous, striate, the mar¬ gins finely ciliate or glabrous. Oral setae 3-4 mm long, straight, white, extremely deciduous. Inner ligule papery, finely ciliate on the margin, 0.5 mm long. Outer ligule a salient rim. Pseudopetiole gla¬ brous, dark, flat, 2.5-4 mm long. Blades 3-20 cm long, 5-20 mm wide, lanceolate or narrowly lan¬ ceolate, attenuate to a very narrow, filiform point, the base rounded to cuneate, slightly asymmetrical, subglabrous on both surfaces except for a small area on the abaxial surface near the base, bearing short, white, dense hairs mixed with wax and for 2-3 lateral nerves on the adaxial surface near one mar¬ gin, bearing parallel rows of antrorse, strigose, hy¬ aline prickle hairs. Inflorescence 5—7(—9) cm long, straight or subfalcate, secund, terminal on a leafy branch, usually fully exserted from the uppermost sheath when mature. Peduncle of the inflorescence and the rachis densely strigose, the hairs white. Volume 5, Number 1 1995 Sendulsky Merostachys from Brazil 89 Figure 7. Merostachys multiramea Hackel. —a. Part of a branch complement with some young inflorescences, —a,. Mature inflorescence. —b. Part of the spike. —c. Lower glume. —d. Upper glume. —e. Lemma. —f. Palea, abaxial surface. —g. Palea, adaxial surface. —h. Spikelet, glumes removed. —i. Bisexual flower. — j. Lodicules. — k. Base of a leaf, adaxial surface. —1. Caryopsis, hilum view. —m. Caryopsis, lateral view. —n. Caryopsis, embryo view. —o. Base of a leaf, adaxial surface, showing ligule and a few oral setae. —p. Culm leaf. All illustrations based on Dutra 518 , except a,, which is based on Dutra 517. Spikelets 2-flowered, 11-12 mm long, grayish green, with wrinkled surface, both florets bisexual, 7-9 mm long, 3 mm wide, laterally strongly compressed when immature, prominently keeled, imbricate at an angle of 45°. Pedicel ca. 1 mm long, densely hispid. Glumes 2, unequal. Lower glume 2.5-3 mm long, 1 mm wide, lanceolate, 1-nerved, the abaxial surface pu- berulent, the margins densely ciliate. Upper glume 90 Novon 7- 9 mm long, ca. 5 mm wide, half as long as upper floret, carinate, apiculate, asymmetrical, 9-nerved, the nerves anastomosing; abaxial surface finely hir- sute-appressed, the margins finely ciliate. Rachilla segment between the lower and the upper florets 2- 3.5 mm long, flattened, narrow below, dilated at the apex, finely appressed-pubescent. Lemma 8 mm long, 5 mm wide, carinate, adaxially slightly dark-spotted, 9-11-nerved. Palea 8 mm long, curved, bicarinate, broadly canaliculate between the finely ciliate keels, 8- nerved, the veinlets anastomosing in the upper part. Prolongation of the rachilla (on the upper floret only) bristle-like, flattened, half as long as the palea or a little over, bearing a minute rudiment. Lodicules 3, membranous, ca. 0.7 mm long, ciliate at the apex. Stamens 3, the anthers 4-5.5 mm long. Ovary oblong, subcylindrical; style single; stigmas 2, plu¬ mose. Caryopsis 6-7 mm long, 3 mm wide, conical, attenuate at the apex, rostrate, dark brown, nearly black, shiny; embryo small, basal; hilum linear, but the sulcus and the embryo weakly manifested ex¬ ternally. Phenology. Flowering collections are known from 1876 (Dutra, 1938), 1906-1907, 1914-1915, 1937-1939, and 1974-1978. Distribution. Southern Brazil. The description of the culm leaves of Merosta- chys multiramea is based on Prance et al. 6887, the only collection in which these leaves are present. This material is not very typical of the species; together with Gibbs et al. 6672 and Ihering s.n. (NY, SP 10315) it makes up a group of three similar specimens, all collected in the State of Sao Paulo. If these specimens can be admitted as good M. multiramea, the description of the culm leaves is valid. The only difference between these collections and the others is that the spikelets are not wrinkled and the short hairs that cover the spikelets are black and not so worn out and colorless as on the spikelets of the specimens from the State of Rio Grande do Sul. Additional specimens examined. BRAZIL. Parana: Ipiranga, Dusen 14367, 5 Jan. 1914 (fl) in US sheet, and 15 Jan. 1915 (fl) in WIS sheet. Rio Grande do Sul: Valle do Rio Taquari, Sao Leopoldo, 1907 (fr), Dutra s.n. (NY, SP 10313, US 1503502); Suburbios, Oct. 1906 (fl), Dutra 151 (RB); Bom Jesus, Fazenda Carauna, 1907 (fl and fr), Dutra 517 (BLA); Sao Leopoldo, Quilombo, Nov. 1937 (fl), Dutra 1634, (ICN); Panambi, 4 Oct. 1970 (fl), Pott s.n. (BLA 8363). Sao Paulo: Reserva Carlos Boteiho, entre S. Miguel Arcanjo e Sete Barras, elev. ca. 800 m, 31 Jan. 1978 (fl), Prance et al. 6887 (RB, US); Rodovia Jaquia a Piedade, 29 Oct. 1977 (fl), Gibbs et al. 6672 (SP); Sao Bernardo, Oct. 1914 (fl), Ihering s.n. (NY, SP 10315). Merostachys pilifera Sendulsky, sp. nov. TYPE: Brazil. Rio Grande do Sul: Mun. Sao Francisco de Paula, Primeiro Distrito, Sao Francisco de Paula, Morro da Pera (uppermost part of Serra Geral), 29°27'S 50°35'W, alt. 970 m, 11 May 1972 (fl), Soderstrom & Sucre 1987 (holotype, RB). Figure 8. Culmi usque 20 m longi, fistulosi ad extremitates solidi. Culmi internodia dense pilosa supra et infra nodos, pilis translucidis, appressis, ca. 1 nun longis. Folia culmorum ignotae. Crista supranodalis tubiformis. Rami in comple- mento 50 ad extremitates culmorum. Foliorum ramorum laminae lanceolatae, 2-8 cm longae, 4-12 mm latae. Inflorescentia spiciformis, pectinata. Spiculae 1-flosculata, bisexualis, late elliptica, 13-14 nun longa; gluma infera 2.5 mm longa; gluma supera 9 mm longa; 11-nervia; lemma 10 mm longum, 15-nervium; palea fusiformis, 11 mm longa, 10-12-nervia, anguste sulcata. Caryopsis non visa. Woody, according to the collectors’ note, culms erect, up to 20 m tall, 5 cm diam., closely clumped and forming a thicket; internodes fistulose, 60-65 cm long at base, with thin walls, becoming shorter, successively thinner and solid above; the tips whip¬ like, arching. New culm intemodes densely pilose above and below the nodes, the hairs sericeous, translucent, appressed, thick, ca. 1 mm long, these substituted in some occasionally hairless parts of the intemode areas by dense, sharp, retrorse prickle hairs with globose, yellowish bases, together with some scattered sericeous hairs. The old thick culms still display prickles and areas with worn and dam¬ aged hairs. Nodes salient, supranodal ridge tubiform, especially evident in new culms, prenodal zone with dense, appressed, retrorse hyaline hairs. Culm leaves not seen. Branch complement from tip of the culm (intemode 5 mm diam.) with 50 branches, these 15-20 cm long, ca. 0.5 mm thick, glabrous, smooth, 1- 2-noded before the leaves. Branch leaves 2-3 per complement: sheaths tight, finely puberulent; the overlapping margin glabrous; adaxial surface shiny, golden, dark-spotted; oral setae fine, ca. 5- 6 mm long, straight; inner ligule short, finely ciliate; outer ligule an inconspicuous rim; pseudopetiole dark, finely puberulent, 2-2.5 mm long; blades lanceolate, 2- 8 cm long, 4-12 mm wide, L:W = 6.2-6.6, glabrous, the abaxial surface with a very few fine hairs. Inflorescences terminal on leafy branches, 2- 3.5 cm long, spikelike, pectinate. Rachis and ped¬ icels pubescent. Spikelets solitary, 1-flowered, bi¬ sexual, broadly ellipsoid, somewhat inflated, 13-14 mm long, 4 mm wide. Glumes 2, unequal. Lower glume 2.5 mm long, ca. 1 mm wide, shortly awned, 1-nerved. Upper glume broadly lanceolate, 9 mm long, 4 mm wide, shortly awned, 11 -nerved, some nerves anastomosing, the adaxial surface dark-spot- Volume 5, Number 1 1995 Sendulsky Merostachys from Brazil 91 Figure 8. Merostachys pilifera Sendulsky. —a. Flowering branch complement. —b. Lodicules. —c. Hermaphroditic flower. —d. Spikelet. —e. Apical portion of the inflorescence. —f. Culm hairs. —h. Lower glume. —i. Upper glume. —j. Lemma. —k. Palea. —1. Upper portion of the branch leaf sheath and base of the blade (abaxial surface). Based on Soderstrom & Sucre 1987. ted. Lemma very broadly lanceolate, deeply con¬ cave, enclosing a palea, 10 mm long, 7 mm wide, 15-nerved, finely puberulent abaxially, the adaxial surface dark-spotted, the margins densely ciliate upward. Palea fusiform, 11 mm long, 2.5 mm wide (folded), 10-12-nerved, 2-keeled, narrowly sulcate, the keels finely ciliate, abaxially puberulent at the apex and along the sulcus; some nerves anastomos¬ ing; adaxial surface shiny. Lodicules 3, subequal, ca. 2.5 mm long, finely ciliate. Stamens 3, the 92 Novon anthers dark, ca. 10 nun long. Ovary elongate, style single, stigmas 2. Caryopsis not seen. Phenology. The only known specimen of this species is the holotype, collected in 1972. Distribution. Known only from the type locality. Merostachys pilifera is known only from the type locality in the State of Rio Grande do Sul. The description presented above was based solely on Soderstrom & Sucre 1897, 1972. The label states that the entire colony of this bamboo flowered after this date and is represented by Soderstrom 2040, 2 Feb. 1973 (US). Merostachys pilifera is distin¬ guished by its pilose culm internodes, the character from which the name is derived. It is somewhat similar to M. abadiana in the details of the spikelet structure. Merostachys scandens Sendulsky, sp. nov. TYPE: Brazil. State of Sao Paulo: Municipio de Sao Paulo, 10 km S of center of city of Sao Paulo, in grounds of the Instituto de Botanica, Parque do Estado, 6 Dec. 1973 (fl), Sendulsky 1319 (holotype, SP; isotypes, BLA, BR, IBGE, ICN, K, MO, NY, P, R, RB, UB, UEC, US, W, WIS). Figure 9. Culmi ca. 6 m longi, 1-3 cm diametro; internodia fistulosa, subtiliter rugulosa; nodi pilis densis, appressis, retrorsis. Foliorum culmorum laminae angustae, 8-12 cm longae, 4-5 mm latae. Rami in complemento 40-60(- 100). Foliorum ramorum laminae lanceolatae, glauco- virides, 3.5-10 cm longae, 8-14 mm latae. Inflorescen- tiae spiciformes, 5-6 cm longae, ramorum foliorum ter- minantes. Spiculae ellipticae, 1-flosculatae, 13-14 mm longae, distichae, lateraliter dispositae; gluma infera 4.5- 5 mm longa; gluma supera 9-10 mm longa, 15-nervia; lemma 11 mm longum, 19—23-nervium, ad marginem superiorem strigosum; palea fusiformis, 11 mm longa, 12- nervia, anguste sulcata. Caryopsis ovoidea ad apicem sub¬ tiliter attenuata, 5.5-6.5 mm longa, cinereo-lutea. Caespitose, woody bamboo. Culms ca. 6 m long, 1-3 cm diam., flexuous, with long, attenuate, threadlike, pendent or clambering tips. Internode fistulose, finely rugose, dark gray, with dull green stripes. Culm walls ca. 2 mm thick. Nodes salient, prenodal zone with dense appressed whitish, retrorse hairs. Culm leaves deciduous: sheaths 15-25 cm long, 4-8 cm wide, narrowing toward the summit, fringed at the apex with 1-cm-long, golden reddish, distally crisped hairs; abaxial surface glabrous, densely nerved; adaxial surface somewhat shiny or dark, apically tessellate; margins glabrous; inner lig- ule 1 mm long, finely ciliolate; outer ligule a salient rim; blades reflexed, deciduous, narrow, 8-12 cm long, 4—5 mm wide, attenuate, forming a fine point. Branch complement with 40-60(-100) branches. these 30-60 cm long, 1-1.5 mm thick, grayish, glabrous or puberulent, 2-3-noded before the leaves, nodes glabrous. Branch leaves 5-6 per complement: sheaths tight, glabrous, the intercostal zone finely transversely rugose, the overlapping margin gla¬ brous; oral setae fine, deciduous, ca. 3 mm long; inner ligule ca. 0.8 mm long, finely ciliolate; outer ligule a salient rim; pseudopetiole flat, slightly pu¬ berulent, 2.5 mm long; blades 3.5-10 cm long, 8- 14 mm wide, L: W = 4.4-7.1, lanceolate, glaucous- green, adaxially glabrous, 2-3 marginal nerves on one side with rows of very fine prickle hairs; abaxial surface glabrous except for finely strigose base, the margins and the apex very finely scabrous. Inflo¬ rescences terminal on leafy branches, spikelike, 5- 6 cm long, with 17-18 spikelets, these distichously arranged, distinctly separate and laterally outspread along the narrow, obscurely triquetrous, slightly sin¬ uous, densely pilose rachis or rarely pectinate. Spike- lets solitary, 1-flowered, bisexual, ellipsoid, 13-14 mm long, 3 mm wide. Glumes 2, unequal. Lower glume 1 -nerved, carinate, attenuate toward the apex, finely puberulent, 4.5-5 mm long, ca. 1 mm wide at the base. Upper glume 9-10 mm long, 4 mm wide, broadly lanceolate, carinate, finely puberulent, 15-nerved, some nerves anastomosing, the apex sca¬ brous, aristulate, ca. 1 mm long; adaxial surface shiny, dark-spotted; margins finely ciliate. Lemma 11 mm long, 5-5.5 mm wide, lanceolate, 19-23- nerved, the nerves anastomosing; abaxial surface somewhat strigose, the hairs appressed, longer and denser toward the upper margins, these densely ciliate; adaxial surface shiny, dark-spotted. Palea fusiform (folded), 11 mm long, ca. 4 mm wide, glabrous, opaque, 2-keeled, narrowly sulcate abax- ially, 12-nerved, the nerves obliquely anastomosing in the upper part; keels finely ciliate, the hairs dark red; adaxial surface shiny. Rachilla extension bristle¬ like, 9 mm long, with a minute rudiment. Lodicules 3, subequal, 2.5 mm long, finely ciliate at the apex. Stamens 3, the anthers 12 mm long, dark. Ovary elongate, the style single, the stigmas 2, plumose; stigmatic papillae sharply pointed. Stamens 3, the anthers 12 mm long, dark. Caryopsis 5.5-6.5 mm long, ca. 3 mm wide, ovoid, apically slightly atten¬ uate, rostrate, grayish yellow, smooth, apically slightly attenuate, the embryo and the hilum scarcely man¬ ifested externally. Phenology. The flowering collections are known from 1941 and 1972-1974, thus suggesting a flow¬ ering cycle of 31-33 years. Distribution. Southern Brazil. Merostachys scandens was in bloom at the same time as M. skvortzovii (described later in this paper) Volume 5, Number 1 1995 Sendulsky Merostachys from Brazil 93 Figure 9. Merostachys scandens Sendulsky. —a. Flowering branch complement. —b. Gynoecium with two lodicules. —c. Stigmatic papillae. —d. Spikelet in bloom (flowering cycle 1973). —e. Lodicules. —f. Apical part of the inflorescence. —g. Culm leaf. —h. Caryopsis, hilum view. —i. Caryopsis, embryo view. —j. Lower glume. —k. Upper glume. —1. Lemma. —m. Palea. —n. Upper portion of the branch leaf sheath and base of the blade (abaxial surface). Based on Sendulsky 1319. and in the same forest of the Biological Reserve, on the grounds of the Institute de Botanica, Sao Paulo. The plants did not form dense clumps, but were slender and leaning on vegetation. The glaucous- green leaves are distinct from the green leaves of M. skvortzovii. Because the plant was deep in the shaded forest, the visiting insects were not observed (see the discussion in M. skvortzovii). The habit of the inflorescence, its widely separated distichous spreading spikelets, is of some interest. This orga- 94 Novon nization of the inflorescence is quite rare in the genus and contradicts the significance of the name Mer- ostachys, formed from the Greek meros, part (par¬ tial or incomplete), and stachys, spike, alluding to the secund (one-sided) profile of the inflorescence (McClure, 1973). In M. scandens the inflorescence may sometimes also be pectinate, when for some unknown reason the very narrow rachis bends lon¬ gitudinally over the ventral surface, bringing to¬ gether its margins and the spikelets of two opposite rows. Paratypes. BRAZIL. Sao Paulo: Municipio de Sao Paulo, 10 km S of center of city of Sao Paulo, in grounds of the Instituto de Botanica, Parque do Estado, 6 Dec. 1973 (fl), Sendulsky 1317 (SP), 6 Dec. 1973 (fl), Sen- dulsky 1320 (SP, US), 17 Dec. 1973 (fl), Sendulsky 1323 (SP, US); Reserva Biologica, Parque Estadual das Fontes do Ipiranga, 8 Mar. 1974 (fr), Sendulsky 1365 (SP); Sao Paulo, nativa no Jardim Botanico, 1 Dec. 1941 (fl), O. Handro s.n. (SP sheet 79756). Merostachys skvortzovii Sendulsky, sp. nov. TYPE: Brazil. Estado de Sao Paulo: Sao Paulo, Parque do Estado e Jardim Botanico, 20 Oct. 1973, Skvortzov s.n. (holotype, SP 120954). Figure 10. Culmi 4-6 m longi, 2-3 cm diametro; internodia fis- tulosa, scabra; nodi subpilosi, pilis albis. Foliorum cul- morum laminae lanceolatae, 6-10 cm longae, 10 mm latae. Rami in complemento 150-200. Foliorum ramo- rum laminae lanceolatae, 3-11 cm longae, 6-16 mm latae. Inflorescentia spiciformis, pectinata, 1.5-3 cm lon- ga, ramorum foliorum terminans. Spiculae 10-12 mm longae, fusiformes, 1-flosculatae; gluma infera 1.5 mm longa; gluma supera 8-9 mm longa, 9-11-nervia, aris- tulata; lemma ca. 9 mm longum, 16-18-nervium, pub- erulum, ad apicem strigosum; palea ca. 10 mm longa, 13-14-nervia, anguste sulcata. Caryopsis ovoidea, ad ap¬ icem globosa, 5 mm longa et ca. 3 mm lata, cinereo-latea. Caespitose, woody bamboo, forming dense clumps. Culms 4-6 m long, 2-3 cm diam., erect except for the flexuose tips. Internodes fistulose, scabrous, with dark green stripes. Culm walls of the middle inter- nodes 1-1.5 mm thick. Nodes salient with a touch of whitish pubescence above and below, younger nodes with short, tubiform supranodal ridge. Culm leaves deciduous: sheath 15-20 cm long, 5-6 cm wide, auriculate, fringed at the apex with a few hairs, abaxial surface glabrous, densely nerved, dark; ad- axial surface opaque, dark, striate, tessellate above; inner ligule 1 mm long, finely ciliate at the margin, the cilia white; outer ligule a short inconspicuous rim; margins glabrous; blades lanceolate, reflexed, deciduous, 6-10 cm long, 10 mm wide. Branch complement with 150-200 slender branches, these 20-50 cm long, 1 mm thick, glabrous or finely puberulent, the intemodes ca. 4-6 cm long, 2-3- noded before the leaves. Branch leaves 5-7 per complement: sheaths tight, glabrous, striate; oral setae fine, 5-6 mm long; inner ligule a finely ciliolate membrane, ca. 1 mm long; outer ligule a short rim; pseudopetiole flat, 3 mm long, finely puberulent; blades lanceolate, 3-11 cm long, 6-16 mm wide, L:W = 5-6.9, adaxially glabrous, slightly pilose abaxially, densely pilose at the base; when old, gla¬ brous on both surfaces except at the base abaxially; adaxial surface with 3-4 marginal nerves on one side with rows of fine prickle hairs; margins finely scabrous. Inflorescences terminal on leafy branches, spikelike, pectinate, 1.5-3 cm long, with 16-18 spikelets. Rachis and pedicels pilose, some of the pedicels present at base on inner side small, swollen, whitish, rugose projections. Spikelets 10-12 mm long, 3 mm wide, solitary, 1-flowered, acuminate, fusiform. Glumes 2, unequal. Lower glume 1-nerved, carinate, acuminate, finely puberulent, 1.5 mm long, 0.8 mm wide at the base. Upper glume 8-9 mm long, 3.5 mm wide, broadly lanceolate, finely pu¬ berulent, 9-11 -nerved, the nerves anastomosing in the upper part, the apex scabrous, aristulate, ca. 1 mm long; adaxial surface shiny, dark-spotted. Upper glume sometimes subtending a rudimentary bud (vestige of a depauperate floret attached to rachilla segment). Lemma ca. 9 mm long, 6 mm wide, broad¬ ly lanceolate, 16-18-nerved, the nerves obliquely anastomosing in the upper part; abaxial surface pu¬ berulent, somewhat strigose in the upper part; ad¬ axial surface dark-spotted; margins ciliate. Palea ca. 10 mm long, 5 mm wide, glabrous, 13-14-nerved, 2-keeled, narrowly sulcate abaxially, the keels finely ciliate; adaxial surface shiny. Rachilla extension bris- tle-like, 8 mm long, with a minute rudiment. Lod- icules 3, subequal, 3.2 mm long. Ovary elongate; stigmas 2, plumose, the stigmatic papillae elongate, rounded at the tips; stamens 3, the anthers 6 mm long, dark. Caryopsis 5 mm long, ca. 3 mm wide, ovoid, apically rounded, rostrate, grayish yellow, smooth; hilum linear, the embryo hardly visible. Phenology. Flowering collections are known from 1941 and 1972-1974, suggesting a flowering cycle of 31-33 years. Distribution. Southern Brazil. Most collections of Merostachys consist of frag¬ mentary flowering branches. The vegetative parts of the plants are nearly always omitted. Conse¬ quently, the descriptions and measurements of the leaves, branch complements, culm and culm leaves, when available, are based on flowering material; usually they are gathered from the easily reachable upper branches of the plant, which are always small¬ er in size and therefore less representative of the Volume 5, Number 1 1995 Sendulsky Merostachys from Brazil 95 Figure 10. Merostachys skvortzovii Sendulsky. —a. Flowering branch complement. —b. Gynoecium with two lodicules. —c. Stigmatic papillae with pollen grains. —d. Bisexual flower with one lodicule. —e. Lodicules. —f. Spikelet in bloom (flowering cycle 1973). —g. Apical portion of the inflorescence. —h. Lower glume. —i. Palea. —j. Lemma. — k. Upper glume. —1. Rudimentary bud attached to rachilla segment. —m. Upper portion of the branch leaf sheath and base of the blade (abaxial surface). —n. Caryopsis, embryo view. —o. Caryopsis, hilum view. — p. Culm leaf. Based on the type specimen Skvortzov s.n. (SP 120954). plant as a whole. Deciduous and decaying culm leaves from basal or midculm nodes, originated dur¬ ing the initial growing period, are lost by the time of flowering. The plant stops producing any new vegetative organs and turns to flowering. When it is possible to find the vegetative, but complete col¬ lection, made from the same plant, the branch leaves are twice as large and the culm leaves are remark¬ ably larger. Unfortunately, it is very seldom that a correctly identified collection displays both vegeta- 96 Novon tive and flowering material. The vegetative mate¬ rials, Soderstrom 2033 (US 2909096) and Sod- erstrom 2035 (US 2909100), show large culm leaves, probably of M. skvortzovii, under the name M. multiramea. The culm leaf sheaths are 25-30 cm long, 18-19 cm wide, culm leaf blades 8.5 cm long, 25 mm wide. During the 1972-1974 flowering cycle, the flow¬ ers showed signs of being of interest to a variety of insects. This species is named in honor of the late Boris V. Skvortzov, Professor of Botany, Chinese Forest Academy, Harbin, Manchuria, who dedicated his later years to the study of the Brazilian flora, col¬ lector of the type material, and the beloved father of the author. Paratypes. BRAZIL. Parana: Municipio Palmas, Es¬ trada Palmas-Ponte Serrado, 5 Dec. 1971 (fl), Hatsch- bach et al. 28267 (NY, SP). Sao Paulo: Sao Paulo, Reserva Biologica, Parque Estadual das Fontes do Ipi- ranga, 7 fev. 1974 (fl), Sendulsky 1341 (SP, WIS); 10 km S of center of city of Sao Paulo, in grounds of the Instituto de Botanica, Parque do Estado, 10 Oct. 1973 (fl), Sendulsky 1314 (SP), 20 Oct. 1973 (fl), Sendulsky 1315 (SP, WIS), 6 Dec. 1973 (fl), Sendulsky 1318 { SP, WIS), 17 Dec. 1973 (fl), Sendulsky 1321 (SP, WIS), 17 Dec. 1973 (fl), Sendulsky 1322 (SP, WIS), 23 Dec. 1973, Sendulsky 1324 (SP, WIS); nativa no Jardim Botanico, 21 Oct. 1941 (fl), Gehrl. s.n. (SP 46002). Rio Grande do Sul: Municipio Colorado, km 251-252 on road between Carazinho and Bonambi, near Carazinho, 555 m, 30 Jan. 1973 (fl), Soderstrom 2029 (US); Depois de Caracol, ca. 10 km N de Canela, 17 dez. 1972, Porto, Girardi & Lindeman s.n. (ICN); Municipio tenente Por- tela, 3 km S of Derrobadas, 31 Jan. 1973, Soderstrom 2033 (US); Reserva Florestal do Turvo, 31 Jan. 1973, Soderstrom 2035 (US). Acknowledgments. This study has been sup¬ ported by the Conselho Nacional e Desenvolvimento Cientifico e Tecnologico (CNPq). I thank the staff of the U.S. National Herbarium, Smithsonian In¬ stitution, for assistance and for making their collec¬ tions available for study, Tarciso S. Filgueiras (IBGE) for the Latin diagnoses, A. G. Burman for interest and collaboration in the initial phase of this study, Fabio de Barros for helpful comments on the manu¬ script, and the anonymous reviewers for their critical readings of the manuscript. Special gratitude is ex¬ pressed to Marshall R. Crosby for helpful sugges¬ tions, and to Amy Scheuler McPherson (MO) and Lynn G. Clark, Department of Botany, Iowa State University, for kind assistance and revision of the original manuscript. The plates were drawn by the author. Literature Cited Dutra, J. 1938. Les Bambusees de Rio Grande du Sud. Revista Sudamer. Bot. 5(5/6): 145-152. Filgueiras, T. S. 1988. Bambus nativos do Distrito Federal, Brasil (Gramineae: Bambusoideae). Revista Brasil. Bot. 11: 47-66. Hackel, E. 1909. Gramineae Novae. VI. Repertorium Specierum Novarum Regni Vegetabilis 7: 311-327. Martius, C. F. P. von. 1840-1906. Flora Bras- iliensis. . .14 volumes (in 40), published in 130 fas¬ cicles. Munich, Vienna. [Bambuseae by J. C. Doell, 1880: volume 2(3): 161-219 + plates 44-56.] McClure, F. A. 1973. Genera of bamboos native to the New World (Gramineae: Bambusoideae). [Edited by T. R. Soderstrom.] Smithsonian Contr. Bot. 9: 1- 148. -& L. B. Smith. 1967. Bambuseae. Pp. 1-78 in P. R. Reitz (editor). Flora Ilustrada Catarinense, Pt. 1 Gramineas—Suplemento. Herbario “Barbosa Rodrigues,” Itajai, Santa Catarina, Brasil. Munro, W. 1868. A monograph of the Bambusaceae, including descriptions of all the species. Trans. Linn. Soc. London 26: 1-157, 6 plates. Smith, L. B., D. C. Wasshausen & R. M. Klein. 1981. Gramineas. In: P. R. Reitz (editor), Flora Ilustrada Catarinense, Vol. 1. Herbario “Barbosa Rodrigues,’ Itajai, Santa Catarina, Brasil. New Combinations in North American Schoenoplectus, Bolboschoenus, Isolepis, and Trichophorum (Cyperaceae) S. Galen Smith Department of Biology, University of Wisconsin-Whitewater, Whitewater, Wisconsin 53190, U.S.A. Abstract. Correct names are provided for Vol¬ ume 11 of the Flora of North America for taxa segregated from Scirpus sensu lato. Schoenoplectus validus is placed in synonymy under Sch. taber- naemontani. New combinations made herein are: Sch. acutus var. occidentalism Sch. x steinmetzii (pro sp.; = Sch. heterochaetus x tabernaemon- tani); Sch. pungens var. longispicatus; Sch. pun- gens var. badius (Scirpus americanus subsp. mon- ophyllus var. monophyllus, misapplied); Sch. x contortus (= Sch. americanus x pungens); Sch. hallii; Bolboschoenus glaucus (Lamarck) (Scirpus tuberosus, misapplied); Isolepis molesta; Tricho¬ phorum dementis; and T. clintonii. The following new combinations, synonymies, and typifications provide correct names in Schoeno¬ plectus, Bolboschoenus, Isolepis, and Trichopho¬ rum, all segregated from Scirpus sensu lato (Cy¬ peraceae) for the upcoming Volume 11 of the Flora of North America (Morin et al. [in prep.]). Descrip¬ tions and keys for identifying the taxa included here will be provided in the upcoming Flora. Types ex¬ amined for this study by A. E. Schuyler, who pro¬ vided me with information on them, are indicated by !Schuyler. Types that I have seen only as pho¬ tocopies or photographs provided by herbarium cu¬ rators or microfiches in libraries are indicated by Iphoto. 1. Schoenoplectus acutus (G. H. E. Muhlenberg ex J. Bigelow) A. Love & D. Love, Bull. Torrey Bot. Club 81: 33. 1954. Scirpus acutus G. H. E. Muhlenberg ex J. Bigelow, FI. Boston.: 15. 1814. Schoenoplectus lacustris L. subsp. acu¬ tus A. Love & D. Love, Taxon 30: 849. 1981. TYPE: U.S.A. Massachusetts: Fresh Pond [near Cambridge], July 1853, Herb. William Boott, s.n. (neotype, GH, selected here). I have selected as neotype the only known spec¬ imen of Scirpus lacustris as traditionally defined that was collected at Fresh Pond. No specimens from the Boston area that could have been seen by Bigelow have been located (Pamela White, GH, pers. comm.). Also, there is no direct evidence that Big¬ elow saw the Muhlenberg herbarium specimens; these add further confusion because each of the two sheets (labeled maculatus) in Folder 63 bears a mixed collection of seven or eight culms of Schoenoplectus acutus, Sch. tabernaemontani, and putative hy¬ brids as described by me (Smith, 1969). Schoenoplectus acutus is part of a difficult world¬ wide group that includes Sch. lacustris (L.) Palla, Sch. tabernaemontani (C. C. Gmelin) Palla, and Sch. heterochaetus (Chase) Sojak. Koyama (1962) treated the entire complex as the single species Scirpus lacustris and placed Scirpus acutus in syn¬ onymy under Scirpus lacustris subsp. glaucus (J. E. Smith) C. J. Hartman (see Sch. tabernaemontani below). Two varieties are here recognized for North America: a. Schoenoplectus acutus var. acutus. Scirpus lacustris L. var. condensatus C. H. Peck, Rep. (Annual) Regents, Univ. State New York, New York State Mus. 53: 853. 1900. Syn. nov. Scirpus val¬ idus Vahl var. condensatus (C. H. Peck) Beetle, Univ. Wyoming Pubis. 13: 6. 1948. TYPE: U.S.A. New York: [Cattaraugus Co.], Lime Lake, 5 and 20 Aug. 1899, F. E. Fenno 17 (holotype, NYS, ’Schuy¬ ler). Distribution. Eastern North America. b. Schoenoplectus acutus var. occidentalis (S. Watson) S. G. Smith, comb. nov. Basionym: Scirpus lacustris var. occidentalis S. Watson, Bot. California 2: 218. 1880. Scirpus occi¬ dentalis (S. Watson) Chase, Rhodora 6: 68. 1904. Scirpus acutus var. occidentalis (S. Watson) A. A. Beetle, Univ. Wyoming Pubis. 13: 5. 1948. TYPE: U.S.A. New Mexico: 1851-1852, C. Wright 1940 (mature achenes) (lectotype, NY, selected here; isolectotype, UC). Novon 5: 97-102. 1995. 98 Novon Scirpus malheurensis L. F. Henderson, Rhodora 32: 20. 1930. Scirpus acutus var. malheurensis (L. F. Hen¬ derson) A. A. Beetle, Univ. Wyoming Pubis. 13: 5. 1948. TYPE: U.S.A. Oregon: Harney Co., in water or on wet shores of Malheur Lake, 15 July 1927, L. F. Henderson 8655 (holotype, ORE, ISchuyler; isotype, CAS). Scirpus rubiginosus A. A. Beetle, Amer. J. Bot. 28: 697. 1941. Syn. nov. Schoenoplectus rubiginosus (A. A. Beetle) Sojak, Cas. Nar. Mus., Odd. Prir. 140: 127. 1972. TYPE: U.S.A. California: Marin Co., along tide flat N of Stinson Beach, 9 May 1941, Beetle <£ Constance 2686 (lectotype [imm.], UC, selected here, with specimen of Sch. acutus x cal- ifornicus on sheet designated by Beetle as holotype; isolectotypes [imm.], CAS, GH, NY—on sheets with specimen of Sch. acutus x californicus). [Other sheets from the type collection at CAS, F, KSC, RSA, and US are Sch. acutus x californicus .] Chase (1904: 68)included all North American Scir¬ pus acutus in Scirpus occidentalis (S. Watson) Chase. I have seen nine specimens that Watson annotated as Scirpus lacustris var. occidentalis. The two not cited by Chase (1904: 69) are: (1) the lectotype of Schoenoplectus acutus var. occidentalis, and (2) U.S.A. California: Sacramento Valley, Wilkes Ex¬ pedition [1842], (no collector given) #1788 (NY). Of all, only the Wright and Wilkes Expedition spec¬ imens bear achenes. I selected the Wright 1940 specimen as the lectotype because it is typical of Sch. acutus var. occidentalis throughout its range in its lenticular achenes and mostly bifid styles, whereas the mostly trigonous achenes and mostly trifid styles of the Wilkes Expedition specimen are known only from low elevations in California. The following collections from the type locality closely match the type collection of Scirpus rubi¬ ginosus: U.S.A. California: Marin Co., S end of Bolinas Lagoon ca. V 2 mi. N of village of Stinson Beach, fresh water seepages at edge of tidal salt marsh. Schoenoplectus acutus var. occidentalis (achenes present): 20 Aug. 1969, S. G. Smith 4713 (MO, PH, UC, US), 24 July 1973, S. G. Smith 5557 (GH, NY). Schoenoplectus acutus x cali¬ fornicus (achenes lacking): 20 Aug. 1969, S. G. Smith 4714 (PH, UC, US, UW), 24 July 1973, S. G. Smith 5556 (GH, MO, NY). For the lectotype of Scirpus rubiginosus I se¬ lected the specimen of Schoenoplectus acutus rath¬ er than the hybrid because: (1) Beetle’s protologue clearly indicates that his name applies to plants correctly placed in Sch. acutus rather than in the hybrid; and (2) Beetle described the achenes, which are lacking in the type specimens collected in May but are present in my collections of Sch. acutus , cited above, collected in July and August. 2. Schoenoplectus tabernaemontani (C. C. Gmelin) Palla, Verh. K. K. Zool.-Bot. Ges. Wien 38, Sitzb.: 49. 1888. Basionym: Scirpus ta¬ bernaemontani C. C. Gmelin, FI. Bad. 1: 101. April or May 1805. Scirpus lacustris L. subsp. tabernaemontani (C. C. Gmelin) Syme, Engl. Bot. ed. 10, 3: 63. 1870. Scirpus lacustris var. tabernaemontani (C. C. Gmelin) Doell, Rhein. FI. 165. 1943. Schoenoplectus lacus¬ tris subsp. tabernaemontani (C. C. Gmelin) Love & Love, Folia Geobot. Phytotax. 10: 275. 1975. TYPE: Europe: in Rheni vivinia (holo¬ type, KR, Iphoto, det. on sheet by Schuyler in 1974). Scirpus validus Vahl, Enum. PI. 2: 268. Oct. 1805 or later. Scirpus lacustris var. validus (Vahl) Kuken- thal, Feddes Repert. 23: 200. 1926 or 1927. Schoenoplectus validus (Vahl) A. Love & D. Love, Bull. Torrey Bot. Club 81: 33. 1954. Scirpus la¬ custris subsp. validus (Vahl) T. Koyama, Canad. J. Bot. 40: 927. 1962. Schoenoplectus lacustris (L.) Palla subsp. validus (Vahl) T. Koyama, Occas. Pap. Pauahi Bishop Mus. 29: 128. 1989. TYPE: e Car- ibaeis ded. Dr. Banks, Herb. Vahl s.n. (holotype, C, Iphoto, ISchuyler). Scirpus glaucus J. E. Smith, Engl. Bot. 33, pi. 2321. 1811. nom. illeg., not Scirpus glaucus Lamarck, Tabl. Encycl. 1: 142. 1791. Scirpus lacustris subsp. glaucus (J. E. Smith) C. J. Hartman, Sv. Norsk Exc.-Fl.: 10. 1846. Schoenoplectus lacustris (L.) Palla subsp. glaucus (J. E. Smith) Becherer, Feddes Repert. 25: 11. 1928. TYPE: Scotland. Galloway, on the sides of a saltmarsh on the W of Ardbigland, 1800, J. Mackay 6 (lectotype, LINN—Herb. Smith 105.25, selected here, Iphoto, det. by Clive Jermy (BM)). Scirpus tabernaemontani forma luxurious Miquel, Ann. Mus. Bot. Lugd.-Batav. 2: 143. 1865. Scirpus la¬ custris subsp. validus var. luxurious (Miquel) T. Koyama, Canad. J. Bot. 40: 927. 1962. Scirpus lacustris subsp. creber var. luxurians (Miquel) T. Koyamya, Phytologia 8: 458. 1963. Schoenoplec¬ tus validus subsp. luxurians (Miquel) Sojak, Cas. Nar. Mus., Odd. Pfir. 141: 62. 1972. TYPE: Japan. In ripa fluminis Jahaki, Gawa ins Nippon, #633 (lectotype, L, selected here—not seen, ISchuyler). Scirpus validus var. creber Fernald, Rhodora 45: 283. 1943. Schoenoplectus validus (M. Vahl) A. Love & D. Love var. creber (Fernald) A. Love & D. Love, Bull. Torrey Bot. Club 81: 33. 1954. Scirpus val¬ idus subsp. creber (Fernald) T. Koyama, Phytologia 8: 458. 1963. Schoenoplectus lacustris subsp. cre¬ ber (Fernald) A. Love & D. Love, Taxon 30: 849. 1981. TYPE: U.S.A. New York: Fisher’s Island, 10-15 Aug., H. St. John 2581 (holotype, GH). The typification of Scirpus glaucus J. E. Smith is important because it is the basionym for the above widely used names Scirpus lacustris subsp. glaucus and Schoenoplectus lacustris subsp. glaucus. Nich- Volume 5, Number 1 1995 Smith North American Cyperaceae 99 olas Turland (BM, pers. comm. 1994) provided the following information: Neither the collection by Bor- rer from Sussex, England, used to produce the il¬ lustration of Scirpus glaucus in English Botany, nor any material from Norfolk, England, as men¬ tioned in the protologue to plate 2321, appears to exist at BM. The only other specimen mentioned in the protologue is Mackay 6, which was examined for me by Clive Jermy (BM); it has bifid styles and dark brown papillae on the floral scales and is a typical example of northwestern European S. ta- bernaemontani. My studies and those of A. E. Schuyler (pers. comm.) indicate that Schoenoplectus tabernaemon- tani and Sch. validus are definitely conspecific. They differ by very few characteristics and inter¬ grade completely in both North America and Eur¬ asia. When Raymond (1957: 140) united these two species he expressed doubts: “1 have tentatively referred [a single specimen from Indo-China] to S. tabernaemontani. . . . S. tabernaemontani and S. validus are very closely related if not conspecific.” Koyama (1962) combined European S. tabernae¬ montani with the North American S. acutus Muhl¬ enberg ex Bigelow in S. lacustris L. subsp. glaucus (J. E. Smith) C. J. Hartman, placed Western Hemi¬ sphere and Hawaiian plants in S. lacustris L. subsp. validus (M. Vahl) T. Koyama var. validus, and placed Asian S. tabernaemontani in S. lacustris subsp. validus var. luxurious (Miquel) T. Koyama. Later, however, Koyama (1978: 209) treated S. tabernaemontani and S. validus as separate but doubtfully distinct species each occurring in both North America and Eurasia. The known distribution of Sch. tabernaemontani as here treated is the boreal to tropical zones of most of Eurasia and North America, the Caribbean, parts of Africa and tem¬ perate South America, the Hawaiian and other Pa¬ cific Islands, Australia, and New Zealand. Variation in Sch. tabernaemontani sensu lato is complex, and my attempts to clearly delimit infraspecific taxa have led me to the opinion that delimitation of infraspecific taxa is premature. Northwestern European plants have densely to sparsely reddish scabrous floral scales, especially near the bases of the spikelets; I have seen very similar plants from scattered localities in northern Canada, along both Atlantic and Pacific coasts of North America, and southern Africa. Scir¬ pus validus var. creber is distinguished only by its short floral scales, which partially expose the achenes, and intergrades with forms with longer scales over most of North America, and I have seen similar forms from northwestern Europe. 3. Schoenoplectus x steinmetzii (Fernald) S. G. Smith (pro sp.), comb. nov. Basionym: Scirpus steinmetzii Fernald, Rhodora 45: 286 and plate 766. 1943. TYPE: U.S.A. Maine: Penobscott County, bank of Passadumkeag Stream [ca. 2 mi. E of the village of], Passadumkeag, 12 Aug. 1937, F. H. Steinmetz 355 (holotype, GH; isotypes, MAINE, NY, US). I consider Schoenoplectus x steinmetzii to be a hybrid between Schoenoplectus heterochaetus (Chase) Sojak and Schoenoplectus tabernaemon¬ tani (C. C. Gmelin) Palla. Koyama (1962: 927) placed Scirpus steinmetzii in synonymy under S. lacustris subsp. validus (Vahl) T. Koyama var. validus and treated S. hetero¬ chaetus Chase as S. lacustris subsp. lacustris var. tenuiculmis (Sheldon) T. Koyama. Scirpus steinmetzii has previously been reported only from Passadumkeag Stream in the immediate vicinity of the type locality, where the plants in the several known colonies are very similar to each other and are intermediate between Schoenoplectus het¬ erochaetus and Sch. tabernaemontani in leaf, culm air cavity, floral scale, style, achene, and perigonial bristle characters. The protologue of S. steinmetzii erroneously describes the styles as bifid, the peri- gonial bristles as 0-2, and the achenes as plano¬ convex. The plants at the type locality produce very few achenes, in which the seeds are often aborted, and most of the pollen grains are abnormal when stained by cotton blue in lacto-phenol (confirmed by Arthur Haines, MAINE, pers. comm. 1994). Al¬ though Sch. tabernaemontani is common in the immediate vicinity of the type locality, the closest known stations of Sch. heterochaetus are each ca. 360 km distant (Lake Champlain and vicinity, where it is common and hybridizes with Sch. acutus, and eastern Massachusetts, where it may be extirpated). The region of the type locality, however, includes vast roadless areas with numerous wetlands that have been little explored botanically. Other specimens from Ontario, Quebec, Iowa, Massachusetts, New York, Minnesota, and Wiscon¬ sin are similar to specimens from the type popula¬ tion, but identification of most is uncertain because they lack achenes. The following collection from a population I have studied is nearly indistinguishable from specimens from the type population except that it produces many achenes containing fully formed seeds: U.S.A. Wisconsin: Rock Co., ca. 1.5 mi. SW of Lima Center along Wis. Hwy. 59, T4N, R14E, section 29, many clones in ca. 1 m of water. 100 Novon in undrained depression with Schoenoplectus het- erochaetus and Sch. tabernaemontani, 26 Aug. 1993, S. G. Smith 5208 ( GH, MAINE, NU, WIS). 4. Schoenoplectus pungens (M. Vahl) Palla, Bot. Jahrb. Verh. K. K. Zool.-Bot. Ges. Wien 38, Sitzb.: 49. 1888. Basionym: Scirpus pungens M. Vahl, Enum. PI. 2: 255. 1805. TYPE: Europe?: Erhart misit, herb. Vahl s.n. (holo- type, C, Iphoto, ISchuyler). Schoenoplectus pungens was long known by the name Scirpus americanus Persoon, misapplied (Schoenoplectus americanus (Persoon) Volkart ex Schinz & R. Keller). The type of Scirpus ameri¬ canus Persoon is conspecific with plants long known by the later synonym Scirpus olneyi A. Gray but correctly known as Sch. americanus (Schuyler, 1974). I agree with Koyama (1963) that North American and European Schoenoplectus pungens can be di¬ vided into three varieties, although the character¬ istics of these are somewhat different from those he gave, and in my opinion the differences are too small for the recognition of subspecies. 4a. Schoenoplectus pungens var. pungens. Synonymy and distribution as given by Koyama (1963: 1117) for Scirpus americanus Persoon subsp. americanus var. americanus. 4b. Schoenoplectus pungens var. longispi- catus (N. L. Britton) S. G. Smith, comb. nov. Basionym: Scirpus americanus Persoon var. longispicatus N. L. Britton, Trans. New York Acad. Sci. 11: 78. 1892. Scirpus longispi¬ catus (N. L. Britton) Smyth, Trans. Kansas Acad. Sci. 16: 163. 1899. Schoenoplectus americanus subsp. longispicatus (N. L. Brit¬ ton) Sojak, Cas. Nar. Mus., Odd. Prir. 141: 62. 1972. Scirpus pungens var. longispicatus (N. L. Britton) Cronquist, Intermountain Flora 6: 74. 1977. Schoenoplectus pungens subsp. longispicatus (N. L. Britton) A. Love & D. Love, Taxon 30: 849. 1981. TYPE: U.S.A. New Mexico: 1851 or 1852, C. Wright 1963 (lectotype, NY, selected here, ISchuyler; iso- lectotypes, PH, US, GH 2 sheets). Scirpus solispicatus Lunell, Amer. Midi. Naturalist 4: 230. 1915. TYPE: U.S.A. North Dakota: Benson Co., Butte, 18 Aug. 1907, J. Lunell s.n. (lectotype, MIN, selected here, ISchuyler). The lectotype sheet of Schoenoplectus pungens var. longispicatus , and the isolectotype sheets at US and GH, all bear mixed collections of Schoen¬ oplectus pungens and putative Sch. americanus x pungens hybrids; and sheet #28000 at GH also bears a specimen of Sch. americanus (which a note handwritten on the sheet suggests belongs to a col¬ lection from Nevada distributed by Wheeler in 1871). I selected the specimens of Sch. pungens as the lectotypes because they are typical of most western North American Sch. pungens and bear mature achenes, which are described in the original de¬ scription, whereas the putative hybrid specimens are intermediate between Sch. americanus and Sch. pungens and lack achenes. The other collection (Colorado: Platte R., Dr. Geo. Smith) that Britton cited in the protologue has not been located. Three other specimens of Scirpus solispicatus collected by Lunell at Butte in 1906 (GH, MIN, NY) are very similar to the lectotype of Scirpus solispicatus , which I chose because of its mature achenes with intact bristles. Schoenoplectus pungens var. longispicatus is composed of at least the North American plants included by Koyama (1963: 1118) in Scirpus amer¬ icanus Persoon subsp. americanus var. longispi¬ catus (N. L. Britton) T. Koyama. 4c. Schoenoplectus pungens (M. Vahl) Palla var. badius (K. Presl) S. G. Smith, comb, et stat. nov. Basionym: Scirpus badius K. Presl, Reliq. Haenk. 1: 193. 1828. TYPE: U.S.A. California: hab. ad Monte-Rey, Haenke s.n. (holotype, PR, ISchuyler). Schoenoplectus pungens var. badius is com¬ posed of at least the North American plants included in Scirpus americanus subsp. monophyllus (K. Presl) T. Koyama var. monophyllus (Koyama, 1963: 1118). My examination of the types and original de¬ scriptions indicate that Scirpus badius K. Presl rather than Scirpus monophyllus K. Presl is the correct basionym for a taxon including the North American Pacific Coastal plants of Schoenoplectus pungens. Koyama (1963) placed Scirpus badius in synonymy under Sch. americanus subsp. mon¬ ophyllus var. monophyllus , based on Scirpus mon¬ ophyllus K. Presl, Reliq. Haenk. 1: 193. 1828, TYPE: Peru: Montana Peruvia, Haenke s.n. (ho¬ lotype, PR, ISchuyler). All three specimens on the type sheet of S. monophyllus (photo in Koyama, 1963) are typical of Sch. americanus rather than Volume 5, Number 1 1995 Smith North American Cyperaceae 101 the Pacific Coastal North American Sch. pungens in their leaves, floral scales, achenes, and perigonial bristles. The styles are a mixture of bifid and trifid, which can occur in either species. The culms, how¬ ever, have more-or-less flat sides and are very firm, retaining their trigonous shape on pressing, which is fairly common in Sch. pungens but rare in Sch. americanus, in which the culm sides are typically deeply concave and readily flattened on pressing. Except for the style number, the pertinent parts of the original description of Scirpus monophyllus, “Colmo triquetro inferne monophyllo, . . . stylo bi- fido, . . . caryopsidem obovatum plano-convex- am . . .” agree with the type. Thus, the name Scir¬ pus monophyllus K. Presl applies to plants belong¬ ing to Sckoenoplectus americanus rather than to Sch. pungens. In the case of Scirpus badius, the type is a fine specimen with a rhizome and four complete shoots bearing inflorescences with some intact styles and mature achenes. It is an excellent example of the North American Pacific Coastal plants of Schoen- oplectus pungens. The original description of S. badius, which reads in part: “Colmo triquetro fol- iato, . . . stylo trifido, . . . Setae sex retrorsum sca- brae, caryopsidae una quarta parte breviores. Car- yopsis obovata trigona. . . ,” agrees well with the type. 5. Schoenoplectus x contortus (E. H. Eames) S. G. Smith, comb. nov. Basionym: Scirpus olneyi A. Gray var. contortus E. H. Eames, Rhodora 9: 220. 1907. Scirpus x contortus (E. H. Eames) T. Koyama, Canad. J. Bot. 41: 1123. 1963. TYPE: U.S.A. Connecticut: Mil¬ ford, plentiful in a brackish marsh at Pond Point, 28 Sep. 1907, E. H. Eames 5847 (ho- lotype, GH, ex herb. Eames; isotype, MO). 6. Schoenoplectus hallii (A. Gray) S. G. Smith, comb. nov. Basionym: Scirpus hallii A. Gray, Man. ed. 3, addend.: xcvii. 1862. Scirpus su- pinus L. var. hallii (A. Gray) A. Gray, Man. ed. 5: 563. 1867. Scirpus uninodis (Defile) Boissier var. hallii A. A. Beetle, Amer. J. Bot. 29: 656. 1942. TYPE: U.S.A. Illinois: Mason Co. or Menard Co., wet banks of pond, [Aug. 1860 or 1861], E. Hall [& D. Vasey & M. S. Bebb?] s.n. (holotype, GH). Raynal (1976: 141) placed Schoenoplectus hal¬ lii in synonymy under Sch. erectus (Poiret) Palla ex J. Raynal. The type locality, date, names of collectors, and number of collections are all in doubt, as indicated by differences among the protologue, Gray’s an¬ notation on the holotype, labels on the holotype and several possible isotypes (F, NY, US), and a letter from Gray [to Torrey?] at NY. 7. Bolboschoenus glaucus (Lamarck) S. G. Smith, comb. nov. Basionym: Scirpus glaucus La¬ marck, Tabl. Encycl. 1: 142. 1791. Scirpus maritimus L. [var.] glaucus (Lamarck) Nees in Wight, Contr. Bot. India 111. 1834. TYPE: Senegal, herb. Lamarck sheet 673/14, Rous¬ sillon s.n. (holotype, P, Iphoto, '.Schuyler; det. as Scirpus maritimus L. by J. Raynal in 1966). Examination of types and protologues indicates that Scirpus glaucus Lamarck is the correct name for the species to which Beetle (1942: 85) and Koyama (1962: 936) misapplied the later name Scirpus tuberosus Desfontaines, FI. Atlant. 1: 50. 1798; synonym: Scirpus maritimus L. [var.] tuber¬ osus (Desfontaines) Roemer & Schultes, Syst. Veg. 2: 139. 1817; Reigera maritimus [var.] tuberosus (Desfontaines) Opiz, Seznam: 83. 1852; Bolbo¬ schoenus tuberosus (Desfontaines) Hadac, Bull. Coll. Sci. Baghdad 6: 13. 1961. The type of S. tuberosus, [Algeria]: habitat in lacubus prope La Calle, Herbier de la Flore Atlantique donne au Museum, par M. Desfontaines, s.n. (holotype, P, Iphoto; det. as Scir¬ pus maritimus L. by A. E. Schuyler in 1966), is probably S. maritimus or a close relative in having most of the spikelets in a sessile mass, bifid and trifid styles, and lenticular achenes. The protologue, which reads in part: “spicufis congestis. . .sessifibus aut brevio pedunculatis,” agrees with the type. How¬ ever, the type of 5. glaucus is typical of the entity sometimes known as S. tuberosus in its achenes and trifid styles (Schuyler, pers. comm.) and its panic¬ ulate inflorescence with most of the spikelets on rays. The protologue of S. glaucus, which reads in part “umbella composita subpaniculata, spicufis pedicel- latis,” agrees with the type. Bolboschoenus glaucus is widely confused with Scirpus maritimus L. (Schoenoplectus maritimus (L.) Lye, Bolboschoenus maritimus (L.) Palla). Its achenes were described by Browning & Gordon- Gray (1993) under “Bolboschoenus maritimus en¬ tity 2.” It is widespread in the warmer parts of the Old World and sparingly introduced into North America. 8. Isolepis molesta (M. C. Johnston) S. G. Smith, comb. nov. Basionym: Scirpus molestus M. C. Johnston, Southw. Naturalist 9: 310. 1964. TYPE: U.S.A. Texas: Walker Co., Huntsville, 27 Apr. 1944,5. R. Warner 20,000 (holotype, TEX not seen, ISchuyler). 102 Novon 9. Trichophorum dementis (M. E. Jones) S. G. Smith, comb. nov. Basionym: Scirpus demen¬ tis M. E. Jones, Contr. W. Bot. 14: 21. 1912. TYPE: U.S.A. California: Rae Lake, King’s River, alpine, 20 July 1910, Mrs. Clements s.n. (lectotype, selected here, POM 111694, ISchuyler; isolectotypes, DS, POM 111695). Scirpus yosemitanus F. J. Smiley, Univ. Calif. Publ. Bot. 9: 108. 1921. TYPE: U.S.A. California: Yosemite National Park, Tuolumne Meadows, soda springs, 8 Aug. 1898, /. 11. Congdon s.n. (holotype, GH). 10. Trichophorum clintonii (A. Gray) S. G. Smith, comb. nov. Basionym: Scirpus clintonii A. Gray, Amer. J. Sci., ser. 2, 38: 290. 1864. TYPE: U.S.A. New York: between Buffalo and Williamsville, June 1864, G. W. Clinton s.n. (holotype, GH, ISchuyler; isotypes, NY, ISchuyler; US, not seen; K not seen, ISchuyler). Two possible isotypes at F bear insufficient data. Acknowledgments. Alfred E. Schuyler (PH) pro¬ vided much essential information and counsel over many years. Fred R. Barrie (MO), Jane Browning (NU), K. D. Gordon-Gray (NU), Hugh H. litis (WIS), Dan H. Nicolson (US), Anton A. Reznicek (MICH), Mark T. Strong (US), and many other colleagues generously gave help and advice. Nicholas Turland and Clive Jermy (BM), Alicia Lourteig (P), G. Thijsse (L), and Pamela White (GH) kindly provided infor¬ mation on types. Curators of the following herbaria made specimens available for study, and many of them were very helpful during my visits to their herbaria: ALA, ALU, ARIZ, ASC, BHO, CAN, CAS, CLEMS, DAV, F, FLAS, FTU, GA, GH, ISC, JEPS, K, KANU, KSC, LAF, LL, LSU, MAINE, MICH, MIN, MO, NEB, NFLD, NLU, NMW, NO, NY, OS, OSC, PH, PR, RM, RSA, SD, TAES, TEX, TRT, UC, UCSB, UNA, UNM, USCH, US, USF, UTC, WIS, WS, WTU. My work was greatly aided by a Smithsonian Institution research fellowship, by research grants from the Flora of North America Project, the Jepson Herbarium, and the University of Wisconsin, and by the facilities generously pro¬ vided since my retirement by the Department of Biology, University of Wisconsin-Whitewater. Literature Cited Beetle, A. A. 1942. Studies in the genus Scirpus L. IV. The section Bolboschoenus Palla. Amer. J. Bot. 42: 82-88. Browning, J. & K. D. Gordon-Gray. 1993. Studies in the Cyperaceae in southern Africa. 21: The taxo¬ nomic significance of the achene and its embryo in Bolboschoenus. S. African J. Bot. 59: 311-318. Chase, A. 1904. The North American allies of Scirpus lacustris. Rhodora 6: 65-71. Koyama, T. 1962. The genus Scirpus Linn. Some North American aphylloid species. Canad. J. Bot. 40: 913-937. -. 1963. The genus Scirpus Linn. Critical spe¬ cies in the section Pterolepis. Canad. J. Bot. 41: 1107-1131. -. 1978. Cyperaceae. In: Hui-Lin Liu et al. (editors), Flora of Taiwan. Vol. 5: 191-372. Epoch Publishing, Taipei, Taiwan. Morin, N. R., Convening Editor, and Flora of North America Editorial Committee. [In prep.] Flora of North America. Vol. 11. Oxford Univ. Press, New York. Raymond, M. 1957. Some new or critical Scirpus from Indo-China. Le Naturaliste Canadien 84: 111 149. Raynal, J. 1976. Notes Cyperologiques: 26. Le Genre Schoenoplectus II. L’amphicarpie et la sect. Supini. Adansonia, ser. 2, 16: 119-155. Schuyler, A. E. 1974. Typification and application of the names Scirpus americanus Pers., S. olneyi Gray, and S. pungens Vahl. Rhodora 76 (805): 51-52. Smith, S. G. 1969. Natural hybridization in the Scirpus lacustris complex in North Central United States. Pp. 175-199 in J. E. Gunckel (editor), Current Topics in Plant Science. Academic Press, New York and London. A New Nerine Species (Amaryllidaceae tribe Amaryllideae) from the Koup Karoo, South Africa D. A. Snijman Compton Herbarium, National Botanical Institute, Private Bag X7, Claremont 7735, South Africa Abstract. Nerine marincowitzii is a rare new species from the Koup Karoo, Cape Province, South Africa. Most closely related to the eastern Cape N. huttoniae Schonland, N. marincowitzii is distin¬ guished by hysteranthous leaves, a sigmoid scape that detaches at ground level when dry, flowers that turn brown at senescence, and a wind-dispersed in- fructescence. It grows on exposed seasonal washes and flowers at the end of the autumn rainfall season in May. Nerine Herbert (1820), comprising approxi¬ mately 22 species, is a horticulturally important southern African genus of Amaryllideae (Dahlgren et al., 1985). Of the ten genera in this mainly sub- Saharan African tribe, Nerine is most closely allied to Brunsvigia Heister (1755), sharing with it apo- morphic fasciculate stamens. Traub (1967) has sug¬ gested that the two genera should be united. Until strong phylogenetic evidence for this becomes avail¬ able, however, Nerine is retained as a distinct genus. Nerine is distinguished from Brunsvigia by some¬ what smaller flowers, mostly with undulate tepal margins, and distintegrating, globose, membranous capsules with bulging, exposed seeds (Dyer, 1976). Traub (1967) recognized four sections in Nerine. Section Laticomae Traub, the most widespread in southern Africa, comprises species with short, stout scapes whose length more or less approximates the width of the flower cluster. According to Obermeyer (1993) the section includes two species, N. laticoma (Ker-Gawler) Durand & Schinz and N. huttoniae Schonland. The newly described N. marincowitzii (Fig. 1) adds a further species to section Laticomae. Nerine marincowitzii was collected southeast of Leeu-Gamka in the Koup Karoo, Cape Province, widely disjunct from N. laticoma and N. huttoniae (Fig. 2). Unlike the highly seasonal summer rainfall region occupied by these two species, the semiarid area occupied by N. marincowitzii has an extended rainfall season, and although annual precipitation in the area is low and highly variable it benefits from summer and autumn rainfall (Zucchini & Adamson, 1984). Nerine marincowitzii is distinguished from N. laticoma and N. huttoniae by hysteranthous, nar¬ row leaves (3-4 mm wide), a scape that is elliptical rather than compressed in transverse section, and by plane-edged tepals. The new species is most sim¬ ilar to N. huttoniae and shares with it a many- flowered inflorescence (up to 40 flowers) and rigid, widely radiating pedicels, which bestow a hemi¬ spherical shape on the flower head. Nerine marin¬ cowitzii is unique in the genus in having an apo- morphic curved scape and tepals that turn brown and remain outspread with age. These specialized floral changes also occur in the closely allied Hessea Herbert and differ from the usual pattern of senes¬ cence in Amaryllideae, where the floral pigmentation becomes accentuated and the tepals ultimately col¬ lapse (Snijman, 1994). The wind-dispersed infructescence of Nerine marincowitzii is a further specialization not before known in Nerine. When dry, the scape detaches at ground level, the curvature of the scape confers a somewhat spherical configuration on the infructesc¬ ence, and the entire structure tumbles away in the wind. This derived mechanism, known as anemo- geochory and reported to occur in arid areas world¬ wide (Van der Pijl, 1982), is also well developed in Brunsvigia, Boophone Herbert, and Cybistetes Milne-Redhead & Schweickerdt in Amaryllideae (Snijman & Williamson, 1994). Nerine marincowitzii Snijman, sp. nov. TYPE: South Africa. Cape Province: 3222 (Beaufort West) Koup Karoo, farm Vredendal, SE of Leeu-Gamka, in washes along Kleinwaterval- rivier (CD), 19 May 1990, Snijman 1245 (ho- lotype, NBG; isotypes, MO, PRE). Figure 1. Inter species sectionis Laticomarum Traub inflores- centia multiflora (floribus usque 40) et pedicillis rigidis radiatim ad Nerine huttoniam Schonland accedens sed foliis hysteranthis, scapo sigmoideo, floribus brunneis ae- tate provecta et infructescentia anemodispersa ab ea re- cedens. Plants small, to 20 cm high. Bulb solitary, de¬ pressed-globose, 45-55 mm across, covered with Novon 5: 103-105. 1995. 104 Novon Figure 1. Nerine marincowitzii Snijman. —1. Inflorescence. —2. Mature leaves. —3. Mature bulb. —4. Flower at antbesis. 5. Anther after dehiscence. —6, 7. Dorsifixed anthers before dehiscence. —8. Mature seeds in disintegrating capsule. Scale bars: 1-3, 8 = 2 cm; 4 = 7 mm; 5-7 = 3 mm. Original painting by E. Ward-Hilhorst from Vlok s.n. Volume 5, Number 1 1995 Snijman Nerine marincowitzii 105 Figure 2. Known distribution of Nerine sect. Laticom- ae: N. laticoma (A), N. huttoniae (★), N. marincowitzii (•). thinly fibrous tunics, cream-colored and fleshy with¬ in, extended into a slender neck to 60 mm long. Foliage leaves 5-6, absent at anthesis, distichous, suberect to recurved, narrowly ligulate, tapering gradually distally, canaliculate, somewhat succulent, to 30 cm x 3-4 mm, bright green, glabrous. In¬ florescence 20-40-flowered, hemispherical, com¬ pact, 10-16 cm across; scape sigmoid, rigid, ellip¬ tical in cross section, 11-14 cm x 2.5-3.5 mm, reddish brown, breaking at ground level when dry; spathe valves 2, narrowly lanceolate, 35-55 x 3- 5 mm, papery, wine-red; bracteoles filiform, to 10 mm long; pedicels radiating widely, straight, terete, 40-75 mm long, shiny green, turning straw-colored and rigid in fruit. Perigone strongly zygomorphic, pink, initially with whitish throat, aging to brown, scentless; tepals shortly connate proximally, oblong- lanceolate, falcate and upturned at anthesis, 17-22 x 4-6.5 mm, plane; filaments slightly declinate, filiform, 13-17 mm long, expanded and connate proximally into a short tube to 0.25 mm long; an¬ thers dorsifixed, ca. 5 mm long and wine-red before opening; pollen whitish. Ovary dark wine-red; ovules 1 or rarely 2 per locule, absent from innermost flowers; style slender, declinate, ca. 20 mm long; nectar present around style base; stigma shortly trilobed. Capsule small, papery, loculicidally dehis¬ cent, rapidly outgrown by seeds. Seeds ellipsoidal, succulent, 6-7 mm across, wine-red; testa with sto¬ mata. Phenology. Flowering is limited to approximately three weeks in May depending on the advent of rain in autumn (probably until the end of April). The leaves appear in spring when growth is rapid and die back during a dry spell in summer. Distribution and habitat. Nerine marincowitzii is known from a single population about 40 kilo¬ meters southeast of Leeu-Gamka, near the Klein- watervalrivier, a tributary of the Gamka River. The population, comprising approximately 400 plants, is confined to sparsely vegetated, seasonal washes with a north aspect, where the bulbs are lodged in fight clay soil, covered with sharp blue-gray slate chips. Etymology. The specific epithet honors C. P. Marincowitz of the farm Kleinsleutelfontein, Koup Karoo, whose original knowledge and appreciation of this remarkable species led to its identification as new. Paratype. SOUTH AFRICA. Cape: 3222 (Beaufort West) Koup Karoo, farm Vredendal, SE of Leeu-Gamka (CD), 10 May 1992, Vlok s.n. (NBG). Acknowledgments. I thank Jan Vlok of Cape Nature Conservation and Museums and Colin Pat- erson-Jones for their assistance in the field, and Adela Romanowski of PRE and artist Ellaphie Ward- Hilhorst for the botanical illustration. Literature Cited Dahlgren, R. H. T., H. T. Clifford & P. F. Yeo. 1985. The Families of the Monocotyledons. Springer-Ver- lag, Berlin. Dyer, R. A. 1976. Amaryllidaceae. Pp. 947-954 in R. A. Dyer (editor). The Genera of Southern African Flowering Plants 2. Department of Agricultural Tech¬ nical Services, Pretoria. Heister, L. 1755. Beschreibung eines neuen Ges- chlechts. Waysenhause, Braunschweig. Herbert, W. 1820. Nerine rosea. Rose-coloured Nerine in J. Sims (editor), Bot. Mag. 47: 2124. Sherwood, Neely & Jones, London. Obermeyer, A. A. 1993. Nerine Herb. P. 162 in T. H. Arnold & B. C. De Wet (editors), Plants of Southern Africa: Names and Distribution. Mem. Bot. Survey S. Africa 62. Snijman, D. A. 1994. Systematics of Hessea, Stru- maria and Carpolyza (Amaryllideae: Amaryllida¬ ceae). Contrib. Bolus Herb. 16: 1-162. - & G. Williamson. 1994. A taxonomic re-as- sessment of Ammocharis herrei and Cybistetes lon- gifolia (Amaryllideae: Amaryllidaceae). Bothalia 24: 127-132. Traub, H. P. 1967. Review of the genus Nerine Herb. PL Life 23: 3-32. Van der Pijl, L. 1982. Principles of Dispersal in Higher Plants. Springer-Verlag, Berlin. Zucchini, W. & P. J. Adamson. 1984. The Occurrence and Severity of Droughts in South Africa. Water Research Commission Report No. 91/1/1984. De¬ partment of Civil Engineering, University of Stellen¬ bosch. Two New Species of Combretaceae from Puntarenas, Costa Rica Clive A. Stace Department of Botany, University of Leicester, Leicester LEI 7RH, England ABSTRACT. Two new species of Combretaceae are described from Costa Rica. Combretum graciliflo- rum is related to C. cacoucia Exell and is only the second species of section Cacoucia (Aublet) Engler 6 Diels known in America. Buchenavia costari- censis is the second species of the genus from Cen¬ tral America and is evidently not closely related to the other (B. tetraphylla (Aublet) R. A. Howard). Combretum assimile Eichler is also reported as new to Costa Rica, from the same area as the above two new species (Osa Peninsula, Puntarenas). Among specimens collected in Costa Rica in 1988- 1991 and recently sent to me for naming by the Missouri Botanical Garden are three taxa new to Costa Rica, two of them new to science. The two latter are the first new Central American Combre¬ taceae to be discovered for many years, indicating the need for further detailed studies of the area in question (Osa Peninsula, Puntarenas). Combretum graciliflorum Stace, sp. nov. TYPE: Costa Rica. Puntarenas: Canton de Osa/Bue- nos Aires, W part of main ridge of Fila Retinto, along and near trail from Palmar Norte to Jal¬ isco, 8°59.5'N, 83°28’W, 780-960 m, 9 Dec. 1988, Grayum & Herrera 9131 (holotype, MO). A speciebus consectionibus floribus extra dense pubes- centibus minoribus angustioribusque differt. Pedicelli 3- 7 mm; hypanthium inferius (cum ovario) 3-4 mm; hy- panthium superius anguste curvato-infundibuliforme 10- 17 mm longum (sepalis 2.5-3.5 mm longis inclusis), 3.5- 6 mm latum; petala usque ad 3 mm exserta, ca. 7x5 mm, late elliptica. Liana. Minute stalked glands frequent on leaves and inflorescences. Leaves opposite, 3.5-11.5 x 2.5-6 cm, ovate-oblong, abruptly and shortly acu¬ minate at apex, cordate to rounded-cordate at base, with 5-6 pairs of strongly curved lateral veins, densely reddish brown appressed-pubescent on veins, otherwise sparsely pubescent. Petiole 3-6 mm, densely reddish brown appressed-pubescent. Inflo¬ rescence a simple, terminal raceme 6-14.5 cm (not fully elongated). Young stems, rachis, bracts, and outside of flowers densely reddish brown appressed- pubescent. Lowest bracts ca. 14 x 5 mm, narrowly Novon 5: 106-108. 1995. elliptic, decreasing in size acropetally. Flowers (from base of pedicel to tip of petals, excluding stamens and style) 18-28 mm. Pedicel 3-7 mm. Lower hypanthium (including ovary) 3-4 mm. Upper hy¬ panthium (including sepals) 10-17 mm, narrowly curved-infundibuliform, gradually widening from base, 3.5-6 mm across at base of sepals. Sepals 2.5-3.5 x 2-3 mm, triangular, acute at apex. Petals ca. 7 x 5 mm, broadly elliptic, pubescent, subacute at apex, exceeding sepals by up to 3 mm. Stamens exserted (but broken off). Style exserted at least 10 mm when mature. The label describes the plant as a coarse liana swooping low in forest, with yellowish green petals. However, the petals have dried very dark, and they and the stamens may have been red, as in most members of the same section; the collectors might have been referring to the sepals. I have seen only one sheet of this species, but it is clearly a member of section Cacoucia (Aublet) Engler & Diels (Exell, 1953). This section contains one American species (C. cacoucia Exell) and prob¬ ably four species in West Africa (Fig. 1). Combretum graciliflorum is readily separable from C. cacoucia because of its much narrower flowers, and comes closer to some of the West African species in flower shape, notably C. mooreanum Exell, C. stenopterum Exell, and some plants of C. bracteatum (M. A. Lawson) Engler & Diels (especially plants formerly segregated as C. gentilii De Wildeman). However, it is distinct from all of these in details of flower structure. The flowers of section Cacoucia are the largest and most highly developed in the genus, with a long curved upper hypanthium, attaining 4.5 cm long in some plants of C. bracteatum (e.g., the type specimen of C. bucciniflorum Exell). They are most¬ ly red or orange in color, have long-exserted stamens and style, and are probably bird-pollinated. The flow¬ ers of the new C. graciliflorum are the smallest in the section. The fruits of the West African species of the genus are 5-winged, the wings being wide and thin in C. bracteatum (including C. gentilii and C. buc¬ ciniflorum) and C. platypterum (Welwitsch) Hutch¬ inson & Dalziel, and very narrow and corky in C. stenopterum and C. mooreanum. In contrast, the Volume 5, Number 1 1995 Stace Combretaceae from Costa Rica 107 Figure 1. Distribution of Combretum sect. Cacoucia. African range covers four species; it coincides closely with the original area of tropical rainforest. American range covers C. cacoucia and (•) C. graciliflorum sp. nov. Continents are drawn to the same scale and correct relative latitudes. fruits of C. cacoucia are larger (5-8 x 1.7-3 cm) and unwinged, with five strong longitudinal ridges. It is thus very desirable that fruiting material of C. graciliflorum be discovered. If it is a riverine spe¬ cies, like the three last-mentioned, then it too prob¬ ably has ridged or very narrowly winged fruits. Buchenavia costaricensis Stace, sp. nov. TYPE: Costa Rica. Puntarenas; Reserva Forestal Golfo Dulce Aguabuena, Sector Cuenca Oeste, 8°42'2'N, 83°31'30'W, 50-150 m, 28 Sep. 1991, Aguilar 502 (holotype, MO; isotypes, CR, LTR). Ab aliis speciebus petiolis longis (1.3-3.5 cm longi) gracilibus (laminae saepe solum duplo longae), inflores- centiae elongato-spicatae, fructibus glabris (apice suba- cuto ad rotundato), hypanthiis superis extus glabris, foliis maturis glabris, hypanthiis inferis rhachidibusque sparse pubescentibus differt. Tree to 35 m, with long, thin, straight buttresses. Leaves spirally arranged, crowded on branchlet tips; lamina 2.5-13 x 1.5-5 cm, obovate or narrowly so, with obtuse or rounded apex, with acute decur¬ rent base, with pubescent prominent midrib and lateral veins on lower surface at flowering, otherwise glabrous; petiole 1.3-3.5 cm, slender, eglandular. Inflorescence an elongated spike; peduncle and ra- chis 6-12 cm, slender, sparsely pubescent at flow¬ ering. Flowers 4-5 mm from base of ovary to tip of calyx, yellowish green; lower hypanthium 2.5-3 mm, slender, puberulous in lower ovary-bearing part, glabrous in upper part; upper hypanthium 1.5-2 x 4-5 mm, glabrous on outside, pubescent on inside, ± entire at margin; stamens 10, exserted up to 1.5 mm; style exserted up to 1.5(-2) mm. Fruit 2-3 x 1.2-1.9 cm, ellipsoid or obovoid, subacute to rounded at apex, yellowish green when ripe, gla¬ brous; endocarp irregularly but strongly longitudi¬ nally furrowed, slightly compressed. The holotype is a tree 20 m x 40 cm DBH in fruit. Buchenavia costaricensis is recognizable on the basis of its leaves with long thin petioles (many about half as long as the lamina); glabrous fruits with obtuse to rounded apex; glabrous upper hypanthia and mature leaves; and leaves at flowering time, inflorescence axis, and lower hypanthium only sparsely pubescent. The leaf shape, texture, and venation is also quite characteristic. This is only the second species of Buchenavia to be found outside mainland South America (Exell & Stace, 1963). Buchenavia tetraphylla (Aublet) R. A. Howard (B. capitata (Vahl) Eichler) occurs in the West Indies (Cuba to Trinidad) and in Costa Rica and Panama, as well as widely throughout South America, where a further 20 species occur 108 Novon (Fig. 2). Buchenavia tetraphylla is distinct from B. costaricensis in its thicker shorter petiole (0.4- 1.5(-2) cm) and capitate inflorescence. Paratypes. COSTA RICA. Puntarenas: Canton de Osa, Playa Campanario o San Josesito, Sierpe, primary forest, 8°39'39"N, 83°42'50‘ , W, 1-10 m, 5 Jan. 1991 (fl), Harmon 192 (CR, LTR, MO); Canton de Golfito Jimenez, Alto de la Carbonera, camino a Cerro de Osa, 8°25'30"N, 83°19'00"W, 200-350 m, 18 Sep. 1990 (fr), Chacon 1046 (CR, LTR); primary forest, Parque Nac. Manuel Antonio, 9°23'N, 84°09'W, 1-10 m, 11 Apr. 1990 (fr), Harmon 95 (CR, LTR); Canton de Golfito La Palma, en pequenas areas de bosque alrededor del pueblo, 8°38'30"N, 83°23'00"W, 50 m, 14 Sep. 1992 (fr), Aguilar 1302 (CR, LTR). The following two specimens have been brought to my attention by Barry Hammel. COSTA RICA. Puntarenas: Parque Nac. Manuel Antonio, woods near ANEP road, Punta Catedral, Puerto Escondido, and Lotz Road, 9°04'N, 84°10'W, 1-100 m, 3 Jan. 1990 (in bud), Harmon 40 (CR not seen); Canton de Osa, Rincon de Osa, entrada a Chocuaco, 8°43'52"N, 83°27'15"W, 100-200 m, 27 Aug. 1992 (fr), Zamora, Aguilar, Ramirez & Martin 1864 (CR not seen). COMBRETUM ASSIM1LE ElCHLER Combretum assimile is a member of section Com- bretum, represented mainly by C. frulicosum (Loe- fling) Stuntz and C. farinosum Kunth in Central America (including Costa Rica). Combretum assi¬ mile differs from these two species in its larger flowers and fruits and in its dark red scales on the flowers, fruits, inflorescence axis, leaves, and young stems (golden in C. fruticosum, gray in C. farinos¬ um). It is primarily a plant of the Amazon basin, especially on the Peru-Brazil border. Combretum sambuense Pittier was described from the old Pan¬ ama Canal Zone, and there are also records from Colombia and Venezuela (Exell, 1953), but in my opinion that species should be included in C. assi¬ mile. The following collection therefore extends the known distribution of C. assimile westward into Cos¬ ta Rica: COSTA RICA. Puntarenas: Reserva For- estal Golfo Dulce Aguabuena, Sector Sur, 8°42'20"N, Figure 2. Distribution of Buchenavia. Continuous line: B. tetraphylla. •: B. costaricensis. Broken line: 20 other species. 83°31'30"W, 50-150 m, 26 Dec. 1991, Aguilar & Herrera 793 (LTR, MO). Combretum fruticosum occurs in the same area, but the southern limit of C. farinosum is in northern Costa Rica. Acknowledgments. I am grateful to Michael Gra- yum (MO) for information concerning the holotype of Combretum graciliflorum, to Barry Hammel (MO) for data on and specimens of Buchenavia costari¬ censis, and to Carel Jongkind (WAG) for valuable data on the African species of section Cacoucia. Literature Cited Exell, A. W. 1953. The Combretum species of the New World. J. Linn. Soc. (Bot.) 55: 103-141. - & C. A. Stace. 1963. Revision of the genera Buchenavia and Ramatuella. Bull. Brit. Mus. (Nat. Hist.), Bot. 3: 3-46. A New Combination in Merremia (Convolvulaceae) for the Flora of China George W. Staples Department of Natural Sciences, B. P. Bishop Museum, Honolulu, Hawaii 96817-0916, U.S.A. Fang Rhui-cheng Kunming Institute of Botany, Academia Sinica, Heilongtan, Kunming, Yunnan, People’s Republic of China Abstract. On the basis of pollen morphology and floral characters, Ipomoea caloxantha is trans¬ ferred to Merremia. While revising the Convolvulaceae for the Flora of China, the name Ipomoea caloxantha Diels came to our attention. Fang & Huang (1979) listed /. caloxantha at the end of their treatment for Ipom¬ oea with a Chinese translation of Diels’s original description, but did not include it in their key to species of the genus nor give it any further discus¬ sion. Based on Diels’s description of the corolla as “intense lutea” and “flowers bright yellow,” one of us (GWS) suspected this to be a Merremia, in which genus yellow corollas are fairly common, whereas very few Ipomoea species have them. The holotype of I. caloxantha was examined, and the suspicion proved correct. A quick check of the pollen grains under the dissecting microscope showed them to be nonspinulose, more or less spherical, and apparently 12-rugate. This conforms to Type 3 pollen described for Merremia in the terminology of Ferguson et al. (1977). A new combination is therefore necessary. Merremia caloxantha (Diels) Staples & R. C. Fang, comb. nov. Basionym: Ipomoea calox¬ antha Diels, Notes Roy. Bot. Gard. Edinburgh 5: 203. 1912. TYPE: China. Yunnan (west¬ ern): valley of the Mekong, at the crossing of the Teng-Yueh-Talifu road, lat. 25°18'N, 4200 ft., Sep. 1903, G. Forrest 1111 (holotype, E). An amplified description of Merremia caloxantha will be published in the Flora of China. We maintain this species until further study can be devoted to Merremia in Asia, where the species are inade¬ quately known, at best, and it is quite possible that other species remain to be described, or may have already been described under Ipomoea. Acknowledgments. We thank the Keeper of the Herbarium at the Royal Botanic Garden, Edinburgh, for the loan of the holotype and Ihsan Al-Shehbaz for expediting the publication of this brief note. Literature Cited Fang Rhui-cheng & Huang Shu-hua. 1979. Convol¬ vulaceae. In: Wu Cheng-yi (editor), FI. Reipubl. Po- pularis Sin. 64(1): 1-153. Ferguson, I. K., B. Verdcourt & M. M. Poole. 1977. Pollen morphology in the genera Merremia and Op- erculina (Convolvulaceae) and its taxonomic signif¬ icance. Kew Bull. 31: 763-773. Novon 5: 109. 1995. Miconia amplinodis (Melastomataceae: Miconieae), una Nueva Especie para Costa Rica Gina Umaha Dodero Herbario Nacional de Costa Rica, Museo Nacional de Costa Rica, Apartado 749-1000, San Jose, Costa Rica Frank Almeda Department of Botany, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118-4599, U.S.A. RESUMEN. Se describe y se ilustra Miconia am¬ plinodis, una nueva especie de Costa Rica. Su ca- racteristica mas sobresaliente es el poseer unas pro- yecciones nodales tanto en tallos como en inflores- cencia. Se comenta su relacion con otras especies del section Cremanium, al cual se adscribe. ABSTRACT. Miconia amplinodis, a new species from Costa Rica, is described and illustrated. The most outstanding feature of this species is its suc¬ culent auriculate projections at branch and inflo¬ rescence nodes. The relationships of M. amplinodis to other species in section Cremanium are also dis¬ cussed. Miconia, el genero mas grande de la familia Me¬ lastomataceae, se encuentra distribuido en la region neotropical con aproximadamente 1000 especies. Estas han sido agrupadas en 11 diferentes secciones en base a la gran variedad presentada en la mor- fologia del androceo. En Costa Rica, este genero se halla representado por unas 100 especies aproximadamente; numero que se ha incrementado durante los ultimos aiios debido a una mayor actividad de colecta en bosques de mediana elevation, de donde se han descrito varias especies nuevas (Almeda & Umana, 1993). La especie que aqui se describe es conocida de la Cordillera de Cuanacaste, Cordillera de Tilaran y la Cordillera Volcanica Central en elevaciones de 1100 a 2000 m.s.n.m. Miconia amplinodis Umana & Almeda, sp. nov. TIPO: Costa Rica. Alajuela: Bajos del Toro Amarillo, 10°12'10'N, 84°18'43"W, 1800 m, 15 jun. 1992, Umana et al. 495 (holotipo, CR; isotipos, CAS, COL, F, MEXU, MO, US, USJ). Figura 1. Novon 5: 110-113. 1995. Frutex 0.5-2 m altus. Rami ramulique sicut petioli inflorescentiaque glabri, nodis tumidis et auriculis 2.5- 3.5 mm longis infra petiolorum insertiones armatis. Petioli 1.3-4.8 cm longi; lamina 8-11.7 x 3.3-5.7 cm elliptica, apice breviter hebeti-acuminato basi acuta vel obtusa, 3- nervata. Panicula 7-17 cm longa multiflora; bracteolis 1-2 x 0.75-1 mm deciduis. Hypanthium (ad torum) 2- 2.5 mm longum; calycis tubus 0.20-0.25 mm longus, lobis usque ad 0.5 mm longis et 0.75 mm latis. Petala 1.5-2 x 1-1.5 mm. Stamina 10, isomorphica glabra; filamenta 2 mm longa; antherarum thecae 1-1.4 x 0.25- 0.50 mm obovato-oblongae 4-porosae, connectivo paulo (0.4 mm) prolongato. Stigma truncatum; stylus 3-3.5 x 0.2 mm glaber; ovarium % inferum, 3(?)-loculare, apice truncato-conico pilis paucis glanduliferis. Arbusto de 0.5-2 m de altura; entrenudos, ramas, peciolos e inflorescencia glabros; nudos del tallo e inflorescencia con unas proyecciones persistentes, glabras, redondeadas y carnosas de 2.5-3.5 mm de largo; hojas de un par iguales en tamano, elipticas, algo coriaceas, margen aserrado, 8-11.7 cm de largo y 3.3-5.7 cm de ancho, apice corto acumi- nado, base anchamente aguda a obtusa; 5 nervios (incluyendo los dos nervios laterales tenues); peciolos 1.3-4.8 cm de largo. Inflorescencia una panicula terminal de muchas flores, 7-17 cm de largo; brac- teas pares, ovadas, 2-6 mm de largo y 2.5 mm de ancho, horde irregular, glabras, sesiles, caducas; bracteolas pares, ovadas, 1-2 mm de largo y 0.75- 1 mm de ancho, borde irregular, glabras, sesiles, caducas; pedicelos glabros, 0.25-0.5 mm de largo; hipantio (en la floracion), campanulado, glabro, con 10 costillas, 2-2.5 mm de largo hasta el toro; tubo del caliz 0.20-0.25 mm de largo; lobulos del caliz glabros, anchamente redondeados, 0.5 mm de largo y 0.75 mm de ancho; dientes externos persistentes, sobrepasados en tamano por los lobulos del caliz; petalos 5, glabros, concavos, suborbiculares, blanco- verdosos, isomorficos, 1.5-2 mm de largo y 1-1-5 mm de ancho; filamentos glabros, distalmente ge- niculados, 2 mm de largo; anteras 1-1.4 mm de Volume 5, Number 1 1995 Umana & Almeda Miconia amplinodis 111 Figura 1. Miconia amplinodis Umana & Almeda. —A. Habito. B. Superficie abaxial de la hoja. C. Detalle de las proyecciones nodales. —D. Flor. —E. Petalo. —F. Estambres, vista ventral (izquierda), vista dorsolateral (centro) y vista lateral (derecha). —G. Bractea. —H. Fruto. I. Semillas. (A - F de Umana et al. 495; H, I de Umana et al. 465.) 112 Novon largo, cuneadas, distalmente truncadas con 4 poros terminales ventralmente inclinados; conectivo en- grosado dorsalmente y prolongado bajo las tecas 0.4 mm. Ovario 3(?)-loculare, infero, estilo glabro, recto, 3-3.5 mm de largo; estigma truncado. Fruto una baya globosa, 6 mm de diametro, morado cuando inmaduro, bianco en la madurez. Semillas nume- rosas, cafe-rojizas, ovoides, 1.3 mm de largo, la testa papilosa. Distribution y fenologia. Se conoce de la Cor¬ dillera de Guanacaste, Cordillera Volcanica Central (Alajuela: Bajos del Toro Amarillo, Fila Volcan Viejo cercanias del Volcan Poas; Heredia: Parque Nacio- nal Braulio Carrillo), y de la Cordillera de Tilaran, Reserva de Bosque Nuboso Monteverde, en la parte noroeste de Costa Rica en elevaciones de 1100- 2000 m. Esta especie ha sido colectada en sitios abiertos y soleados, en orillas de caminos y en po- treros (Fig. 2). Los especimenes con flor han sido colectados en los meses de abril, mayo, junio, julio y septiembre; muestras con fruto en febrero, abril, septiembre, noviembre y diciembre. Miconia amplinodis fue colectada por primera vez en Monteverde en 1979; esta muestra presen- taba solamente frutos lo mismo que posteriores co- lectas hechas en la misma localidad y en Fila Volcan Viejo y Bajos del Toro. No fue hasta 1992 cuando se logro realizar una colecta de material plenamente florecido. Miconia amplinodis se caracteriza por ser un arbusto glabro, con hojas glabras de 5 nervios con borde ligeramente aserrado de color morado, y con flores de 5 partes y anteras cuneiformes de 4 loculos. Su caracteristica mas sobresaliente, y unica dentro de las especies de Miconia conocidas para el pais, es el poseer unas proyecciones persistentes, glabras, redondeadas y carnosas a nivel de nudos tanto en tallos como en inflorescencias. Estas proyecciones nodales engrosadas son raras en Miconia y en la familia Melastomataceae en general, y hasta el mo¬ menta no se conoce su significancia funcional. Cu- riosamente estas estructuras presentan un color di- ferente dependiendo de si estan expuestas o no a la luz solar; cuando es asi, el color varia entre rojo y morado, cuando se hallan en sitios sombreados estas presentan un color verde. Generalmente se presen¬ tan 2 por nudo con excepcion de una muestra que presentaba 3 de estas estructuras por nudo. Esta especie, asignable a la seccion Cremanium Volume 5, Number 1 1995 Umana & Almeda Miconia amplinodis 113 en virtud de sus anteras cuneiformes de 4 loculos, se halla relacionada cercanamente a Miconia re- sima Naudin, especie conocida solamente para Co¬ lombia. Las caracteristicas que ambas comparten son el poseer flores de 5-partes y androceos muy similares. A diferencia de Miconia amplinodis, M. resima carece de las proyecciones nodales de la que parten los peciolos y las ramificaciones de las inflo- rescencias; posee hojas de mayor tamano (15-20 x 6-7 cm), y presenta un estigma subpeltado. Entre las especies de Miconia, dentro de la sec- cion Cremanium, que poseen proyecciones anula- riformes conspicuas en cada nudo, M. amplinodis es tambien similar a M. manicata Cogniaux & Glea¬ son de Colombia, en el tamano de las hojas, mor- fologia del hipantio, y forma de los petalos. Miconia manicata difiere de nuestra nueva especie en poseer lamina foliar ciliada (los cilios de approx. 1 mm de largo) y base largo cuneada, tecas de las anteras de 4 poros incompletos con el conectivo basalmente trilobulado, ovario medio infero, y estigma capitado. El nombre dado a esta especie hace alusion a las proyecciones conspicuas que presente a nivel de nudos en tallos e inflorescencias, caracteristicas que la hace facil de reconocer. Paratipos. COSTA RICA. Alajuela: Volcan Poas, faldas del Cerro innominado ubicado al N del crater, 5 mayo 1992, Ballestero 148 (CR); San Carlos, Fila Volcan Viejo, 1800-2000 m, 11-14 feb. 1986, Gomez-Laurito 11021 (CR); San Carlos, Fila Volcan Viejo, 1800-2000 m, 13 nov. 1986, Gomez-Laurito 11251 (CR); Bajos del Toro Amarillo, 10°12'10'N, 84°18'43"W, 1800 m, 10 abr. 1989, Umana etal. 276 (CR), 28 abr. 1992, Umana et al. 465 (CR), 14 mayo 1992, Umana et al. 484 y 488 (CR). Alajuela/Puntarenas/Guanacaste: Cor¬ dillera de Tilaran, Reserva Biologica Monteverde, 1540- 1600 m, 21 feb. 1986, Almeda et al. 4999 (CAS, CR); Reserva Biologica Monteverde, 10°18'N, 84°46'W, 1100 m, 1 jul. 1989, Bello 996 (CAS, CR); Reserva Biologica Monteverde, 1500-1600 m, Haber ex Bello 5190 (CAS, CR, MO); Reserva Biologica Monteverde, 10°18'N, 84°47'W, 1500-1600 m, 16 jul. 1990, Haber & Zu chowski 10014 (CAS, CR, MO); Reserva Biologica Mon¬ teverde, 10°18'N, 84°47' r W, 1500-1600 m, 31 die. 1990, Haber & Zuchowski 10378 (CAS, CR, MO); Re¬ serva Biologica Monteverde, 10°18'N, 84°47'W, 1520- 1600 m, 14 sep. 1991, Haber & Zuchowski 10872 (CAS, CR, MO); Reserva Biologica Monteverde, 10°18'N, 84°48'W, 1600 m, 17 jun. 1992, Haber et al. 11206 (CR); Reserva Biologica Monteverde, 3 jul. 1979, Lumer 1086 (CAS). Heredia: Parque Nacional Braulio Carrillo, San Isidro, Cerro Zurqui, 10°03'30*’N, 84°01T5*’W, 1800 m, 16 jul. 1992, Ballestero et al. 177 (CR). Guana- caste: Parque Nacional Rincon de la Vieja, Sector Las Pailas en la cima del Volcan, 10°49'30*’N, 85°19 / 30"W, 1700 m, 9 enero 1994, Morales 2243 (CR). Agradecimientos. Queremos hacer publico nues- tro agradecimiento a Pablo Sanchez V. y a Jorge Gomez-L. por su colaboracion en la descripcion la- tina de la especie que aqui se describe; a Jorge Gomez-L., Miguel Ballestero y William Haber por el comentario sobre las colectas hechas de la especie en los diferentes lugares; y a Rafael Chacon por su colaboracion en las giras realizadas para la colecta del material. Literatura Citada Almeda, F. & G. Umana D. 1993. New species of Miconia (Melastomataceae) from Costa Rica. Novon 3: 5-10. Justicia nevlingii (Acanthaceae), a New Species from Mexico Dieter C. VTasshausen Department of Botany, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Thomas F. Daniel Department of Botany, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118, U.S.A. ABSTRACT. A new species of Justicia with heter- omorphic bracts, J. nevlingii , is described from southern Mexico. The species is compared to two other heteromorphically bracted species from Mex¬ ico and Central America, J. chol and J. costaricana. It differs from these in several features, including its 4-aperturate pollen. In 1974, Larry Nevling of the Field Museum of Natural History called to the attention of the senior author an unusual collection of Acanthaceae from Veracruz representing an unknown species. Other collections resembling this species from Veracruz and Chiapas have since been assembled and studied. They are described below as a new species of Jus¬ ticia with heteromorphic bracts. Justicia nevlingii Wasshausen & T. F. Daniel, sp. nov. TYPE: Mexico. Veracruz: Hidalgoti- tlan, 0-2 km del Plan de Arroyo-Alvaro Obre- gon, 17°01'N, 94°40'W, 14 Apr. 1974, J. Dorantes et al. D-2799 (holotype, US; isotype, F). Figures 1, 2. Herba perennis usque ad 1.5 m alta. Folia petiolata, laminae ovatae, 6.5-18.5 cm longae, 3-10.4 cm latae. Spicae terminales et axillares, densae, pedunculis et rhach- idibus dense pilosis. Bracteae heteromorphae; bracteae fertiles petiolatae, obovatae vel ellipticae, 5.5-12 mm longae, 2.7-6.5 mm latae; bracteae steriles lineares-lan- ceolatae vel lineares vel subulatae, 2-5 mm longae, 0.3- 0.5 mm latae. Calyx 5-lobus. Corolla alba vel flavida vel cremea, 9-10 mm longa, extus pubescens trichomatibus eglandulosis. Thecae antherarum 2, superpositae, glabrae, theca inferiore basi calcarata; pollen 4-apertum. Capsula 7-8.5 mm longa, pilosa. Semina 2. Erect, perennial herbs to 1.5 m tall. Stems subquadrangular, quadrifariously pilose with tri- chomes eglandular, appressed and spreading, 0.7- 2 mm long, multi-septate with maroon septae. Leaves petiolate, petioles 1.5-7.2 cm long, densely pilose, blades ovate to elliptic, 6.5-18.5 cm long, 3-10.4 Novon 5: 114-117. 1995. cm wide, short-acuminate at apex, obtuse or rounded to subattenuate at base, entire to sinuate, upper surface glossy, sparingly pilose with trichomes ap¬ pressed, lower surface more densely pilose (especial¬ ly the central and lateral veins) with trichomes ap¬ pressed and spreading, to 1 mm long. Inflorescence of (axillary and) terminal dense panicles of spikes to 15 cm long and 1.5-4.4 cm diam., erect or ascending, rachises densely pilose with trichomes sordid, erect, to 1 mm long. Floral bracts hetero¬ morphic; fertile bracts petiolate, obovate to elliptic, 5.5-12 mm long, 2.7-6.5 mm wide, acute to trun- cate-apiculate at apex, abaxial surface pubescent with antrorse-appressed eglandular trichomes 0.1- 0.5 mm long and glandular trichomes to 0.2 mm long, margin ciliate with cauline type trichomes; sterile bracts linear-lanceolate to linear to subulate, 2-5 mm long, 0.3-0.5 mm wide. Bracteoles oblan- ceolate, 4-9 mm long, 1.2-2.2 mm wide. Calyx deeply 5-lobed, 3-5 mm long, lobes lanceolate, 2.5- 4 mm long, 0.5-0.6 mm wide, puberulous. Corolla white to yellowish to cream, 9-10 mm long, exter¬ nally pubescent with eglandular trichomes 0.2-0.5 mm long; tube 5-5.5 mm long, 0.8 mm diam. at base, 3 mm diam. at apex; limb 2-lipped, upper lip erect, rugulate, ovate-lanceolate, 3-4.5 mm long, 2 mm wide, 2-lobed at apex, lobes ca. 0.25 mm long and wide, rounded, lower lip ± spreading, 3- lobed, 3.5-6 mm long, 3 mm wide at base of lobes, lobes obovate, 1.2-1.5 mm long, 1.2-1.5 mm wide, rounded at apex. Stamens inserted near apex of corolla tube, 4 mm long, filaments glabrous, thecae superposed, glabrous, upper theca ca. 1 mm long, lower thecae ca. 0.8 mm long, calcarate with blunt appendage ca. 0.3 mm long; pollen prolate 4-aper¬ turate, apertures flanked on each side by a ± con¬ tinuous band of exine and a pseudocolpus (Fig- 2); 2 staminode-like protrusions of corolla present near base of filaments, 0.4-1.2 mm long, densely villous. Style 6.5 -7 mm long, glabrous, stigma 0.2-0.3 mm long. Capsules 7-8.5 mm long, pilose. Seeds 2, Volume 5, Number 1 1995 Wasshausen & Daniel Justicia nevlingii 115 ED C Figure 1. Justicia ne vlingii Wasshausen & T. F. Daniel (Dorantes et al. D-2799). A. Habit. B. Bract. C. Bracteoles. — D. Calyx lobes. — E. Corolla and stamens. —F. Expanded corolla, stamens, and staminode-like protrusions. 116 Novon Figure 2. Scanning electron (SEM) photomicrographs of pollen of Justicia nevlingii (Dorantes et al. D-2799). — A. Equatorial view, x2800. —B. Portion of aperture, x7000. spherical to discoid, flattened, 2.1-2.8 mm long, 2- 2.5 mm wide, smooth. Distribution. Southern Mexico (Chiapas and Ve¬ racruz); plants occur in lowland and lower montane rainforests at elevations from 130 to 470 m. Paratypes. MEXICO. Veracruz: Mpio. Hidalgotitlan, 0-2 km road Plan de Arroyo-Arroyo Alegre, 17°15'N, 94°35'W, 23 Apr. 1974, Jesus Dorantes et al. D-2964 (F, US); Mpio. Hidalgotitlan, between Hnos. Cedillo-Au¬ gustin Melgar, 17°13'30' r N, 94°35'W, 26 Apr. 1974, Mario Vazquez et al. 411 (F, US); Las Choapas, Las Cruces, 14 July 1970, A. Gomez-Pompa & L. Nevling 1522 (F, MEXU); Mpio. Hidalgotitlan, La Laguna, Uxpanapa, 29 Mar. 1974, A. Gomez-Pompa & L. Nevl¬ ing 5183 (F). Chiapas: Mpio. Ixtacomitan, above Ixta- comitan, 10 May 1973, D. Breedlove 35058 (DS, MEXU); St. Bartolo, Apr. 1840, J. Linden s.n. (G, P); along Rio Pichucalco, 3 km NE of Pichucalco toward Villahermosa, 17°32'N, 93°04'W, 2 July 1969, B. Marchs & C. Marcks 913 (LL, WIS); 37 km NW of Bonampak, 6 km NNW of Nuevo Guerrero, G. Davidse et al. 20534 (BM, LL, MEXU); between Pichucalco and Rayon near Solosuchiapa, cuadrante 57-A, J. Chavelas P. et al 442 (MEXU). Justicia nevlingii resembles both J. chol T. F. Daniel (Chiapas) and J. costaricana Leonard (Costa Rica) by its heteromorphic bracts and staminode- like protrusions of the corolla. It may be distin¬ guished from these species by the following key: la. Bracts eciliate or inconspicuously ciliate with trichomes to 0.2 mm long; calyx 5-7 mm long; corolla externally glandular; cauline trichomes lacking maroon septae; cloud forests at 900- 1550 m in Costa Rica. J- costaricana lb. Bracts conspicuously ciliate with trichomes to 2 mm long; calyx 2.5-5 mm long; corolla ex¬ ternally eglandular; cauline trichomes with con¬ spicuous maroon septae; lowland to lower mon¬ tane rainforests at 50-470 m in Mexico. 2a. Young stems with trichomes concentrated in 2 lines; inflorescence of axillary spikes; bracteoles 0.4-1 mm wide; style pubes¬ cent; seeds 4, 1.3-1.6 mm long, covered with low rounded papillae or ridges; pollen 3-aperturate. J- c h°l 2b. Young stems with trichomes restricted to 4 lines; inflorescence of (axillary to) ter¬ minal panicles of spikes; bracteoles 1.2- 2.2 mm wide; style glabrous; seeds 2, 2.1 - 2.8 mm long, smooth; pollen 4- aperturate. J. nevlingii Graham (1988) stated that the pollen of Justicia is either 2- or 3-aperturate but that 4-aperturate pollen (not confirmed by her study) had been re¬ ported in a few New World species. Daniel (1990, Volume 5, Number 1 1995 Wasshausen & Daniel Justicia nevlingii 117 1993) documented 4-aperturate pollen in several species of Justicia (i.e., J. angustijiora D. N. Gib¬ son, J. tabascina T. F. Daniel, and J. valvata T. F. Daniel) and has since observed it in several others (e.g., J multicaulis Donnell Smith, J. silvicola D. N. Gibson, and /. warnockii B. L. Turner). Graham (1988) also noted that all the species of Justicia that she had observed have an androecium of two stamens and no staminodes. Justicia nevlin¬ gii and the two other heteromorphically bracted species of Justicia noted above exhibit two stami- node-like protrusions near the base of the filaments. It is not known whether these protrusions technically are staminodes or whether they are merely thick¬ enings of the corolla near the point of insertion of the filaments. Whatever their origin, they appear to be present in species of Justicia with hetero- morphic bracts. Heteromorphically bracted species of Justicia are now known from Mexico and Central America, and they undoubtedly occur in South America as well. Determining whether they consti¬ tute a monophyletic infrageneric taxon within Jus¬ ticia deserves additional study. Acknowledgments. We thank Alice Tangerini for skillfully preparing the line drawings, and Stephen F. Smith and the staff of the National Museum of Natural History SEM Laboratory for providing the high-quality pollen photomicrographs. Literature Cited Daniel, T. F. 1990. New and reconsidered Mexican Acanthaceae. IV. Proc. Calif. Acad. Sci. 46: 279- 287. -. 1993. New and reconsidered Mexican Acan¬ thaceae. V. Contr. Univ. Michigan Herb. 19: 271- 291. Graham, V. A. 1988. Delimitation and infra-generic classification of Justicia (Acanthaceae). Kew Bull. 43: 551-624. Errata Croat, T. B. & M. H. Grayum. 1994. Philodendron Subgenus Pteromischum (Araceae) from Saul, French Guiana. Novon 4: 211-219. On page 216, second column, third paragraph, in the discussion of Pteromischum placidum, the following should be changed: “It [ Pteromischum placidum ] is very easily confused with P. rud- geanum Schott, which differs in climbing directly up the side of the tree, and in having blades that are matte rather than glossy on the lower blade surfaces, a slender spadix, and in generally flowering ahead of P. placidum. It was rarely at anthesis during mid-February, whereas many plants of P. placidum were in flower.” It should read: “It [ Pter¬ omischum placidum ] is very easily confused with P. rudgeanum Schott. . . and in generally flowering later than P. placidum. Philodendron placidum was rarely in flower in mid-February, whereas many plants of P. rudgeanum were in flower.” On page 219, the first paragraph of the second column, the following should be changed: “In ad¬ dition, Philodendron placidum flowers later than P. rudgeanum , which was only rarely in flower in mid-February when P. placidum was in full flower." It should read: “In addition, P. rudgeanum flowers later than P. placidum, which was only rarely in flower in mid-February when P. rudgeanum was in full flower.” Volume 5, Number 1, pp. 1-118 of NOVON was published on 27 March 1995. Novon 5: 118. 1995. Volume 5 Number 2 1995 NOVON MISSOURI BOTANICAL SEP 1 2 1995 GARDEN LIBRARY A Note on the Kazakh Endemic Lap pula zaissanica (Boraginaceae) N. K. Aralbaev Main Botanical Garden, Timiryazev St., 480070 Almaty, Republic of Kazakhstan Abstract. Studies of the Kazakh endemic Lap- pula coronifera (M. Popov) subsp. zaissanica Ar¬ albaev show that it is a distinct species. The new combination Lappula zaissanica (Aralbaev) Aral¬ baev is proposed. Recent collections of an interesting Lappula from the Zaissan depression of eastern Kazakhstan were referred to Lappula coronifera by Kudabaeva & Aralbaev (1989). Subsequent collections showed that the plants represent a distinct subspecies, L. coron¬ ifera subsp. zaissanica (Aralbaev, 1990). Further studies and critical comparison with authentic plants of L. coronifera, as well as additional fieldwork, revealed that the plants in question represent a distinct species quite common in the Zaissan de¬ pression. A new combination is herein proposed. Lappula zaissanica (Aralbaev) Aralbaev, comb, nov. Basionym: Lappula coronifera M. Popov subsp. zaissanica Aralbaev, Bot. Zhurn. 75: 707. 1990. TYPE: E Kazakhstan. Northern Zaissan: above Schubarschilik, sandy-pebbled desert, 8 June 1984, Aralbaev s.n. (holotype, Altay Botanical Garden, Leninogorsk (AIC); iso¬ type, LE). Herbs annual, 7-15(-20) cm tall. Stems 1 or several, branching at base or apex. Basal leaves rosulate, dried at fruiting time; cauline leaves sessile, lanceolate, 9-12 mm long, 1-2.5 mm wide, some¬ what longitudinally folded, sparsely tomentose and greenish adaxially, moderately tomentose abaxially. Bracts shorter than fruits, lowermost oblanceolate, upper ones narrowly lanceolate. Pedicels tomentose, as long as or slightly longer than calyx, slightly reflexed in fruit. Calyx tomentose, with bristly tu¬ bercles; lobes in fruit narrowly lanceolate, 3.5-4 mm long, slightly reflexed. Corolla blue, 3 cm long, with broadly campanulate limb. Fruit broadly ovoid; nutlets similar, with a smooth crownlike cup turning from yellow to brown as nutlets mature, each side with 6 long, glochidiate spines shorter than or nearly as long as the cup; disc not keeled, smooth, bright; gynophore 2-2.5 mm long; style exserted from nut¬ lets by 0.2-0.3 mm. Lappula zaissanica, which is a Kazakh endemic known thus far only from the Zaissan depression, is most closely related to L. coronifera, from which it differs by having nutlets with smooth, not keeled, bright-colored discs and broader wings with shorter teeth. Additional specimens examined. KAZAKHSTAN. N Zaissan Region: in valleys above Kiin-Kerish, 18 May 1983, Aralbaev s.n. (paratype; AIC); Kara-Biryuk, peb¬ bled desert, 12 June 1993, Aralbaev s.n. (Main Botanical Garden, Almaty). Acknowledgments. I am grateful to Ihsan Al- Shehbaz and Larissa A. Karavaeva for their help with the manuscript. Literature Cited Aralbaev, N. K. 1990. A note on Lappula coronifera (Boraginaceae). Bot. Zhurn. (Moscow & Leningrad) 75: 707-708. Kudabaeva, G. M. & N. K. Aralbaev. 1989. Genus Lappula Moench. in flora of north part of Zaissan’s depression. Mat. Kazakh. Acad. Bot. Inst. Herb. Alma-Ata 1989: 34-42, Novon 5: 119. 1995. New Species of Gibsoniothamnus (Bignoniaceae: Schlegelieae) Kerry Barringer Brooklyn Botanic Garden, 1000 Washington Avenue, Brooklyn, New York 11225-1099, U.S.A. Alwyn Gentryf Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. ABSTRACT. Three new species of Gibsoniotham¬ nus L. O. Williams, G. grandiflorus, G. stellatus, and G. versicolor, are described as new. Gibsoniothamnus L. 0. Williams is a genus of about 14 species of epiphytic shrubs, variously clas¬ sified in the Bignoniaceae or Scrophulariaceae. This unusual group of Central American epiphytes is most closely related to the neotropical genus Schlegelia Miquel, currently classified in the Bignoniaceae tribe Schlegelieae. Undescribed species have been filed with both Scrophulariaceae and Bignoniaceae, so a collaborative project was begun to identify and de¬ scribe the new species. Collaborative work was cut short by the untimely death of A1 Gentry. The fol¬ lowing species were discovered during the study. Gibsoniothamnus grandiflorus A. Gentry & Barringer, sp. nov. TYPE: Panama. Veraguas: ridge of Cordillera de Tute along trail to Cerro Tute, ca. 3-4 km past Escuela Agricola Alto de Piedra, just W of Santa Fe, 8°32'N, 81°07'W, 20 Mar. 1982, Knapp & Kress 4389 (holotype, MO; isotype, PMA). Figure 1. Species fioribus grandis, a Gibsoniothamno truncato lobis calyce triangulatis longe acuminatis, pedicellis lon- gioribus differt. Epiphytic shrub to 3 m tall; bark gray-brown; stems angled, glabrous, to 5 mm diam., with raised leaf bases at the nodes. Leaves opposite, subequal to equal across the node; petioles 5-14 mm long, glabrous or sparsely pubescent; blade ovate to ob- ovate or lanceolate, 6.5-14.5 cm long, 3.0-6.2 cm wide, coriaceous, glabrous, shiny above, the base cuneate, the apex acuminate, secondary veins 3 or 4 per side, impressed above, prominent below, glan¬ dular pits in fields between the veins below, pocket domatia occasionally present in the axils of the mid¬ vein and secondary veins toward the base of the blade. Flowers in congested, axillary panicles; pri¬ mary peduncle to 1.5 cm long; bracts narrowly Novon 5: 120-124. 1995. triangulate, 3-5 mm long, pubescent; the secondary peduncles to 7 mm long; bracteoles to 3 mm long, ovate, acuminate; pedicels 2.5-3.5 cm long, thicker above; calyx funnelform, the tube 5-9 mm long, red to red-purple, the lobes narrowly triangulate, unequal, 7-13 mm long, long-acuminate, glabrous to minutely ciliate; corolla cylindric, slightly ampliate above, 3.5-4.0 cm long, 6-7 mm diam., reddish pink to red, fleshy, the upper lobes rounded, 3-4 mm long and wide, the lateral lobes rounded to ligulate, 4-5 mm long, 4 mm wide, the median lobe ligulate, 4-6 mm long, 3-4 mm wide; stamens at¬ tached 6-9 mm from the base of the corolla tube, filaments 20-26 mm long, glabrous; staminode 10- 20 mm long, filiform, the apex clavate; pistil gla¬ brous, the ovary globose to oblate, 2-3 mm diam., the style 30 mm long, the stigma slightly bilobed. Berry globose, 8-10 mm diam., green. This species is most easily distinguished by its large, slightly ampliate flowers. It appears to be most closely related to Gibsoniothamnus truncatus and G. latidentatus . It differs from G. truncatus by its larger, ampliate corollas, pedicels more than 2 cm long, and narrowly triangulate calyx lobes. It differs from G. latidentatus by its corollas more than 3 cm long, reddish calyx, short staminode, and calyx lobes 7-13 mm long. This species has an interesting disjunct distribution: the plants from Panama, col¬ lected along the El Llano-Carti road, have narrower, more acuminate leaves than those from Code, most¬ ly collected on Cerro Tute. Paratypes. PANAMA. Code: vicinity of Escuela Agricola Alto de Piedra, 3200-3400 ft., 3 Apr. 1980, Antonio 3961 (MO); about 16 km NW of Santa Fe on descent to Rio Caloveborita, 650 m, 4 Sep. 1975, Dress¬ ier 5142 (MO); near third branch of Rio Santa Maria, 10-14 km NW of Santa Fe, 650-750 m, 10-11 Oct. 1975, Dressier 5171 (MO); shoulder of Cerro Tute, 25 May 1977, Folsom & Edwards 3345 (MO); ridge of Cerro Tute, along trail to Cerro Tute, 3-4 km past Escuela Agricola Alto de Piedra, 8°32'N, 81°07'W, 800-1400 m, 20 Mar. 1982, Knapp & Kress 4363 (MO); Cerro Tute, E slopes, 1 km beyond Escuela Agricola Alto Piedra Volume 5, Number 2 1995 Barringer & Gentry New Species of Gibsoniothamnus 121 above Santa Fe, 900-1200 m, 14 May 1981, Sytsma & Andersson 4655 (MO, PMA). Panama: El Llano- Carti road, 10 km N of Pan Am Highway, 340 m, 15 Feb. 1975, Gentry & Mori 14216 (F, MO); 6-7 mi. from PanAmerican highway on El Llano-Carti road, 9°12'N, 79°00'W, 400 m, 26 Feb. 1982, Knapp & Mallet 3829 (MO, PMA); 8.2 mi. from Panamerican highway on the El Llano-Carti rd„ 9°14’N, 79°00'W, 450 m, 24 Mar. 1982, Knapp & Huft 4415 (MO, PMA); toad between El Llano and Carti-Tupile road, 12 mi. above Pan American highway, 200-500 m, 30 Mar. !973, Liesner 1321 (F, MO, NY, PMA); El Llano-Carti td-, 10.8 km from interamerican highway, 1100-1200 ft-, 27 Dec. 1974, Mori, Kallunki & Hansen 4121 (MO); El Llano-Carti road, 21.6 mi. from Inter-American high¬ way, 350 m, 20 Mar. 1975, Mori & Kallunki 5104 (MO, PMA); El Llano-Carti road, 9 mi. from Pan Amer¬ ican highway, 350—400 m, 28 Apr. 1981, Sytsma 4146 (NY). Gibsoniothamnus stellatus A. Gentry & Barrin¬ ger, sp. nov. TYPE: Panama. Chiriqui; Cerro Colorado, along road to copper mine 33.1 km beyond the bridge over the Rio San Felix, 13.1 km beyond turnoff to Escopeta, 1400 m, 15 July 1976, Croat 37200 (holotype, MO; iso¬ type, MO). Figure 2. 122 Novon A Gibsoniothamno alato foliis minoribus, petiolis 3- 5 ram longis, tubis calyce 6-8 mm longis, staminibus 17— 18 mm longis differt. Epiphytic shrub to 1.5 m tall; bark pale gray- brown; stems angled, glabrous, the leafy portions to 2 mm diam., the older portions with raised leaf bases. Leaves opposite, unequal to subequal; petioles 3-5 mm long, glabrous, unwinged; lamina ovate-lanceo¬ late, coriaceous, glabrous, gland-dotted, the base cuneate, the apex acuminate to acute, the midvein impressed above, prominent below, the secondary veins not impressed above, prominent below, 2-3 per side, domatia occasionally found on the under¬ side of the lamina near the base, the larger leaves 15-25 mm long, 9-16 mm wide, the smaller leaves 5-12 mm long, 4-8 mm wide. Flowers borne near the tips of the branches, solitary, axillary; pedicels 10-12 mm long, thick above, bracteolate at base, glabrous; calyx campanulate, purple, the tube 6-8 mm long, the lobes 8-9 mm long, triangulate, the wings broad, 3-4 mm wide; corolla tubular, white. Volume 5, Number 2 1995 Barringer & Gentry New Species of Gibsoniothamnus 123 2.0-2.5 cm long, 4-5 mm diam., the upper and lateral lobes rounded to ligulate, 3-4 mm long and wide, the margin ciliate, the median lobe ligulate, 3 mm long, 1.5-2.0 mm wide; stamens attached 5 mm above the base of the corolla tube, with a ring of trichomes at the point of insertion, the filaments 17-18 mm long; staminode 5-6 mm long; pistil glabrous, the ovary globose, 1.5 mm diam., the style 14 mm long, the stigma capitate. Immature berries globose, green. This species is distinguished by its small leaves and large, winged calyx lobes. It appears to be most closely related to Gibsoniothamnus alatus, native 124 Novon to southern Darien and adjacent Choco, because both species have strongly alate calyces and white corollas. It can be distinguished from that species by its smaller leaves, calyx tube more than 5 mm long, and stamens less than 18 mm long. Paratypes. PANAMA. Bocas del Toro: region of Cerro Colorado, 7.5 mi. from Chami camp, 8°35'N, 81°45'W, 1220-1250 m, 13 Apr. 1986, McPherson 8882 (MO, NY, PMA). Gibsoniothamnus versicolor A. Gentry & Bar¬ ringer, sp. nov. TYPE: Panama. San Bias: Playon Chico, Rio Ukupseni, Campamento Neba Dummat hasta las cascada, 50-100 m, 9°15'N, 78°15'W, 30 Oct. 1991, H. Herrera, J. Mor¬ ris & J. Mojica 1022 (holotype, MO; isotypes, BKL, PMA). Figure 3. A Gihsoniothamno allenii foliis majoribus anguste el- lipticis, petiolis ciliatis, floribus paniculatis, lobis calyce majoribus differt. Epiphytic shrub or climber; bark gray-brown; stems angled, glabrous to sparsely pilose, with raised leaf bases at the nodes. Leaves opposite, equal to subequal; petioles 5—12 mm long, pilose, unwinged; blade narrowly elliptic, 4-11 cm long, 1.7-4.0 cm wide, subcoriaceous, dull olive gray above, tannish below, sparsely pilose above, with scattered tri- chomes along the midvein below, strongly lepidote- glandular below, domatia absent. Flowers in terminal panicles; pedicel 2-3 cm long, sparsely pilose, slight¬ ly thickened above; calyx campanulate, pink to mauve, the tube 3—5 mm long, the lobes nearly linear, 7-12 (-15 in fruit) mm long, porrect, then reflexed after anthesis; corolla tubular, white or red¬ dish violet, 2 cm long, 4 mm wide at the base, 3 mm wide near the apex, the lobes ovate to round, 1.5-3.0 mm long; stamens inserted 4-5 mm from the base of the corolla tube, with a basal ring of trichomes, the filaments 1 cm long; staminode 5 mm long, filiform; ovary depressed-globose, 2 mm long, glabrous, the style 15 mm long, the stigma subcapitate. Young fruit green, subglobose, 6 mm long. This species differs from Gibsoniothamnus alatus A. Gentry, the only other species in this genus known from eastern Panama, by its linear, unwinged calyx lobes. It appears to be most closely related to G. allenii A. Gentry, which grows in the hills around El Valle. This species differs from G. allenii by its longer calyx lobes, 7-15 mm long, and its larger, subcoriaceous leaves. D’Arcy (1979) listed Croat 27281 under G. cornutus (Donnell Smith) A. Gen¬ try, but that species differs by its short, triangulate calyx lobes and coriaceous leaves. Paratype. PANAMA. Darien: headwaters of the Rio Tuquesa, gold mining camp of Tyler Kitteridge, ca. 2 air km from the continental divide, 26 Aug. 1974, Croat 27281 (MO). Acknowledgments. I thank the curators of F, MO, NY, and US for allowing us to study their collections. Literature Cited D’Arcy, W. G. 1979. Scrophulariaceae. In: R. E. Woodson & Shery, Flora of Panama. Ann. Missouri Bot.Gard. 77:173-274. [Gibsoniothamnus pp. 220- 227.] Two New Species of Weinmannia (Cunoniaceae) from the Venezuelan Guayana Paul E. Berry and Jason Bradford Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A. Abstract. Two new species of Weinmannia (Cu¬ noniaceae) from the highlands of southern Venezuela are described and illustrated. Weinmannia ilute- puiensis P. E. Berry & J. Bradford is a simple¬ leaved, serrate-margined species known only from Ilu-tepui in Estado Bolivar, and Weinmannia coro- coroensis J. Bradford & P. E. Berry is a pinnate¬ leaved species with tiny, revolute-margined leaflets currently known only from Cerro Coro Coro in Es¬ tado Amazonas. While preparing the floristic treatment of Cu¬ noniaceae for the Flora of the Venezuelan Guayana and as part of a broader study of neotropical Wein¬ mannia by the second author, two distinctive new species of Weinmannia were found among the col¬ lections examined from the tepuis, or tabletop moun¬ tains, of southern Venezuela. Both species were col¬ lected from relatively poorly explored mountains, one from Ilu-tepui in Estado Bolivar, and the other from Cerro Coro Coro in Estado Amazonas. These species are described and illustrated below. Weinmannia ilutepuiensis P. E. Berry & J. Bradford, sp. nov. TYPE: Venezuela. Estado Bolivar: Ilu-tepui, lower plateau, with varied habitats, rocky, boggy, and short forest to 5 m tall, 05°25'03"N, 60°29'W, 2500 m, 16 Apr. 1988, Liesner 23413 (holotype, MO; isotypes, NY, US, VEN). Figure 1A-D. Frutex vel arbor l-5(-10)-metralis, foliis simplicibus, laminis foliorum infra dense ochraceo-lanatis, ellipticis, coriaceis, marginibus serratis, 1. 5-6.0 x 0.8-3. 5 cm; pseudoracemis 4-6 cm longis, lanatis, pedicellis sub an- thesi 1-2.5 mm longis post anthesin 2.5-4 mm longis; capsulis villoso-lanatis, 3-3.5 mm longis. Shrub or small tree 1 -5(-10) m tall; young stems lanate, nodes hirsute. Leaves opposite, simple, ellip¬ tical, subcoriaceous, 1.5-6.0 cm long, 0.8-3.5 cm wide, broadly acute to rounded at the apex, broadly acute at the base, sparsely pilose to glabrescent and dark green on the upper surface, light brown and densely lanate-villous on the lower surface; second¬ ary veins 10-15 per side, impressed on the upper surface and prominent on lower surface, subparallel from midvein until divaricating within ca. 4 mm of the margin with each branch joining an adjacent branch at a marginal sinus; margins serrate, planar to revolute, with 10-15 glandular teeth per side between successive sinuses or adjacent to a sinus; petioles 3-4 mm long, lanate; stipules broadly ellip¬ tical to suborbicular, 3-6 mm long, 3-5 mm wide, hirsute on the outside, dark brown and glabrous adaxially. Pseudoracemes in pairs at the branch apex, exceeding the leaves, 4-6 cm long, axis lanate. Flowers arranged in fascicles along the inflorescence axis with 1-several flowers per fascicle; bracteoles subtending fascicles, linear to obovate, 1.5-2 mm long, with long hairs on the dorsal surface; pedicels 1.0-2.5 mm long in flower, 2.5-4.0 mm long in fruit, lanate; sepals four, elliptic to narrowly deltoid, acute, 1-1.5 mm long, lanate abaxially, glabrous adaxially, persistent; petals four, narrowly obovate, 1.5-2.5 mm long, ca. 1 mm wide, glabrous, off- white; filaments eight, 2.5-4.0 mm long, anthers oblong, ca. 0.4 mm wide, glabrous; ovary lanate, ca. 1.5 mm long, the 2 styles divergent and each 1.5 mm long, basally pubescent but glabrous toward the tips. Capsules 3.0-3.5 mm long (not including style), lanate-villous; seeds oblong, ca. 1.0 mm long, sparsely covered with simple hairs 0.5 mm long. Ecology and distribution. Occurring in patches of short forest on lower summit plateau overlying sandstone, known only from Ilu-tepui in the eastern tepui chain of Estado Bolivar, Venezuela, between 2500 and 2630 m elevation. In the size and shape of leaves, inflorescences, flowers, and fruits, Weinmannia ilutepuiensis most closely resembles the Andean W. rollottii Killip, especially variety testudineata (Cuatrecasas) Ber¬ nards Both species belong to series Dryadifoliae Bernardi (Bernardi, 1963). However, W. ilute¬ puiensis differs in its dense, lanate-villous indumen¬ tum, including the calyx and the capsule (which are glabrous in W. rollottii). On the same helicopter trip to Ilu-tepui during which Ronald Liesner collected the type specimen, he also made three other collections of Weinmannia (numbers 23422, 23417, and 23366, all at MO). None of these additional collections fits easily into Novon 5; 125-127. 1995. 126 Novon D 8 Detail of ^r‘»7T a i au ‘:f ul(,n r P , E - Berr y & J - Bradf °rd- -A. Habit. -B. Flower. -C. Fruit. - LctLL'f Sft.SE? E BSrirSStf ^ 2 f 38 ^nmannia fruit. Drawn from Huber 12296. Ha b.t. F. Deta.1 of leaf pa.r and stipule. -G. Flower. -H. Young Volume 5, Number 2 1995 Berry & Bradford Weinmannia from Venezuela 127 any known species, and they appear to be inter¬ mediates between W. ilutepuiensis and W. bra- chystachya Willdenow ex Engler. Since these two species are so different phenotypically ( W. bra- chystachya is characterized by small, compound, glabrous leaves with pseudo-umbellate inflores¬ cences), the apparent hybrids were only recognized when all Liesner collections from the area were seen together. Weinmannia brachystachya is not yet known from Ilu-tepui, but it does occur on the near¬ by Roraima-tepui. Paratypes. VENEZUELA. Bolivar: Cumbre del Ilu- (Uru-)tepui, sector centro-meridional, vegetacion herba- ceo-fruticosa sobre superficies poco disectadas, 5°25'N, 60°59'W, 2630 m, 29 Apr. 1984, Huber 9520 (MYF, US); Ilu-tepui, lower plateau, with varied habitats, rocky, boggy, and short forest to 5 m tall, 05°25'03"N, 60°29'W, 2500 m, 15 Apr. 1988, Liesner 23338 (MO, US, VEN); Ilu-tepui, cumbre slopes, saddle between N peak and central plateau, 7000-7800 ft., 15 Mar. 1952, Maguire 33426 (NY, US); Ilu-tepui, slopes below uppermost W-facing escarpment, 7500-8000 ft., Maguire 33508 (NY, US). Weinmannia corocoroensis J. Bradford & P. E. Berry, sp. nov. TYPE; Venezuela. Estado Amazonas: cumbre del Cerro Coro-Coro, en las cabeceras nor-occidentales del Rio Manapiare (sector NW de la Serranxa Yutaje), 05°46'N, 66°12'W, 2200 m, 12 Nov. 1987, Otto Huber 12296 (holotype, MO; isotypes, MYF, VEN). Figure 1E-H. A Weinmannia guyanensi Klotzscb foliolis subintegris valde revolutis, foliolo terminali lateralibus simili, inffo- rescentiis congestioribus, pubescentia densiore longio- reque differt. Shrub 0.5-1.5 m tall; young stems villous, the hairs 1-1.5 mm long. Leaves opposite, imparipin- nate, rachis winged, rigid-coriaceous, leaflets 1- or 2-paired (occasionally unifoliate), the terminal and lateral leaflets elliptical to obovate, similar in size and shape, 5—12 mm long, ca. 3—5 mm wide when flattened but lateral margins strongly inrolled, lateral margins entire, apex generally with three small teeth, upper surface glabrous with hairs occasional along midvein, lower surface glabrous except for strigose hairs 1-1.5 mm long along midvein especially to¬ ward the base; secondary veins 6—9 per side, barely visible below, subparallel from midvein until divar¬ icating within ca. 1 mm of the margin with each branch joining an adjacent branch and together forming a vein which extends to the margin; petioles 1-2 mm long, densely villous; stipules obovate, cil- iate, hairs sparse on outer surface, glabrous on inner surface, 2-4 mm long, 1.5-3.5 mm wide. Pseu¬ doracemes in axillary pairs near branch tips, ex¬ ceeding the leaves, 20-30 mm long in flower, axis villous, the basal 8—12 mm sterile, the apical portion densely covered with flowers. Flowers arranged in fascicles along the inflorescence axis with 1-several flowers per fascicle; bracteoles subtending fascicles, rectangular, apex blunt, ciliate, 0.7-1 mm long, 0.5-0.8 mm wide; pedicels 1.5-3.0 mm long in flower, with sparse, shaggy hairs; sepals four, nar¬ rowly deltoid, acute, 1.0-1.5 mm long, hairs on midvein and apex; petals four, obovate, 1.5-2.0 mm long, ca. 1 mm wide, glabrous, white, with a prominent midvein divaricating near the apex; fil¬ aments eight, 2.0-2.8 mm long, anthers oblong, ca. 0.4 mm long, glabrous; gynoecium 1.5-1.8 mm long, glabrous, the two styles divergent, glabrous, ca. 1 mm long. Mature fruits not seen, recorded on labels as being red. Ecology and distribution. Locally frequent in tepui scrub communities overlying sandstone and in high-tepui meadows along streams; known only from the summit of Cerro Coro Coro in Estado Amazonas, Venezuela, at 2200 m elevation. Weinmannia corocoroensis resembles W. guy- anensis Klotzsch, especially variety quinata Cua- trecasas, in its habit and leaf venation pattern, but differs in having denser, longer pubescence along the stems, nodes, and inflorescence structures; smaller, more revolute and nearly entire leaflets; terminal and lateral leaflets of the same size; and shorter, more congested inflorescences. This species was noted in the field by Otto Huber as being com¬ mon, while another specimen ( Huber 12332, MO) from the same location and apparently belonging to W. guyanensis was rare. Weinmannia corocoroen¬ sis is also similar to W. brachystachya Willdenow ex Engler in the few, apical teeth of the leaflets and in its shrubby habit, but differs in its more elongate inflorescence, denser pubescence (on stems, mid¬ veins of leaves, inflorescence axis, and petioles), more regularly elliptical leaflets, and more congested nodes on the stem. Acknowledgments. This work was supported in part by National Science Foundation grant DEB- 9201044. We thank Jose Cuatrecasas for sharing his knowledge of the genus Weinmannia with us and first suggesting that W. ilutepuiensis was a new species. Bruno Manara made the line drawings of the two species, and Roy Gereau kindly reviewed the Latin diagnoses. Literature Cited Bernardi, L. 1963. Revisio generis Weinmannia. Pars II: Sectio Simplicifoliae. Candollea 18: 285-334. New Combinations in Eastern North American Elymus (Poaceae) Julian J. N. Campbell The Nature Conservancy, 642 W. Main St., Lexington, Kentucky 40508, U.S.A. ABSTRACT. Two new varietal combinations are made in the grass genus Elymus: E. glabriflorus (Vasey) Scribner & Ball var. australis (Scribner & Ball) J. J. N. Campbell, and E. villosus Muhlenberg var. arkansanas (Scribner & Ball) J. J. N. Campbell. The following new combinations have been found necessary in preparing the treatment of eastern Ely¬ mus species for the forthcoming Manual of North American Grasses (M. E. Barkworth & K. Capels, editors, in prep.). Elymus glabriflorus (Vasey) Scribner & Ball var. australis (Scribner & Ball) J. J. N. Campbell, comb. nov. Basionym: Elymus australis Scrib¬ ner & Ball, U.S.D.A. Div. Agrostol. Bull. 24: 46. 1901. TYPE: U.S.A. North Carolina: Bilt- more Herbarium 411b (holotype, US). Elymus villosus Muhlenberg var. arkansanas (Scribner & Ball) J. J. N. Campbell, comb. nov. Basionym: Elymus arkansanas Scribner & Ball, U.S.D.A. Div. Agrostol. Bull. 24: 45. 1901. TYPE: U.S.A. Arkansas: Harvey 7 in part (holotype, US). In most recent treatments, these taxa have gen¬ erally been recognized at the forma level or not at all. The morphological differences are largely a mat¬ ter of presence or absence of pubescence in the spike. However, each taxon is locally more frequent than typical plants of the species, and there appear to be some associations with different habitats and some geographic trends. Novon 5: 128. 1995. Festuca diclina (Poaceae), a New Species from Northwestern Mexico Stephen J. Darbyshire Centre for Land and Biological Resources Research, Agriculture and Argi-Food Canada, Wm. Saunders Bldg., Central Experimental Farm, Ottawa, Ontario, K1A 0C6, Canada ABSTRACT. A series of collections from Guadalupe y Calvo in southern Chihuahua State, Mexico, is described as a new species, Festuca diclina (Po¬ aceae), and compared with similar species from Cen¬ tral America. Festuca diclina S. J. Darbyshire, sp. nov. TYPE: Mexico. Chihuahua: Mpio. Guadalupe y Calvo, Cerro de Mohinora, S de Guadalupe y Calvo, 3300 m, alpina-subalpina, en ladera rocosa, 27 Aug. 1987, McDonald & Martinez 2392 (ho- lotype, DAO 661232; isotypes, TEX (ex DAO 591464), MEXU (ex DAO 592610)). Fig¬ ure 1. Plantae perennes caespitosae, innovationibus intrava- ginalibus praeditae. Vaginae foliorum ima basi tantum integrae, vertice scabrae; ligulae 0.2-0.5 mm longae; laminae foliorum plicatae vel convolutae, 1.2-1.5 mm latae, abaxialiter ima basi scabrae, adaxialiter sparsim scabrae, 5-7-costatae, fasciculis sclerenchymaticis cum nervis infra et saepe supra conjunctis, non confluentibus. Culmi 30-75 cm longi, glabri. Paniculae 8-15 cm longae, plus minusve laxae, pyramidales, axe et ramulis glabris vel trichomatibus sparsis, ramulis interdum tortuosis. Spic- ulae 8-12 mm longae, (2-)3-4(-5) flosculis; glumae in- feriores 2.4-4 mm longae, 1-nerviae; glumae superiores 3.5-5 mm longae, (l-)3-nerviae; rachillae trichomatibus 0.1-0.3 mm longae praeditae; lemmata lanceolata, glabra vel apice scabra, l-3(-5)-nervia, 6-8 mm longa, inter¬ dum arista ad 1.0 mm longa; paleae 5.5-7 mm longae, bidentatae, apice piloso-hirsutae inter nervos, nervis cum trichomatibus 0.1-0.2 mm longis prope apicem; antherae interdum abortivae praesertim flosculis inferioribus, 3- 4.2 mm longae; ovariorum apex dense pubescens. Caespitose, loosely tufted perennial (Fig. 1A); shoots intravaginal. Vegetative shoots to 25 cm tall, light green (not glaucous); sheaths rapidly decaying to fibers, the margins closed only at the base, the upper parts of the sheaths, collars, and lower abaxial blade surface retrorsely scabrous; auricles absent; ligules short, 0.2-0.5 mm long, acute to truncate, sometimes lacerate, puberulent apically and abaxi- ally, membranous; blades plicate to convolute, adax- ially sparsely scabrous, 1.2-1.5 mm wide on flat¬ tened blades (about 0.4-0.8 mm in the longest di¬ mension of a cross section), with 7(-9) veins and 5-7 adaxial ribs; sclerenchyma in discrete fascicles opposite veins abaxially and usually adaxially (Fig. 2A-C), the fascicles usually joining veins abaxially and sometimes adaxially thus forming a girder be¬ tween the epidermises through the veins (Fig. 2A). Culms 30-75 cm tall, glabrous; nodes black or brown, glabrous. Flag leaf sheath sometimes loose. Panicle open, pyramidal, 8-15 cm long; rachis glabrous below and sparsely scabrous on the angles above; branches ± flexuous and sometimes sinuously un¬ dulating, 1 or 2 per node, glabrous or sparsely scabrous, the lowest 5-9 cm long. Spikelets 8-12 mm long, distal on the branches, with (2-)3-4(-5) florets; glumes chartaceous, glabrous or scabrous on the keel (especially apically), ovate-lanceolate to lan¬ ceolate with wide hyaline margins; lower glume 2.4- 4 mm long, 1-nerved; upper glume 3.5-5 mm long, (l-)3-nerved; rachilla with hairs 0.1-0.3 mm long (Fig. 1C); lemmas 6-8 mm long, narrowly lanceolate (Fig. IB), glabrous or scabrous-hirsute apically, 1- 3(-5)-nerved (the lateral ones faint), the apex at¬ tenuate, sometimes with a terminal awn or mucro to 1.0 mm long; paleas subequal to the lemmas, 5.5-7 mm long, bidentate, apically pilose-hirsute between the nerves and with hairs 0.1-0.2 mm long on the nerves (Fig. ID, E); anthers sometimes abor¬ tive especially in lower florets, 3-4.2 mm long when developed; ovary apex densely pubescent (Fig. IF); styles 2. Caryopses not seen. Specimen label data from the type collection de¬ scribe the culms as up to 2 m high, although all specimens examined were much less. Distribution. Collections of Festuca diclina are known only from the vicinity of Guadalupe y Calvo, Sierra Madre Occidental, Mexico, in montane pine forests at altitudes of 2950-3300 m. Festuca diclina belongs to Festuca subg. Festuca and is similar to F. hintoniana E. B. Alexeev (Figs. 1G-K, 2D) but differs primarily in the loose tufts with non-persistent sheaths decaying to separate fi¬ bers (densely tufted with persistent chartaceous sheaths in F. hintoniana), scabrous upper sheaths, collars, and lower leaf blades (not glabrous), nar¬ rower leaf blades (1.6-1.9 mm on flattened blades and 0.7-1.0 mm in longest dimension of transverse section in F. hintoniana ), blade sclerenchyma in discrete fascicles (more or less continuous abaxial Novon 5: 129-132.1995. 130 Novon Figures A, G Figures B, D, H, J Figures C, E, F, I, K 1 cm 1 mm 1 mm Volume 5, Number 2 1995 Derbyshire Festuca diclina 131 C D Figure 2. Camera lucida drawings of cross sections of mature leaves from vegetative shoots. A C. Festuca diclina S. J. Darbyshire. —D. F. hintoniana E. B. Alexeev. (A based on McDonald & Martinez 2392 (TEX), B based on McDonald & Nesom 2475 (ARIZ 285274), C based on Nesom & McDonald 6473 (ARIZ 281336), D based on Hinton 17243 (K).) Figure 1. Festuca species. A-F. F. diclina S. j. Darbyshire. —A. Habit. —B. Lower floret. —C. Rachilla. —D. Palea. — E. Palea apex. —F. Ovary. G K. F. hintoniana E. B. Alexeev. —G. Habit. —H. Lower floret. —I. Rachilla. —J. Palea. —K. Palea apex. (A composite, B-F based on McDonald & Nesom 2489 (DAO 592613), G- K based on Hinton 17243 (K).) 132 Novon band in F. hintoniana; Fig. 2D), which extend in¬ ward to meet vascular bundles, sclerenchyma fas¬ cicles of the blades visible from the abaxial surface (invisible or very faint in F. hintoniana ), longer rachilla hairs (0.05-0.07 mm in F. hintoniana; Fig. 11), longer hairs on the palea veins (scabrous prickles 0.02-0.05 mm in F. hintoniana; Fig. IK) and pilose-hirsute apically between the palea veins (gla¬ brous in F. hintoniana). The Mexican species F. coahuilana Gonzalez-Ledesma & S. D. Koch is dis¬ tinguished from F. diclina by the longer ligules (to 1 mm), which form erect auricular projections, wider leaf blades (1.5-2.2 mm) with 8-11 nerves, erect inflorescence branches, shorter lemmas (5.0-6.2 mm), and shorter anthers (2.5-2.7 mm). Two taxa recently described from Costa Rica, F. herrarae Davidse and F. talamancensis Davidse, are both readily distinguished from F. diclina by their gla¬ brous ovaries and shorter anthers (less than 2.1 mm long). Festuca diclina is known from the Sierra Madre Occidental of northwestern Mexico, while both F. hintoniana and F. coahuilana are known only from the Sierra Madre Oriental in northeastern Mexico. This species is so named because the lower florets within a spikelet frequently contain abortive anthers, while the upper florets frequently contain apparently abortive ovaries. Paratypes. MEXICO. Chihuahua: Mpio. Guadalupe y Calvo, NW side of Cerro Mohinora, ca. 13 mi. SW of Guadalupe y Calvo, 25°57'N, 107°03'W, open pine-fir woods with scattered spruce, top and near top of summit, open rocky slopes at lip of precipice with scattered krum- holz [sic] pines, 3250 m, 21 Aug. 1988, McDonald & Nesom 2489 (ARIZ 285277, DAO 592612, 592613); NW side of Cerro Mohinora, ca. 13 mi. SW of Guadalupe y Calvo, 25°57’N, 107°03’W, open pine-fir woods with scattered spruce, top and near top of summit, open rocky slopes at lip of precipice with scattered krumholz pines, 3250 m, 21 Aug. 1988, McDonald & Nesom 2475 (ARIZ 285274, DAO 592611); N side of Cerro Mohi¬ nora, ca. 13 mi. SW of Guadalupe y Calvo, 25°57'N, 107°03'W, open pine-fir woods with scattered spruce, nearly vertical N-facing rock wall, very moist with many bryophytes and rich herbaceous flora, 2950 m, 20 Aug. 1988, Nesom & McDonald 6473 (ARIZ 281336, TEX). Acknowledgments. I am grateful to Andy McDonald for sending material from his collections, J. Cayouette and John and Charlotte Reeder for useful comments, James E. Darbyshire for assistance with the Latin description, Marcel Jomphe for the illustrations, and herbarium curators at ARIZ, DAO, K, and TEX for making specimens available for study. Three New Species of Rustia (Rubiaceae, Condamineeae) from Panama and Ecuador Piero G. Delprete Botany Department, The University of Texas at Austin, Austin, Texas 78713, U.S.A. Abstract. Three new species of Rustia are de¬ scribed, two from Ecuador ( R. alba and R. viri- diflora) and one from Panama (R. dressleri ). Illus¬ trations of the three species are provided. RESUMEN. Se describen tres especies nuevas de Rustia, dos de Ecuador (R. alba y R. viridiflora) y una de Panama (R. dressleri). Se incluyen ilus- traciones de las tres especies. Rustia (Cinchonoideae, Condamineeae; Hooker, 1873; Robbrecht, 1988; Delprete, in prep.) is a genus of trees and shrubs that was founded by Jo¬ hann Friedrich Klotzsch, curator of the Berlin her¬ barium in the mid-1800s. Klotzsch (1846) separated Exostema formosum Chamisso & Schlechtendal to establish Rustia, in honor of Dr. Rust (“Dem An- denken des verstorbenen Prasidenten Dr. Rust gew- idmet”). Rustia, along with Tresanthera H. Kar- sten, is unique in the woody Rubiaceae in having poricidal anthers and pellucid-punctate leaves. Rus¬ tia comprises approximately 15 species endemic to tropical America, from Nicaragua to southern Brazil. The only previous work on Rustia was published by Donald R. Simpson (1976), but this was restricted to only a few species of western South America. During preparation of a monographic treatment of the tribe Condamineeae (Delprete, in prep.), I examined several herbarium specimens from poorly collected areas of Central and South America, many of them unidentified or incorrectly identified. Among these collections, I encountered three species new to science, two from Ecuador and one from Panama. In the following descriptions I use the abbrevia¬ tions L/W to represent the length to width ratio of the leaf blades, and BA to represent the basal angle from the leaf margin to the midrib. Rustia alba Delprete, sp. nov. TYPE: Ecuador. Carchi: environs of Maldonado, wet montane forest, 1450-1650 m, 31 June 1978, M. T. Madison 4808 with T. C. Plowmann, H. A. Kennedy & L. Besse (holotype, AAU; isotypes, F, QCA, SEL not seen, US). Figure 1. Arbores ve! frutices glabri; stipulae interpetiolares cad- ucae. Foliorum laminae 23-28 cm longae, 13-16 cm latae, glanduli-punctatae; domatia desunt. Inflorescentia 40-46 cm longa; alabastra floralia cylindrica aestivatione valvati-reduplicata; calyx cupulatus ac membranaceus 1.5- 3.0 mm longus; corolla alba 1.5-1.9 cm longa ad basim constricta lobis % longitudinis corollae; antherae exsertae dorsifixae concavae poricidales, thecae ad basim subae- qualis; filamentis basiliter dilatatis ascendenti-strigosisque. Capsulas non visi. Shrub or tree, with several straight trunks 6 m tall (probably stump sprouts). Young branches smooth, glabrous, terete; older branches rugose, te¬ rete, grayish to pale brown and with sparse lenticels. Stipules interpetiolar, free at base, present only in bud and overlapping, narrowly triangular, glabrous outside, glabrous and with few basal colleters inside, 2.8-3.7 mm long, 7-9 mm wide at base, reddish green, leaving a linear scar ca. 0.5 mm wide of the same color as the stem. Leaves petiolate, petioles 2-3 cm long, 3.0-4.5 mm thick, adaxially narrowly concave to flattened, glabrous; basally thickened, not pulvinate; blade 23-28 cm long, 13-16 cm wide, L/W = 1.6:1 to 1.7:1; widely elliptic to widely obovate, acute to rounded at base (BA = 30-45°), obtuse and short-acuminate at apex; the acumen up to 1 cm long; blade when fresh pale green above and yellowish green below, subcoria- ceous, drying pale green, glabrous above and below, pellucid punctate (see Fig. 1). Primary and second¬ ary veins glabrous and prominent below, secondary veins 16-21 each side; tertiary veins starting sub¬ parallel and openly reticulate in the center, faintly evident above and below; domatia absent. Inflores¬ cences terminal, laxly paniculate and pyramidal, 40-46 cm long, basal branches 22-30 cm long, lateral branches 6-8 pairs, decussate; basal portion of axis not branched, 7-11 cm long. Rachis terete, rachis and branches glabrous to minutely hirtellous; flowers on distal branches in cymules (rarely alter¬ nate). Distal bracts 4-5 mm long, 4-5 mm wide, deltoid; bracteoles subtending the flowers 2-3 mm long, 2-3 mm wide, deltoid, glabrous. Flowers pro- tandrous, pedicellate, pedicel 4-6 mm long, glabrous to minutely puberulent; hypanthium narrowly ob- conical, 4-6 mm long, 2-3 mm wide, glabrous. Calyx cupular with small lobes, membranous and Novon 5: 133-139. 1995. / cm 134 Novon Figure 1. Rustia alba Delprete. —A. Habit of inflorescence with mature leaf. —B. Abaxial view of blade showing peUueid punctation. —C. Flower bud showing cupular calyx. —D. Anther. —E. Detail of anther external surface. — F. Style. (Drawn from type: Madison et al. 4808.) 2 mm Volume 5, Number 2 1995 Delprete Rustia from Panama and Ecuador 135 glabrous, 1.5-3.0 mm long, calyx lobes 5, 1.0-2.5 mm long and 1.5-2.0 mm wide at base, glabrous and ciliolate; calyx persistent. Corolla tubular, 1.5- 1.9 cm long, with a basal constriction and reflexed lobes, flower bud cylindrical and pointed at apex, aestivation valvate with contact zone, white, and semifleshy when fresh; tube subcylindrical with basal constriction, 5-6 mm long, ca. 2 mm wide at base and 3.5-4.0 mm wide at the orifice; glabrous and minutely verrucose outside; microscopically (40 x) granulose inside, without a ring of hairs; corolla lobes 5, 9-11 mm long, ca. 3 mm wide, narrowly tri¬ angular and reflexed, glabrous and minutely ver¬ rucose outside; densely ascending short-strigose at base, minutely papillose throughout and microscop¬ ically (40 X) papillose at margins inside. Lobes % of corolla length. Stamens 5, fully exserted, subequal, filaments attached 4-5 mm from the base of the tube, anthers cylindrical and subsagittate at base (see Fig. 2), rectilinear to slightly curved, 6.5-7.0 mm long, ca. 1.5 mm wide, dorsifixed near the base, the thecae subequal at base, papillose throughout, dehiscing by two pores at apex; filaments ca. 3 mm long, flattened and widened at base, with a basal tuft of ascending strigose hairs 0.3-0.5 mm long. Style exserted, 16-20 mm long, glabrous, style branches ovate, 3.0-3.5 mm long and 1.0-1.3 mm wide at base, stigmatic surface papillose. Ovary two- celled, placentation axile, turbinate to narrowly ob¬ long, glabrous, with many ovules in each locule horizontally inserted. Capsule not seen, but probably obconical to obovoid, like the hypanthium. Rustia alba is similar to R. formosa (Chamisso & Schlechtendal) Klotzsch but differs mainly in hav¬ ing a bigger inflorescence (exceptionally up to 36 cm long in R. formosa), smaller flowers; calyx cu¬ pular, more expanded and membranous; shorter styles; and longer membranous style lobes (ca. 3 mm long in R. alba vs. ca. 1 mm long in R. formosa ). Rustia alba is known only from the type, which was collected in the cloud forest of northern Ec¬ uador, around Maldonado; in contrast, R. formosa is endemic to central and southern Brazil. Rustia alba is easily distinguishable from all other species of Rustia in western South America by its white erect flowers (hence the epithet) and by its widely obovate leaves. Rustia viridiflora Delprete, sp. nov. TYPE: Ec¬ uador. Morona-Santiago: Achutza, Jibaro set¬ tlement in the vicinity of Macuma, ca. 50 km N of Macas, 23 Mar. 1973, H. Lugo 3674 (holotype, GB sheets A and B). Figure 2. Arbores usque ad 16 m altae; stipulae interpetiolares caducae. Foliorum laminae 34-62 cm longae, 9.5-20.5 cm latae, glandulis punctatae; domatia desunt. Inflores- centia 22-54 cm longa, paniculata et secundiflora; ala- bastra floralia cylindrica acuminata, aestivatione valvati- reduplicata; calyx valde redactus 0.3-0.5 mm longus truncatus; corolla viridis 1.9-2.2 cm longa, lobis longitudinis corollae ad basim bulbosa; antherae rectae pilosae inclusae poricidales basaliter subcaudatae dorsifix- ae prope basim; filamenta tereta ad basim aurei-pilosa. Capsulae angustissime obconicae 18-23 mm longae disco conico. Tree 6-16 m tall. Young branches glabrous to minutely puberulent, terete; older branches becom¬ ing glabrous, rugulose, terete, grayish to pale brown and with sparse punctiform to linear lenticels, up to 1.0 cm long. Stipules not seen, probably interpe- tiolar and free at base (observed from stem scars), leaving a linear scar 1.0-1.5 mm wide, of the same color as stem. Leaves petiolate, petiole 4.5-7.0 cm long, 3-5 mm thick, subterete, glabrous; basally thickened, not pulvinated; blade 34-62 cm long, 9.5-20.5 cm wide, L/W 3:1 to 3.5:1; lanceolate to elliptic, acute to obtuse (BA = 35-65°) at base, acute to attenuate at apex, sometimes tapering to an acumen 2-3 cm long; blade when fresh dark green above and dull green below; drying pale brown, stilf-chartaceous; glabrous above and below, pellucid punctate (see Fig. 2). Primary and secondary veins glabrous and prominent below, secondary veins 19- 25 each side; tertiary veins starting subparallel and openly reticulate in the center, faintly evident above and fairly evident below; domatia absent. Inflores¬ cences terminal, 22-54 cm long, 9-22 cm wide at base, laxly paniculate and pyramidal, with decussate racemoid lateral branches; lateral branches 3-7 pairs, basal portion of axis not branched, 10-20 cm long. Rachis terete to decussately obtuse-compressed, ra- chis and branches glabrous to minutely puberulent; flowers on distal racemoid branches or on racemoid cymules. Bracts subtending lateral branches 1-3 mm long, 2-3 mm wide, deltoid; bracteoles sub¬ tending the flowers 0.5-1.0 mm long, 0.7-1.0 mm wide, deltoid, glabrous. Flowers protandrous, pedi¬ cellate, pedicel 6-9 mm long, glabrous to minutely puberulent; hypanthium narrowly obconical, 4-6 mm long, 2-4 mm wide, glabrous to minutely golden hirtellous. Calyx reduced to a wavy margin with barely distinguishable lobes, 0.3-0.5 mm long and persistent. Corolla tubular with bulbous base and small constriction just above, 1.9-2.2 cm long, flow¬ er bud cylindrical, aestivation valvate with contact zone, color green throughout (rarely reddish with green lobes), semifleshy when fresh; tube subcylindr¬ ical and striate, 1.4-1.8 cm long, 4-5 mm wide at base and 2.0-2.5 mm wide at the orifice, glabrous 136 Novon show n. f v " ld jfi° ra Delprete. -A Habit of inflorescence with mature leaves. -B. Abaxial view of blade showing 7 C ['?' er bu d showing bulbous base. -D. Anther. -E. Style. -F. Mature capsule OTmT " ° type: 80 3674 ' SheetS A and B; capsule drawnfr ° m paratype: F. Prieto Volume 5, Number 2 1995 Delprete Rustia from Panama and Ecuador 137 or golden hirtellous outside; ascending strigose at base inside, without a ring of hairs, the remaining medial and superior zone glabrous or with sparse hairs; corolla lobes 5, 3-5 mm long, ca. 2.5 mm wide at base, triangular to ovate, glabrous outside, glabrous and with minute glandular hairs (hairs up to 0.5 mm long) at margins inside. Lobes V t to % of corolla length. Stamens 5, included (only the very tip exserted), subequal, filaments attached 3.5-5.5 mm from the base of the tube, anthers elongated and rectilinear, 11-12 mm long, 1.2—1.3 mm wide, dorsifixed near the base, base subcaudate, with sharp ridges, papillose-echinate to ascending-strigulose and descending glandular-pilose at base; dehiscing by two pores at apex; filaments 6-7 mm long, sparse as¬ cending-pilose and with a tuft of golden pilose hairs at base. Style exserted, 22-28 mm long, glabrous and minutely verrucose throughout, style branches ovate and acute at apex, 1.0-1.3 mm long, stigmatic surface smooth to minutely papillose. Ovary two- celled, placentation axile, turbinate to narrowly ob- conical, glabrous, with many ovules in each locule horizontally inserted. Immature fruits green to red¬ dish. Capsules very narrowly obconical (see Fig. 2), apex shortly conical, (14—)18—23 mm long, 4-6 mm wide, black and without lenticels, capsule and disk black and glabrous, disk obviously exceeding the calyx; dehiscing loculicidally, disk-septicidal de¬ hiscence absent even in old capsules. Rustia viridiflora is unique in the genus in having basally bulbous green flowers and capsules very nar¬ rowly obconical with a conical disk. It is known from the eastern slopes of the southern Ecuadorian Andes (Prov. Morona-Santiago), at the western margin of the Amazon forest. Paratype. ECUADOR. Morona-Santiago: low hills W of Rio Chupiangas, 20-22 Nov. 1944, F. Prieto CHUP-20 (NY [2], US). Rustia dressleri Delprete, sp. nov. TYPE: Pan¬ ama. Panama: El Llano-Carti Highway, 8-10 km N of El Llano, 31 Aug. 1974, R. Dressier 4703 (holotype, MO). Figure 3. Arbores vel frutices; stipulae interpetiolares caducae. Laminae 30-45 cm longae, 9.5-13.0 cm latae, glanduli- punctatae; domatia desunt vel tricomata pauca brevia praesentia. Inflorescentia 12-21 cm longa paniculata; alabastra floralia clavata, aestivatione valvati-reduplicata; calyx valde redactus ca. 0.5 mm longus truncatus; corolla alba 1.2-1.4 cm longa lobis l / 2 longitudinis corollae; an- therae exsertae poricidales concavae basaliter rotundatae dorsifixae prope basim; filamenta basaliter dilatata ad bas- im descendens-pilosa. Capsulas non visi. Shrub or tree (specimen collected from fallen branches). Young branches grayish, glabrous, te¬ rete; older branches glabrous, rugose, terete, pale brown-grayish, lenticels absent; leafy branchlets semisucculent, 7-9 mm wide. Stipules not seen, interpetiolar, free at base (observed from stem scars), leaving a linear scar ca. 1 mm wide, of the same color as stem. Leaves petiolate, petioles 28-32 mm long, ca. 3.0 mm thick, adaxially concave to flat¬ tened, glabrous; thickened and becoming corky at base; blade 30-45 cm long, 9.5-13.0 cm wide, L/ W 3:1 to 4:1; narrowly elliptic to oblanceolate, cuneate (BA = 20-25°) at base, acute and acu¬ minate at apex, the acumen ca. 1 cm long; blade when fresh shiny grass-green above and pale grayish green below, subcoriaceous; drying olive-green, semi- coriaceous; glabrous above and below; evidently pel¬ lucid punctate (see Fig. 3). Primary and second¬ ary veins glabrous and prominent below, secondary veins 24-30 each side, subparallel; tertiary veins starting subparallel and openly reticulate in the cen¬ ter, very evident above and faintly evident below; domatia absent or a tuft of few hairs. Inflorescences terminal, 12-21 cm long, reduced panicles with opposite decussate branches; basal branches 6.5- 11.0 cm long, lateral branches 4-5 pairs, basal portion of axis not branched, 0.9-2.0 cm long. Rachis decussately compressed and terete at base, rachis and branches semisucculent, glabrous throughout and moderately puberulent at nodes; flowers in opposite cymules on rachis and lateral branches. Distal bracts subtending lateral branches 2.5-3.0 mm long, 3-4 mm wide, deltoid, glabrous; bracteoles subtending the flowers 1.0-1.5 mm long, 1-2 mm wide, deltoid, glabrous. Flowers protan- drous, subsessile or pedicellate, pedicel 0.5-4 mm long, glabrous; hypanthium obconical, 2.5-3.5 mm long, 2.0-2.5 mm wide, glabrous. Calyx extremely reduced, with a wavy margin or with barely distin¬ guishable lobes, ca. 0.5 mm long, sometimes cilio- late, persistent. Corolla tubular with spreading lobes, 1.2-1.4 cm long; flower bud clavate, ca. 2.5 mm at base and 3.5-4.0 mm at bulge; aestivation valvate with contact zone, color white, semifleshy when fresh; tube 7-8 mm long, 2.5 mm wide at base and 3.5- 4 mm wide at the orifice, glabrous outside, glabrous inside and with a ring of white-pilose retrorse to erect hairs, 5-6 mm from the base, at the point of attachment of the filaments, which is also the base of the corolla lobes; corolla lobes 5, 5-6 mm long, ca. 3 mm wide, triangular, semifleshy, glabrous out¬ side, white-strigose at base and margins microscop¬ ically (40 x) papillose inside. Lobes ca. V 2 of corolla length. Stamens 5, partially exserted because of narrowly spreading lobes, subequal, filaments at¬ tached 5-6 mm from the base of the tube, anthers concave, yellow and banana-shaped, 4-5 mm long. 138 Novon Figure 3. Rustia dressleri Delprete. —A. Habit of inflorescence with mature leaf. —B. Abaxial view of blade showing pellucid punctation. C. Detail of open flower showing callous base and descending pilose tufts. —D. Flower bud. (Drawn from holotype: Dressier 4703.) 1.8-2.0 mm wide, dorsifixed near the base, bases of thecae rounded and subequal, microscopically (40 x) papillose-echinate throughout, dehiscing by two pores at apex; filaments ca. 2 mm long, distally terete and basally flattened, adnate to tube, glabrous throughout and barbate at base, with a tuft of de¬ scending white-pilose hairs 0.1-0.2 mm long (see Fig. 3). Style exserted, 13-16 mm long, glabrous, style branches ovate to narrowly ovate, ca. 1.5 mm long, stigmatic surface smooth. Ovary two-celled, placentation axile, obconical, with many ovules in each locule horizontally inserted. Capsule not seen. Volume 5, Number 2 1995 Delprete Rustia from Panama and Ecuador 139 Rustia dressleri is very closely related to R. occidentalis (Bentham) Hemsley in having fleshy clavate buds, but differs in having white flowers (flesh-red to purple in R. occidentalis), filaments with basal tuft of descending white-pilose hairs (pu- berulent in R. occidentalis), larger inflorescences (up to 11 cm long in R. occidentalis), young bran- chlets thick and semisucculent, the whole plant gla¬ brous, leaf blades 30-45 cm long (exceptionally up to 30 cm in R. occidentalis), and with 24-30 secondary veins each side (11-16 in R. occiden¬ talis). In his treatment of Rubiaceae for the Flora of Panama, Dwyer (1980) erroneously cited the para- type of Rustia dressleri (Dressier 4749) as R. panamensis Dwyer. The latter was subsequently reduced to synonymy under the new combination R. costaricensis (Standley) Lorence (Burger & Tay¬ lor, 1993). Rustia dressleri differs from R. costaricensis in its inflorescence 12-21 cm long (vs. 20-33 cm long in R. costaricensis), twice compound (vs. thrice compound in R. costaricensis), with 4-5 pairs of lateral branches (vs. 8-11 pairs in R. costaricensis), flower bud clavate (vs. pointed in R. costaricensis), corolla tubular (vs. cupular in R. costaricensis) and 12-14 mm long (vs. 3-4 mm long in R. costari¬ censis). Known only from two collections, this species was first encountered by Robert L. Dressier as fallen branches along a stream. Dressier is a well-known orchidologist, and I am pleased to honor him with this eponym. Paratype. PANAMA. Panama: El Llano-Carti High¬ way, 10-12 km N of El Llano, 12 Sep. 1974, R. Dressier 4749 (MO). Acknowledgments. I am grateful to the directors and staff of the following institutions for the loan of material and for help in fieldwork: AAU, B, BR, CAS, CHOCO, COL, CR, DAV, DS, F, G, GB, GH, JBGP, JEPS, K, LL, MO, NY, P, QCA, QCNE, S, TEX, UC, US. I thank Lennart Andersson, John Dwyer, and Charlotte Taylor for the stimuli toward my research. I especially thank Billie Turner and Guy Nesom for reviewing the manuscript, and David Lorence for helpful comments. Guy Nesom also helped me with the Latin descriptions. The illustra¬ tions are provided by the author. Literature Cited Burger, W. & C. M. Taylor. 1993. Rubiaceae. In: Flora Costaricensis. Fieldiana, Bot. 33: 306-307. Dwyer, J. D. 1980. Rubiaceae. In: Flora of Panama. Ann. Missouri Bot. Gard. 67: 483-485. Hooker, J. D. 1873. Rubiaceae. In: G. Bentham & J. D. Hooker, Genera Plantarum 2: 7-151. Klotzsch, J. F. 1846. In: F. G. Hayne, Getreue Dar- stellung und Beschreibung der in der Arzneykunde Gebraiichlichen Gewachse. 14: 14-15. Nicolai, Ber¬ lin. Robbrecht, E. 1988. Tropical Woody Rubiaceae. Opera Bot. Belgica 1: 1-271. Simpson, R. D. 1976. Studies in neotropical Rubiaceae. Rustia. Phytologia 33: 4-8. New Orchid Species from Costa Rica Robert L. Dressier Missouri Botanical Garden. Mailing address: Rt. 2, Box 565C, Micanopy, Florida 32667, U.S.A. Abstract. Five new orchid species from Costa Rica are described and illustrated from living or liquid-preserved material: Encyclia ortizii, Malaxis monsviridis, M. talamancana, Sobralia dissimilis, and S. doremiliae. The orthography of Sobralia “corazoi ” is discussed, and S. carazoi is thought to be correct. Costa Rica has a long tradition of resident nat¬ uralists and orchidophiles collecting and pressing plants for study. The orchid flora of Costa Rica is thus much better sampled than those of neighboring countries. Even so, the preparation of the orchid treatment for Manual de la Flora de Costa Rica continues to reveal unnamed species, especially among terrestrials and smaller-flowered epiphytes. Five of these are here named and described. Encyclia ortizii Dressier, sp. nov. TYPE: Costa Rica. Prov. Alajuela: Reserva San Ramon, aprox. 30 km NNO de San Ramon, 8, 9 die. 1984, R. L. Dressier y Biologta 350 no. 288 (holotype, USJ). Figure 1. Pseudobulbi pyriforme-ovoidei unifoliati; folia oblan- ceolato-oblonga; sepala oblongo-oblanceolata acuta; petala anguste oblanceolata; labello unguiculato trilobato lobi laterales parvi oblongi lobo intermedio subquadrato-ovato callo alto trigono carinato ornato. Epiphytic, caespitose; pseudobulbs unifoliate, narrowly pyriform-ovate, somewhat flattened, 3.7 x 0.8-1.2 cm; leaves oblanceolate-oblong, nar¬ rowed basally, obtuse to subacute, 8.2-10 x 2.3- 2.6 cm; inflorescence racemose, 12-24 cm, bracts deltoid, acute, 3-4 x 1.8-2 mm; pedicel with ovary 10-12 mm; sepals oblong-oblanceolate, acute, 9 x 2.8-3 mm; petals narrowly oblanceolate, 8x2 mm; lip unguiculate, claw ca. 4 mm, basally adnate to column, blade 3-lobed, lateral lobes oblong, por- rect, clasping column apex, 2 x 1.3 mm; midlobe at right angle to column, ovate, 4 x 3.8 mm, callus prominent, trigonous, 1.5 x 1.5 mm, with laminar keel ca. 1.3 mm high; column ca. 7 mm, dorsal profile concave, midtooth ovate-elliptic, 1.7 x 0.8 mm, surpassing anther and lateral teeth, lateral teeth deltoid, 0.8 x 0.6 mm. Encyclia ortizii is a member of subgenus Os- mophyta and is closely allied to E. campylostalix (Reichenbach f.) Schlechter, E. fortunae Dressier, and E. glauca (Knowles & Westcott) Dressier & Pollard. The relatively slender column resembles that of E. campylostalix, while the sharply defined, rounded lateral lobes of the lip resemble those of E. glauca and E. fortunae. It is quite distinctive in the high, keeled callus and the orientation of the lip. The epithet honors Rodolfo Ortiz, director of the San Ramon nature reserve, in whose company the type material was collected. Malaxis monsviridis Dressier, sp. nov. TYPE: Costa Rica. Puntarenas: Monteverde Reserve, near Alajuela boundary, Cerros Centinelas, elev. 1550-1600 m, 10°18'N, 84°47'W, 19 Aug. 1984, M. H. Grayum & P. Sleeper 3842 (holotype, MO). Figure 2A, B. Herba perennis erecta; pseudobulbis epigaeis ovato- conicis; folia duo late ovata breviter acuminata; inflores- centia subumbellata; bracteae deltoideo-lanceolatae; se¬ pala oblongo-ovata obtusa; petala filiformia; labello subquadrato profunde hastato apice subquadrato trilob- ulato profunde excavato. Pseudobulbs above ground, 1.5-3 x 0.6-1.2 cm, approximate or up to 3.5 cm apart on rhizome. Leaves 2, petioles 5-6 cm, blades broadly ovate, short-acuminate, 5-7.5 x 3.2-5 cm. Inflorescence subumbellate, scape 10-15 cm, floral bracts deltoid- lanceolate, to 1.5 mm; pedicel with ovary 6.5-10 mm; sepals oblong-ovate, obtuse, dorsal sepal 3-3.5 x 1.3-1.6 mm, lateral sepals 2.7-3 x 2mm; petals filiform, 2-3 mm; lip hastate, subquadrate, 2.5-3 x 2-2.6 mm, basal auricles subdeltoid, retrorse, prominent; cavities 2, about half length of blade, cavity apically projecting under blade of lip; apex of lip 3-lobulate, lobules subequal, lateral lobes rounded, midlobule rounded or deltoid; column about 1 mm long; color: pale green or lip brownish green. Flowering July through September. Malaxis monsviridis may be recognized by the prominently hastate lip with short, deep cavities and wide, trilobulate apex. Both M. carpinterae (Schlechter) Ames and M. hastilabia (Reichenbach Novon 5: 140-145. 1995. Volume 5, Number 2 1995 Dressier Orchids from Costa Rica 141 view. B no. 288). —A. Flower. —B. Column and lip, lateral lateral view. 142 Novon f.) Kuntze have lips with retrorse lobules and tri- lobulate apices, but the apices are much narrower and the cavity either narrower or longer. Paratypes. COSTA RICA. All from the area of Monteverde: J. T. Atwood 89-319 (SEL), K. Barrin¬ ger,, E. Christenson & B. Perez 4168 (F), V. J. Dryer 699 (F), W. A. Haber & E. Bello C. 2423 (MO), W. Haber & W. Zuchowski 9315 (MO), ft. 0. Lawton 1329 (F). Malaxis talamancana Dressier, sp. nov. TYPE: Costa Rica. San Jose: Villa Mills, Cordillera de Talamanca, ca. 3000 m, 28 mar. 1993, R. L. Dressier & D. E. Mora s.n. (holotype, USJ). Figure 2C, D. Herba perennis, erecta; pseudobulbis epigaeis, conicis; folia duo deltoideo-ovata cuneata vel subcordata breviter acuminata; inilorescentia subumbellata; bracteae delto- ideo-lanceolateae; sepala ovata vel elliptica obtusa; petala sublinearia; labello deltoideo-ovato hastato ad apicem min¬ ute tridentato auricula retrorsa deltoidea vel denticulata. Terrestrial, pseudobulbs above ground, conic, 2- 4.5 x 0.8-2 cm, to 2.5 cm apart on rhizome. Leaves 2, petioles 5-7 cm; blades deltoid-ovate, cuneate to subcordate, apices short-acuminate, 5.5- 8 x 3-4.2 cm. Inflorescence subumbellate, scape 15-20 cm, bracts deltoid-lanceolate, ca. 2 x 1 mm, ovary with pedicel 17-21 mm; sepals ovate to el¬ liptic, narrowly obtuse, 3-6 X 1.5-2.3 mm; petals sublinear, 3-4 x 0.6 mm; lip 3-5 x 2-2.5 mm (in natural position), hastate, deltoid-ovate, apex mi¬ nutely 3-dentate, lateral lobules folded upward, au¬ ricles retrorse, rounded, deltoid or denticulate, 0.8- 1 x 0.8 mm; basal cavity deep, 1.5-2 mm, with rounded median keel, distal margin emergent. Malaxis talamancana may be recognized by the deltoid-ovate, hastate lip minutely 3-dentate api- cally. Superficially, the flowers are similar to those of M. brachyrrhynchos (Reichenbach f.) Ames, but that species has subterranean corms and the details of the lip are quite distinct. Paratypes. COSTA RICA. San Jose: trail from Ca¬ naan to Cerro Chirripo, between 2000 and 3000 m elev., 20 & 25 Aug. 1971, W. C. Burger & L. Gomez P. 8358 (F). Limon: Cordillera de Talamanca, Atlantic slope, between Rio Terbi and the Rio Sini, elev. 2400- 2750 m, 13 Sep. 1984, G. Davidse , G. Herrera Ch. & M. H. Grayum 29048 (MO). Sobralia dissimilis Dressier, sp. nov. TYPE: Cos¬ ta Rica. Puntarenas: Monteverde Reserve, TV towers at 1750 m elev., 26 June 1989, J. T. Atwood 89-243 (holotype, MO; isotypes, CR, FLAS, SEL). Figure 3. Herba epiphytica; folia elliptica; flores e bracteis erectis imbricantibus nonullis erumpentis; sepala ovato-oblonga obtusa mucronata; petala ovato-oblonga; labellum ovatum margine fortiter undulatum; discus carinis biformis un- decim ornatus. Epiphytic, caespitose, stems stout, 35-75 cm; sheaths glabrous, verruculose. Leaves several, ellip¬ tic, acuminate, 13-15 x 4.5-5.2 cm, thin, not markedly corrugated. Inflorescence bract cluster el¬ lipsoid, 5-6 cm, loose, glabrous, with 2 flowers at once; sepals ovate-oblong, obtuse, mucronate, 3.1- 3.3 X 1.4 cm, lateral sepals basally connate for 6- 7 mm; petals oblong-ovate, 3.1 x 1.3 cm; lip ovate, about 3 x 2.3 cm, tube about 1 cm, then flaring, exposing column, with 2 fleshy calli about 8 mm from base, with 11 keels from near center, outer 6 keels high, curved, not undulate, outermost short, next intermediate, then reaching to within 5 mm of apex, inner 5 keels strongly undulate, reaching 2- 5 mm from apex; column 2.5 cm, basally slender, column wings narrowly deltoid, 4.5 x 3 mm, an- trorse. The epithet dissimilis , or different, refers both to the fact that the species is strikingly distinct from all others and to the two different types of keel on the Up. Sobralia dissimilis is clearly a close ally of S. undatocarinata C. Schweinfurth, of which there is an excellent photograph in Orquidea 5: 209, 1975 (as S. mucronata Ames & C. Schweinfurth; Horich, 1975). Sobralia dissimilis is quite distinct in the shorter, rounded limb of the lip (not pandurate or deeply bifid), in the wider, antrorse column wings, and in the two sharply different types of keel on the lip. Sobralia doremiliae Dressier, sp. nov. TYPE: Costa Rica. Cartago: slopes and steep roadside banks along the Rio Grande de Orosi about 11 km SSE of bridge at Tapanti, elev. about 1500 m, 10 July 1977, R. L. Wilbur, F. Almeda ni Service Forestier de Madagascar SF 27670, and G and H from Service Forestier de Madagascar SF 10027. ) Species apice foliolorum contracto, foliis et leguminibus glabris, et inflorescentiis cum axibus secundariis brevibus et gracilibus pares apicales florum gerentibus distincta. A deciduous tree 4-8 m tall, flowering on young, developing shoots; [DBH not known]; twigs glabrous wih raised lenticels and small, spherical dormant buds. Leaves with 5-11 opposite leaflets; rachis glabrous; stipels absent. Leaflets ovate, 25-55 x 15-40 mm, obtuse to rounded basally, the apex acuminate and tapering into a short to long drip tip, finally obtuse and mucronate, entirely glabrous, pal¬ er beneath. Pseudoracemes delicate, ca. 4-6 cm king, from the leaf axils near the base of new growths, the flowers toward the base of the inflorescence paired on short, slender secondary axes up to 6 mm lung; pedicels with a pair of minute, filiform brac- teol B s pear the middle which are caducous before the flower opens. Flowers ca. 12-15 mm long, violet. Calyx 3~4 mm long, subglabrous with a minutely c iliate margin, with 5 short, broadly triangular teeth, Volume 5, Number 2 1995 Labat & Du Puy Millettieae from Madagascar 173 the upper pair smallest and close together. Standard limb ca. 10 x 10 mm, cuneate basally, finely and sparsely appressed pubescent behind. Wings as long as the keel. Staminal sheath ca. 9 mm long. Pods obovate to oblong-obovate, flat, 65-80 x 15-20 mm, tapering toward the base, the apex rounded and beaked, glabrous, splitting into 2 twisted, woody valves, with 1-4 seeds. Seeds oblong-discoid, ca. 10 x 10 mm, chestnut brown, the aril extended into a short flap on one side. Millettia capuronii is recognizable by its tapering leaflet apices, its glabrous leaves and pods, and its distinctive inflorescences with short, slender sec¬ ondary axes bearing apical pairs of flowers. Distribution. Eastern Madagascar, collected in¬ frequently but in widespread localities along the east¬ ern coastal region including Fort Dauphin (Taolan- aro), Farafangana, and Sambava. Habitat. Lowland evergreen forest near the coast, at low altitudes. Flowering time. Only recorded in April. Vernacular name. Sikidihazo. This species is dedicated to Rene Capuron, col¬ lector of the type specimen, in recognition of his outstanding contributions to the knowledge of the forest flora of Madagascar. Paratypes. MADAGASCAR. Prov. de Toliara, Pre¬ fecture Taolaiiaro (Fort Dauphin), foret a 5 km de Ste. Luce, au N de Maliaforaky, 24°46'S, 47°09'E, 0-10 m, 28 Avr. 1989(st), Rabevohitra, Dumetz & Gereau 1946 (MO, P, TAN); Prov. de Toliara, Prefecture de Fort Dauphin, foret ombrophile de Manatantely, 50-250 m, 9 Nov. 1990 (jfr), Rabevohitra 2426 (MO, P, TAN); Dct. de Farafangana, Andafia, 20 Avr. 1953 (fr), Service Forestier de Madagascar 10027-SF (P, TEF). Millettia hitsika Du Puy & Labat, sp. nov. TYPE: E Madagascar. Foret d’Analalava, Fenoarivo, 4 km W de Foulpointe, 20—50 m, 3 Dec. 1958 (fl). Service Forestier de Madagascar 20154- SF, R. Capuron (holotype, P; isotypes, K, L, MO, P, TEF). Figure 2. Species habitu tenui et delicato et foliolis cum apice acuto, supra glabris et subtus appresso-pubescentibus dis- tincta; a M. orientalis differt foliis, foliolis, inflorescentibus at floribus magnopere parvioribus. A deciduous tree or shrub to 10 m tall, flowering on new, developing shoots soon after they emerge from the buds; DBH 18 cm (one record only); twigs brownish pubescent when young, glabrescent with rounded dormant buds when mature. Leaves with 9-13 opposite leaflets; rachis pubescent when young; stipels absent. Immature leaflets narrowly elliptic, [mature leaflets not known], acute and mucronate apically, glabrous except for the midvein above, densely covered in short, appressed hairs beneath. Flowers in delicate pseudoracemes 4-9.5 cm long from the lowest nodes and leaf axils on very young shoots with developing leaves, the flowers solitary or paired on very short side spurs up to 1.5 mm long; pedicels with a pair of very small, filiform bracteoles up to 3 mm long which are caducous before the flower opens. Flowers 10-12 mm long, [pink or violet]. Calyx 2.5-4 mm long, brownish pubescent, with 5 short, broadly triangular teeth, the upper pair smallest and close together. Standard limb ca. 10 x 10 mm, thinly pubescent behind. Wings as long as the keel. Staminal sheath 8-9 mm long. [Pods and seeds not known.] Millettia hitsika is characterized by its fine, del¬ icate habit and by its narrow (immature) leaflets with acute apices, which are glabrous above and ap- pressed-pubescent beneath. It is only known in flow¬ er with immature foliage, in which it somewhat re¬ sembles Pongamiopsis pervilleana (Baillon) R. Vi- guier. It is easily distinguished from M. orientalis, the only other Millettia species that occurs in the humid, eastern forests, a much more robust species with much larger leaves, leaflets, inflorescences, and flowers. Distribution. Eastern Madagascar, on the coastal plain, from Ambila-Lemaitso to Foulpointe. Habitat. In evergreen forest on sand, at up to ca. 50 m altitude. Flowering time. October-March. Local uses. The wood is used in construction. Vernacular name. Hitsika. The species name is derived from the vernacular name for this species. Paratypes. MADAGASCAR. Ambila Lemaitso, 13 Mar. 1951 (fl), Service Forestier de Madagascar 3260-SF (K, P, TEF); Ambila Lemaitso, 15 Jan. 1953 (fl). Service Forestier de Madagascar 7257-SF (P, TEF); Dct. de Brickaville, Ambila Lemaitso, sous parcelle F2K, 20 Fev. 1958 (fl), Service Forestier de Madagascar 19002-SF (P, TEF). Millettia nathaliae Du Puy & Labat, sp. nov. TYPE: N Madagascar. Track to Irodo along the northern ridge above the Irodo River, ca. 5 km E of Route Nationale 6 to Antsiranana (Diego-Suarez), the nearest village Andrano- mena, Sarainana River, 12°37'S, 49°26'E, ca. 235 m, 26 Nov. 1992 (fl, fr), D. J. Du Puy, G. Lewis & B. Schrire M564 (holotype, K; isotypes, P, MO, TAN). Figure 3. 174 Novon Figure 2. Millettia hitsika Du Puy & Labat. —A. Habit. —B. Leaflet upper surface. —C. Leaflet lower surface. —D. Flower. —E. Calyx. (A-E drawn from Service Forestier de Madagascar SF 20154.) Species M. richardiana affinis, inflorescentia elongata cum fasciculis pluribus florum, floribus minoribus, pedi- celbs multo brevioribus, et leguminibus glabris. A shrub or small tree ca. 3-15 m tall, flowering on new shoots as they emerge from the buds and whilst the leaves are still very immature, DBH up to 40 cm; bark smooth with shallow vertical fissures, dark gray; young shoots densely golden pubescent; mature twigs glabrous, gray, with pale lenticels and large spherical dormant buds. Leaves with (13-) 15- 25 opposite leaflets; rachis slightly pubescent; stipels absent. Leaflets narrowly oblong-elliptic, 13-37 x 6-11 mm, with a long apical mucro (up to ca. 2 mm long), subcoriaceous and with decurved mar¬ gins, densely golden pubescent when young, becom¬ ing glabrous when mature except for the midvein beneath. Pseudoracemes l-3.5(-5.5) cm long, from the axils of very immature leaves, golden-velvety pubescent when still in flower, the axis often angular. the flowers in 1-3 clusters, often paired, but the bottom group may have 2-4 flowers; pedicels short, 2-4 mm long, with a pair of linear bracteoles ca. 3 mm long, which are caducous as the flowers open. Flowers 8-10 mm long, bright fuchsia pink fading to pale mauve, the standard pale green at the base. Calyx 4.5-5 mm long, pale pubescent with occa¬ sional brown hairs, purple, with 3 narrowly trian¬ gular lower teeth with slender, tapering tips, as long as or shorter than the calyx cup, and with 2 small, very closely spaced upper teeth. Standard limb ca. 6-8 x 8-10 mm, shallowly notched at the apex, white pubescent behind. Wings longer than the keel, both slightly pubescent on the outer surfaces. Sta- minal sheath ca. 7 mm long. Pod obovate to oblong, flat, 40-70 x 1 6- 1 8 mm, tapering toward the base, rounded and with a decurved beak apically, gla¬ brous, splitting into two woody valves with l - 3 seeds. Seeds disc-shaped, 8-9 x 8—9 mm, finely Volume 5, Number 2 1995 Labat & Du Puy Millettieae from Madagascar 175 D M-leUa a*rtdM. v A Figure 3. Millettia nathaliae Du Puy & Labat. —A. Flowering stem. —B. Leaf. —C. Flower. —D. Calyx. E. Pod. —F. Seed. (A-F drawn from Du Puy , Lewis & Schrire M564.) mottled dark brown, the aril extended into a short flap on one side. Millettia nathaliae is closely related to M. ri- chardiana but is distinguished by its elongated in¬ florescence with several clusters of flowers, its small¬ er flowers, its much shorter pedicels, and its glabrous pods. It also closely resembles Pongamiopsis amyg- dalina (Baillon) R. Viguier when in flower and before 176 Novon the leaflets develop, but its much smaller flowers are characteristic. Distribution. Northern Madagascar, Ankarana Massif, around Diego-Suarez and Sahafary forest north of the Irodo River. Habitat. In deciduous woodland and shrubland, especially on limestone, but also on sand, at ca. 200-250 m altitude. Flowering time. Only recorded in November. This species is dedicated to Nathalie Labat. Paratypes. MADAGASCAR. Ankarana Massif, SE margin, about 25 km NE of Ambilobe, near the village of Ambilomagodro, 13 o 01'S, 49°07'E, 200 m, 17 Nov. 1992 (fl), Du Puy, Lewis & Schrire M559 (K, MO, P, TAN); Foret d’Andranomavo, Canton d’Anamakia, Dis¬ trict de Diego-Suarez, 24 Juin 1951 (fr), Service Forestier de Madagascar 3596-SF (K, P, TEF); Bassin de la Saharaina, foret de Sahafary, 27 Nov. 1958 (fl), Service Forestier de Madagascar 20112-SF, Capuron (K, MO, P, TEF). Millettia orientalis Du Puy & Labat, sp. nov. TYPE: E Madagascar. Baie d’Antongil, Nosy Mangabe, 5 km from Maroansetra, 15°33'S 49°46'E, 0-330 m, 9 Jan. 1989 (fl), G. Schatz & J. Miller 2495 (holotype, MO; isotypes, K, P, TAN). Figure 4. Species habitu robusto ramulis crassis et gemmis mag- nis, foliis longis (ca. 25-30 cm), foliolis numerosis (11- 21) grandibus et inflorescentiis longis (9-20 cm) densis floribus numerosis grandibus (15-20 mm longis) distincta. A large, deciduous tree ca. 15-30 m tall, flow¬ ering on new, developing shoots; [DBH not known]; young shoots densely velvety-pubescent, becoming glabrous and gray, the older twigs thick and blunt- ended with large, subspherical dormant buds. Stip¬ ules very large and conspicuous on emerging shoots, obovate, ca. 20-25 x 8-10 mm, soon caducous. Leaves large, ca. 25-30 cm long, with (11—)15— 21 opposite or subopposite, subequal leaflets; rachis velvety-pubescent when young; stipels absent. Leaf¬ lets very large, oblong-elliptic to obovate, usually narrow, 60-110 x 20—35 mm, acuminate apically, obtuse to rounded basally, glabrous above, densely velvety beneath especially while young. Flowers in robust pseudoracemes 9-20 cm long from the axils of developing leaves near the base of new shoots, the flowers often paired and the lower ones on short secondary axes up to 5 mm long; pedicels with 2 slender, soon caducous bracteoles ca. 4 mm long. Flowers 15-20 mm long, pink fading to pale lav¬ ender or mauve, the standard green at the base, sweetly fragrant. Calyx 4-7 mm long, densely pu¬ bescent, purplish, with 3 broadly triangular lower teeth, the upper pair short and much reduced. Stan¬ dard limb 11-15 X 12-14 mm, pubescent behind. Wings as long as the keel, finely pubescent outside. Staminal sheath 14-17 mm long, the free portion of the filaments rather long. Pods narrowly oblong to obovate, flat, ca. 70-150 x 20-30 mm, slightly thickened along both margins, velvety pubescent, splitting into 2 twisted, woody valves with 2-6 seeds. Seeds oblong to discoid, flat, 13-16 x 8-11 mm, chestnut brown, the aril extended into a short flap on one side. Millettia orientalis is the only Millettia that forms a canopy tree in the humid eastern forests. It is characterized by its robust habit with thick twigs and large buds, its long leaves with many large leaflets, and its long, dense inflorescences with nu¬ merous large flowers. Distribution. Known from around the Bay of Antongil and in the region of Taolaiiaro (Fort Dau¬ phin). Habitat. In lowland humid evergreen forest, up to ca. 300 m altitude. Flowering time. October-January. Vernacular name. Fanamoakondro. The specific epithet indicates the provenance of this species, in the eastern forests of Madagascar. Paratypes. MADAGASCAR. Prov. of Toliara, Fort Dauphin [Taolaiiaro] Region, NW of town near road to Andohahela Reserve, along trail W of road, 24°46'S, 46°53'E, 100-200 m, 11 Nov. 1989 (fl), McPherson 14456 (K, MO, P, TAN); Emafotra, Mahatalaky, Fort Dauphin, 27 Mai 1954 (fr). Service Forestier de Mad¬ agascar 10084-SF (K, MO, P, TEF); Beandri, Maha¬ talaky, Fort Dauphin, 23 Oct. 1954 (st), Service Forestier de Madagascar 11139-SF (P, TEF); Amboangy, Ma¬ hatalaky, Fort Dauphin, 20 Oct. 1955 (fl), Service Fo¬ restier de Madagascar 14861-SF (K, MO, P, TEF); E Madagascar, environs de la Baie d’Antongil, massif de Farankaraina, entre Navana et Andranofotsy, 0-150 m, 16 Sep. 1957 (fr). Service Forestier de Madagascar 18307-SF, Capuron (K, P, TEF). Millettia taolanaroensis Du Puy & Labat, sp. nov. TYPE: SE Madagascar. SW de Fort Dau¬ phin [Taolaiiaro], Baie de Ranofotsy, colline pres d’ltaly, 10 Jan. 1963 (fl). Service Fores¬ tier de Madagascar 22367-SF, R. Capuron (holotype, P; isotypes, K, MO, P, TEF, WAG). Figure 5. Species foliolis subglabris nitidis obtusis ad rotundatis subtus cinerascentibus, pubescentia aperta pilis nitidis cu- preis brevibus praesertim in rachidi, petiolulis, axe inflo- rescentiae, pedicellis et calycibus, et leguminibus glabris secus marginem superiorem incrassatis distincta. Volume 5, Number 2 1995 Labat & Du Puy Millettieae from Madagascar 177 Figure 4. MiUettia orientals Du Puy & Labat. -A. Habit. -B. Leaflet. -C. Leaflet undersurface. -R Young stem with stipules, leaf and inflorescence. —E. Flower. —F. Calyx. —G, H. Pod. I. Seed. (A, E, and F drawn from Schatz & Miller 2495; B and C from Service Forestier de Madagascar SF 18307; D from Service horestier de Madagascar SF 11139; G-I from Service Forestier de Madagascar SF 10084.) 178 Novon Figure S. Millettia taolanaroensis Du Puy & Labat. —A. Habit. —B. Petiole and first leaflet pair. —C. Leaflet apex. —D. Leaflet undersurface. —E. Flower. —F. Calyx. —G, H. Pod. —I. Seed. (A, C-F drawn from Service Forestier de Madagascar SF 22367 ; B and G from Phillipson & Milijaona 3526 ; H and I from Service Forestier de Madagascar SF 8503.) A deciduous shrub or small tree 4-6 m tall, flowering on young shoots with fully expanded leaves; twigs sparsely pubescent when young, glabrescent, with small dormant buds. Leaves with 7-11 opposite leaflets; rachis thinly coppery-pubescent; stipels ab¬ sent. Leaflets elliptic, 12-30(-36) x 9-19 mm, rounded basally, obtuse or rounded to notched and mucronulate apically, dark green, glossy and gla¬ brous above, gray-green and almost glabrous with very scattered, short brown hairs beneath. Flowers in delicate pseudoracemes ca. 4-6 cm long from the axils of leaves near the base of new shoots, the flowers solitary or paired on small knobs; pedicels with 2 slender, soon caducous bracteoles near the middle. Flowers 10-12 mm long, mauve or violet. Calyx 3-4 mm long, densely coppery-pubescent, with 5 short, broadly triangular teeth, the upper pair smallest and close together. Standard limb ca. 9x9 mm, cuneate basally, finely pubescent behind. Wings as long as the keel. Staminal sheath ca. 8 mm long. Pods obovate to oblong, flat, (40-)50-90 x (12—)14— 19 mm, tapering basally, the apex Volume 5, Number 2 1995 Labat & Du Puy Millettieae from Madagascar 179 rounded and beaked, slightly thickened along the upper margin, glabrous or with very scattered brown hairs, splitting into 2 twisted, woody valves, with 1- 5 (usually 1-3) seeds. Seeds oblong-discoid, 9-10 x 7-8 mm, chestnut brown, the aril extended into a small flap on one side. Millettia taolanaroensis is recognizable by its subglabrous, glossy, obtuse to rounded leaflets, which are grayish beneath, by its thin pubescence of lus¬ trous, copper-colored short hairs most noticeable on the leaf rachis and petiolules, the inflorescence axis and pedicels and the calyces, and by its glabrous pods thickened along the upper margin. It appears to flower when the leaves are almost full-sized rather than as new growth emerges from the buds. Distribution. Southeastern Madagascar, only known from around Taolafiaro (Fort Dauphin), on the coastal plains to the north and on the lower slopes of the mountains to the west (including An- dohahela, parcelle 3). Habitat. In coastal, evergreen forest on sand, and in transitional, deciduous woodland often near the coast. Flowering time. Only recorded in January. Local uses. Used as a fish poison. Vernacular name. Anakaraka. The specific epithet indicates the provenance of this species, in the southern region of Madagascar, near the town of Taolafiaro. Paratypes. MADAGASCAR. Maravato pres de Fort Dauphin, Avr. 1972 (fr), Boiteau s.n. (P); Prov. de Fort Dauphin, Andrahomana, 24 Juin 1926 (fr), Decary 3993 (P); Baie des Galions (Ranofotsy) au SW de Fort Dauphin, 1-100 m, 18-21 Fev. 1955 (fr), Humbert & Capuron 29019 (L, MO, P); Falaise cohere NW du Cap Sainte- Marie, vers Lavanono, 1-150 m, 8 Mar. 1955 (fr), Humbert & Capuron 29323 (K, P); Fort Dauphin region, W of town along highway on southern boundary of Re¬ serve Integrale no. 11, Andohahela, parcel 3, dry forest W of Col des Ranopiso, 120-140 m, 27 Jan. 1990 (fr), McPherson & Pigeon 14911 (K, MO, P, TAN); 5 km S of Manambaro, 23 km W of Fort Dauphin, 25°05'S, 46°49'E, 150 m, 30 Mar. 1991 (fr), Miller & Randri- anasolo 6 224 (K, MO, P, TAN); Reserve d'Andohahela, parcelle 3, 25°02'S, 46°40'E, 100 m, 17 Feb. 1990 (fr), Phillipson & Milijaona 3526 (K, MO, P, TAN); Pre¬ fecture de Fort Dauphin, Petriky, 25°05'S, 46°51'E, 0- 10 m, 16 Oct. 1989 (fr), Rabevohitra 2018 (K, MO, P, TAN); Mandena, Fort Dauphin, 8 Avr. 1953 (fr). Service Forestier de Madagascar 8172-SF (P, TEF); Bords d'un cours d'eau. aux environs de Ranopiso, route Ambovombe-Fort Dauphin, 23 Sep. 1953 (fr). Service Forestier de Madagascar 8503-SF, Capuron (K, MO, P> TEF); entre km 454 et 455 de la route de Fort Dauphin, 19 Sep. 1954 (fr), Service Forestier de Mad¬ agascar 10911-SF (P, TEF). Millettia aurea (R. Viguier) Du Puy & Labat, comb. nov. Basionym: Neodunnia aurea R. Viguier, Notul. Syst. (Paris) 14: 73, 1950. TYPE; (based on three collections) W Mada¬ gascar. Ambongo, Manongarivo [S of Soalala], Oct. 1903 (fl, jfr), H. Perrier de la Bathie 1617 (lectotype, chosen here, P; isolectotypes, K, L, MO, P, TAN). R. Viguier cited three syntypes in the original description, including H. Perrier de la Bathie 4831 and 4879, in addition to the lectotype selected here. Millettia aurea may be confused with M. ri- chardiana, which differs in its much shorter inflo¬ rescences with a single cluster of (larger) flowers, its smaller and less pubescent calyx with shorter teeth, especially the upper pair, which are very short and connate to near the tip, its much less pubescent petals, and its usually smaller leaflets. Millettia aurea is an uncommon species of west¬ ern Madagascar, with a restricted distribution in the Boina and the Ambongo, south to Maintirano, most frequently recorded from the region of Ankara- fantsika, and from Soalala to Mitsinjo. It occurs in deciduous woodland, on sand, often near the coast but also inland at up to ca. 300 m altitude. Millettia richardiana (Baillon) Du Puy & Labat, comb. nov. Basionym: Mundulea richardiana Baillon, Bull. Soc. Linn. Paris 1: 389-390, 1883, pro parte. TYPE: (based on three col¬ lections) N Madagascar. [Diego-Suarez (Antsi- ranana)], Baie des Amis (11), C. Richard 158 (lectotype, chosen here, P). Mundulea hysterantha Baker, J. Linn. Soc., Bot. 25: 309, 1890. Syn. nov. TYPE: W Madagascar. An- drona (fl, jfr), R. Baron 5444 (holotype, K; isotype, P). Millettia baroni Drake in Grandidier, Hist. Phys. Nat. Pol. Madagascar, Plantes, Texte 1: 141, 1903. Syn. nov. TYPE: Madagascar (fr), R. Baron 4952 (ho¬ lotype, P; isotypes, K, P). Neodunnia atrocyanea R. Viguier, Notul. Syst. (Paris) 14: 72, 1950. Syn. nov. TYPE: NW Madagascar. Riviere de Mananjeba, Oct. 1909 (fl), H. Perrier de la Bathie 4845 (holotype, P; isotype, P). Neodunnia edentata R. Viguier, Notul. Syst. (Paris) 14: 73, 1950, pro parte. Syn. nov. TYPE: (based on four collections) NW Madagascar. Majunga, Dec. 1924 (fl), H. Perrier de la Bathie 16800 (lectotype, chosen here, P; isolectotypes, K, P, MO). H. Baillon cited three syntypes in the original description of Mundulea richardiana, including Boivin 2712 and 2713, in addition to the lectotype selected here. Of the two Boivin specimens (both mounted on the same herbarium sheet) only 2712 180 Novon corresponds with M. richardiana; Boivin 2713 is Pongamiopsis amygdalina (Baillon) R. Viguier. In the original description of Neodunnia eden- tata, R. Viguier cited four syntypes: H. Perrier de la Bathie 1601 is a specimen of Pongamiopsis pervilleana (Baillon) R. Viguier; H. Perrier de la Bathie 4076, 16800, and Louvel 53 are all spec¬ imens of Millettia richardiana. H. Perrier de la Bathie 16800 is composed of two elements: the first was collected in December 1924, in flower, and is selected here as the lectotype; the second was col¬ lected in January 1926, in fruit, and is considered here to be a separate specimen. Millettia richardiana is especially variable in the number of leaflets, the form and pubescence of the leaflets, and the pubescence of the pods. Variants can be constant within certain localities. However, none of these variants seem distinct enough to be recognized as separate taxa. Millettia richardiana is very widely distributed throughout western Madagascar from Antsiranana (Diego-Suarez) to Toliara (Tulear) and southwestern Madagascar, including the forest of Zombitsy, the Bemaraha Massif, the Ambongo, the Boina and the Ankarana Massif. It occurs in deciduous woodland, mainly on calcareous or sandy soils, on limestone, sand over limestone or on coastal dunes, at up to ca. 300 m altitude. Pongamiopsis viguieri Du Puy & Labat, sp. nov. TYPE: W Madagascar. Tsingy de Bemaraha (Reserve Naturelle no. 9), environs d’Antsa- lova, 5 km a l’E de Bevitika, 18°38'S, 44°43'E, 150-250 m, 30 Nov. 1992 (fr), J.-N. Labat, R. Edmond & O. Laivao 2243 (holotype, P; isotypes, K, MO, P, TAN). Figure 6. A P. amygdalina differt inflorescentiis brevibus con- densatis, dentibus calycis latioribus, apicibus tenuibus des- titutis, vexillo, alis et carina dense pubescentibus, vagina staminea breviori, et leguminibus plerumque elongatiori- bus seminibus numerosioribus, A deciduous tree (2-4-)10 m tall, flowering on new shoots as they emerge from the buds and whilst the leaves are still very immature; young shoots densely pubescent, glabrescent; twigs with large leaf scars and large, spherical dormant buds. Leaves becoming very large, with 7-11 opposite leaflets; stipels absent. Leaflets oblong, oblong-elliptic or ob- long-obovate, 50-200 x 20-80 mm, with a broad, V-shaped apical notch with a mucro at the base of the notch, silky pubescent at first but becoming glabrous above and pubescent mainly on the veins beneath, the veins strongly raised beneath, the straight, parallel secondary veins extending to the slightly thickened margin. Racemes produced in the axils of the lowest leaves of developing shoots, short and condensed, the axis ca. 1-3 cm long, with numerous flowers in closely spaced clusters of 2 or 3 on short spurs; pedicels with a pair of small, caducous bracts toward the apex. Flowers 13-15 mm long, white washed with pale lilac, the standard with a pale green basal eye. Calyx 6-8 mm long, densely appressed-hairy, with 3 triangular, acute lower teeth, the upper pair partially connate but with free, widely separated, recurved tips. Standard limb 12-14 x 12-13 mm, densely silky pubescent behind. Wings about as long as the keel, narrow, both appressed-pubescent outside and upcurved api- cally, the keel shortly beaked. Staminal sheath 12- 15 mm long. Style short-hairy, hooked apically. Pod strongly inflated, ellipsoidal to oblong-ellipsoidal, 45- 70 x 25-35 x 23-35 mm, obliquely beaked api¬ cally, the upper margin broad and thickened, vel¬ vety-pubescent, wrinkled, thick and woody, inde- hiscent or eventually splitting into 2 valves, usually with 3-5 seeds (occasionally some pods with fewer seeds). Seeds subspherical, compressed, ca. 13 mm diam., becoming dark gray, the hilum raised and surrounded by a narrow aril. Pongamiopsis viguieri is closely related to P. amygdalina, with similar leaves, pod type, and growth habit, but differs in its short, condensed inflorescences, its broader calyx teeth lacking the very slender tips, its densely pubescent standard, wings, and keel, its shorter staminal sheath, and its usually more elongated pods with more numerous seeds. It occurs in western rather than northern Madagascar, which is the range of P. amygdalina. Distribution. Western Madagascar, uncommon, mainly from the Bemaraha Massif, but also collected at Amboanio south of Mahajanga (Majunga) and at Andranomavo near Namoroka. Habitat. In deciduous woodland, always over limestone outcrops or “tsingy,” from near sea level to ca. 500 m altitude. Flowering time. October. This species is dedicated to Rene Viguier, in rec¬ ognition of his extensive research on the Legumi- nosae of Madagascar. Paratypes. MADAGASCAR. Vallee de la Tsiribihina dans le Bemaraha, 20-350 m, Oct. 1933 (fl, jfr), Hum¬ bert 11480 (K, MO, P); Antsingy de Bemaraha, vers Anbodiriana (E d’Antsalova), NE du pont, 100-150 m, 10 Dec. 1952 (fr), Leandri, Capuron & Razafindrakoto 2102 (BR, K, L, MO, NY, P, PRE, WAG); Antsingy de Bemaraha, vers Anbodiriana (E d’Antsalova), 100-150 m, 21-27 Jan. & 8-11 F6v. 1960 (fr), Leandri & Saboureau 3071 (K, L, MO, NY, P, PRE); Ambongo, environs d’Andranomavo, Jan. 1905 (jfr), Perrier de la Volume 5, Number 2 1995 Labat & Du Puy Millettieae from Madagascar 181 Figure 6. Pongamiopsis viguieri Du Puy & Labat. A. Habit with mature leaf and pod- j j Habit with leaf and flowers. —C. Leaflet undersurface. —D. Leaflet apex. —E. Flower. —F. Calyx. — G. Pod. — H. seed (A, C, and D drawn from Labat, Edmond & Laivao 2243; B, E, and F drawn from Humbert 11480; G and H from Leandri & Saboureau 3071.) 182 Novon Bathie 1801 (P); Rochers calcaires du bord de mer a Amboanio pres de Majunga, Fev. 1901 (fr), Perrier de la Bathie 1828 (P); Antsingy de Bemaraha, route An- tsalova-Tsiandro, aux environs de la clairiere d’Ambo- diriana, 10 Dec. 1952 (fr), Service Forestier de Mad¬ agascar 6808-SF, Capuron (K, L, MO, P, TEF). Acknowledgments. We thank Frangoise Theu- reau and Margaret Tebbs for the illustrations and R. Hoogland for assistance with the Latin descrip¬ tions. D. Du Puy thanks the Royal Society for the opportunity to undertake collaborative research in the Laboratoire de Phanerogamie, Paris, and the Weston Foundation for the support of his continued research in Madagascar and Kew. We thank the Directors and staff of the Laboratoire de Phanero¬ gamic, Paris, the Herbarium, Royal Botanic Gar¬ dens, Kew, the Parc de Tzimbazaza, Antananarivo, and the Centre National de la Recherche Ecologique, Antananarivo. We are also grateful to the National Geographic Society for supporting our recent field¬ work in Madagascar. Literature Cited Geesink, R. 1981. Tephrosieae. Pp. 245-260 in R. M. Polhill & P. H. Raven (editors), Advances in Legume Systematics, Part 1. Royal Botanic Gardens, Kew. -. 1984. Scala Millettiearum. A survey of the genera of the tribe Millettieae (Legum.-Pap.) with methodological considerations. E. J. Brill/Leiden Univ. Press, Leiden Botanical Series 8. Hutchinson, J. 1964. The Genera of Flowering Plants. Vol. 1. Oxford Univ. Press, Oxford. Another New, Nearly Extinct Species of Hibiscadelphus (Malvaceae) from the Hawaiian Islands David H. Lorence National Tropical Botanical Garden, P.0. Box 340, Lawai, Hawaii 96765, U.S.A. Warren L. Wagner Department of Botany, National Museum of Natural History, Smithsonian Institution, MRC- 166, Washington, D.C. 20560, U.S.A. ABSTRACT. Four individuals of a new species of Hibiscadelphus (Malvaceae: Hibisceae) have been discovered on Kaua'i, the oldest of the major Ha¬ waiian Islands. Hibiscadelphus woodii is described and illustrated, and its status and basal relationship within the genus are discussed. The imperiled status of the other six species is reviewed, and a key to the genus is given. Hibiscadelphus Rock consists of seven species endemic to the Hawaiian Islands (Bates, 1990), in¬ cluding the one described herein. The genus was originally established by Joseph F. Rock (Radlkoffer & Rock, 1911) to accommodate three species of shrubs or small trees that he had collected from dryland forests: two species on the island of Hawai‘i, H- giffardianus Rock (Mauna Loa) and H. huala- laiensis Rock (Hualalai), and H. wilderianus Rock from Auwahi on East Maui. Four additional species were subsequently described: H. bombycinus C. Forbes from the Kohala Mountains on the island of Hawai‘i (Forbes, 1920), H. distans L. Bishop & Herbst from the Koai‘e Canyon of Kaua'i (Bishop & Herbst, 1973), H. crucibracteatus Hobdy from Puhielelu Ridge on Lana'i (Hobdy, 1984), and the new species, H. woodii, described herein from Ka- lalau Valley, Kaua'i. Hibiscadelphus is a member of the tribe Hibis¬ ceae, represented in the Hawaiian Islands by the indigenous genus Hibiscus L., following Fryxell’s (1968) classification. Although closely related to Hi¬ biscus (the generic name means “brother of Hibis- cus ), Hibiscadelphus is clearly a distinct mono- phyletic group based on its curved and narrowly convolute zygomorphic corollas with the lower two lobes shorter than the other three, calyces that are circumscissily deciduous in fruit, and relatively thick, woody capsules (Hobdy, 1984; Bates, 1990). The green, yellow, or magenta flowers with tubular, curved corollas are presumably adapted to pollina¬ tion by native Meliphagid and Drepanidine birds. known as the honeyeaters and honeycreepers (Hob¬ dy, 1984). The Hawaiian name for members of the genus is hau kuahiwi, meaning upland or mountain hau, because of their resemblance to the common lowland hau, Hibiscus tiliaceus L. (Rock, 1913; Bates, 1990). Due to the destruction and severe alteration of the diverse dry and mesic forest habitats and loss of all of the Meliphagid birds in the Hawaiian Islands (Hobdy, 1984), five of the seven Hibiscadelphus species have become extinct in the wild, although at least two of these are maintained through culti¬ vated plants: H. giffardianus and H. hualalaiensis. Only H. distans, a U.S. federally fisted endangered species, and the new species described in this paper have naturally occurring wild populations (Table 1; Bates, 1990). The status of the genus is extremely precarious at present. In March 1991 an unusual species of Hibisca¬ delphus not corresponding with any of the described taxa was discovered on Kaua'i by Kenneth R. Wood, staff member of the National Tropical Botanical Garden, along with M. Query and S. Montgomery. Four plants were found growing on a steep slope below the rim of Kalalau Valley in northwestern Kaua'i (K. R. Wood, 1992, unpublished). Study of flowering specimens has revealed that they represent a new species. Hibiscadelphus woodii Lorence & W. L. Wag¬ ner, sp. nov. TYPE: Hawaiian Islands. Kaua'i: Hanalei District, Kalalau Rim, N of Kahuama'a Flat, steep diverse lowland mesic forest, 990- 1020 m, 3 Mar. 1991, K. R. Wood, M. Query & S. Montgomery 629 (holotype, PTBG, a flower also in spirit collection; isotypes, BISH, K, MO, NY, US). Figure 1. Species foliis glabratis vel venis sparsim stellato-pub- erulis, pagina abaxiali laminae axillis venarum trichom- atibus stellatis instruetis, calyce trichomatibus stellatis cum radiis 5-8 ornatis, corolla cerina cupreo-suffusa atropur- purescentia distinguibilis. Novon 5: 183-187. 1995. 184 Novon Table 1. Status of species of Hibiscadelphus (D. Ragone, unpublished observations, 1993). DOFAW — Division of Forestry and Wildlife, Department of Land and Natural Resources, State of Hawaii (plantings at Pu‘uwa‘awa a, Kona, Hawaii Island); HVNP = Hawaii Volcanoes National Park (planting at Kipuka Puaulu, Hawaii Island); NTBG = National Tropical Botanical Garden (plantings at (plantings at Haleiwa, 0‘ahu). Lawai and Limahuli Gardens, Kaua‘i); WA = Waimea Arboretum Species Distribution Wild populations Wild Known individuals Cultivated H. bombycinus Hawaii Extinct — — (Kohala Mts.) H. crucibracteatus Lana‘i Extinct — — (Puhielelu Ridge) H. distans Kaua‘i 2 137 39 NTBG (Koai‘e Canyon) 14 WA H. giffardianus Hawai‘i Extinct — 7 HVNP (Mauna Loa) 1 NTBG 8WA H. hualalaiensis Hawai‘i Extinct — 3 NTBG (Hualalai) 20-30 DOFAW H. wilderianus East Maui Extinct — — (Auwahi) H. woodii Kaua‘i i 4 — (Kalalau Valley) Small branched trees 2.5-5 m tall with dense rounded crown, trunk with DBH 10-20 cm, the twigs glabrate. Leaves with petioles 2.8-5.8 cm long bearing scattered stellate trichomes when young, glabrate; stipules subulate, 3-5 mm long, sparsely stellate-pubescent; blade broadly ovate to very broadly ovate, 7-9 cm long, 6.5-8.4 cm wide, thin- coriaceous, drying chartaceous; adaxially glabrate or with scattered small, translucent stellate tri¬ chomes on principal veins, abaxially with few scat¬ tered stellate trichomes on principal veins or gla¬ brate, with sparse tufts of stellate trichomes in major vein axils, when young with scattered stellate tri¬ chomes along margins and at junction with petiole, the base deeply cordate, occasionally shallowly so, the apex acute to weakly acuminate or broadly ob¬ tuse, the principal veins 5 (7), the margins irregu¬ larly and coarsely crenate-dentate. Flowers solitary, the peduncles 1.4-2.1 cm long, sparsely stellate pubescent; involucral bracts 4-6, free essentially to their base, 11-15 mm long, 1.8-4 mm wide, linear- oblong; calyx tubular, 1.3-1.5 cm long (up to 1.7 cm when fresh), green, shallowly 5-lobed, the lobes 2-3 mm long, sometimes in age splitting further, moderately densely pubescent with stellate tri¬ chomes composed of 5-9 horizontal radiating arms free to their base; corolla 45-47 mm long, stellate pubescent, yellow with coppery tinge when fresh, rapidly becoming purplish maroon, the lobes 35- 40 mm long, 10-12 mm wide, with conspicuous veins, moderately stellate pubescent with trichomes becoming restricted to veins in inner % of lobes abaxially; staminal column exserted ca. 7 mm be¬ yond corolla lobes, stigmas ca. 1 mm long, subglo- bose, reddish brown, villosulous, exserted 14 mm beyond corolla lobes; style villous; ovary 7 mm long, 4-5 mm wide, 5-angled, stellate pubescent, endo- carp segments apparently 10 (in cross section). Fruit unknown. Distribution and habitat. The new species is known only from a population of four trees at the type locality—accessible only with climbing ropes. It occurs on a steep slope of predominantly basalt talus with patches of brown soil and leaf litter in remnant diverse lowland mesic forest. Associated native taxa include Melicope pallida (Hillebrand) T. Hartley & B. Stone, Nototrichium sp. nov., Hed- yotis sp. nov., Chamaesyce sp. nov., Lysimachia glutinosa Rock, L. kalalauensis Skottsberg, Poa mannii Munro ex Hillebrand, Stenogyne campan- ulata Weller & Sakai, and Lobelia niihauensis St. John. The alien daisy fleabane, Erigeron karvin- skianus DC., is an extremely invasive perennial herb that threatens to overrun and smother the native cliff habitat in this region. Accessible areas above these cliffs have been severely degraded by feral goats and pigs. The Hibiscadelphus woodii plants are also susceptible to damage by falling rocks dis¬ lodged by these animals, which aggravate erosion. Reproduction and conservation. Flowering ma¬ terial has been collected in March, April, and Sep- Volume 5, Number 2 1995 Lorence & Wagner Hibiscadelphus 185 Figure 1. Hibiscadelphus woodii Lorence & W. L. Wagner. Habit. Scale 1 cm. tember, but no fruit set has been observed in spite of efforts to manually outcross and bag the flowers. Introduced birds, i.e., the Japanese white eye (Zos- terops japonicus), were seen piercing the corollas above the calyx, presumably robbing nectar (note on Wood et al. 640). A liquid-preserved flower of Wood 726 contained three adult Nitidulidae beetles probably representing an endemic species (D. Ja¬ mieson, pers. comm.). The flower also contained two half-grown larvae of the endemic moth Crocidosema marcidella (Walsingham) (Lepidoptera: Tortrici- dae), which had completely eaten the style down to the ovary. Larvae of Hawaiian Crocidosema species are known to feed on indigenous Malvaceae fruits and foliage (Zimmerman, 1978), and damage by these larvae may be responsible for the observed 186 Novon lack of fruit set in H. woodii. Attempts to propagate the new species by air layering, cuttings, grafts, and tissue culture thus far have failed. We take pleasure in naming this new species for Kenneth R. Wood, in recognition of his continued efforts to understand and protect the rare and en¬ dangered plants of the Hawaiian Islands, especially those of Kalalau Valley. Paratypes. HAWAIIAN ISLANDS. Kaua'i: Hanalei District, Kalalau Rim, N of Kahuama'a Flat, steep diverse lowland mesic forest, 990-1000 m, 6 Mar. 1991, K. R. Wood et al. 640 (BH), 15 Apr. 1991, K. R. Wood et al. 726 (PTBG), 23 Sep. 1991, K. R. Wood & S. Perl¬ man 1600 (PTBG). Affinities. Hibiscadelphus woodii differs from its congeners in having leaves that are glabrate except for rare stellate trichomes adaxially and sparse stel¬ late trichomes on the veins and in the principal vein axils abaxially, stellate trichomes on the calyx with rays free to the base, and yellow corollas with a coppery tinge, rapidly becoming purplish maroon with age. Hibiscadelphus distans differs in its in- volucral bracts connate for about one-third of their length, multi-rayed calyx trichomes of two sizes, a smaller yellowish green corolla rapidly becoming dull reddish, and densely stellate pubescence on the co¬ rolla with the trichomes becoming restricted to the veins on the inner one-third of the lobes abaxially. Hibiscadelphus giffardianus differs from If. woodii in having filiform involucral bracts up to 1.1 mm wide, whereas those of H. hualalaiensis are short, triangular teeth only 0.5-3 mm long. Hibiscadel¬ phus crucibracteatus differs in having longer in¬ volucral bracts (20-30 mm), whereas H. wilderi- anus has a longer calyx (2.3-2.5 cm). Hibisca¬ delphus bombycinus differs from H. woodii in hav¬ ing leaves with the lamina densely pubescent beneath and a shorter corolla (3.3 cm). Fruiting material, unknown for H. woodii, is required to further assess its affinities. Examination of transverse hand-cut sections of a flowering ovary of H. woodii suggests the endocarp will separate into ten segments in fruit as in H. bombycinus, H. crucibracteatus, H. gif¬ fardianus, H. hualalaiensis, and H. wilderianus, rather than into five boat-shaped segments as in H. distans. The phylogenetic relationships of this unique Ha¬ waiian endemic genus were recently analyzed (Funk & Wagner, 1995). In the single cladogram resulting from their morphological study the position occupied by Hibiscadelphus woodii is basal. Because fruit characters are among the most important ones for determining the tree topology, the placement of H. woodii is in doubt. Their analysis estimated that the fruit character of H. woodii is plesiomorphic (i.e., chartaceous, not woody) as in H. distans. In order to investigate the stability of the basal position of H. woodii in the phylogeny they recoded the missing character as apomorphic. Even with this change, however, H. woodii remained in the basal position. Species of Hibiscadelphus may be separated by characters given in the following key (adapted from Bates, 1990). Key to the Species of Hibiscadelphus la. Involucral bracts connate ca. V, of their length; mesocarp weakly developed and usually adnate to exocarp; endocarp segments 5, boatlike; Koai‘e Canyon, Kaua'i. II. distans lb. Involucral bracts free or connate slightly at base; reticulate mesocarp strongly developed; endocarp segments 2a. Involucral bracts filiform or obsolete, up to 1.1 mm wide. 3a. Involucral bracts 0.5-2(-3) mm long; corolla yellowish green, fading to purplish within, 2-5(-5.5) cm long; Hualalai, Hawai'i. H. hualalaiensis 3b. Involucral bracts 18-35 mm long (hybrids between H. giffardianus and other species tend to have bracts 5-15 mm long); corolla grayish green externally, dark magenta within, (5-)6-7 cm long; Mauna Loa, Hawai'i . H. giffardianus 2b. Involucral bracts linear to spatulate, 1-7 mm wide. 4a. Involucral bracts 20-30 mm long, 3-7 mm wide; calyx 4.2-4.9 cm long; corolla 5-6.5 cm long; Puhielelu Ridge, Lana'i. H. crucibracteatus 4b. Involucral bracts 9-18 mm long, 1.5-4 mm wide; calyx 1.3-2.5 cm long; corolla 3.3-4.7 cm long. 5a. Leaf lamina adaxially glabrate or with minute, scattered, stellate trichomes only on principal veins, abaxially glabrate or with scattered stellate trichomes only on principal veins, the trichomes sparsely tufted in principal vein axils; Kalalau Valley, Kaua'i . H. woodii 5b. Leaf lamina adaxially sparsely stellate pubescent, abaxially sparsely to densely stellate pu¬ bescent, the trichomes densely tufted in principal vein axils; Mauai, Hawai'i. 6a. Involucral bracts 9-10 mm long; calyx ca. 1.2 cm long; Kohala Mts., Hawai'i . . H. bombycinus 6b. Involucral bracts 11-18 mm long; calyx 2.3-2.5 cm long; Auwahi, East Maui ...... . H. wilderianus Volume 5, Number 2 1995 Lorence & Wagner Hibiscadelphus 187 Acknowledgments. We thank Anna Asquith for preparing the skillfully rendered illustration. Dean Jamieson and Laura Ishi kindly identified the insect specimens. We are also grateful to Kenneth R. Wood and Diane Ragone for providing data and field observations, Robynn Shannon for a careful review of a draft of the manuscript, and to Loyal Mehrhoff and an anonymous reviewer for valuable comments on the manuscript. Literature Cited Bates, D. M. 1990. Malvaceae. Pp. 868-903 in W. L. Wagner, D. R. Herbst & S. H. Sohmer, Manual of the Flowering Plants of Hawaii. Univ. Hawaii Press & Bishop Museum Press, Honolulu. Bishop, L. E. & D. R. Herbst. 1973. A new Hibis¬ cadelphus (Malvaceae) from Kauai. Brittonia 25: 290-293. Forbes, C. N. 1920. New Hawaiian plants. Occas. Pap. Bernice Pauahi Bishop Mus. 7(3): 33-39. Fryxell, P. A. 1968. A redefinition of the tribe Gos- sypieae. Bot. Gaz. 129: 296-308. Funk, V. A. & W. L. Wagner. 1995. Biogeography of seven ancient Hawaiian plant lineages. In: W. L. Wagner & V. A. Funk (editors), Hawaiian Bioge¬ ography: Evolution on a Hot Spot Archipelago. Smithsonian Institution Press, Washington, D.C. Hobdy, R. W. 1984. A re-evaluation of the genus Hibiscadelphus (Malvaceae) and the description of a new species. Occas. Pap. Bernice P. Bishop Mus. 25(11): 1-7. Radlkoffer, L. & J. F. Rock. 1911. New and noteworthy Hawaiian plants. Hawaiian Board of Agriculture and Forestry Botanical Bull. 1: 1-15. Rock, J. F. 1913. The Indigenous Trees of the Hawaiian Islands. Charles E. Tuttle, Rutland, Vermont. Wood, K. R. 1992. New Hibiscadelphus found on Kauai. Hawaii’s Forests and Wildlife 7(1): 15-117. Zimmerman, E. C. 1978. Insects of Hawaii. Vol. 9, Microlepidoptera, Part 1. The Univ. Press of Hawaii, Honolulu, Hawaii. A New Species of Tournefortia (Boraginaceae) from La Planada, Colombia James S. Miller Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. ABSTRACT. Tournefortia restrepoae J. S. Miller, sp. nov., is described from the Reserva Natural La Planada in Narino, Colombia. The species is distinct in its evidently and evenly pustulate leaf surface with depressed tertiary venation and elevated are- oles, and in its widely separated petiolate flowers. It is currently known only from the type locality. The genus Tournefortia comprises about 150 species distributed throughout tropical and subtrop¬ ical parts of the world (Miller, 1988). The vast majority of the species occur in the Neotropics, and the genus is only sparsely represented in the Old World. South America, particularly the northern Andes, is the region with the greatest number of species and morphological diversity (Killip, “The Andean species of Tournefortia ,” Smithsonian In¬ stitution Archives, unpublished; Miller, 1988). The species described below is one of many narrowly distributed endemics in mid-elevation Andean for¬ ests. Tournefortia restrepoae J. S. Miller, sp. nov. TYPE: Colombia. Narino: Mpio. de Ricaurte, Reserva Natural La Planada, elev. 1800 m, 18 Nov. 1993 (young fruits), Carla Restrepo 778 (holotype, MO 4619857; isotype, PSO not seen). Figure 1. Frutex sciophilus. Folia alterna; lamina elliptica ad el- liptico-oblonga vel leviter obovata, 12-24 cm longa, 6- 11 cm lata, apice acuminata, base acuta, supra hirsuta, infra strigosa, adaxialiter manifeste aequaliter pustulata, nervis tertiariis depressis, areolis elevatis; petiolo 1.7-5 cm longo, supra canaliculate, dense brunneo-pubescenti. Inflorescentia intemodalis, cymosa; pedicellis 1.5-2(-4) mm longis inter se 1-7 mm distantibus. Flos sepalis lan- ceolatis, 4-6 mm longis, pubescentibus; corolla tubiformi, ca. 6 mm longa, lobulis anguste ovatis, ca. 2 mm longis; staminibus ignotis; stylo 4 mm longo. Fructus late ovoi- deus, albus, glaber. Understory shrub. Leaves alternate, blades ellip¬ tic to elliptic-oblong or slightly obovate, 12-24 cm long, 6-11 cm wide, the apex acuminate, the base acute, sometimes asymmetrically so, the margin en¬ tire, somewhat undulate, the adaxial surface prom¬ inently and evenly pustulate with the tertiary ve¬ nation depressed and the areoles elevated, hirsute with several erect, siliceous hairs arising from each areole, the abaxial surface strigose with the slightly appressed hairs mostly restricted to the veins, each areole with one to several sunken pits, the venation brochidodromous with 9-16 secondary veins; peti¬ oles 1.7-5 cm long, canaliculate on the adaxial surface, densely brown pubescent. Inflorescence in- temodal, a dichotomously branched cyme, the branches 6-10 cm long, the peduncle 7.5 cm long, both densely brown pubescent. Flowers on 1.5- 2(-4)-mm-long pedicels, 1-7 mm apart, bisexual; sepals green, lanceolate, ± equal, 4-6 mm long, 0.4-1.3 mm wide, sparsely but obviously pubescent; corolla tubular, ca. 6 mm long, the lobes narrowly ovate, ca. 2 mm long; stamens unknown; ovary ovoid, ca. 1 mm long, ca. 0.5 mm broad, borne on an annular nectariferous disk, the style ca. 4 mm long, the stigma conical. Fruits white, drupaceous, the exocarp glabrous, the mesocarp thin, the en- docarp distinctly 4-parted, broadly ovoid, ca. 5 mm long, ca. 4 mm wide. Distribution. Tournefortia restrepoae is known only from the type collection from cloud forest at 1800 m at La Planada in Narino, Colombia. Tournefortia restrepoae is a distinctive member of Tournefortia sect. Tournefortia, which is by far the largest section of the genus, containing the ma¬ jority of the neotropical species, and is characterized by having free anthers and fruits that are unlobed and lack a corky mesocarp. E. P. Killip’s unpub¬ lished treatment of the Andean species of the genus recognized 38 species in the section, none of which appear very close to T. restrepoae. The species is distinct in its alternate, distinctly pustulate leaves and widely spaced, pedicellate flowers. The leaf sur¬ face is unique in the genus. The type collection is mostly in young fruit with only the old remains of a few floral parts, thus better knowledge of rela¬ tionships will require the collection of better flow¬ ering material. The species is named in honor of Carla Restrepo, who has done much to improve our knowledge of the plants of the Andean Cordillera in Colombia. Novon 5: 188-189. 1995. Volume 5, Number 2 1995 Miller Tournefortia restrepoae 189 Figure 1. Tournefortia restrepoae J. S. Miller (Restrepo 778). —A. Fruiting branch. B. Section of infructescence showing fruiting calyx. —C. Lateral and apical view of fruit. —D. Upper leaf surface. E. Lower leaf surface. Paratypes. COLOMBIA. IMarino: Mpio. de Ricaurte, Reserva Natural La Planada, elev. 1800 m, Restrepo 730 (PSO not seen). Acknowledgments. I thank John Myers for the illustration, Joan W. Nowicke and the Smithsonian Institution for making E. P. Killip's unpublished manuscript available, Roy E. Gereau for help with the Latin description, and Gordon McPherson for review comments. Literature Cited Miller, J. S. 1988. A revised treatment of Boraginaceae for Panama. Ann. Misssouri Bot. Gard. 75: 456- 521. Aristida tuitensis (Poaceae: Aristideae), a New Species from El Tuito, Jalisco, Mexico Jorge G. Sanchez-Ken and Patricia Davila A. Instituto de Biologla, U.N.A.M., Depto. de Botanica, Herbario Nacional, Apartado Postal 70- 233, 04510 Mexico, D.F., Mexico Abstract. A new species, Aristida tuitensis from El Tuito, Jalisco, Mexico, is described and illustrat¬ ed, and its distribution and ecology are discussed. It belongs to Aristida sect. Streptachne and is dis¬ tinguished from A. schiedeana by its spreading in¬ florescence, shorter spikelets, and the lack of dentate glumes, and from A. hintonii by its inflorescence shape, the internode, and sheath length. A table summarizing the differences and similarities among the three species is presented. As part of research to revise the genus Aristida L. section Streptachne (R. Brown) Domin, some unusual specimens of Aristida were collected in the area of El Tuito, Jalisco, Mexico, in early 1993. The material was initially identified as A. schie¬ deana, a common grass in the area. However, with additional studies of these specimens, as well as other collections from IBUG and COCA, it was realized that these specimens represent a new species. Aristida tuitensis Sanchez-Ken & P. Davila, sp. nov. TYPE: Mexico. Jalisco: Municipio El Tui¬ to, El Tuito, 9 a 10 km sobre la brecha a las Minas del Cuale (E de El Tuito), elev. 1900 m, 13 ene. 1991, J. Sanchez-Ken, M. May- field & B. Westlund 500 (holotype, MEXU; isotypes, ENCB, IBUG, MICH, MO). Figure 1. Gramen perenne caespitosum; internodia ramosa, 9- 15 infra inflorescentiam; vaginae coriaceae internodiis longiores; ligulae externae presentes; spiculae effusae bin- atae; glumae infernae glumis supernis longiores; glumae infernae no dentatae, 3-nervatae, nervis centrales con- spicui, nervi laterales inconspicui; calli 0.7-1.0 mm longi; lemmata glumis breviores vel eas aequantes raro; col- umnae lemmatum 6-10 mm longae; aristae laterales ab- sentes; antherae 3-4 mm longae. Perennial tufted grass, erect; pseudorhizome present, 80-120 cm long. Internodes 2.0- 9.0(-12.0) cm long, 1-2 mm diam., glabrous, straight, terete, bamboo-like, branched, with 9-15 intemodes below the inflorescence; nodes glabrous, inconspicuous. Leaf sheaths 3-9 cm long, 4-6 mm wide, longer than the internodes, strongly overlap¬ ping, glabrous, not striate, strongly coriaceous, in- Novon 5: 190-192. 1995. eluding the margins; inner ligule ciliate, 0.2-0.3 mm long; auricles lacking, sometimes a tuft of lateral hairs 2 mm long instead of the auricles in juvenile stages; outer ligule or contra ligule present; blade 10-50 cm long, 2-4 mm wide, linear, flat, straight to slightly flexuous when dry, adaxially scaberulous with scattered hairs toward the base, abaxially smooth or rarely scaberulous, with 7 main conspicuous nerves, but with 21, 23, or 25 nerves in total. Inflorescence an open panicle, 14-35 cm long, 12- 20 cm wide; basal internode 18-70 cm long; pe- duncule 11-30 cm long, glabrous or scaberulous; branches 9— 13(— 15) cm long, branching until 5th order, ascendant, strongly spreading, with the spike- lets located at the apex; pulvinus present in the axils of all branches and pedicels, glabrous; pedicels 5- 18 mm long. Spikelets usually paired, 20-28 mm long from the base of glumes to the tip of the awn, spreading; glumes varying in size, usually the first larger than the second one, sometimes the second glume larger, persistent; first glume 4-8 mm long, to 1 mm wide, 3-nerved, the two lateral nerves inconspicuous, glabrous, keeled and scabrous in the keel, apically acute and awned, not dentate, awn 0.5-2.0 mm long; second glume 6.0-7.5 mm long, to 1.0 mm wide, 1-nerved, keeled and glabrous in the keel, apically obtuse and awned, not dentate, the awn 0.5-1.0 mm long; calluses 0.7-1.0 mm long; lemma shorter than or sometimes as long as the glumes, 4.5-6.0 mm long without the column, 1.0-1.3 mm wide, convolute, 3-nerved, terete, sca¬ berulous in the upper third, not articulate with the column, not articulated; column 6-10 mm long, twisted, abaxially scaberulous, adaxially muricate, basally geniculate; awn 1, 8-10 mm long, straight, basally geniculate, thin; lateral awns lacking; palea 0.5-0.7 mm long, not keeled, smooth; lodicules 2, 1.0-1.2 mm long, hyaline; stamens 3; anthers 3- 4 mm long; stigma 1-2 mm long. Caryopsis not seen. Distribution. Aristida tuitensis is endemic to the area of El Tuito in the state of Jalisco, northeast of Chamela. It occurs in a pine and oak forest that also contains some elements that are characteristic Volume 5, Number 2 1995 Sanchez-Ken & Davila Aristida tuitensis 191 Figure 1. Aristida tuitensis Sanchez-Ken & P. Davila. -A. Habit. -B. Stem. -C. Adaxial and abaxial view of the leaf sheath. —D. Inflorescence. —E. Spikelet (all from Sanchez-Ken et al. 500). of a tropical rainforest. Altitudinally, it ranges from 1300 to 1900 m. The soils are reddish and usually sandy to clayish. Flowering occurs between Septem¬ ber and January. According to Henrard’s (1929) criteria, this spe¬ cies belongs to Aristida sect. Streptachne. As shown by the herbarium annotations, this species was con¬ fused with A. schiedeana in Flora Novo-Galiciana 192 Novon Table 1. Morphologic comparison among Aristida tuitensis, A. hintonii, and A. schiedeana. Character A. tuitensis A. hintonii A. schiedeana Branching basal, middle basal basal Number of internodes 9-15 2-10 2-7 Stem shorter longer longer Sheath length compared with internode length longer shorter shorter Leaf shape straight to flexuous straight to flexuous flexuous to coiled Inflorescence few-flowered flowered few-to many-flowered Inflorescence branches strongly spreading spreading appressed Pulvinus all branches usually all branches usually only first-order branches Spikelet usually paired paired and triads paired and triads Larger glume usually the first usually the second second or first Glumes not dentate dentate dentate Lateral awns lacking 0.1-1.0 mm 0.1-2.0 mm Anthers 3-4 mm 1.5-2.0 mm 1.5-2.0 mm (McVaugh, 1983). However, A. tuitensis has reli¬ able diagnostic characters by which it can be dis¬ tinguished from A. schiedeana and A. hintonii, probably the two most closely related species (Table 1). A notable character of A. tuitensis is the rela¬ tively long anthers compared to the lemma body. To our knowledge, such long anthers have not been reported in any other species of the genus. Artificial Key to Three Related Species of Aristida Sect. Streptachne la. Anthers 3-4 mm long; sheaths not striate; in¬ florescence branches strongly spreading. . A. tuitensis lb. Anthers 1.5-2 mm long; sheaths striate; inflo¬ rescence branches slightly spreading or ap- pressed. 2a. Spikelets usually spreading; pulvinus usu¬ ally in ail branches; leaf blades straight to flexuous; lateral awns up to 1.0 mm long . A. hintonii 2b. Spikelet appressed to branches; pulvinus in the branches of 1st order; leaf blades flexuous to coiled; lateral awns up to 2.0 111111 l° n 8 .. A. schiedeana The specific epithet refers to the area in which the species was discovered, El Tuito. Many other taxa have recently been discovered in this region. Paratypes. MEXICO. Jalisco: Mpio. Cuautitlan, Cer- ca de 10 km al sur-suroeste de las Joyas, Sierra de Manantlan, R. Guzman 6120 (COCA); Mpio. El Cuale, Lado este de las Minas de Zimapan, brecha a Cuale, R. Guzman M. 6101a (IBUG); Mpio. El Tuito, 30 mi. S of Puerto Vallarta, A. A. Beetle et al. M-3610 (COCA); Al sur-sureste de Puerto Vallarta, 14 km por la brecha a Minas de Zimapan, 1 km al oeste de Providencia por la brecha El Tuito-El Cuale, R. Guzman M. 6094 (COCA); Km 2 por la brecha a Minas de Zimapan, F. J. Santana M. 1196 (IBUG); Km 14 de la brecha El Tuito-Minas Zimapan, 1 km al oeste del Aserradero Provincia, F. J. Santana M. 1204 (IBUG); Mpio Talpa, Entre Cuale y Talpa, J. de la Torre V. s.n. (IBUG). Acknowledgments. We thank Jose Luis Villa¬ senor and Oswaldo Tellez for reviewing the manu¬ script. We thank the McArthur Foundation, the Consejo Nacional de Ciencia y Tecnologia (CONACyT CPECBN-021759), and the Direccion General de Asuntos del Personal Academico (D.G.A.P.A.- U.N.A.M.) for financial support for fieldwork and botanical research in different parts of Mexico. A special thanks to Eloisa Duarte for the elaboration of the drawing. Finally, we thank the Instituto de Biologia, U.N.A.M., where this research was carried out. Literature Cited Henrard, T. 1929. A monograph of the genus Aristida. Meded. Rijks-Herb. Leiden no. 58: 1-156. McVaugh, R. 1983. Gramineae. In: Flora Novo-Gali- ciana: A Descriptive Account of the Vascular Plants of Western Mexico. Vol. 14. The Univ. Michigan Press, Ann Arbor. Novelties in the Neotropical Genus Ouratea Aublet (Ochnaceae) Claude Sastre Laboratoire de Phanerogamic, Museum National d’Histoire Naturelle, 16 rue Buffon, F-75005, Paris, France ABSTRACT. A new section of Ouratea, section Cad- vcae Sastre, and eight new species of Ouratea from Venezuela and adjacent countries are described: O. pendulosepala Sastre (sect. Cardiocarpae), O. oli- gantha Steyermark ex Sastre (sect. Cardiocarpae), 0. apurensis Sastre (sect. Ouratea), 0. paratatei Sastre (sect. Ouratea), O. cidiana Sastre (sect. Our- atella), O. guriensis Sastre (sect. Ouratella), O. pseudoguildingii Sastre (sect. Ouratella), and O. squamata Sastre (sect. Ouratella). During preparation of the Ochnaceae treatment for the Flora of the Venezuelan Guayana, approx¬ imately 20 undescribed species of Ouratea Aublet from southern Venezuela and adjacent areas have been found. In order to properly place these new species within the subgeneric classification, a brief review and update of the previous system (Sastre, 1988) is needed. In the previous classification, I recognized six sections: Kaieteurea (Dwyer) Sastre, Cardiocarpae (Engler) Sastre, Polyouratea (Tiegh- em) Sastre, Persistens Sastre, Ouratella (Tieghem) Sastre, and Ouratea, which is the type section of the genus (with O. guianensis Aublet as the type). The new system also has six sections, but section Persistens is now placed in synonymy of section Ouratea, and section Caducae is newly described below. Aublet’s (1775) description of Ouratea guianen¬ sis was based on a flowering sheet (in BM), illustrated by plate no. 152, and all additional material of this species studied for my 1988 classification was either in flower or old fruit, making it impossible to assess the structure of the carpids (fruiting carpels) with precision. Recent study of herbarium specimens of O. guianensis from CAY, MO, NY, and VEN clearly shows, however, that its carpids are similar to those of the species previously included in section Per¬ sistens. This necessitates the transfer of section Persistens into the type section, and the recognition of a new section to include species previously (and erroneously) thought to be closely related to 0. guianensis, but whose fruit structure is now seen to be different. Ouratea sect. Caducae Sastre, sect. nov. TYPE: Ouratea longifolia (Lamarck) Engler in Mart., FI. Bras. 12(2): 316. 1876. Ochna longifolia Lamarck, Encycl. Meth. 4: 511. 1798. Inflorescentia terminalis, flores 5 sepalis, 5 carpellis, fructus 5 sepalis caducis, 1-3 carpellis verticalibus. Inflorescence terminal, flowers with 5 sepals and 5 carpels, sepals caducous in fruit, fruiting carpels 1-3, vertical. Key to the Sections of Ouratea 1. Flowers with 2-4 sepals united in 2-3 parts .. .1. sect. Kaieteurea 1. Flowers with 5 distinct sepals . 2 2(1). Fruits with the carpels borne horizontally .. .2. sect. Cardiocarpae 2. Fruits with the carpels borne vertically . 3 3(2). Flowers with (5)6-10 carpels. .3. sect Polyouratea 3. Flowers always with 5 carpels . 4 4(3). Fruits with persistent sepals ... 4. sect. Ouratea 4. Fruits without sepals . 5 5(4). Inflorescences axillary, rarely terminal .... .5. sect. Ouratella 5. Inflorescences always terminal 6. sect. Caducae Description and Nomenclature of the Sections 1. Ouratea Aublet sect. Kaieteurea (Dwyer) Sastre, Bull. Mus. Natl. Hist. Nat. 4°. ser. 10, sect. B Adansonia 1: 50. 1988. Kaieteurea Dwyer, Bull. Torrey Bot. Club 70: 50. 1943. TYPE: Ouratea gillyana (Dwyer) Sandwith & Ma¬ guire. Inflorescence terminal or axillary; flowers with 2-4 sepals united in 2-3 parts, and 5 (-7 in O. articulata Sastre) carpels; fruits with sepals cadu¬ cous or persistent and the carpels vertical. 2. Ouratea Aublet sect. Cardiocarpae (Engler) Sastre, Bull. Mus. Natl. Hist. Nat. 4°. ser. 10, sect. B Adansonia 1: 55. 1988. Ouratea sect. Eugomphia (Planchon) Engler, ser. Cardi¬ ocarpae Engler in Mart., FI. Bras. 12(2): 307. 1876. TYPE: Ouratea cardiosperma (DC.) Engler. Novon 5; 193-200. 1995. 194 Novon Diouratea Tieghem, Ann. Sci. Nat. Bot. ser. 8, 16: 227. 1902. Inflorescence terminal; flowers with 5 sepals and 5 carpels; fruit with the carpels horizontal. 3. Ouratea Aublet sect. Polyouratea (Tieghem) Sastre, Bull. Mus. Natl. Hist. Nat. 4°. ser. 10, sect. B Adansonia 1: 55. 1988. Polyouratea Tieghem, Ann. Sci. Nat. Bot. ser. 8, 16: 268. 1902. TYPE: Ouratea hexasperma (A. St. Hilaire) Baillon. Inflorescence terminal; flowers with 5 sepals and 6-10 carpels; fruit with the carpels vertical. 4. Ouratea Aublet sect. Ouratea. TYPE: Ouratea guianensis Aublet. Ouratea sect. Persistens Sastre, Bull. Mus. Natl. Hist. Nat. 4°. ser. 10, sect. B Adansonia 1: 59. 1988. Inflorescence terminal; flowers with 5 sepals and 5 carpels; fruits with sepals persistent and the carpels vertical. 5. Ouratea Aublet sect. Ouratella (Tieghem) Sastre, Bull. Mus. Natl. Hist. Nat. 4°. ser. 10, sect. B Adansonia 1: 59. 1988. Ouratella Tieghem, Ann. Sci. Nat. Bot. ser. 8, 16: 289. 1902. TYPE: Ouratea mexicana (Humboldt & Bonp- land) Engler. Inflorescence axillary, rarely terminal, spicate or paniculate; flowers with 5 sepals and 5 carpels, fruit with carpels vertical. 6. Ouratea Aublet sect. Caducae Sastre. TYPE: Ouratea longifolia (Lamarck) Engler. Ouratea Aublet sect. Ouratea, pro parte fide Sastre (1988) excluding O. guianensis. See description above. Inflorescence terminal; flowers with 5 sepals and 5 carpels; fruit with carpels vertical. Following are descriptions of eight new species in sections Cardiocarpae (two species), Ouratea (two species), and Ouratella (four species). The descrip¬ tion of new species in section Caducae will be pre¬ sented in a future paper. Ouratea Section Cardiocarpae Ouratea pendulosepala Sastre, sp. nov. TYPE: Brazil. Amazonas: Mun. Pres. Figueredo, 8 km da Vila Residencial Atrari, Canteiro de obras de Balbina. 1°30'-2°S, 59°30'-60°W, mata de terra firme, solo argiloso, 19 sep. 1986 (fr), C. A. Cid Ferreira et al. 8219 (holotype, P; is¬ otypes, INPA not seen, NY not seen). Figure 1 . Ab Ouratea cerebroidea Sastre, foliis nervis non im- pressis, paucis visibilibus, margine sub-integra, sepalis 12 x 2 mm, persistentibus pendusque in fructu, staminibus papillatis, stylo 10 mm longo, differt. Tree 12 m tall, branches glabrous. Leaves cori¬ aceous, 12-16 cm long, 5-6 cm wide, obovate, apex acuminate, base attenuate, margin undulate, subentire, principal vein salient on lower surface, lateral veins faintly evident. Inflorescence panicu¬ late, terminal, 14 cm long. Peduncle 13-15 mm long, weakly papillate; sepals 5, 10-11 x 2 mm, ovate; petals 5, obovate, 15-16 x 6-7 mm; sta¬ mens 10, papillate, 10 mm long. Young fruits with peduncle 15 mm long, sepals 5, pendent and per¬ sistent in older fruits, 12 x 2 mm, ovate. Fruits with 1-4 horizontal carpels, the carpophore sub- cylindric. Until now, section Cardiocarpae was restricted to the Guianas and had only two species: Ouratea cardiosperma (DC.) Engler and O. cerebroidea Sastre. The pendent, persistent sepals are charac¬ teristic of the new species and are unknown in the other species of Ouratea. Paratypes. BRAZIL. Amapa: Camipi, EMBRAPA reserve and vicinity, 00°10'N, 51°37'W, 6 Sep. 1983, S. Mori et al. 15864, 15865, 15868, 15869, 15874 (MG not seen, NY not seen, P). Amazonas: Mun. Bar- celos, 00°30'N, 63°30'W, rio Araca, 3 jul. 1985, E. Sette Silva et al. 214 (INPA not seen, NY not seen, P). Para: Mun. de Almeirin, Mte. Dourado, estrada MTD- oeste em dire$ao a serra da mina de bauxita, 17 sep. 1985 (fl), M. J. P. Pires et al. 627 (K not seen, P). Ouratea oligantha Steyermark ex Sastre, sp. nov. TYPE: Venezuela. Bolivar: Alto Rio Cuyuni, SE escarpment of Cerro Pitin, Cordillera Epi- cara, 400 m, B. Maguire, J. A. Steyermark & C. Maguire 53644 (holotype, NY). Figure 2 . Ab Ouratea pendulosepala Sastre, inflorescentiis par¬ vis (4-10 cm longis), laminis parvis (7-9 cm longis) et sepalis caducis in fructus, differt. Shrub 1.5—4 m tall, branches glabrous. Petiole 1 cm long, leaves coriaceous, 7-9 x 3-3.5 cm. TYPlj inntnit mil rwsimn CU-tiM-h^- f+-d~Ln[A. £»£>«. Figures 14. — 1 . Ouratea pendulosepala Sastre; Cid Ferreira et al. 821 9 (holotype, P). —2. Ouratea oligantha '^mark ex Sastre; Maguire et al. 53644 (holotype, NY). —3. Ouratea apurensis Sastre; Steyermark et al. Oh 62 (holotype, P). —4. Ouratea paratatei Sastre; Blanco 525 (holotype, P). 196 Novon elliptic, apex acuminate, base obtuse, margins toothed, undulate, principal vein salient on lower surface, with 8-12 unpaired lateral veins on each side of the principal vein, ascending, salient on lower surface between the small, subparallel veins, a few of which are also salient. Inflorescences usually ax¬ illary, sometimes terminal, spicate or sometimes pa¬ niculate, 4-10 cm long. Pedicel 12 mm long, sepals 5, 7 x 2 mm, oblong; petals 5, 15 x 6 mm, ovate; stamens 10, 1 cm long, carpels 5, horizontal; style single, 5.5 mm long. Young fruit with peduncle 15 mm long, sepals late-caducous, carpels 1-3, hemi¬ spheric, carpophore conical. Distribution. Known only from the type locality. The type and paratypes cited here were annotated by Steyermark at different times as “O. oligantha sp. nov.” and “O. soderstromii Sastre.” I have chosen to use Steyermark’s name for the new spe¬ cies. The material cited clearly does not belong to Ouratea soderstromii; the new species belongs to section Cardiocarpae, while O. soderstromii is a member of section Ouratella. The name used by Steyermark is validated here. Ouratea oligantha is most closely related to O. pendulosepala, described above, but differs by the short inflorescences (4-10 cm), that are usually spicate (vs. largely paniculate), short leaves (7—9 cm long), and the sepals caducous in fruit. Ouratea pendulosepala has longer inflo¬ rescences (14 cm long), longer leaves (12-16 cm long), and persistent sepals. Ouratea oligantha is the first species of section Cardiocarpae known in Venezuela. Paratypes. VENEZUELA. Bolivar: Alto Rio Cuyuni, Rio Chicanan, Cerro Pitin, Cordillera Epicara, 3 sep. 1962, Maguire et al. 53568 (NY), 9-11 sep. 1962, Maguire et al. 53679 (NY). Ouratea Section Ouratea Ouratea apurensis Sastre, sp. nov. TYPE: Ven¬ ezuela. Apure: Reserva Forestal San Camilo, selva siempreverde a lo largo de la Quebrada de la Cristalina hasta la vega del Cerro la Nulita, 250-280 m, 2 apr. 1968, Steyermark et al. 101762 (holotype, P; isotypes, NY, VEN). Fig¬ ure 3. Ab Ouratea marahuacensi Maguire et Steyermark, folia petiolo 2-3 mm longo (non 5-9 mm) nervis later- alibus prominentibus, sepalis 8 x 2-3 mm, petalis 10 x 5 mm, staminibus 7.5 mm longis, stylo 7.5 mm longo, differt. Tree 7 m tall, branches glabrous. Petioles 2-3 mm long, leaf blades elliptic, undulate, 6.5-9 x 2- 2.5 cm, apex acuminate, base attenuate, margin crenulate, principal vein salient on the lower surface, lateral vein prominent, transversal tertiary veins parallel. Inflorescence paniculate, terminal, 8-11 cm long. Peduncle 5-6 mm long, with short hairs 0.2-0.3 mm long; sepals 5, oblong-lanceolate, 8 x 2-3 mm; petals 5, obovate, 10 x 5 mm; stamens 10, 7.5 mm long; style 7.5 mm long. Young fruits with sepals persistent. Both Ouratea marahuacensis Maguire & Stey¬ ermark and O. apurensis occur in Venezuela, the former on Cerro Marahuaca, Estado Amazonas, at 1150 m elevation, the latter in the Reserva Forestal San Camilo, Estado Apure. Ouratea apurensis dif¬ fers from 0 . marahuacensis by the shorter petiole (2-3 mm long vs. 5-9 mm long), salient lateral veins, longer sepals (8 mm long vs. 6-7.5 mm long), longer petals (10 mm long vs. 6-7.5 mm long), and longer stamens (7.5 mm long vs. 5-6 mm long). Ouratea apurensis is known only from the type collection. Ouratea paratatei Sastre, sp. nov. TYPE: Ven¬ ezuela. Bolivar: Campamentos “El Paraiso” y “La Yagua” a 48 km NE del caserio Las Rosas, este ultimo a 17 km de Upata (carretera nueva Upata-San Felix), jun. 1966, Blanco 525 (ho¬ lotype, P; isotypes, NY, VEN). Figure 4. Ab Ouratea tatei Gleason, lamina elliptica viridi, nervo principali prominenti pagina superiore, nervis lateralibus non impressis, differt. Ab O. pseudotatei Maguire et Stey¬ ermark, lamina nervo principali prominenti pagina su¬ periore, nervis tertiaris paucis visibilibus (non impressis), fructus sepalis persistentibus, differt. Tree 7 m tall, branches glabrous. Petiole 1 cm long, leaf blade coriaceous, elliptic-ovate, 14-19 x 4.5-7 cm, apex acuminate, base obtuse to round, margin undulate-denticulate, principal vein raised on upper surface, lateral veins subequal, tertiary veins anastomosing, faintly evident. Inflorescence paniculate, terminal, 12 cm long. Flowers not known. Fruits with 5 persistent sepals, these 7 x 2.5 mm, lanceolate, acuminate, the carpels 1-3, obovate, 8- 9 x 5.6-6 mm, carpophore conical, 5-6 mm long, peduncle 1 cm long. Ouratea tatei Gleason, O. pseudotatei Maguire & Steyermark, and O. paratatei are similar, but the fruits of O. pseudotatei do not have persistent sepals and so belong to section Caducae; O. pseu¬ dotatei is endemic to Guyana. Ouratea tatei and O. paratatei are Venezuelan species, the former occurring in the highlands of Estado Bolivar (Ro- raima-tepui, Chimanta-tepui), the latter from the Volume 5, Number 2 1995 Sastre Ouratea 197 lowlands of Estado Bolivar in the Sierra de Imataca. The differences of the Venezuelan species in section Ouratea are outlined in the key below. Key to the Venezuelan Species of Ouratea Sect. Ouratea 1. Leaf blades with 2 types of secondary veins: large ascending ones terminating along the blade margin, and between these and the midvein, some smaller, horizontal, ± parallel veins . 2 1. Leaf blades with subequal secondary veins .... 4 2(1). Leaves 18-22 cm long, sepals 9-10 mm long . 0. guianensis Aublet 2. Leaves 5-9 cm long, sepals 5-8 mm long ... 3 3(2). Petiole 2-3 mm long; sepals 8 mm long; petals 10 mm long. O. apurensis Sastre 3. Petiole 5-9 mm long; sepals 6-7.5 mm long; petals 5-6 mm long . .... 0. marahuacensis Maguire & Steyermark 4(1). Leaves 12-19 cm long . 5 4. Leaves 3-12 cm long . 6 5(4). Leaf blades oblong, veins on the lower surface of the blade impressed . 0. tatei Gleason 5. Leaf blades elliptic-ovate; veins on the lower surface of the blade not impressed . . O. paratatei Sastre 6(4). Leaf margin clearly undulate and toothed .. . 0. superba Engler 6. Leaf margin few-toothed . 7 7(6). Leaf blades 3-6 x 1. 7-2.7 cm . . O. longistyla Maguire & Steyermark 7. Leaf blades 6-12 x 3-5 cm. . 0. spruceana Engler Ouratea Sect. Ouratella Ouratea cidiana Sastre, sp. nov. TYPE: Brazil. Para: Mun. Oriximina, Rio Trombetas a mon- tante do Cachoeira Portura prox. as ilhas Res- posta, 00°52'S, 57°03'W, 22 aug. 1986, Cid Ferreira et al. 7971 (holotype, P; isotypes, INPA not seen, NY not seen). Figure 5. Ab 0. soderstromii Sastre basi foliae obtusa vel ro- tundata, margine serrata, inflorescentia 2-8 cm longa differt. Shrub 3 m tall, stems glabrous. Leaves coria¬ ceous, elliptic, 6-9.5 x 1.5-4 cm; base rounded; margin undulate and denticulate; apex acuminate; principal vein slightly prominent above, with 9-11 unpaired, arcuate lateral veins on each side of prin¬ cipal vein, with fine and parallel intersecondaries; petiole 5-6 mm long. Inflorescences axillary, spi- cate, 2-8 cm long, the peduncle 4 mm long. Sepals 5, obovate, 6x3 mm; petals 5, obovate, 7x4 mm; stamens 10, anthers sessile, 5 mm long; style 6 mm long. Fruit not seen. Distribution. Known only from the type locality, in Rio Trombetas, Para, Brazil. Ouratea cidiana differs from O. soderstromii by its rounded leaf bases (vs. obtuse); serrate margins (vs. denticulate), and inflorescences 2-8 cm long (vs. 8-10 cm). Paratype. BRAZIL. Para: Rio Trombetas, near Cach¬ oeira Porteira, 28 May 1974, Campbell et al. P.22383 (COL, INPA not seen, NY). Ouratea guriensis Sastre, sp. nov. TYPE: Ven¬ ezuela. Bolivar: Represa Guri, islands and W side of river, 1 Apr. 1981, Liesner & Gonzalez 11081 (holotype, VEN; isotypes, MO, NY). Figure 6. Ab 0. soderstromii Sastre nervis II et III anastomo- santibus, inflorescentiis 2-8 cm longis, carpophoro fructi discoideo 3 mm alto (non conico 10-13 mm alto) differt. Shrub 1.5 m tall, stems glabrous. Leaves coria¬ ceous, unequal, elliptic, 6-9.5 x 1.5-4 cm; base obtuse or rounded, margin undulate and serrate, apex acuminate; principal vein slightly prominent on lower surface, the secondary veins subequal and anastomosing with the tertiary; petiole 5-7 mm long. Inflorescences axillary, spicate, 2-8 cm long with 1-3 leaves in the basal part, the peduncle 6 mm long. Sepals 5, coriaceous, triangular, 7x3 mm; petals 5, obovate, 9-10 x 1.5-2 mm; stamens 10, 6 mm long. Fruits with carpophore discoid, 3x8 mm, carpels 7-9 x 4-5 mm. Distribution. Known only from the type locality in Estado Bolivar near the Guri Dam. Ouratea guriensis differs from 0. soderstromii by its anastomosed secondary and tertiary veins, inflorescence 2-8 cm long (vs. 8-10 cm), and car¬ pophore 3 mm long and discoid (vs. 10-13 mm long and conic). It differs from O. cidiana by its anastomosed secondary and tertiary veins. Paratype. VENEZUELA. Bolivar: Represa Guri, ca. 0.5 km SSW of dam, on a steep forested slope with open rocks at top, ca. 250-350 m, 7°46'N, 63°00'W, 31 Mar. 1981, Liesner & Gonzalez 11034 (MO, NY, VEN). Ouratea pseudoguildingii Sastre, sp. nov. TYPE: Venezuela. Bolivar: Rio Horeda, 1-4 km above mouth, 90 m, 15 Dec. 1955, Wurdach & Monachino 39901 (holotype, P; isotype, NY). Figure 7. Ab O. guildingii (Planchon) van Tieghem, foliis apice obtusa (non acuta), nervis II subparallelis (et non inae- qualibus cum 8-12 magnis inter nervis subparallelis ten¬ ues transversales), differt. Ab O. grosourdyi Van Tiegh¬ em, foliis apice obtusa (non acuta et mucronata), nervis II et III anastomosantibus, inflorescentiis paniculatis (et non spiciformibus), differt. Figures 5-8. —5. Ouratea cidiana Sastre; Cid Ferreira et al. 7971 (holotype, P). —6. Ouralea guriensis Sastre; Liesner & Gonzalez 11081 (holotype, VEN). —7. Ouratea pseudoguildingii Sastre; Wurdack & Monachino 39901 (holotype, P). —8. Ouratea squamata Sastre; Steyermark et al. 123963 (holotype, VEN). Volume 5, Number 2 1995 Sastre Ouratea 199 Tree 3-5 m tall, stems glabrous. Leaves coria¬ ceous, elliptic, 3-5.5 x 1.5-2.5 cm; base rounded; margin denticulate; apex obtuse; principal vein prominent below, the secondary veins subparallel, the secondary and tertiary veins anastomosed; pet¬ iole 2 mm long. Inflorescence axillary, paniculate, 5-10 cm long, the peduncle 7-8 mm long. Sepals 5, coriaceous, obovate, 8x2 mm; petals 5, spa- thulate, 9x6 mm; stamens 10, a few ornate, anthers subsessile, 5 mm long; carpels 5, style 5 mm long. Fruit not seen. Ouratea pseudoguildingii differs from O. guil- dingii by its obtuse leaf apices (vs. acute) and sub¬ parallel venation. It differs from O. grosourdyi by its obtuse leaf apices (vs. acute and mucronate), secondary and tertiary venation anastomosed (vs. not anastomosed), and paniculate inflorescence (vs. spicate). Distribution. Middle Rio Orinoco, Venezuela. Paratypes. VENEZUELA. Bolivar: Puerto Ordaz, San Felix, Apr. 1964, Aristiguieta 5280 (NY, VEN). Guarico: Dto. Infante, Municipio Cabrita, laguna del Caiman, Finca los Becerros, 21 mar. 1990, S. Tillelt el al. SPB 279 (P, UCU). Ouratea squamata Sastre, sp. nov. TYPE: Ven¬ ezuela. Bolivar: Distr. Heres, cerro Marutani, cumbre, afloramiento de piedra arenisca en la altiplanicie a lo largo del Rio Carla, afluente de las cabeceras del Rio Paragua, 3°50'N, 62°15'W, 1200 m, 11-14 jan. 1981, Stey- ermark et al. 123963 (holotype, VEN; isotype, NY). Figure 8. Ab O. poeppigii van Tieghem foliis inaequalibus, nervis lateralibus pagina superiore impressis, pagina inferiore non visibilibus, nervis II et III anastomosantibus, differt. Tree, 10 m tall, young stems with scales. Leaves coriaceous, unequal, elliptic, sometimes folded, 10- 14 x 4-6 cm; base rounded; margin denticulate; apex acuminate; principal vein prominent below, secondary veins subparallel, impressed above, not apparent below, secondary and tertiary veins anas¬ tomosed, petiole 3-4 mm long. Inflorescence axil¬ lary, paniculate, 16-26 cm long with 1-3 leaves at the basal part, axes with scales. Flowers not seen. Fruit with conical carpophore, 5-6 x 5 mm, carpels 5-6 x 4 mm. Distribution. Known only from the type locality on Cerro Marutani, Bolivar, Venezuela, near the headwaters of the Rio Paragua. Known only from the type specimen; Ouratea squamata differs from O. poeppigii, an Amazonian Brazil species, by its unequal leaves, lateral veins impressed above and not apparent below, and the secondary and tertiary venation anastomosed. Key to the Species of Ouratea Sect. Ouratella from Venezuela and Adjacent Areas 1. Inflorescence spicate, not or few-branched .... 2 1. Inflorescence paniculate .. 10 2(1). Inflorescence 11-30 cm long . 3 2. Inflorescence 2-10 cm long. 4 3(2). Leaves 10-15 x 4-6.5 cm. . 0. croizatii Steyermark & Maguire 3. Leaves 6-10 x 3-6 cm . . O. grosourdyi (Tieghem) Steyermark 4(2). Leaf blade with the margin rolled and few¬ toothed ... O. yapacana Sastre 4. Leaf blade flat with the margin toothed . 5 5(4). Leaves 20-60 cm long . 6 5. Leaves less than 15 cm long . 7 6(5). Leaf blade oblong, 30-60 cm long . . O. ramiflora Sastre 6. Leaf blade ovate, 20-35 cm long . . O. ornata Maguire & Steyermark 7(5). Leaf blade 3-5 x 2-5 cm. .O. guaiquinimensis Sastre 7. Leaf blade 6-15 x 1.5-7 cm. 8 8(7). Leaf blade membranous, inflorescences 8-10 cm long . O. soderstromii Sastre 8. Leaf blade coriaceous, inflorescences 2-8 cm long . 9 9(8). Secondary and tertiary leaf venation reticu¬ late . O. guriensis Sastre 9. Secondary and tertiary leaf venation not re¬ ticulate . O. cidiana Sastre 10(1). Leaf blades with length/width ratio 1-2.5 11 10. Leaf blades with length/width ratio > 2.5 14 11(10). Leaf blades 3-5.5 x 1.5-2.5 cm . . O. pseudoguildingii Sastre 11. Leaf blades 6-15 x 3.5-6 cm. 12 12(11). Leaf blade with 2 types of lateral veins; inflorescence 4-7 cm long. . O. caracasana (Planchon) Engler 12. Leaf blade with lateral veins subequal, in¬ florescence 7-17 cm long . 13 13(12). Venation of the leaf blade not impressed, inflorescence 7-10 cm long . . 0. ramossisima Maguire & Steyermark 13. Venation of the leaf blade impressed, in¬ florescence 16-17 cm long . .O. timehriensis Sastre 14(10). Inflorescences 3-5 cm long. . O. guildingii (Planchon) Urban 14. Inflorescences > 5 cm long . 15 15(14). Leaf blade with lateral veins subequal, sec¬ ondary and tertiary veins anastomosed .. 16 15. Leaf blade with 2 types of lateral veins, secondary and tertiary veins not anasto¬ mosed . 1 ‘ 16(15). Leaf blades 5.5-8 x 2-3 cm; young stems without scales; inflorescence axis papillate . O. papillata Maguire & Steyermark 16. Leaf blades 10-14 x 4-6 cm; young stems with scales; inflorescence axis not papillate .... O. squamata Sastre 200 Novon 17(15). Leaf blades coriaceous . . O. duidae Steyermark 17. Leaf blades membranaceous . . O. poeppigii Van Tieghem Ouratea cidiana, 0. ramiflora , and O. poep¬ pigii occur in Amazonian Brazil, and 0. timehrien- sis occurs in Guyana. Acknowledgments. I thank Bruce K. Holst for assistance with the manuscript. Literature Cited Aublet, F. 1775. Histoire des plantes de la Guyane fran£aise 1: 1-261. Paris. Engler, A. 1876. Ochnaceae. In: Martius, Flora Bras- iliensis 12(2): 301-332, pi. 62-77. Monachii. Kanis, A. 1968. A revision of the Ochnaceae of the Indo-Pacific area. Blumea 16: 1-82. Sastre, C. 1988. Synopsis generis Ouratea Aublet (Och¬ naceae). Bull. Mus. Natl. Hist. Nat. Paris 4°. ser., sect. B Adansonia 1: 47-67. Tieghem, P. van. 1902. Sur les Ochnacees. Ann. Sci. Nat. Bot., ser 8, 16: 161-416. New Species and Combinations in Rubiaceae from Costa Rica and Panama Charlotte M. Taylor Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. ABSTRACT. The new combinations Psychotria ne- bulosa (Dwyer) C. M. Taylor and P. roseocrema (Dwyer) C. M. Taylor are made, and the following new species are described: Chiococca caputensis D. H. Lorence & C. M. Taylor, Manettia longi- pedicellata C. M. Taylor, Pentagonia lobata C. M. Taylor, Psychotria burgeri C. M. Taylor, Psy¬ chotria saltatrix C. M. Taylor, and Rudgea laevis C. M. Taylor. During review of Rubiaceae materials for prep¬ aration of the Manual to the Plants of Costa Rica, the following new species and taxonomic problems were discovered. These comments supplement the treatment by Burger & Taylor (1993). Chiococca caputensis D. H. Lorence & C. M. Taylor, sp. nov. TYPE: Panama. Panama: Cer- ro Jefe, NE of Panama City, 9°15'N, 79°30'W, 12 July 1986, G. McPherson 9744 (holotype, MO 3624428; isotype, PMA not seen). Figure IB, C. Chiococcae belizensi Lundell similis, sed ab ea corollis albis etiam nervis secundariis brochidodromis semel ar- cuatis differt. Glabrous low shrubs or usually woody vines. Leaves 2-8 cm long, 1.2-4 cm wide, elliptic, at apex acute to slightly acuminate or sometimes ob¬ tuse, at base cuneate, coriaceous; secondary veins 3-5 pairs, brochidodromous, looping once to inter¬ connect, above and below plane, with the costa plane or prominulous on both surfaces; petioles 4— 10 mm long; stipules interpetiolar, triangular, 2-3 mm long, acuminate to usually apiculate. Inflorescences ax¬ illary, 1 per axil, racemose to sometimes 1-2 times branched, 3-6 cm long; peduncles 5-15 mm long; bracts subtending pedicels deltoid to narrowly tri¬ angular, 0.5-1.5 mm long; pedicels 1-4 mm long; calyx glabrous, green, with hypanthium ellipsoid, somewhat flattened laterally, 2-2.5 mm long, limb 1-1.3 mm long, divided for ca. 14 its length, lobes deltoid, entire to ciliolate; corolla glabrous through¬ out, white becoming cream to pale yellow, funnel- form, tube 6.5-7 mm long, ca. 1 mm diam. at base and 4-5 mm diam. at apex, lobes narrowly trian¬ gular, 3-4 mm long, acute; stamens included, fil¬ aments ca. 1 mm long, pilose, anthers very narrowly oblong, 3.5-4 mm long; style 11-11.5 mm long, twisted, white, papillose throughout, at apex clavate. Fruit ellipsoid, strongly laterally compressed, 8-12 mm diam., white, fleshy. Habitat and phenology. Known only from wet forest on the upper slopes and summit of Cerro Jefe in central Panama, at 550-1000 m. Collected in flower in July, in mature fruit September through February. Chiococca caputensis is similar in its included stamens and climbing habit to C. alba (L.) A. S. Hitchcock, which can be distinguished by its pa¬ pyraceous leaf blades and usually smaller mature fruits, 4-8 mm in diameter. This new species has also been confused with C. belizensis Lundell, which has coriaceous leaves, included stamens, and fruit size similar to that of C. caputensis, but which can be distinguished from C. caputensis by its lemon yellow corollas and secondary leaf veins connected by one to three additional smaller loops outside the principal loops. Additionally, C. belizensis is found from sea level to 1500 m but has been collected predominantly below 400 m elevation. Chiococca caputensis has been confused with C. durifolia Dwyer, a synonym of C. belizensis, and also with C. jefensis Dwyer, which is a synonym of Elaeagia nitidifolia Dwyer. The specific epithet refers to the type locality, Cerro Jefe, which trans¬ lated from the Spanish means “chief’ mountain. Several other species of Rubiaceae, for example, Palicourea tubuliflora Dwyer and Psychotria ol- gae Dwyer & M. V. Hayden, appear to be endemic to this mountain, and C. caputensis may be a similar local endemic. Paratypes. PANAMA. Panama: beyond Goofy Lake on road to Cerro Jefe, Correa & Dressier 467 (MO), Croat 15235 (MO); Cerro Jefe, NE of Panama City, Correa et al. 4576 (MO, PMA), Croat 35865, D’Arcy 12192 (MO), Elias & Hayden 1799 (MO), Folsom & Harp 1373 (MO), Folsom 3854 (MO), Gentry 2878 (MO), Gentry & Dwyer 5540 (MO), Gentry 6133 (MO), Kirkbride & Crobbs 9 (MO), McDonagh et al. 47 (BM, MO), McPherson 9733 (MO, PMA), 10012 ( MO, PMA), 10032 (MO, PMA, PTBG), 15855 (MO, PMA), Mori Novon 5: 201-207. 1995. 202 Novon 6527 (MO), 7994 (MO), Sytsma 1490 (MO), Tyson et al. 3207 (MO), Tyson 3374 (MO), Witherspoon 8554 (MO). Manettia longipedicellata C. M. Taylor, sp. nov. TYPE: Costa Rica. Limon: Canton de Matina, Cordillera Talamanca, 200 m aguas abajo de la confluencia de la quebrada Canabral con el rio Barbilla, 10°00'10''N, 83°25'30"W, 5 Nov. 1988, G. Herrera 2290 (holotype, CR; isotype, MO 3649361). Figure 1A. Flores in cymas paucifloras pedunculis 1-2.5 cm longis insidentes aggregati, pedicellis 1-2.5 cm longis; lobis ca- lycinis 4, 2 mm longis; corolla viridula ad rosea, tubo 6- 10 mm longo, lobis 4, 3-4 mm longis. Glabrescent to retrorsely hispidulous, extensively twining, herbaceous to suffrutescent vines. Leaves elliptic to ovate or broadly lanceolate, 3-12 cm long, 1.5-5 cm wide, acuminate at apex, cuneate to trun¬ cate at base, membranaceous; secondary veins 4- 6 pairs; petioles 5-15 mm long; stipules interpetio- lar, triangular, 1 mm long, with 4-8 cartilaginous teeth or triangular appendages, these often caducous and the stipules appearing truncate. Inflorescences axillary and terminal, cymose, with 1-4 flowers; peduncles 1-2.5 cm long; bracts triangular, to 2.5 mm long, scarious; pedicels 1-2.5 cm long, often thickened near apex and flowers and fruits appearing stipitate; calyx glabrescent, limb 2-4 mm long, di¬ vided to base, lobes 4, narrowly triangular, acute, reflexed; corolla tubular, externally glabrous, inter¬ nally pilose, green to white or usually pink with exserted yellow trichomes in throat, tube 6-10 mm long, lobes ovate, 3-4 mm long, acute. Capsules obconic, 5-9 mm long, frequently somewhat dicoc- cous, compressed laterally; seeds orbicular to some¬ what irregular, ca. 2 mm diam. Habitat and phenology. Wet Caribbean slopes of Nicaragua and Costa Rica at 100-1100 m, most frequently collected as a vine climbing on thickets along streams below 300 m elevation. Collected in flower and fruit usually concurrently, February to May and in December. This new species is similar to Manettia reclinata L., which can be distinguished in southern Central America by its six to eight calyx lobes 4-7 mm long, dark red corollas, and pedicels usually 1 cm long or shorter, in contrast to four calyx lobes 2- 4 mm long, green to white or pink corollas, and pedicels 1-2.5 cm long in M. longipedicellata. Plants of M. reclinata from Mexico and northern Central America may have peduncles and pedicels to 4 cm long, but differ in the other characters. Paratypes. COSTA RICA. Alajuela: Piedades de San Ramon, Brenes 4555 (CR). Cartago: Florencia Norte, CATIE, Turrialba, Poveda 3109 (CR). Heredia: Finca La Selva, the OTS field station near Puerto Viejo de Sarapiqui, near the junction of the Rios Puerto Viejo and Sarapiqui, Almeda & Anderson 5130 (F), Folsom 8851 (DUKE), 10032 (DUKE), Grayum 1144 (DUKE), 2711 (DUKE), Hammel 8099 (DUKE), 11752 (DUKE), Opler 1645 (CR), Wilbur 28181 (DUKE), 28308 (DUKE), 38626 (DUKE), 39485 (DUKE). Limon: Canton de Pococi, along road between Mata de Limon (on quadrangle map as “Milloncito”) and Millon, 10°24'30"N, 83°37'W, Grayum & Hammel 8407 (CR, MO). NICARAGUA. Zelaya: Cano Monte Cristo, “Las Faldas,” 11°36'N, 83°51'W, Moreno 14776 (MO); Montecristo, al N de Barra Punta Gorda, 11°34'N, 83°48'W, Sandino 2232 (MO); Cano Costa Riquita, ca. 1.8 km SW of Colonia Naciones Unidas, above road between there and Colonia Nueva Leon, 11°43'N, 83°18'W, Stevens 4942 (MO). Pentagonia lobata C. M. Taylor, sp. nov. TYPE: Costa Rica. Puntarenas: Canton de Golfito, Re- serva Natural de Vida Silvestre Golfito, final del camino hacia las torres, 8°39'52"N, 83°10'00"W, 7 Dec. 1992, B. Hammel 18619 (holotype, MO 4355639; isotypes, CR, F, MO 4355638). Figure ID. A speciebus ceteris Pentagoniae Bentham foliis pin- natifidis etiam calyce spathaceo differt. Monocaulous shrub or small tree to 7 m tall; stems lenticellate. Leaves 62-99 cm long, 60-90 cm wide, deeply pinnatifid, with sinuses extending to within 3-5 cm of costa, with lobes 4-6 on each side, these acute to somewhat acuminate, 7-14 cm wide, the basalmost lobes usually ca. % as long and wide, the entire blade chartaceous to subcoriaceous, glabrous and smooth adaxially, abaxially glabrescent to usually spreading-puberulous or shortly pilosulous on principal veins and sometimes also on lamina, abaxially lamina also finely striate, costa and longer secondary veins prominent above and below, shorter secondary veins slightly thickened, minor venation not visible; petioles 12-22 cm long; stipules ligu- late, 6-8 cm long, 2.5-3.5 cm wide, coriaceous, acute, with costa thickened near base, finely ap- pressed-puberulous to sericeous. Flowers hermaph¬ roditic, glomerulate in groups of ca. 12-20 in leaf axils, each inflorescence enclosed by 1 bract resem¬ bling a stipule, these bracts 8-9 cm long, 3-7 cm wide, each flower with a stipe 1-3 mm long and subtended by 1 floral bract, these floral bracts 15- 25 mm long, 6-12 mm wide, ovate, acute; hypan- thium glabrous, cylindrical to ellipsoid, 3—5 mm long, 2-3 mm diam.; calyx limb red, coriaceous, glabrous, 2.5-4 cm long, spathaceous, splitting for ca. % of its length, acute at apex; corolla green to cream, funnelform, carnose, fibrous, glabrous Volume 5, Number 2 1995 Taylor Rubiaceae from Costa Rica and Panama 203 Figure 1. —A. Manettia longipedicellata C. M. Taylor (from Grayum 1144, DUKE), habit. B, C. Chiococca caputensis C. M. Taylor & D. H. Lorence. —B. Habit (from McPherson 10012, MO). —C. Flower (from McPherson 9744, MO). —D. Pentagonia lobata C. M. Taylor, flower (from Hammel 18619, MO). —E. Psychotria burgeri c. M. Taylor, habit (from Haber 9778, F). F, G. Rudgea laevis C. M. Taylor. —F. Flower (from Haber 452, MO). —G. Habit (from Haber 1070, MO). A, B, D, E, G to 5-cm scale; C, F to 5-mm scale. 204 Novon throughout, tubes 38-46 mm long, 3-5 mm diam. at base, 10-15 mm diam. at apex, curved, lobes 5, deltoid, 4-8 mm long; stamens 5, included, fila¬ ments attached ca. V 3 of length of tube above base, ca. 15 mm long, swollen and densely pilose to villous at attachment point, glabrous otherwise, anthers narrowly oblong, ca. 3.5 mm long; style included, ca. 2 cm long, stigmas 1, lanceolate, ca. 2 mm long; disk bilobed, ca. 1.5 mm high. Fruit not seen. Habitat and phenology. Moist forest in the vi¬ cinity of Golfito and on the Osa Peninsula in the Golfo Dulce region of Costa Rica at 300-500 m. Collected in flower in July and December. This species is included in Pentagonia based on its fleshy monocaulous habit (characteristic though not exclusive in this genus), relatively large, firm- textured leaves with the abaxial surface finely stri¬ ate, and five-merous, relatively large, fleshy flowers borne in contracted axillary inflorescences. The fine¬ ly striate texture of the abaxial leaf surfaces with the minor venation not at all visible seems to be unique to this genus. Pentagonia lobata is distinguished within Pen¬ tagonia by its combination of pinnatifid leaves and a spathaceous calyx. Pinnatifid leaves also charac¬ terize P. tinajita Seemann of Costa Rica and Pan¬ ama (here circumscribed to include P. alfaroana Standley and P. gymnopoda Standley), which can be separated by its regularly lobed calyx limb. A spathaceous calyx also characterizes P. spathicalyx K. Schumann of Amazonian Brazil, Ecuador, and Peru, which has elliptic entire leaves. Pentagonia is poorly known taxonomically and ecologically. Individuals of most species are appar¬ ently infrequent, or at least infrequently collected, and good specimens are difficult to make due to the large fleshy leaves and flowers. Some species cannot be separated vegetatively. Flowering characteristics appear to be reliable for separating species, but flowering specimens are sometimes difficult to relate to fruiting specimens (the majority of collections). The range of size and shape variation seen in her¬ barium material is notable in some species, to 100% in some features (see descriptions in Burger & Tay¬ lor, 1993): this may be real morphological variation, or it may reflect variable shrinkage or other artifacts of the preparation of dried specimens. Some species of Pentagonia were originally separated based on details of leaf shape such as acute versus rounded leaf bases, but with more material available these conditions often show continuous variation and are not adequate to distinguish species. The deeply pin¬ natifid leaves found in some Pentagonia species are here regarded as a consistent character at the spe¬ cies level, although this has not been verified with field observations. Good taxonomic knowledge of Pentagonia will require extensive field-based study. Paratypes. COSTA RICA. Puntarenas: Canton de Osa, Reserva Forestal Golfo Dulce, Rancho Quemado, 8°42'00"N, 83°35'40”W, Hammel et al. 18309 (CR, F, MO). Psychotria burgeri C. M. Taylor, sp. nov. TYPE: Costa Rica. Puntarenas: Canton de Puntarenas, Monteverde, along Quebrada Maquina, 10°18’N, 84°48’W, 12 Mar. 1990, W. Haber 9778 (holotype, CR; isotypes, F, MO 4619858, US). Figure IE. A Psychotria pleuropoda J. Donnell Smith foliis sub- sessilibus basim rotundatis ad truncatis differt. Small trees to 3 m tall, glabrous. Leaves narrowly elliptic to oblanceolate, 7-13 cm long, 1.4-3.6 cm wide, at apex acute to slightly acuminate, at base truncate to rounded, chartaceous, glabrous above and below; secondary veins 10-12 pairs, looping to interconnect in a rather straight submarginal vein, plane above, prominulous below, without domatia; petioles stout, 2-3 mm long, glabrous; stipules unit¬ ed around the stem, deciduous below distalmost 1- 3 nodes, ovate, split along one side, 6-15 mm long, acute or frequently bilobed with the lobes acuminate, sometimes strongly costate, glabrous, ciliate. Inflo¬ rescences terminal, pyramidal; peduncles 2-4.5 cm long; panicles 2-3 cm long and wide, branched once, with bracts 3-4 mm long, triangular to ovate, ciliate, acute; flowers sessile in 3 glomerules of 3-5; hy- panthium not seen; calyx limb green, glabrous, 1- 1.5 mm long, lobed for ca. 14 °f its length, lobes deltoid, acute, ciliolate; corollas not seen. Infruc- tescences similar to inflorescences; fruit ellipsoid, 8-11 mm long, 6-12 mm diam., becoming red; pyrenes with 4-5 low rounded longitudinal ridges. Habitat and phenology. Wet premontane forest at ca. 1400 m in the Cordillera de Tilaran, Costa Rica. Collected in flower and fruit in March. Psychotria burgeri is placed in subgenus Psy¬ chotria based on its deciduous stipules, red fruits, and distinctive gray drying color. Subgenus Psy¬ chotria is well known in Central America (Hamilton, 1989), and even though its flowers have not been seen, this new species is clearly undescribed. It is similar to P. pleuropoda J. Donnell Smith of north¬ ern Central America, which can be separated by its leaves with acute to tapered bases and secondary veins 15-18 pairs, petioles 2-10 mm long, and elevational range of 50-500 m; P. pleuropoda is so far not known from Costa Rica. Range disjunc- Volume 5, Number 2 1995 Taylor Rubiaceae from Costa Rica and Panama 205 tions and disjunct species pairs between northern Central America and the Cordillera de Tilaran in Costa Rica are frequent in Costa Rican Rubiaceae (Burger & Taylor, 1993), although few are accom¬ panied by habitat shifts such as this. The specific epithet honors William C. Burger, a long-time stu¬ dent of the flora and phytogeography of Costa Rica. Psychotria nebulosa (Dwyer) C. M. Taylor, comb, nov. Basionym: Coussarea nebulosa Dwyer, Ann. Missouri Bot. Gard. 67: 131. 1980. TYPE: Panama. Chiriqui: Monte Rey near Bo- quete, 20 July 1971, T. B. Croat 15868 (ho- lotype, MO 2162999; isotype, MO 4043108). Rudgea chiriquiensis Dwyer, Ann. Missouri Bot. Gard. 67: 476. 1980. Syn. nov. Not Psychotria chiri¬ quiensis (Standley) C. M. Taylor & W. C. Burger, 1993. TYPE: Panama. Chiriqui: Cerro Colorado, along road above San Felix 29 km above bridge over Rio San Felix (7.9 km above turnoff to Escopeta), 14 July 1976, T. B. Croat 37071 (holotype, MO 2798805; isotype, MO 2389189). Study of additional material available since the publication of Coussarea nebulosa and Rudgea chi¬ riquiensis shows that they refer to the same species, and that this species is best placed in Psychotria. The inflorescences on the type specimens of R. chiriquiensis are older and consequently larger and relatively more open than those on the type speci¬ mens of C. nebulosa, but additional material shows this to be a developmental series, and no other features separate these specimens. The mature fruits are fleshy drupes 12-14 mm long with two hard, longitudinally ridged pyrenes. This fruit structure is characteristic of both Psychotria and Rudgea but excludes this species from Coussarea, which has smooth, thin-walled, usually solitary seeds in fleshy to leathery fruits. Rudgea is separated from Psy¬ chotria by its glandular-fimbriate stipules. The stip¬ ules of this species are shortly bidentate and not at all glandular, and it is here transferred to Psycho¬ tria. Psychotria nebulosa belongs to subgenus Het- eropsychotria Steyermark, which is characterized by green or gray-green drying color, persistent usu¬ ally bidentate stipules, and blue or black mature fruits. The mature fruit color of P. nebulosa is not known. Based on inflorescence, stipule, and fruit morphology it seems closely related to P. eurycarpa Standley, which can be separated by its smaller leaves that are usually shiny on the adaxial surfaces and pyrenes that are longitudinally angled rather than ridged, and to P. umbelliformis Dwyer & M. V. Hayden, which is densely pilosulous throughout. Psychotria nebulosa is also similar in general aspect to Coussarea austin-smithii Standley, which can be distinguished by its subglobose to ellipsoid fruits with smooth to slightly angled, usually solitary seeds. Psychotria nebulosa appears to be restricted to Chiriqui and Veraguas provinces in western Panama. Psychotria roseocrema (Dwyer) C. M. Taylor, comb. nov. Basionym: Coussarea roseocrema Dwyer, Ann. Missouri Bot. Gard. 67: 133. 1980. TYPE: Panama. Panama: El Llano- Carti highway, 6-10 km N of El Llano, 13 Apr. 1973, R. L. Dressier 4334 (holotype, MO 2126258). Additional material collected since the publication of this name shows that the fruits of this species are fleshy and blue-black at maturity with two hard, longitudinally ridged pyrenes. As discussed above, this fruit structure is characteristic of Psychotria but is not found in Coussarea, and accordingly this species is here transferred to Psychotria. Psycho¬ tria roseocrema belongs to subgenus Heteropsy- chotria. Psychotria roseocrema is similar in general as¬ pect to P. umbelliformis, which can be distinguished by its pilosulous pubescence throughout, and to P. eurycarpa and Coussarea austin-smithii, both of which lack the strong looping submarginal leaf veins found in P. roseocrema. Psychotria saltatrix C. M. Taylor, sp. nov. TYPE: Costa Rica. Cartago: Refugio Nacional Tapanti, por la orilla del Rio Grande de Orosi, 9°45'N, 83°47'W, 14 Aug. 1991, B. Hammel et al. 18328 (holotype, CR; isotypes, F, MO 4619859). Figure 2. A speciebus ceteris Psychotriae L. subg. Psychotriae foliis stipulis bracteis calycibusque majoribus etiam in- fructescentiis ut videtur axillaris differt. Soft shrubs to 2 m tall; stems puberulous to hir- tellous becoming glabrescent with age. Leaves ellip¬ tic, 18-32 cm long, 6.5-13.5 cm wide, at apex acute to acuminate with the tip ca. 1 cm long, at base acute to usually tapered, drying membrana¬ ceous, above glabrous, below glabrous on lamina but moderately to usually densely puberulous to pilo¬ sulous on veins; secondary veins 18-24 pairs, di¬ verging from costa at 80-90° or more, usually loop¬ ing to interconnect in an undulating to rather straight submarginal vein, with 1-2 secondary loops as well, plane above, prominulous below, without domatia; petioles 1.5-8 cm long; stipules interpetiolar, de¬ ciduous after 1-2 distal nodes, obovate, 25-35 mm long, 12-16 mm wide, acute, basal and central 206 Novon Figure 2. Psychotria saltatrix C. M. Taylor (from Hammel 18030 , MO). portion chartaceous to subcoriaceous and smooth, marginal portions membranaceous and with fine par¬ allel veining, both portions puberulous or pilosulous becoming glabrescent. Inflorescences terminal, py¬ ramidal; peduncles 3-60 mm; panicles 2-10 cm long, 4-12 cm wide, with branches spreading at ca. 80-90° or occasionally more, with 2 branches at the basalmost nodes or on larger inflorescences fre¬ quently 4 branches and then these generally similar in size but 2 spreading and 2 reflexed, puberulous to hirtellous, with bracts lanceolate, membrana¬ ceous, puberulous to glabrescent, ciliolate, acute, 2- 10 mm long, those subtending primary branches 8- 10 mm long and those subtending flowers 2-4 mm long; flowers sessile or very shortly stipitate in glom- erules of 3-5, each subtended by 1-3 bracts; hy- panthium turbinate, 1 — 1.2 mm long, glabrescent to puberulous; calyx limb green, glabrous to puberu¬ lous, 1.5-2 mm long, divided for ca. %-% its length, lobes 5, deltoid to ovate, ciliolate, acute; corolla salverform, white, externally glabrous or usually densely puberulous at top of tube, internally glabrous except densely pilose in 10 tufts at stamen attach¬ ment, tubes 2-3 mm long, 1-1.2 mm diam., lobes 5, ligulate, 2-3 mm long, somewhat thickened at apex in bud, becoming acute at anthesis; stamens 5, exserted, attached just below top of corolla tube, filaments 0.5-0.8 mm long, anthers oblong, ca. 1 mm long; style exserted, 3.5-4.5 mm long, stigma lobes 2, linear, ca. 0.5 mm long; disk annular, smooth, ca. 0.3-0.5 mm long. Infructescences over¬ topped by sympodial stem growth and appearing pseudoaxillary, as fruit matures usually displaced to 3rd or 4th node below apex, more expanded than inflorescences, to 12.5 cm long and 16 cm wide, bracts often deciduous ; fruit with stipe ca. 0.5 mm long, obovoid, 6-6.5 mm long, 4-4.5 mm wide, puberulous to glabrescent, becoming orange to red; pyrenes with 4—5 rather sharp longitudinal ridges. Habitat and phenology. Costa Rica to Colombia, in wet premontane and montane forest at 1250- 2100 m. Collected in flower in May and August, in fruit January, July, September, and October. Psychotria saltatrix is placed in subgenus Psy¬ chotria (Hamilton, 1989) based on its deciduous interpetiolar stipules, orange to red mature fruits, and distinctive gray-green to brown drying color. Within this subgenus it is distinguished by its ap¬ parently pseudoaxillary inflorescences, relatively large leaves and stipules, secondary veins that often diverge from the costa at more than a 90° angle and usually form a distinct submarginal vein, well- developed inflorescence bracts, relatively long calyx limbs, obovoid fruits, and very wet premontane or montane habitats. The inflorescences are terminal in position but the infructescences are typically over¬ topped by sympodial growth from one axil. This growth pattern is seen in other species of subgenus Psychotria, but is rarely as strongly or consistently developed as in this new species. Relatively few other species of subgenus Psychotria inhabit wet pre¬ montane or montane forest. In the two flowering collections seen, both the stamens and the style are Volume 5, Number 2 1995 Taylor Rubiaceae from Costa Rica and Panama 207 exserted in all of the flowers, and P. saltatrix may be homostylous. This condition (if actually present) would be highly unusual in the subgenus. The spe¬ cific epithet refers to the occurrence of this species in widely separated wet cool sites in the Andean Cordillera, western Panama, and Costa Rica. Three fruiting specimens from similar habitats in southern Colombia and Ecuador, de Benavides 9292 (MO), Sparre 17686 (S), and Hoover 1993 (MO), are similar to P. saltatrix but can be separated by their shorter calyx limbs and bracts subtending the flowers, both ca. 1 mm long, secondary leaf veins 14-15 pairs, and stipules 5.5-6 cm long. These specimens may represent a separate, apparently un¬ named taxon, or they may reflect clinal variation in P. saltatrix', no conclusion can be made without flowering material. Paratypes. COLOMBIA. Antioquia: municipio de Valdivia, vereda San Feraiin, sitio “Alto de Ventanas,” 2 km al NE de la via Medellin-Valdivia en la via que conduce al municipio de Briceno, 7°50'N, 75 C 32 W, Cal- lejas et al. 10735 (F, HUA, MO); municipio de Valdivia, vereda San Fermin, cerca 120 km NE de Medellin, en la via a Briceno, finca de Los Restrepo, 07°06'N, 07°32'W, Callejas & Gomez 10913 (HUA, MO); municipio de Urrao, Parque Nacional Natural “Las Orquideas,” cam- ino de Calles hacia Venados arriba. Sector Bellavista, Cogollo et al. 5028 (JAUM, MO). Narino: municipio Ricaurte, Reserva Natural La Planada, Restrepo 637 (MO). COSTA RICA. Cartago: Tapanti, /. Chacon et al. 1432 (CR, MO). San Jose: Parque Nacional Braulio Carrillo, Estacion Zurqui, 10°03'30"N, 84°01'W, Ham- mel et al. 18030 (CR, F, MO). PANAMA. Chiriqui: Cerro Colorado Mine, Antonio 4878 (MO). Darien: Cer- ro Pirre, valley between Pirre and next most southerly peak, Folsom 4441 (MO); Cerro Pirre, top of mountain and ridge just S of Pirre, Folsom 4521 (MO). Rudgea laevis C. M. Taylor, sp. nov. TYPE: Costa Rica. Puntarenas: Monteverde, Bajos de La Lindora, rio Guacimal, 10°18'N, 84°48’W, 19 Feb. 1988, W. Haber & E. Bello 8271 (ho- lotype, CR; isotype, MO 3646480). Figure IF, G. A Rudgea laurifolia (Kunth) Steyermark venulis pau- cioribus calycibus corollisque minoribus differt. Shrubs or small trees to 6 m tall; stems to 15 cm DBH, glabrous. Leaves elliptic, 5.5—12 cm long, 1 -8 5.5 cm wide, at apex acute, at base cuneate to obtuse, drying chartaceous, glabrous above and below; secondary veins 4-7 pairs, not looping to interconnect, above plane to somewhat sulcate, be¬ low prominulous, usually with foveolae in axils; pet¬ ioles glabrous, 4-12 mm long; stipules glabrous, united around stem into a persistent continuous trun¬ cate sheath 1.5—2 mm long bearing on each side 1 caducous deltoid gland ca. 1 mm long and wide. Inflorescences terminal, pyramidal to somewhat rounded, green, pilosulous to glabrescent; peduncles 1.5-3 cm long; panicles 2.5-3 cm long, 4-8 cm wide, with bracts 0.5-1 mm long, deltoid, obtuse; flowers sessile in dichasial cymes of 5-9, with hy- panthium turbinate, ca. 1 mm long, glabrous; calyx limb green, glabrous, 0.3-0.5 mm long, subtruncate to 5-denticulate; corolla salverform, white, glabrous externally and internally except barbate in throat; tube 3-4 mm long, 1-2 mm diam.; lobes 5, ligulate, 3-4 mm long, acute, unappendaged; stamens 5, exserted, attached just below top of corolla tube, filaments 0.8-1 mm long, anthers ca. 2 mm long, narrowly oblong; style exserted, 6-6.5 mm long, puberulous in distal portion, stigma lobes 2, linear, ca. 1 mm long; disk annular, smooth, ca. 0.5 mm high. Infructescences similar in size, proportion, and color to inflorescences; fruit globose, 6-10 mm diam., glabrous, white; pyrenes smooth. Habitat and phenology. Costa Rica, in wet pre- montane and lower montane forest of Cordillera de Tilaran at 1000-1400 m. Collected in flower in February and April, in fruit June, September, and November. This new species is similar in aspect to Rudgea laurifolia (Kunth) Steyermark, which can be dis¬ tinguished from R. laevis by its several to numerous acicular stipular glands, more numerous secondary leaf veins (9-11 pairs), longer calyx limb (2.5-2.7 mm long), longer corolla tube (ca. 7.5 mm long), shortly appendaged corolla lobes, and lowland wet forest habitat. Rudgea laurifolia is known only from the basins of the Rios Negro and Orinoco in South America. Paratypes. COSTA RICA. Alajuela: Reserva Biolo- gica Monteverde, Finca Ston’s, 10°19'N, 84°43'W, Ob- ando 140 (CR, MO). Puntarenas: Monteverde comu- nidad, 10°18'N, 84°48'W, Bello 4 (CR), Haber 452 (MO), 1070 (MO). Acknowledgments. I thank the curators of CR, DUKE, F, and HUA for making specimens available for study; R. Callejas for facilitating fieldwork in Antioquia, Colombia; R. E. Gereau for revision of the Latin diagnoses; and W. C. Burger, M. Grayum, B. Hammel, D. Lorence, and W. D. Stevens for helpful comments. Travel related to this work was supported in part by the National Science Foun¬ dation (BSR 87-00068). Literature Cited Burger, W. B. & C. M. Taylor. 1993. Flora Costari- censis: Rubiaceae. Fieldiana, Bot. n.s. 33: 1-333. Hamilton, C. 1989. A revision of Mesoamerican Psy- chotria subg. Psychotria (Rubiaceae). Ann. Missouri Bot. Card. 76: 67-111, 386-429, 886-916. Erratum Ten, Chi & Jun-liang Yang. 1994. Roegneria triden- tata, a new species of Triticeae (Poaceae) from Qing- hai, China. Novon 4: 310-313. On page 313, first column, four lines from bottom should read “Chromosome 19 has a large satellite, and chromosome 21 has a very short arm connected with a small satellite.” Volume 5, Number 2, pp. 119-208 of NOVON was published on 21 June 1995. Volume 5 Number 3 1995 _-JS0VON $£p 121995 pMtDEN UBRARt, Novelties and Notes in North American Aristida (Gramineae) Kelly W. Allred Range Science Herbarium, Department of Animal & Range Sciences, New Mexico State University, Las Cruces, New Mexico 88003, U.S.A. Jesus Valdes-Reyrm Departamento de Botanica, Universidad Autonoina Agraria “Antonio Narro,” Saltillo, Coahuila 25315, Mexico Abstract. New taxa of Aristida include three re¬ lated to A. schiedeana Trinius & Ruprecht (A. pe- tersonii sp. nov., A. eludens sp. nov., and A. spa- nospicula sp. nov.) and one variety of A. purpurea Nuttall (var. perplexa var. nov.). Two species are re¬ duced in rank (A. schiedeana var. orcuttiana [Vasey] comb. nov. and A. purpurea forma brownii [War- nock] comb, nov.), and one species is reinstated (A. curvifolia Fournier). A key is given for the A. schie¬ deana complex. Resumen. Nuevos taxa de Aristida que incluyen tres especies con las aristas laterales reducidas, re- lacionadas con A. schiedeana Trinius & Ruprecht (A. petersonii sp. nov., A. eludens sp. nov., y A. spa- nospicula sp. nov.) y una variedad con las aristas laterales desarrolladas de A. purpurea Nuttall (var. perplexa var. nov.) son descritos. Dos especies son reducidas en rango (A. schiedeana var. orcuttiana [Vasey] comb. nov. y A. purpurea forma brownii [Wamock] comb, nov.) y una especie es reinstalada (A. curvifolia Fournier). Una clave para el complejo A. schiedeana es presentada. The genus Aristida (Arundinoideae: Aristideae) comprises approximately 300 species found throughout the world, but its members are partic¬ ularly frequent in warm, semiarid environments. Many species have three awns terminating the sin¬ gle floret, but it is not uncommon to find single- awned relatives of three-awned taxa. Species are notorious for their taxonomic difficulty, due partly, we believe, to undescribed variation that obscures the taxonomic boundaries. Major revisionary efforts include those of Henrard (1927, 1928, 1929) and Hitchcock (1924, 1935). Aristida petersonii Allred & Valdes-Reyna, sp. nov. TYPE: Mexico. Oaxaca: 7.7 km NW of Tlaxiaco on road to San Juan Mixtepec, steep slopes with Pinus, Quercus, and Arbutus, elev. 2020 m, 2 Sep. 1990, P. M. Peterson & A. Campos-Villanueva 9731 (holotype, NMCR; is¬ otypes, ANSM, US). Figure 1. Species nova propria vaginis lanosis, glumis glabris et valde inversis (primis secundis longioribus), paniculis an- gustis, rostris tortis lemmatum, et aristis singularibus a congeneribus Americanae borealis diversa. Plants perennial, tufted; culms 50-100 cm tall, erect, unbranched except for basal tillering; inter¬ nodes glabrous, striate, terete. Sheaths striate, shorter than the intemodes, rounded on the back, loosely lanose, the hairs curling and tangled, the upper sheaths sparsely lanose to glabrous; collars often with a tuft of longer cobwebby hairs at the comers; throat glabrous except for the ligule. Lig- ules a minute fringe of hairs less than 0.5 mm long. Blades glabrous, flat when fresh and mature (new leaves convolute), folded upon drying, 15-30 cm long, 2-2.5 mm wide when flat, the lateral nerves generally coalescent and the margin thickened, the Nnvniv 5- 900-999. 1QQ5 210 Novon F**"® 1 - Aristida petersonii Allred & Valdfe-Reyna (Peterson & Campos-Villanueva 9731). —A. Vegetative habit. —B. Sheath pubescence. —C. Inflorescence. —D. Spikelet: glumes left, floret right. Volume 5, Number 3 1995 Allred & Valdes-Reyna North American Aristida 211 Figure 2. Geographic distributions of Aristida petersonii Allred & Valdes-Reyna (stars), A. eludens Allred & Valdes- Reyna (closed circles), and A. spanospicula Allred & Valdes-Reyna (squares). marginal veins ca. 0.5 mm wide. Panicles 20—30 cm long, narrow, contracted, erect, the nodes gla¬ brous; primary branches few, single or paired at the nodes, erect to appressed, 4- to 8-flowered, the low¬ er ones 6—12 cm long; secondary branches weakly developed, 2- to 3-flowered, appressed; pulvini ab¬ sent in the axils of the primary and secondary branches and pedicels, the spikelets thus ap¬ pressed along the branches and main axis. Glumes glabrous, inverse, membranous and sub-hyaline, 1- nerved, light tan to purplish, the apices acute but not awned, the second glume Vi to % the length of the first; first glume (9—)10—13 mm long; second glume 6—7 mm long. Lemmas glabrous, mottled, 7— 8 mm long from the base of the callus to the twisted portion of the beak; beak prominently twisted, (4—) 5—6 mm long; central awn not twisted, geniculate- bent, 7—9 mm long; lateral awns completely re¬ duced, represented by tiny points ca. 0.1 mm long at the end of the beak; callus ca. 0.5 mm long, with straight hairs to 1 mm long. Palea completely en¬ closed by the lemma, 2-nerved, hyaline, obtuse to rounded, 1—1.5 mm long. Lodicules 2, flabellate, 1— 1.5 mm long. Caryopsis fusiform, chestnut-brown, 4.5-5.5 mm long. Flowering known only from September, but pre¬ sumed August-October. Distribution. Pine/oak/ju¬ niper woodlands; Mexico, known only from south- central Oaxaca (Fig. 2). The specific epithet, petersonii, honors the col¬ lector of this species, Paul M. Peterson of the Smithsonian Institution. Aristida petersonii is one of three perennial spe¬ cies in North America with lanate or floccose sheaths, the other two being A. lanosa Muhlenberg ex Elliott and A. scribneriana A. S. Hitchcock. Ar¬ istida lanosa, of the southeastern United States, dif¬ fers in having sheaths mostly longer than the in- temodes, blades 2-6 mm wide, panicles mostly 30- 70 cm long with lanose tufts in the branch axils, longer glumes (though they share the inverse po¬ sition), and three well-developed awns. Aristida scribneriana, of the central highlands of Mexico, differs in having lanose intemodes (including the peduncles) and blades (both surfaces), sparsely vil- 212 Novon lous glumes that are nearly equal in length, and normally three well-developed awns. Other single- awned Aristida with narrow panicles that have gla¬ brous (or very sparsely pilose) sheaths and equal glumes are A. eludens, A. spanospicula, and epul- vinate forms of A. pansa. Aristida petersonii is the only North American Aristida with lanose sheaths, narrow panicles, glabrous glumes in the inverse po¬ sition, and single-awned spikelets. Paratypes. MEXICO. Oaxaca: 24.5 km W of Tlax- iaco and 7.4 km NE of San Juan Mixtepec, gravelly slopes with Pinus, Quercus, Arbutus and scattered Juniperus, elev. 2220 m, 3 Sep. 1990, P. M. Peterson & A. Campos-Vil- lanueva 9746 (ANSM, NMCR, US). Aristida schiedeana Trinius & Ruprecht Complex The Aristida schiedeana complex includes A. schiedeana, A. orcuttiana Vasey, A. laxa Cavanilles, and the two new species A. eludens and A. spano¬ spicula. This complex is characterized by having flat blades often curling like wood shavings, gen¬ erally equal glumes, a twisted lemma beak, and single awns (except A. laxa). Its members are cus¬ tomarily found as sparse understory in pine/oak for¬ ests of the Mexican and Central American central cordilleran mountain ranges. Aristida eludens Allred & Valdes-Reyna, sp. nov. TYPE: Mexico. Coahuila: San Lorenzo Can¬ yon, ca. 8 km S of Saltillo, mountain scrub grassland vegetation with Berberis, Bouteloua, Hilaria, Dasylirion, very rocky sandy loam, flat bajada on N-facing side of canyon, 6000 ft. (1830 m), 3 Sep. 1991, J. Valdes-Reyna 2254 (holotype, ANSM; isotype, NMSU). Figure 3. Species nova laminis ubi vivis planis vel plicatis laxe, paniculis angustis absque pulvinis axillaribus praeter in- terdum ramos infimos, glumis subaequalibus, et aristis singularibus a congeneribus Americanae borealis diversa. Plants perennial, tufted; culms (35-)40-65(-80) cm tall, erect, branching only at the base; inter- nodes terete, glabrous, leaves basal and cauline, light green. Sheaths longer than the intemodes, gla¬ brous or scaberulous except for the summit, round¬ ed on the back; collars with a tuft of cobwebby hairs at the comers (becoming glabrous upwards), some¬ times with a line of minute hairs across the back; throat glabrous or puberulent. Ligules a fringe of hairs 0.2-0.3 mm long. Blades flat or loosely folded when fresh, rolled upon drying, glabrous or sca¬ berulous abaxially, puberulent or glabrate adaxially especially toward the base, 12-35 cm long, 1-1.5 mm wide when flat. Panicles 10-22 cm long and 1— 2 cm wide, with 16 or more spikelets, narrow, contracted, the nodes glabrous; primary branches erect, appressed to the main axis except occasion¬ ally the lowermost branch divaricate, naked at the base, the lower ones 5—9 cm long; pulvini absent, the spikelets appressed, only occasionally present in the axil of the lowermost branch and causing it to spread from the axis. Glumes glabrous, brownish, subequal or the first slightly shorter or longer, 1- nerved, 8-13 mm long, the apices acute and often with a mucro. Lemmas glabrous, mottled, 10-13 mm long from the base of the callus to the diver¬ gence of the awns; beak twisted, (3-)4-5 mm long; central awn not twisted, geniculate-bent, 5-10 mm long; lateral awns highly reduced, erect, 0.1—3 mm long; callus 0.5—1 mm long with stiff hairs ca. 1 mm long. Paleas completely enclosed by the lem¬ ma, 2-nerved, hyaline, rounded, ca. 1 mm long, shorter than the lodicules. Stamens 3, the anthers 2- 2.5 mm long. Lodicules 2, flabellate, 1—1.5 mm long. Caryopses fusiform, chestnut-brown, ca. 6 mm long. Flowering April—May, August—November. Distri¬ bution. Desert scrub/grassland habitats in calcare¬ ous soil, but extending up into the pine/oak zones; Mexico, in the states of Chihuahua, Coahuila, Durango, Guanajuato, Nuevo Leon, Oaxaca, Que- retaro, and San Luis Potosf (Fig. 2). The epithet eludens refers to the tendency of plants of this species to grow only within the branches and foliage of protective shrubs, evading, as it were, grazing animals. Aristida eludens is the desert scrub/grassland representative of the A. schiedeana complex. Aris¬ tida eludens is distinguished most easily from the other species of the complex by possessing con¬ tracted panicles where the primary branches lack pulvini; it also differs by having cauline leaves with non-coiling blades and by the habitat. Aristida elu¬ dens occurs mostly in the desert scrub and semi- desert grasslands of northern Mexico, and only in¬ frequently extends into the lower reaches of adjacent pine/oak communities. It has not been found to grow sympatrically with any of the other species of the complex. Consistent with the other members of the complex, its blades are nearly al¬ ways flat or only loosely folded in the natural state (this inay be obscure in dried specimens), though they do not coil like wood shavings. The flat blades contrast with the tightly involute blades of sympat- ric A. pansa Wooton & Standley and A. purpurea Nuttall s.l. Although the panicles of A. eludens usu¬ ally lack pulvini and are thus narrow and contract¬ ed, it is not uncommon to find an isolated plant or Volume 5, Number 3 1995 Allred & Valdes-Reyna North American Aristida 213 Figure 3. Aristida eludens Allred & Valdfe-Reyna (Valdes-Reyna 2254). -A. Habit. -B. Spikelet: glumes left, floret right. 214 Novon two in each population with only the lowermost branch spreading from a single axillary pulvinus. Only one individual from northern Coahuila (Ibarra s.n.) has been found to be completely pulvinate (pulvini in all axils). Paratypes. MEXICO. Chihuahua: Sierra Carrasco, ca. 31 mi. NW of Julimes, 6000 ft., 15 Sep, 1973, J. Henrickson 12965.5 (TEX/LL); Santa Eulalia Mts., 10 Sep. 1885, C. G. Pringle 386 (TEX/LL); La Campana, 1650 m, 7 Sep. 1973, J. ValcUs-Reyna 148 (TAES). Coahuila: Si¬ erra de la Paila (lado norte), Canada becerros, 1700 m, 13 Oct. 1989, J. A. Villarreal 5469 (ANSM); 12 km al W de Saltillo, carr. 40, 2000 m, 9 July 1983, ,/. Valdds-Reyna & /.. Rodriguez 1522 (ANSM); San Lorenzo Canyon, ca. 8 km S of Saltillo, mountain scmb grassland vegetation with Berber! s, Bouteloua , Hilaria, Dasylirion, very rocky sandy loam, flat bajada on N-facing side of canyon, anthers 1.5 mm, 6000 ft. (1829 m), 3 Sep. 1991, K. W. Allred & J. Valdds-Reyna 5488, 5491, 5492, 5495, 5497 (ANSM, NMCR); Est. Cameros, Cameros Pass, steep hills W of town, very rocky calcareous soil along cobble road to mi¬ crowave tower, desert grassland/scrub vegetation with Yuc¬ ca carnerosana. Agave, Nassella, Aristida curvifolia, 6750 ft. (2057 m), 5 Sep. 1991, K. W. Allred & J. Valdds-Reyna 5527, 5528 (ANSM, NMCR); Sierra Garcia, San Lazaro Pass, desert mountain scrub vegetation with ocotillo, le- chuguilla. Acacia, and Opuntia, limestone N-facing steep slopes, 1200 ft. (366 m), 6 Sep. 1991, K. W. Allred & J. Valdds-Reyna 5540 (ANSM, NMCR); San Lorenzo Canyon, 25 Aug. 1981, S. L. Hatch 4501 (ANSM, TAES); Cameros, camino a torre de microondas, 3 km al poniente de la estacidn, 30 km al S de Saltillo, matorral de Yucca car¬ nerosana, Pinus cembroides, Dasylirion cedrosanum, Cea- nothus greggii, Nassella tenuissima, 17 Oct. 1986, J. Es- pinosa-A. 97, 186 (ANSM, NMCR); San Lorenzo Canyon, 22 Aug. 1980, M. Madrigal-A. s.n. (TAES); Sierra de Par¬ ras en Los Chupaderos, 14 Apr. 1981, A. Rodriguez s.n. (CIID1R-DURANGO); Estacidn Cameros, camino torre de microondas, 2100 m, 24 May 1982, J. Valdds-Reyna 1456 (TAES, US); 18 mi. S of Saltillo on hwy. 54, just W of Est. Cameros, 2250 m, 18 Oct. 1989, P. Peterson 8416 (ANSM, US); Canon San Lorenzo, 1900 m, 22 Aug. 1980, [no collector] (TAES); Mpio. Arteaga, 10 mi. NE of hwy! 57 at Los Alpes, with pinyon. Yucca, Agave, 24 Aug. 1983 S. L Hatch 5022 (ANSM, NMCR, TAES); Mpio. Saltillo’, Cerro del Pueblo, al W de la ciudad de Saltillo, 11 Sep. 1990, J. Valdds-Reyna 2055 (ANSM, NMCR); Mpio. Ocampo, Rancho La Rueda, D. Ibarra s.n. (ANSM). Du¬ rango: 45 km WNW of Huejuquilla El Alto, 1830-2150 m, 23 Oct. 1983, D. E. Breedlove 59212 (CAS, MO). Gua¬ najuato: 30 km SE of San Felipe on hwy. 37 to Ledn, rocky slopes in open grassland, 2290 m, 30 Aug. 1990 P. M. Peterson 9693 (ANSM, NMCR, US); oak slopes ca! 12 mi. on rd. between Guanajuato and Santa Rosa 30 Sep. 1946, H. E. Moore, Jr. 1351 (US). Oaxaca: 19.5 km NW of Tlaxiaco and 12.3 km NE of San Juan Mixtepec, near jet. of road to Yosonama, sandy slopes with pine, oak and manzanita, 2410 m, 3 Sep. 1990, P. M. Peterson 9736 (ANSM, NMCR, US). Nuevo Leon: near Puerto Mexico along hwy. 57, 22 Aug. 1983, S. L. Hatch 5011 (ARIZ, TAES). Queretaro: 5 km al NE de Bernal, sobre la car- retera a Tolim4n, ladera caliza con vegetacidn de matorral xerftfilo, 1950 m, 10 Nov. 1988, Rzedouiski s.n. (ANSM, NMCR). San Luis Potosi: Mpio. Villa de Arriaga, Cerro del Gallo, 2500 m, 3 Oct. 1962, E. Hemdandez-X. 1413 (ANSM); mountain slopes, 13 mi. S of San Luis Potost, 7100 ft., 2 Sep. 1939, F. Shreve 9307 (US); near village of San Francisco in Sierra de San Miguelito, ca. 25 km SW of San Luis Potosf, pine/oak, 22—2400 m, 5 Sep. 1954, E. R. Sohrts 1062 (US); Cnyn. del Muerto, ca. 3 km W of Rodrigo, in Sierra de San Miguelito, 1800-2200 m, 18 Sep. 1954, E. R. Sohns 1317 (US); ca. 5 mi. SW of city in hills, 18 July 1950,,/. R. Reeder 1372 (ARIZ). Several specimens from central Mexico approach Aristida eludens in having single awns and narrow panicles, but differ in being very tall (to 1.5 m) robust plants with knotty bases and curling blades, and in occurring in oak and pine/oak forests. They are here excluded from A. eludens and are tenta¬ tively considered aberrant forms of plants normally with spreading panicle branches. Duplicates of some of these were referred by McVaugh (1983) to A. schiedeana or A. laxa. Excluded from A. eludens: MEXICO. Jalisco: Autldn, 1290 m, 5/XI/1975, C. Castro 84 (MEXU); near Villa Guerrero, Cerro del Aquila, 25 Sep. 1980, A. Beetle 5811 (MEXU); Paso de la Troje, near km 36, SW of Ojuelos on rd. to Aguascalientes, 2100—2300 m, 9-12 Aug. 1958, R. McVaugh 16818 (US); Rio Blanco, 6 Oct. 1886, E. Palmer 476 (MO); Rio Blanco, June—Oct. 1886, /'.. Palmer 517.5 (US); Rio Blanco, June-Oct. 1886, E. Palmer 769 (US); Mpio. Mezquitic, 5 km E of Rancho El Mortero, pine/oak, 2200 m, 5 Nov. 1963, Rzedouiski 17687 (US). Morelos: Alarc an. 31 Aug. 1910, C. Orcutt 3863 (MO). Puebla: 35 km de Tetela, 1550 m, 1/VI11/1981, S. Contreras 534 (MEXU). Aristida spanospicula Allred, Valdes-Reyna & Sdnchez-Ken, sp. nov. TYPE: Mexico. Chihua¬ hua: Sierra Madre Occidental, at Cuesta Prie- ta, along road from San Juanito to Creel, 3.1 mi. S of San Juanito, pine forest with scrub oak understory of Lupinus, Muhlenbergia, and Trisetum, thin forest loam calcareous soil, nu¬ merous clumps among the scrub oak, 7500 ft. (2280 m), 22 Sep. 1992, K. W. Allred & J- Valdds-Reyna 5787 (holotype, NMCR; iso¬ types, ANSM, NMC, TAES). Figure 4. Species nova Aristidae schiedeanae affinis laminis plan¬ ts saepe lorsivis scobiformibus, paniculis angustis absque pulvinis axillaribus, spiculis paucis dispersis late, glumis magnis spadiceis, rostris exsertis tortis lemmatum, et ar- istis singularibus a congeneribus Americanae borealis div- ersa. Plants perennial, tufted; culms (20-)25-55(-60) cm tall, ascending, slender, not branching above the base; internodes terete, glabrous to minutely scaberulous. Leaves basal, light green to yellowish green. Sheaths longer than the intemodes, glabrous to sparsely pilose, rounded on the back; collars gla- brous to sparsely pilose at the comers, lacking a line of hairs across the back; throats glabrous ex- Volume 5, Number 3 1995 Allred & Valdes-Reyna North American Aristida 215 Figure 4. Aristida spanospicula Allred & Valdfc-Reyna (AUred 5787). -A. Habit. -B. Spikelet: glumes left, floret right. 216 Novon cept for the ligule. Ligules a fringe of hairs less than 0.5 mm long. Blades flat to loosely folded when fresh, rolled upon drying, usually coiling like wood shavings in age, glabrous abaxially, glabrous to pu- berulent adaxially, 14-25 cm long, 1.5-2 mm wide when flat, the margins thickened by the coales¬ cence of the outer veins. Panicles (8-)12-22 cm long, 1—2 cm wide, few flowered with 3—8(—12) spikelets, often drooping, narrow but loose, the nodes glabrous; primary branches weakly devel¬ oped, appressed-erect, bearing 1—2(-4) spikelets; pulvini absent. Spikelets few and widely spaced, scarcely overlapping, the pedicels often drooping and somewhat capillary. Glumes glabrous to sca- berulous, mostly 1-nerved, occasionally a short lat¬ eral nerve developed, brownish, subequal or the first 1-2 mm shorter or longer, 9—15 mm long. Lem¬ mas glabrous, (16—)18—22(—25) mm long from the base of the callus to the divergence of the awns; beak twisted, exserted beyond the glumes, (6—)8— 10(—15) mm long; central awn not twisted, genic¬ ulate-bent at ca. 90°, 10-16(—20) mm long; lateral awns highly reduced, erect, less than 0.5 mm long; callus 0.5-1 mm long with stiff hairs less than 1 mm long. Paleas completely enclosed by the lem¬ mas, 2-nerved, membranous, ca. 1 mm long, shorter than the lodicules. Stamens 3, the anthers light brown, ca. 2 mm long. Lodicules 2, flabellate, 1- 1.5 mm long. Caryopses fusiform, brown, 7—9 mm long. Flowering August-October. Distribution. As yet, found only in pine/oak forests of the Sierra Madre Occidentale, often on nearly bare ground, usually rocky or crumbly calcareous soil; Mexico, in the states of Chihuahua, Durango, and Sonora (Fig. 2). We are pleased to cooperate with Jorge SAnchez- Ken, of the Herbario Nacional de Mexico (MEXU), in describing this new species. The epithet spano- spicula alludes to the panicles with few spikelets. Aristida spanospicula differs from others in the complex by the epulvinate panicle, fewer spikelets, and longer lemmas. It is apparently restricted to the pine/oak forests of the Sierra Madre Occidental in northern Mexico. Aristida spanospicula occasion¬ ally grows intermingled with A. schiedeana, and some of the sympatric populations contain inter¬ mediate plants suggesting hybridization or incom¬ plete morphological delineation of the species. The latter would not be uncommon in Aristida. Paratypes. MEXICO. Chihuahua: ca. 6 mi. S of Creel, rocky outcrops near small stream, in ponderosa pine community with 5-needle pine and juniper, sandy loam soil, 22 Aug. 1985, K. W. Allred 3074 (NMCR); Cas- cada Basaseachic Overlook on S side of canyon, very steep N-facing slopes in oak/pine forest, with Quercus crassifol- ia, Arbutus, 6600 ft. (2012 m), 11 Sep. 1987, K. W. Allred 4607 (NMCR); 4 mi. W of Cuesta Blanca in the Sierra Brena, approximately 32 road mi. SW of Col. Juarez, pine- oak woodland with a very sparse understory, rocky gravelly soil, 6500 ft. (1981 m), 21 Sep. 1992, K. W. Allred & J. Valdts-Reyna 5745, 5746 (NMCR); Sierra Madre Occi¬ dental, 11 mi. W of Babicora on hwy. 180 to Madera, pine- oak forest with sparse grass understory of Bouteloua, Muh- lenbergia, Piptochaetium, rocky loam soil, 6800 ft. (2072 m), 22 Sep. 1992, K. W. Allred & J. Valdes-Reyna 5776 (ANSM, NMCR); Sierra Madre Occidental, at Cuesta Prie- ta, along road from San Juanito to Creel, 3.1 mi. S of San Juanito, pine forest with scrub oak understory of Lupinus, Muhlenbergia, and Trisetum, thin forest loam calcareous soil, approaching A. schiedeana , 7500 ft. (2280 m), 22 Sep. 1992, K. W. Allred & J. Valdes-Reyna 5792 (ANSM, NMCR); Cusarare, S of Creel, 14 Sep. 1973, Bye, R. 5047 (TEX/LL); S&nchez, rocky pine woods, 8000 ft., 12 Oct. 1910, A. S. Hitchcock 7671 (US); Chuhuichupa, Aug.-Sep. 1936, Lesueur 6112 (CAS, MO, TEX/LL); Colonia Cum- bres de Majalca, approx. 32 km W of hwy. 45 N of Chi¬ huahua, table rock and rocky slopes with oak, pine, ju¬ niper, and Arctostaphylos, 23 Sep. 1988, P. M. Peterson 5810 (ANSM, NMCR, US); 43.5 km W of Balleza and 51.6 km E of Guachochi, sandy, clay flats with pine, oak, and Arctostaphylos pungens, 2320 m, 18 Sep. 1991, P. M. Peterson et al. 10756 (ANSM, NMCR, US); Parque Na¬ cional Barranca del Cobre, 24.8 km NE of La Bufa on road to Samachic, gentle slopes in forest of pine, oak, and Arbutus, 2440 m, 20 Sep. 1991, P. M. Peterson et al. 10799 (ANSM, NMCR, US); pine-oak region, Sierra Ma¬ dre occidental, W of Casas Grandes, 3 mi. W of Cuesta Blanca, 2073 m, 4 Sep. 1958, J. R. & C. G. Reeder 3214 (ARIZ, US). Durango: Sierra Madre Occidental, moun¬ tains E of Durango City, approximately 30 mi. E of the city at the movie set for “Fat Man and Little Boy,” deep barrancas in pine/oak/manzanita communities, soil chalky, calcareous, very crumbly, 7500 ft. (2286 m), 24 Sep. 1992, K. W. Allred & J. Valdes-Reyna 5831, 5837 (ANSM, NMCR); Sierra Madre Occidental, along hwy. 40 from Du¬ rango City to El Salto, 30 mi. W of Durango, pine/oak/ arctostaphylos woodland with thick grass understory of Muhlenbergia, Trachypogon, slight slope, loam soil, 8400 ft. (2560 m), 24 Sep. 1992, K. W. Allred & J. Valdes- Reyna 5845 (ANSM, NMCR); Sierra Madre along hwy. 40 from Durango City to El Salto, 32 mi. W of Durango, adjacent to Parque El Tecuan, pine savannah parkland with 5-nee¬ dle pine, gently W-facing slope, grass understory of Aris¬ tida, Panicum, forest loam soil, 8000 ft. (2438 m), 24 Sep. 1992, K. W. Allred & J. Valdts-Reyna 5851 (ANSM, NMCR). Sonora: E of Cananea, pine woods. Sierra de los Ajos, 26 Sep. 1981, A. A. Beetle 7859 (MO). The relationship of A. schiedeana and A. orcut- tiana has long been problematical. D4vila & S£n- chez-Ken (1994), Gould & Moran (1981), and Pohl & Davidse (1994) treated the two as conspecific without distinction; Henrard (1927, 1928, 1929) treated the two as distinct species; Hitchcock treat¬ ed them both ways (conspecific in 1924, distinct in 1935); and Beetle (1983) presumably treated the two as conspecific, though he does not mention A. orcuttiana in the text at all, even though the type is from Baja California Norte. Table 1 presents the salient differences between the two taxa. Although Volume 5, Number 3 1995 Allred & Valdes-Reyna North American Aristida 217 the morphological distinctions are incomplete and difficult to interpret, they are separated geograph¬ ically (Fig. 5). Aristida orcuttiana is here relegated to varietal status under A. schiedeana: Aristida schiedeana Trinius & Ruprecht var. or¬ cuttiana (Vasey) Allred & Valdes-Reyna, comb. nov. Basionym: Aristida orcuttiana Va¬ sey, Bull. Torrey Bot. Club 13: 27. 1886. SYN- TYPES: Mexico. Baja California Norte: Han¬ sen’s Ranch, 6000 ft., 30 July 1883, C. R. Orcutt 507 (lectotype, selected here, US). U.S.A. Arizona: M. E. Jones (not seen). Key to the Members of the Aristida schiedeana Com¬ plex: la. Panicles open, all the primary branches with ax¬ illary pulvini and spreading sharply outward from the main axis upon exsertion of the panicle from the sheath and throughout maturity. 2a. Lemmas with three well-developed awns, the lateral awns at least V 4 the length of the cen¬ tral awn. A. laxa Cavanilles (We use the name A. laxa Cavanilles in its traditional sense, although S4nchez-Ken (pers. comm.) informs us that in the type of A. laxa , the lateral awns are nearly absent; there may be difficulties with the application of this name.) 2b. Lemmas with a single well-developed awn, the lateral awns minute or at most V 4 the length of the central awn. . A. schiedeana Trinius & Ruprecht 3a. First glume usually equal to or shorter than the second; glumes usually sca¬ brous; blade above ligule usually with scattered pilose hairs; collar usually with a line of short hairs . .var. schiedeana 3b. First glume usually longer than the sec¬ ond; glumes usually glabrous; blade above ligule usually glabrous; collar usually lacking a line of hairs .... .var. orcuttiana (Vasey) Allred & Vald6s-Reyna lb. Panicles closed, contracted, none of the primary branches (or occasionally only the lowermost) with axillary pulvini and thus not spreading from the axis, though the branch tips or spikelets may droop outward. 4a. Lemmas (16-)18-22(-25) mm long (from the base of the callus to the divergence of the awn); panicles few flowered, with 3—12 loosely disposed spikelets. . A. spanospicula Allred & Valdes-Reyna 4b. Lemmas less than 10-13 mm long (from the base of the callus to the divergence of the awn); panicles densely flowered, with 16 or more congested spikelets. . A. eludens Allred & Valdes-Reyna Aristida purpurea Nuttall Complex The Aristida purpurea complex has tightly invo¬ lute to loosely folded blades, panicles without ax- Table 1. Comparison of Aristida schiedeana Trinius & Ruprecht and A. orcuttiana Vasey. A. schiedeana A. orcuttiana Pilose hairs above Usually present Usually absent ligule Collar hairs Usually present in a line Usually absent Glume pubescence Usually scabrous Usually glabrous Glume 1 length 6—15 usually 6-14 usually (mm) equal to longer than glume 2 glume 2 Glume 2 length 8-14 5—10(—13) (mm) Central awn length (5—)7—15 5-14 (mm) Lateral awn length 0.3-1.5(-4) 0-3(-7) (mm) Distribution (Fig. North-central Southwestern 5) Mexico to United States Nicaragua to northern Mexico illary pulvini (except in two varieties), and strongly unequal glumes, the first much shorter than the second. Its members are common in arid and semi- arid plains and deserts, and include A. purpurea s.l. (Allred, 1984), A. brownii Wamock, and A. cur- vifolia Fournier. Aristida purpurea Nuttall var. perplexa Allred & Valdes-Reyna, var. nov. TYPE: U.S.A. New Mexico: Dona Ana County, USDA Jornada Ex¬ perimental Range, ca. 16 mi. NE of Las Cru¬ ces, pasture 12, ca. 4800 ft., sandy soil, very common, 15 July 1986, K. W. Allred 4035 (ho- lotype, NMCR). Figure 6. Varietas nova Aristidae pansae et A. purpureae similis; ab A. pansa arachnoideis destitutis angulis vaginarum, glumis inaequalibus (primis secundis brevioribus), et ar- istis longis ascendentibus differt; ab A. purpurea paniculis apertis habentibus pulvinos axillares differt. Plants perennial, tufted; culms (30-)40—65 cm tall, erect, simple or sparsely branched at the base; intemodes terete, glabrous to puberulent. Leaves basal and cauline, yellowish to pale green. Sheaths rounded on the back, glabrous to puberulent, with longer hairs at the summit; collars usually with a line of short stiff hairs across the back, the comers glabrous or with a tuft of erect, straight to somewhat crinkly hairs (but not cobwebby) ca. 1 mm long; throats pubemlent in addition to the ligule. Ligules a fringe of hairs ca. 0.5 mm long. Blades rolled when fresh, sometimes only loosely folded, usually glabrous abaxially and pubemlent adaxially, mostly 218 Novon 8-20 cm long and less than 1 mm wide, arcuate to curling upon drying. Panicles (8-)10-26 cm long and (4-)6-12 cm wide, open, pyramidal; primary branches stiffly ascending to widely spreading from axillary pulvini, sometimes somewhat flexuous or capillary, naked at the base, the lower ones 3—6 (-8) cm long; pulvini present in the axils of the primary branches but usually absent from the ped¬ icels, the spikelets thus appressed along the branches. Glumes glabrous, membranous to scari- ous, light tan, strongly unequal and the first Vi to % the length of the second, 1-nerved, the first (4.5-) 5-7(-7.5) mm long, the second 8—11(—12) mm long. Lemmas glabrous to scabrous, often mottled, (8-) 10-12(-13) mm long from the base of the callus to the divergence of the awns; beak usually not twist¬ ed, reaching about to the tip of the second glume or slightly beyond, 2.5-3.5(-4) mm long and 0.1- 0.2 mm wide; awns subequal, mostly spreading 40- 50°, rarely approaching 90° from the vertical, the central awn (1.5-)2-3 cm long, the lateral awns 2- 4 mm shorter; callus 0.5-1 mm long with short stiff hairs ca. 0.5 mm long. Paleas completely enclosed by the lemmas, membranous, acute, 2-nerved, 1— 1.3 mm long. Stamens 3, the anthers 0.7—1.5 mm long. Lodicules 2, flabellate, 1-1.4 mm long. Car- yopses fusiform, light brownish. Flowering July-October. Distribution. Sandy to rocky plains and mesas in desert grassland/scrub communities, often calcareous soil; Mexico, in the states of Chihuahua, Coahuila, and San Luis Potosf; United States, in the states of Arizona, New Mexico, and Texas (Fig. 7). The epithet perplexa refers to the prior confusion of this variety with Aristida pansa. Because of its spreading panicle branches, spec¬ imens of variety perplexa have long been confused with Aristida pansa Wooton & Standley. The new variety differs from this latter species by its lack of cobwebby hairs at the collar (a feature consistent throughout the A. pansa complex), by the acuminate unequal glumes (the first %-% the length of the second), and by the longer awns (mostly 2—3 cm vs. 1.5 cm or less), which generally ascend at an angle of 40-50° rather than being nearly horizontal. Volume 5, Number 3 1995 Allred & Valdes-Reyna North American Aristida 219 Figure 6. Aristida purpurea Nut,all var. perplexa Allred & Valdfe-Reyna (Peterson 10632). -A. Habit. -B. Spikele, glumes left, floret right. 220 Novon The spikelets of variety perplexa are essentially identical to spikelets of A. purpurea var. purpurea. Variety perplexa may grow in homogeneous isolated populations (the type locality) or be sympatric with A. pansa and other varieties of A. purpurea, as is the case at the type locality of A. pansa in Dona Ana County, New Mexico. Variety perplexa is the second variety in the A. purpurea complex with spreading, pulvinate panicle branches. The other, variety parishii (A. S. Hitchcock) Allred, which is restricted to Sonoran Desert habitats, is similar to variety wrightii (Nash) Allred and often lacks the axillary pulvini so that the panicle is narrow or spi- cate. The panicles of variety perplexa, however, are consistently pulvinate. Paratypes. MEXICO. Chihuahua: just S of New Mex¬ ico border near Columbus, rocky hills, 15 Oct. 1983, K. W. Allred 2629 (NMCR); Sierra Las Pampas, N end of the sierra along dirt road from Las Pampas to jet. with road to Camargo, 6.2 mi. SE of jet., Chihuahuan Desert Scrub, rocky calcareous slope, 4500 ft. (1372 m), 23 Sep. 1992, K. W. Allred & J. Valdes-Reyna 5806 , 5807 (NMCR); near Chihuahua, 13 Oct. 1885, C. G. Pringle s.n. (POM/RSA). Coahuila: 16.9 km NE of San Miguel on road towards Boquillas, 1180 m, 13 Sep. 1991, P. M. Peterson 10608 (ANSM, NMCR, US); Sierra El Pino, 9.2 km SW of Ran¬ cho El Cimarron along the eastern slope, calcareous rocky slopes, 1500 m, 14 Sep. 1991, P. M. Peterson 10632 (ANSM, NMCR, US); near La Rosa, 5000 ft., 18 Aug. 1940, F. Shreve & E. R. Tinhham 4593 (ARIZ). San Luis Potosi: along hwy. 49 from Zacatecas to San Luis Potost, 32.4 mi. E of Salina, 21 mi. W of San Luis Potosi, desert thorn scrub community, rocky soil, 6200 ft. (1890 m), 26 Sep. 1992, K. W. Allred & J. Valclts-Reyna 5873 , 5874, 5875 (NMCR); along hwy. 57 at jet. to San Jose del Re¬ fugio, gypsum, 1340 m, 20 Oct. 1978, J. R. Reeder 7052 (ARIZ). U.S.A. Arizona: Cochise Co., along Cascabel to Willcox road, near 8 mi. marker, rocky limestone slope with ocotillo, Agave and grasses, 1450 m, 25 Aug. 1992, J. R. & C. G. Reeder 8878 (ARIZ, NMCR); 1 mi. NW of Naco (Waco?), 6 Oct. 1936, L A. Goodding 389 (ARIZ); Apache Pass between Dos Cabezas and Chiricahua Mts., Siphon Cnyn., 1450 m, 23 Aug. 1993, J. R. Reeder 9020 (ARIZ). Pima Co., Santa Rita Mts., hwy. jet. E of Vail, 3000 ft., 7 Sep. 1938, L. Benson 9121 (ARIZ). New Mex¬ ico: Bernalillo Co., 3 mi. W of Albuquerque, 9 Oct. 1945, R. A. Darrow 3367 (ARIZ). Chaves Co., Hwy. 380 ca. 1^ mi. W of Caprock, 8 July 1982, K. W. Allred 231 (NMCR); 3.5 mi. E of Elkins on hwy. 70, sandy, 03 " Artemisia-grassland, 26 Aug. 1974, L C. Higgins 917 (NMC). Dona Ana Co., Ropes Spring, San Andres Mts^, 17 Aug. 1933, [collector unknown] 89 (NMCR); 7 mi- Volume 5, Number 3 1995 Allred & Valdes-Reyna North American Aristida 221 of Hatch at jet. of hwy. 26 and 85, 2 Oct. 1982, T. Potter 17 (NMCR); Las Cruces, 6 Oct. 1904, D. Griffiths 7417 (NMCR); near Tortugas Mt., 24 Oct. 1904, E. 0. Wooton s.n. (NMC); White Sands Missile Range, 5 km E of head¬ quarters, 200 m S of Nike Ave., granitic sandy bajada, 23 Sep. 1990, R. Spellenberg 10678 (NMC). Eddy Co., abt. 40 mi. E of Roswell, 7 mi. N of hwy. 380, Mather Natural Area, sandy, shin oak, 3 Aug. 1979, R. Spellenberg 5249 (NMC). Guadalupe Co., 2 mi. S of Vaughn, 15 July 1979, L Rockhill 20 (NMCR). Luna Co., Cooke’s Range, Prov- inger Draw, 12 Aug. 1986, ]. T. Columbus 419 (NMCR); 5 mi. W of Columbus, 9 July 1984, J. S. Trent 107 (NMCR). Socorro Co., ca. 12 mi. E of San Antonio along hwy. 380, 5 July 1984, K. W. Allred 2700 (NMCR); road¬ side E of Belen on hwy. 60, 7 Apr. 1952, E. F. Costetter 7962 (NMC). Valencia Co., 20 mi. NE of Suwanee, 1 Sep. 1935, K. W. Parker 633 (NMCR). Texas: Culberson Co., deep sand 21 mi. W of Kent, 4000 ft., 17 Aug. 1950, B. H. Wamock 9317 (SR); dunes near Salt Flats, 4000 ft., 29 Aug. 1965, B. H. Warnock 20678 (SR); at Salt Flat Station, E of Salt Flat, 4000 ft., 29 Aug. 1965, B. H. Warnock 20674 (SR). Pecos Co., limestone hills 20—35 mi. S of Ft. Stockton, along Sanderson hwy., 3100 ft., 1 July 1955, B. H. Warnock 13266 (SR). Aristida brownii Wamock (Wamock, 1982) was described to accommodate single-awned Aristida (the lateral awns highly reduced) otherwise identi¬ cal to A. purpurea var. wrightii. Since its descrip¬ tion from Brewster County, Texas, individuals of this entity have been found from other Texas lo¬ calities, several locations in southern New Mexico, and one locality in Coahuila, Mexico. Plants of A. brownii are always found intermingled with variety wrightii, usually in sparse numbers, and never as isolated homogeneous populations. In addition, it is not uncommon to find in the same population with typical A. brownii plants with lateral awns 14—% the length of the central awn, an attribute intermediate between A. brownii and the sympatric variety wrightii. We find the recognition of this taxon un¬ tenable at the species level. Because the single- awned feature is unique in A. purpurea s.l. and eas¬ ily confused with other taxa, we propose its recognition at the level of forma. We have placed the name in a 3-level hierarchy to indicate its af¬ finities within A. purpurea, i.e., A. purpurea var. wrightii forma brownii. Aristida purpurea Nuttall forma brownii (War¬ nock) Allred & Valdes-Reyna, comb. nov. Bas- ionym; Aristida brownii Wamock, Sida 9: 358. 1982. TYPE: U.S.A. Texas: Brewster County, widespread perennial on limestone hills of Del Norte Mountains, elevation 4600 ft. or more, 27 June 1981, Warnock 141 (holotype, Lajitas Museum, not seen; isotypes, SMU, TEX). Allred (1984) reduced Aristida curvifolia Four¬ nier to a variety of A. purpurea because its strongly unequal glumes are also characteristic of the other varieties of A. purpurea. Further field studies during the subsequent 10 years have shown that disposi¬ tion to be unwarranted, and we propose the rein¬ statement of this taxon at the species rank, as did Divila & Sanchez-Ken (1994). Aristida curvifolia is consistently distinguished from all other desert Ar¬ istida by the following features: blades stiff, tightly involute, yellow-green, the epidermis nearly smooth and the veins hardly noticeable; glumes broad, blunt, unequal; lemma beak (apex) 1-2 mm long, 0.2-0.5 mm wide, straight to only slightly twisted even when mature; panicles spicate, always epul- vinate. Aristida curvifolia often grows with both A. purpurea and A. partsa, and no evidence of hybrid¬ ization or intermediacy has been noted. Indeed, this taxon turns out to be one of the more uniform spe¬ cies of Aristida in North America. It is presently known from 12 states in Mexico: Aguascalientes, Baja California Norte, Chihuahua, Coahuila, Gua¬ najuato, Nuevo Le6n, Oaxaca, Puebla, San Luis Po- tosf, Tamaulipas, Veracruz, and Zacatecas. Acknowledgments. Portions of this research were supported by the New Mexico State University Ag¬ ricultural Experiment Station. We thank Paul Pe¬ terson for sharing his collections, Jorge Sdnchez- Ken for his cooperation and hospitality, Richard Spellenberg for allowing us use of his maps, Miguel A. Carranza-P. for help in fieldwork, Cathy Montes of the Department of Agricultural Communications (New Mexico State University) for digitizing and printing the maps, anonymous reviewers for their helpful comments, and the staffs of the following institutions for the generous use of their specimens and facilities: AR1Z, ASU, CHAPA, COCA, ENCB, JEPS, MEXU, MO, POM/RSA, TAES, TEX/LL, UC, US. The illustrations were rendered by Cuauht¬ emoc Gonzfilez de Le6n. Literature Cited Allred, K. W. 1984. Morphologic variation and classifi¬ cation of the North American Aristida purpurea complex (Gramineae). Brittonia 36: 382—395. Beetle, A. A. 1983. Las Gramfneas de Mexico. Secretaria de Agricultura y Recursos Hidriulicos, Mexico City. D4vila, P. & J. SSnchez-Ken. 1994. Flora del Valle de Tehuacta-Cuicatlan. Fasc. 3: 7-22. Inst. Biol. Univ. Nac. Aut. Mexico, Mexico D.F. Gould, F. W. & R. Moran. 1981. The Grasses of Baja California, Mexico. San Diego Society of Natural His¬ tory, Memoir 12. Henrard, J. T. 1927. A critical revision of the genus Ar¬ istida. Meded. Rijks-Herb. 54: 221-464. -. 1928. A critical revision of the genus Aristida. Meded. Rijks-Herb. 54: 465-701. -. 1929. A monograph of the genus Aristida. Med¬ ed. Rijks-Herb. 58: 1-153. 222 Novon Hitchcock, A. S. 1924. The North American species of Aristida. Contr. U.S. Natl. Herb. 22: 517-586. -. 1935. Aristida. North American Flora 17: 376— 406. McVaugh, R. 1983. Flora Novo-Galiciana, Vol. 14, Gra- mineae. Univ. Michigan Press, Ann Arbor. Pohl, R. W. & G. Davidse. 1994. Aristida. Pp. 253-257 in: G. Davidse, M. Sousa S. & A. O. Chater (editors), Flora Mesoamericana, Vol. 6. Alismataceae a Cypera- ceae. Universidad Nacional Aut6noma de Mexico, Mex¬ ico D.F.; Missouri Botanical Garden, St. Louis; The Nat¬ ural History Museum, London. Wamock, B. H. 1982. A new three-awn grass from trans- Pecos Texas. Sida 9: 358-359. Two New Species of the Genus Sterigmapetalum (Rhizophoraceae) from the Venezuelan and Brazilian Amazonian Region Gerardo A. Aymard C. and Nidia Cuello A. UNELLEZ-Guanare, Programa de R.N.R. Herbario Universitario (PORT), Mesa de Cavacas, Edo. Portuguesa, Venezuela 3323 Abstract. Two new species of Sterigmapetalum subg. Sterigmapetalum are described from Vene¬ zuela and Brazil: 5. chrysophyllum and S. plum- beum. Relationships with related species are dis¬ cussed, and an illustration for each species is provided. Resumen. Se describen dos nuevas especies de Sterigmapetalum subgenero Sterigmapetalum de Venezuela y Brasil: S. chrysophyllum y .S', plum- beum. Se discuten sus relaciones con las especies afines y se presentan ilustraciones de cada una de ellas. Sterigmapetalum Kuhlmann is a small neotropi¬ cal genus with nine species in the Rhizophoraceae. Sterigmapetalum and Cassipourea Aublet were seg¬ regated by Steyermark & Liesner (1983) into a new tribe, the Hypogyneae. Subsequently, Tobe & Ra¬ ven (1987), based on a detailed study of the mor¬ phology of the seed, flower, and fruit, placed these two genera in the Macarisieae, a treatment origi¬ nally employed by Schimper (1893) a century ear¬ lier. Eight of the nine species of Sterigmapetalum oc¬ cur in northern South America (Colombia, Vene¬ zuela, Guyana, and Surinam) and one (5. obovatum Kuhlmann) is restricted to the western and central Amazonian region of Peru and Brazil (Prance et al., 1975). Sterigmapetalum chrysophyllum Aymard & Cuello, sp. nov. TYPE: Venezuela. Amazonas: Dpto. Atabapo, Rio Ventuari, bosques en pla¬ teau, 04°14'N, 65°S8'W, 1250 m, Feb. 1992 (fl), Proyecto Inventario de los Recursos Na- turales de la Guayana Venezolana-CVG-TEC- MIN, A. Chaviel 411 (holotype, PORT; iso¬ types, MO, VEN). Figure 1. Species nova quae a S. exappendiculato Steyermark & Liesner stipulis triangularibus, acutis, 3—5 mm longis; fo- liis coriaceis, 3—8 cm longis, supra glabris, pagina infema tomento flavo omnino occulta; petiolo 2-4 mm longo, ju- ventute dense flavo-tomentoso; petalis extus epapillatis, fimbriis lateralibus munitis differt. Trees 15—20 m high; branches 3-verticillate, te¬ rete, covered completely by a yellow-ocher pubes¬ cence when young, glabrous or sparsely adpressed- pubescent when mature. Stipules triangular, acuminate, 3—5 mm long, with yellow pubescent trichomes on both surfaces, internally glandular at the base. Leaves 3-whorled, the blades coriaceous, elliptical or oblong-elliptical, 3-8 X 1.5-5.5 cm, obtuse to truncated at the base, retuse at apex, mar¬ gin subrevolute, crenate, blades glabrous on upper surface, covered completely by yellow tomentum on under surface, lateral nerves 8-14 pairs, tertiary venation more evident on upper surface than on under surface; petiole 2-A X 1-2 mm, densely yel¬ low-ocher pubescent when young, glabrous or sparsely adpressed-pubescent when mature. Inflo¬ rescences solitary in the leaf axils, 1-2 cm long; 1 to many flowers, peduncle striate, adpressed-pu¬ bescent with yellow trichomes; pedicels 1 mm long, adpressed-pubescent with yellow trichomes on both surfaces; flowers subtended by bractlets, lanceolate, 1-2 mm long, adpressed-pubescent with yellow tri¬ chomes on both sides; staminate flowers with 5- lobulate calyx, obovate, 3-5 X 2-3 mm (calyx and lobes), adpressed-pubescent with yellow trichomes externally, adpressed-pubescent with sericeous tri¬ chomes internally; petals 5, 2-3 mm long, unguic- ulate, externally adpressed-pubescent with yellow trichomes from the middle to the apex, glabrous internally, with three apical appendices and two lat¬ eral ligules, the ligules 1-1.5 mm long, glabrous, 4-5-laciniate; stamens 8-10, united at the base, filaments glabrous, 1-2 mm long, anthers 1 mm long, oblong, glabrous; pistilodium rudimentary, conic, hispid with yellow trichomes; pistillate flow¬ ers not seen. Fruit obovate, 8—12 X 4—7 mm, com¬ pletely covered by an adpressed yellow-ocher in- dument, seeds 4-5 mm long, oblong, glabrous and reticulate. Distribution and common names. Sterigmape¬ talum chrysophyllum is only known from the middle and upper Ventuari River basin, Atabapo Depart¬ ment, Amazonas State, Venezuela, at 1250 m. It is Novon 5: 223-226. 1995. 224 Novon Figure 1. Sterigmapetalum chrysophyllum Aymard & Cuello. —A. Inflorescence with open flowers. —B. Fruiting branch. —C. Flowering branch. —D. Flower with 1 sepal removed leaving a horizontal scar. —E. Young fruit. Scale: A, D = 0.5 cm; B, C = 5 cm; E = 1 cm. A, C, and D drawn from holotype (Chaviel 411); B and E drawn from paratype (Chaviel 406). Volume 5, Number 3 1995 Aymard C. & Cuello A. Amazonian Sterigmapetalum 225 Figure 2. Sterigmapetalum plumbeum Aymard & Cuello. —A. Pistillate flower showing the ligules. -B. Flowering branch. —C. Base of the under surface of the leaf lamina. —D. Pistillate inflorescence. Scale: A, C, and D - 1 cm; B — 5 cm. All from Daly et al. 5608. known by the common names of “kasemani” and “tadiji” (Y’ekuana). Ecology. Mesothermic humid forests of medium height (18-25 m), on sandstone surface outcrop¬ pings. The absence of resinous exudate and the pres¬ ence of 3-whorled leaves and laciniate petal ligules indicate that Sterigmapetalum chrysophyllum be¬ longs to subgenus Sterigmapetalum (Steyermark & Liesner, 1983). This new species is related to 5. exappendiculatum Steyermark & Liesner, but it dif¬ fers in its triangular-acuminate stipules that are 3- 5 mm long (vs. stipules ovate-lanceolate, subob- tuse, and 7-10 mm long); coriaceous leaves that 226 Novon are 3-8 cm long, glabrous or sparsely adpressed- pubescent above and completely covered by yellow indumentum below (vs. leaves subcoriaceous, 7—14 cm long, sparsely pilose on the midrib above, mod¬ erately to densely adpressed-sericeous below); pet¬ ioles 2-4 mm long, completely covered by a yellow- ocher pubescence when young, and glabrous or sparsely adpressed-pubescent when mature; and petals that are not papillate externally, with two lat¬ eral ligules 4—5-laciniate (vs. petiole 5-11 mm long, sericeous-tomentose, and petals that are pa¬ pillate externally and without lateral ligules). Paralype. VENEZUELA. Amazonas: Dpto. Atabapo, Rto Ventuari, bosques en plateau, 04°14'N, 65°58'W, 1250 m, Feb. 1992 (fl), Proyecto Inventario de los Re- cursos Naturales de la Guayana Venezolana-CVG-TEC- MIN, A. Chaviel 406 (MO, PORT, VEN). Sterigmapetalum plumbeum Aymard & Cuello, sp. nov. TYPE: Brazil. Amazonas: Mun. Sao Gabriel da Cachoeira, Upper Rio Negro, near Comunidade Aparecida, opposite mouth of Igarape Tuari, 0°20'N, 67°19'W, 6 Nov. 1987 (fl), D. C. Daly, P. J. Maas & J. Ramos 5608 (holotype, MO; isotypes, INPA, NY). Figure 2. A S. colombiano Monachino stipulis triangularibus, 5- 6 mm longis; foliis coriaceis, oblongis, 8-11 cm longis, 5—7.5 cm latis, utrinque glabris, nervis lateralibus 17-24- jugatis; inflorescentiis 1.4-2.2 cm longis; floribus 4-5 mm longis, sepalis intus indumento grisaceo nitido obtectis, petalis oblanceolatis differt. Trees 8 m high; branches 3-verticillate, lenticel- late, sparsely adpressed-pubescent when young, glabrous when mature. Stipules triangular, 5-6 mm long, densely adpressed-pubescent on both sur¬ faces, barbate and glandular internally at the base. Leaves 3-whorled, the blades coriaceous, oblong, 8-11 X 5-7.5 cm, base rounded, apex emarginate, margins subrevolute, glabrous on both surfaces, lat¬ eral nerves 17—24 on both sides, tertiary venation very evident on both surfaces, petioles 5-7 X 2-3 mm, sparsely adpressed-pubescent on young leaves, glabrous on mature leaves. Inflorescences solitary in the axils, 1.4—2.2 cm long; peduncles adpressed-pubescent, especially at the base; pedi¬ cels 1-2 mm long, adpressed-pubescent, subtended by 3 bracts, lanceolate-triangular, the bracts 3 mm long, sparsely adpressed-pubescent externally, densely adpressed-pubescent internally; pistillate flowers with a 5-lobulate calyx, 4—5 mm long, 3-4 mm wide, glabrous to sparsely pilose externally, densely adpressed-pubescent internally with a sil¬ very gray indumentum; petals 5, white, 5-5.5 mm long, unguiculate, oblanceolate, externally adpres¬ sed-pubescent from the middle to the apex, inter¬ nally glabrous, with five lateral ligules, the ligules 3 mm long, glabrous, 5-laciniate; staminodes 10, united at the base, filaments glabrous, 1 mm long; ovary densely pubescent, style glabrous, 3 mm long, stigma capitate, staminate flowers and fruits not seen. Distribution. Only known in the upper Rio Ne¬ gro region, Municipio San Gabriel de Cachoeira, Amazonas, Brazil. Ecology. Upper Amazonian caatinga, on sandy soil. Sterigmapetalum plumbeum is related to S. col- ombianum Monachino (restricted to mesothermic humid forests in the department of Magdalena, northwestern Colombia; Barbosa, 1988), but it can be distinguished from this species by the triangular stipules, 5-6 mm long, coriaceous leaves, oblong, 8-11 cm long, 5-7.5 cm wide, glabrous on both sides, lateral nerves 17—24 (vs. ovate stipules, 1.5- 2 mm long, subcoriaceous, obovate leaves, 4.5-6.7 cm long, 2-4 cm wide, glabrous to sparsely pilose on the main vein and lateral ones below, lateral nerves 8-12); inflorescences 1.4—2.2 cm long, pis¬ tillate flowers 4—5 mm long, with sepals internally covered by a silvery gray indumentum and petals oblanceolate (vs. inflorescences 8-10 mm long, pis¬ tillate flowers 2-3 mm long with sepals internally sericeous and the petals subobovate). Acknowledgments. We thank Paul Berry and Bruce Holst for their collaboration during our stay at the Missouri Botanical Garden, Roy Gereau (MO) for the verification of the Latin descriptions, Laur¬ ence Dorr (US) for valuable comments on the manuscript, and John Myers for the excellent illus¬ trations. Literature Cited Barbosa, C. 1988. Rhizophoraceae en el Herbario Na¬ tional Colombiano. Trianea 2: 251—263. Prance, G. T., M. Freitas da Silva, B. W. Alburquerque, I. de J. da Silva Araujo, L. M. Medeiros Carreira, M. M. Nogueira Braga, M. Macedo, P. N. Da ConceicSo, P. L. Braga Lisboa, P. I. Braga, R. C. Lobato Lisboa & R. C. Queiroz Vilhena. 1975. RevisSo Taxon6mica das es- pecies amazfinicas de Rhizophoraceae. Acta Amazonica 5: 5-22. Schimper, A. F. W. 1893. Rhizophoraceae. In: A. Engler & K. Prantl (editors). Die Natlirlichen Pflanzenfamilien. Wilhelm Engelmann, Leipzig. 3(7): 42—56. Steyermark, J. A. & R. Liesner. 1983. Revision of the genus Sterigmapetalum (Rhizophoraceae). Ann. Missou¬ ri Bot. Gard. 70: 179-193. Tobe, H. & P. H. Raven. 1987. The embryology and re¬ lationships of Cassipourea and Sterigmapetalum (Rhi¬ zophoraceae—Macarisieae). Opera Bot. 92: 253-264. Taxonomic Changes in Ixonanthaceae from the Venezuelan Guayana Paul E. Berry Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A. Nelson Ramirez Centro de Biologfa Tropical, Universidad Central tie Venezuela, Apartado 20513, San Martin, Caracas, Venezuela ABSTRACT. Cyrillopsis micrantha (Steyermark) P. E. Berry & N. Ramirez is a new combination based on Ochthocosmus micranthm Steyermark and con¬ stitutes only the second known species of Cyrillop¬ sis (Ixonanthaceae). The number of varieties in Ochthocosmus roraimae Bentham is reduced from three to two, and Ochthocosmus parvifolius Hallier f. is treated as a species of uncertain status, with no known extant type material. The family Ixonanthaceae, which is sometimes treated as a subfamily of the Linaceae, includes two genera in the Neotropics, Cyrillopsis and Ochtho¬ cosmus. These two genera are distinguished mainly by fruit and pedicel characters. Ochthocosmus has a 5-locular ovary, the fruit is a prismatically cylin- dric, elongate, 5-valved capsule, and the seeds have a conspicuous, thin, lateral wing. Further¬ more, the pedicels are non-articulated, that is, they abscise flush with the rachis. In contrast, Cyrillopsis has a 2-locular ovary, its fruit is an elongate, slight¬ ly asymmetrical, 2-valved capsule, and the seeds lack wings but are partly covered at the upper end by a 2-lobed membranaceous aril; its pedicels are articulated, with an abscission zone and bracteoles located roughly one-third of the way up the pedicel from the rachis. Until now, Cyrillopsis was considered to have a single species, C. paraensis Kuhlman, and Ochtho¬ cosmus to have eight recognized neotropical species (Steyermark & Luteyn, 1980; Steyermark, 1984, 1988). Although some botanists, such as Mabberley (1990), consider Ochthocosmus to include species both in South America and in Africa, Forman (1965) presented convincing arguments to place the African species in the genus Phyllocosmus Klotzsch, thus leaving Ochthocosmus as a strictly neotropical group. While preparing the treatment of the Ixonantha¬ ceae for the Flora of the Venezuelan Guayana, we examined recently collected fruiting material of Ochthocosmus micranthus Steyermark, a species that was described from flowering specimens only (Steyermark, 1988). The articulated pedicels, aril- late seeds, and 2-locular fruits of these new collec¬ tions place the species clearly in Cyrillopsis rather than in Ochthocosmus. The transfer of O. micran¬ thus to Cyrillopsis brings the number of currently recognized species in Cyrillopsis to two and in Ochthocosmus to seven. We also reduce the number of varieties in O. roraimae from three to two by removing 0. roraimae Bentham var. parvifolius (Hallier f.) Steyermark & Luteyn as a variety of this species. Cyrillopsis micrantha (Steyermark) P. E. Berry & N. Ramirez, comb. nov. Basionym: Ochthocos¬ mus micranthus Steyermark. Ann. Missouri Bot. Gard. 75: 318. 1988. TYPE: Venezuela. Bolivar: Ayavaparu, 10 km SW of Wadacapia- pue, 5°18'N, 60°58'W, 1100 m, 13 Nov. 1986, Herndndez 348 (holotype, VEN; isotype, NY). [Steyermark (1988) was incorrect in citing the holotype at MO; rather, the holotype is at VEN and is so labeled, and there is no type material at MO.] Figure 1. Steyermark described this species based on two flowering specimens that strongly resemble other species of Ochthocosmus from the Guayana region. Three more recent collections from the type locality, however, Huber 11958, Huber & Herndndez 11948, and Herndndez 395, all have nearly mature fruits, and they conform to the characteristics of Cyrillopsis described above (see Fig. 1). Also, the type collection and the other specimens cited below all have the char¬ acteristic articulated pedicels of Cyrillopsis that do not occur in Ochthocosmus. This second species of Cyrillopsis has much smaller leaves than the previously known species, C. paraensis Kuhlman. Whereas C. paraensis is pri- Novon 5: 227-229. 1995. 228 No von Figure 1. Details of an inflorescence and fruit of Cyrillopsis micrantha (Steyermark) P. E. Berry & N. Ramirez. —A. Section of inflorescence after most of the flowers have dehisced. Note the articulated pedicels and the bracteole at the point of articulation. Drawn from Herndndez 348 (VEN). —B. Mature fruit, a somewhat fleshy capsule splitting into two halves. —C. Seed with arillate structure at the apex. B and C drawn from Huber & Herndndez 11948 (MO). marily a lowland species distributed from south¬ eastern Venezuela to French Guiana and northern Brazil, C. micrantha is an upland species endemic to the southeastern part of the Gran Sabana at around 1100 m elevation. It occurs in an unusual shrub savanna (described in Folster, 1992: 37) that is dominated by a sympatric population of Ochtho- cosmus roraimae. Additional specimens examined. VENEZUELA. Boli¬ var: Ayavaparu, 10 km SW of Wadakapiapue, 5°18'N, 60°58'W, 1100 m, 13 Nov. 1986, Herndndez 359 (US, VEN); Ayavapard, 15 km al W de Wadakapiapud-tepui, 5 16 30'N, 61°00'W, 1110 m, 6 Mar. 1987, Herndndez 395 (herbarium at San Ignacio de Yuruant, Venezuela); ca. 10 km SW of Wadakapiapuc-tepui. N of the junction of the Rfo Caraurfn and the Rfo Yuruant, ca. 1100 m, 5 15 N, 60 58 W, 18 Feb. 1987, Huber & Herndndez 11948 (MO, US, VEN); ca. 10 km SW of Wadakapiapud- tepui, N of the junction of the Rto Caraurfn and the Rfo Yuruanf, ca. 1030 m, 5°14'N, 60°58'W, 19 Feb 1987 Huber 11958 (MO, MYF, US—2 sheets, VEN). The label of Herndndez 395 cites “Wontai” as a common name for this species (in Taurepan, a di¬ alect of the Pemon Amerindian group). Ochthocosmus roraimae Bentham in Hooker, Lon¬ don J. Bot. 2: 366. 1843. TYPE: Guyana or Venezuela (“British Guiana” on label), large shrub 12—16 ft. high, banks of rivers, near Mt. Roraima, Oct.-Nov. 1842, Richard Schom- burgk 1037 (holotype, K, photo, MO). This species is widespread over much of the Ven¬ ezuelan Gran Sabana and extends into western Guyana and adjacent Roraima state in Brazil. It occurs between 800 and 1300 m elevation; field observations by Gabriel Pic6n and Nelson Ramfrez (pers. comm.) have shown 0. roraimae to be a vari¬ able taxon occurring in a variety of local habitats, from fairly open shrublands to low or moderately tall and dense forest. Steyermark & Luteyn (1980) recognized three va¬ rieties under 0. roraimae: var. roraimae, var. parvi- folius (Hallier f.) Steyermark & Luteyn, and var. grandifolius (Steyermark) Steyermark & Luteyn. Ochthocosmus roraimae var. grandifolius, which is known only from the type collection, is unique in its large, stiff, and virtually entire-margined leaves, so it is maintained here as a distinct variety. However, Steyermark and Luteyn employed an extremely nar¬ row concept for the typical variety roraimae, char¬ acterizing it by a shiny and reticulately raised- veined upper surface and assigning to it only two additional specimens besides the type (of which they examined a reduced black-and-white photograph). They then relegated to variety parvifolius all remain¬ ing specimens of the species, encompassing a wide range of variation in leaf size, crenulation of the mar¬ gins, and length of the inflorescence. After examin¬ ing a full-sized color Cibachrome print of the type of 0. roraimae, kindly sent by the Kew herbarium. Volume 5, Number 3 1995 Berry & Ramirez 229 Ixonanthaceae from Venezuelan Guayana and studying the full range of variation of the spe¬ cies, there is no consistent way to distinguish the specimens of variety roraimae from variety parvifol¬ ius, and all specimens cited by Steyermark & Luteyn (1980) under variety parvifolius are now treated as belonging to variety roraimae. The status of the bas- ionym of variety parvifolius is discussed following the key to the varieties of O. roraimae. Key to the Varieties of Ochthocosmus roraimae 1. Leaf blades of fertile branches oval, 7-13 X 4—8 cm, margins entire or nearly so; panicles 11—14 cm long, 10-12 cm wide; known only from the upper slopes of Cerro Venamo. . 0. roraimae var. grandifolius 1. Leaf blades of fertile branches mostly obovate, 3— 10 X 2—6 cm, margins usually noticeably crenate or toothed; panicles 3—8 cm long, 2—5 cm wide; widespread throughout much of the Gran Sabana, also in western Guyana and adjacent Brazil . . . . 0. roraimae var. roraimae Ochthocosmus parvifolius Hallier f., Beih. Bot. Cen- tralbl. 39 (2). 15. 1921. Ochthocosmus rorai¬ mae var. parvifolius (Hallier f.) Steyermark & Luteyn, Brittonia 32: 135. 1980. TYPE: Guy¬ ana or Venezuela (“British Guiana” on label), undershrub 6—10 ft. high, sandstone region, Mt. Roraima, Oct.—Nov. 1842, Richard Schom- burgk 1046 (holotype, B destroyed, photo, US, VEN). The holotype of Ochthocosmus parvifolius Hallier f. at B was destroyed in 1943, and there is appar¬ ently no extant duplicate material (inquiries were made at B, BM, G, K, and W). The type photo¬ graphs are dark and much reduced in size from the original specimen, with no evidence of flowers or inflorescences. Steyermark & Luteyn (1980) treated this as a variety of O. roraimae, but the inflores¬ cence, described as just 2.5 cm long, is very short for that species. According to the protologue of 0. parvifolius and Schomburgk’s collection notes in the New York Botanical Garden archives, the type specimen came from a low shrub smaller than the more robust and often tree-like O. roraimae. Ochthocosmus parvifolius was also described as having unequal sepals, which are characteristic of species such as 0. floribundus Gleason, but not of O. roraimae, which has equal-sized, glandular-mar- gined sepals. Ochthocosmus floribundus is not known to occur around Mt. Roraima and is known primarily from areas farther north and west in Es- tado Bolivar such as AuyAn-tepui and Cerro Guia- quinima. The possibility also exists that Ochthocosmus parvifolius is conspecific with a later-described spe¬ cies such as 0. attenuatus Steyermark & Luteyn, which is known so far only from the type locality near San Rafael de Camoirdn in the northern part of the Gran Sabana. Given the current uncertainties concerning the identity of O. parvifolius, and until the issue can be better resolved, we recommend that this species be treated as one of uncertain po¬ sition (incertae sedis). Acknowledgments. We thank Bruno Manara for making the illustrations, the Royal Botanic Gardens at Kew for the Cibachrome type photograph of Ochthocosmus roraimae, The New York Botanical Garden for providing a loan of their specimens and copies from the Schomburgk collection notes, and H. Sipman (B), S. Knapp (BM). F. Jacquemoud (G), G. Lucas (K), J. Pruski (US), and H. Riedl (W) for checking for possible type material of 0. parvifol¬ ius. This work was supported in part by National Science Foundation grant BSR-9201044 to the Missouri Botanical Garden. CVG-EDELCA (Elec- trificacidn del Caronf, C.A.) kindly provided logis¬ tical support for field studies related to this paper. Literature Cited Folster, H. 1992. Holocene autochthonous forest degra¬ dation in southeast Venezuela. Pp. 25-A4 in J. G. Gol- dammer (editor). Tropical Forests in Transition. Birk- hauser Verlag, Basel. Forman, L. 1965. A new genus of Ixonanthaceae, with notes on the family. Kew Bull. 19: 517-526. Mabberley, D. J. 1990. The Plant-Book. A Portable Dic¬ tionary of the Higher Plants. Cambridge Univ. Press, Cambridge. Steyermark, J. A. 1984. Flora of the Venezuelan Guaya¬ na. I. Ann. Missouri Bot. Gard. 71: 297-340. -. 1988. Flora of the Venezuelan Guayana. IV. Ann. Missouri Bot. Gard. 75: 311—351. -& J. L. Luteyn. 1980. Revision of the genus Ochthocosmus. Brittonia 32: 128—143. Three New Combinations in Pfaffia (Amaranthaceae) from the New World Tropics Thomas Borsch Forschungsinstitut Senckenberg, Senckenberganlage 25, 60325 Frankfurt am Main, Germany Abstract. The new combinations Pfaffia aurata (Martius) Borsch, Pfaffia completa (Uline & W. L. Bray) Borsch, and Pfaffia costaricensis (Standley) Borsch are made. In analyzing the pollen morphol¬ ogy and floral structures of the three species, it is shown that they belong to Pfaffia Martius and not to Iresine P. Browne, nom. cons. Pfaffia belongs to Amaranthaceae subfamily Gomphrenoideae (Schinz, 1934; Townsend, 1993). It comprises about 35 species, all characterized by long trichomes arising in tufts from the pedicel or the base of the perianth, a bilobate stigma, and the flowers all being hermaphrodite. The genus is dis¬ tributed in the New World, according to Stiitzer’s (1935) monograph, ranging from Mexico south¬ wards throughout the tropics, including the Amazon basin, to the region of Bahia Blanca, Argentina. During preparation of the Amaranthaceae treat¬ ment for Flora de Nicaragua and subsequent stud¬ ies in the genera Pfaffia and Iresine, three new combinations turned out to be necessary. Pfaffia aurata (Martius) Borsch, comb. nov. Bas- ionym: Trommsdorffia aurata Martius, Nova Genera et Species Plantarum, Vol. 2; 41. 1826. TYPE: Brazil. Provinciae Rio Negro: habitat in argillosis praeruptis oparatis flu[. . .] Japura ripis ad Mar[ibis], Dec.-Jan., C. F. P. v. Mar¬ tius s.n. (holotype, M). Pfaffia completa (Uline & W. L. Bray) Borsch, comb. nov. Basionym: Iresine completa Uline & W 1,. Bray, Bot. Gaz. (Crawfordsville) 21: 349. 1896. TYPE: Honduras. Santa Barbara: San Pedro Sula, 200 m, C. Thieme 338 (ho¬ lotype, US; isotypes, BM, K). Figure 1. Iresine wrightii Standley, Contr. U.S. Natl. Herb. 18: 97. 1916. Syn. nov. TYPE: Nicaragua [Herbarium of the U.S. North Pac. Expl. Exped. under Commanders Ringgold and Rodgers 1853-1856], C. Wright s.n. (holotype, US). Iresine ealea (Ibanez) Standley, Contrib. U.S. Natl. Herb. 18: 94. 1916. Iresine completa Uline & W. L. Bray f. glabriuscula Sues- senguth, Repert. Spec. Nov. Regni Veg. 39: 11. 1935. Syn. nov. TYPE: Mexico. Mazatl&n: Felsige Abhange, Jan.-Mar. 1902, C. A. Purpus 362 (holo¬ type, WU). There has been some confusion about the iden¬ tity of this species because John Donnell Smith re¬ numbered Thieme’s specimen ( Thieme 338) with his own number, 5443. There is a second collection by Thieme from the same locality in Honduras, re¬ numbered 5447. Suessenguth (1934) erroneously cited “Donnell-Smith n. 5447" as the type. This specimen (BM, US) is a functionally male plant of Iresine calea (Ibanez) Standley. It has small pistil- lodes and might have been interpreted as being hermaphroditic for that reason, but the plant lacks the long basal trichomes that in hermaphroditic or functionally female flowers of all species of Iresine serve for wind dispersal. Obviously, this specimen cannot be the type mentioned by Uline & Bray (1896). In line with his misinterpretation, Suessen¬ guth (1935) described Iresine completa Uline & W. L. Bray f. glabriuscula Suessenguth from a speci¬ men that is a functionally male plant of Iresine ca¬ lea. When publishing Iresine completa, Uline & Bray (1896) stressed the complete hermaphroditism of their plant, a remarkable character in contrast to the many species belonging to Iresine which are dioecious or polygamous. It is obvious that they called it “ completa ” for that reason. Pfaffia costaricensis (Standley) Borsch, comb, nov. Basionym: Iresine costaricensis Standley, Contrib. U.S. Natl. Herb. 18: 94. 1916. TYPE: Costa Rica. Tucurrique, Buissons a Las Vuel- tas, 635 m. Mar. 1899, A. Tonduz 13183 (ho¬ lotype, US; isotypes, K, US). Figure 2. Pfaffia aurata, P. completa, and P. costaricensis show the bilobate stigma, which is a distinguishing character of Pfaffia. In P. aurata the lobes have almost the outline of an equilateral triangle. In P- completa the lobes are narrowly triangular, and in P. costaricensis the stigma is shaped more or less Novon 5; 230-233. 1995. Volume 5, Number 3 1995 Borsch New Combinations in Pfaffia 231 Figures 1, 2. Scanning electron (SEM) photomicrograph of air-dried pollen. —1. Pfaffia completa (Thieme 338, BM). —2. P. costaricensis (Tonduz 13183, K). like a mushroom cap and the lobes are very short and broad. All three species have subequal, 3- veined tepals with a tuft of long, straight and stiff simple trichomes arising abaxially at the base of the tepal and from the pedicel. The flowers are al¬ ways hermaphroditic. Duke’s (1961) suspicion that P. completa could be a polygamous species with the pistillate flowers perfect and the staminate flowers bearing pistillodes is a result of the above-men¬ tioned confusion about the species’ identity. In all three species the filaments are united at the base into a shallow cup. Broadly rounded pseudostami- nodia are present, which are distinctly shorter than the free part of the filament. Sometimes they are notched at the apex or they are indistinct and the margin of the cup is only slightly raised. The pollen grains of Pfaffia aurata, P. completa (Fig. 1), and P. costaricensis (Fig. 2) are pantopor- ate, having 12—14, 5-angular lumina and sunken pores. They represent the Gomphrena- type, one of two major pollen types in the Amaranthaceae (Erdt- man, 1952). The Gomphrena-type is placed in a specialized category in Nowicke’s (1975) study of centrospermous pollen. Pfaffia has pollen of the Gomphrena-type, whereas pollen of lresine is sim¬ ilar to the Amaranthus- type of Erdtman, which cor¬ responds to Pollen-Type II of Nowicke (1975). In Pfaffia aurata, P. completa, and P. costaricensis, the tectum appears to cover the columellae completely and the muri have more or less vertical sides. The extraporal sexine bears distinct elongate spinules in all three species. This is exactly the same struc¬ ture as in species of Pfaffia (see illustrations of Pfaffia pollen by Nowicke & Skvarla (1979), Cuad- rado (1988), Eliasson (1988), and Townsend (1993)). There is only variation in aperture number, which is known to occur between species within many genera of Amaranthaceae. Eliasson (1988) showed Pfaffia- pollen to have a row of warts or spi¬ nules on the distal part of their tectum. In P. au¬ rata, a species illustrated by Eliasson as figure 34 under the misapplied name Pfaffia paniculala (Martius) Kuntze, these rows are not very straight. In investigating more specimens of P. aurata from different localities the same observation was made. Sometimes the spinules were in an undulate row, some of them even side by side. The pollen of P. costaricensis (Fig. 2) also has some of the spinules side by side, but there is a certain amount of vari¬ ation, from a side by side arrangement of the spi¬ nules to straight rows, even within a number of grains from a single anther. In contrast to lresine. 232 Novon the spinules are never evenly spread over a more or less convex distal part of the tectum. In Iresine the pores are small and apertures cover a minor part of the surface of the grain. Iresine angustifolia Euphrasen has more specialized pollen with wider pores, compared to the other species of the genus, but the distal parts of the tectum are still convex and the spinules are evenly spread. Pfaffia aurata, P. completa, and P. costaricensis are closely related and belong to a group that was described as Trommsdorffia by Martius (1826). In 1839 Dietrich transferred Trommsdorffia aurata Martius (the type species of Trommsdorffia ), T. ar- gentata Martius, and T. pulverulenta Martius to Ire¬ sine P. Browne. Similarities in habit (all species be¬ ing scandent shrubs or lianas), several times branched racemose inflorescence structures, and the common occurrence of long trichomes serving for dispersal were the decisive factors for merging the two genera. My own studies in connection with a revision of Iresine revealed that Trommsdorffia is different from Iresine not only in pollen morphology but also in floral structure, most strikingly in the morphology of the stigma (two more or less elon¬ gated, filiform branches). Iresine and Trommsdorffia belong to completely different evolutionary lines. This is also supported by the strong trend to the evolution of polygamous or dioecious species in Ire- sine, whereas the relatives of Pfaffia aurata and all other species of Pfaffia are generally hermaphro¬ ditic. Consequently, Trommsdorffia has to be ex¬ cluded from Iresine. Trommsdorffia pulverulenta Martius (= Pfaffia densipellita Borsch), a morpho¬ logically well defined species also allied to P. au¬ rata from Peru, was excluded from Iresine earlier (Borsch, 1993). Pfaffia aurata and its allies form a small group of closely related species within the genus Pfaffia. Differences from the other species of Pfaffia are merely a matter of degree. The occurrence of more or less broadly rounded pseudostaminodia in P. au¬ rata and its relatives is not sufficient to support generic status. Their position within the genus Pfaf¬ fia and the generic concept of Pfaffia will be dis¬ cussed in detail in a separate paper. It is question¬ able whether Trommsdorffia argentata Martius is specifically distinct from Pfaffia aurata, and sev¬ eral other morphologically very similar taxa (e.g., Iresine argentata (Martius) D. Dietrich var. ama- zonica Seubert = Iresine aurata (Martius) D. Die¬ trich var. amazonica (Seubert) Suessenguth) are be¬ ing studied more thoroughly before a taxonomic decision is made as to their status among the rel¬ atives of P. aurata. Pfaffia aurata has tepals that are ovate-oblong. 3-veined, 1.3-1.8 mm long and stigma lobes that have more or less the outline of an equilateral tri¬ angle. Pfaffia completa is characterized by its te¬ pals, which are narrowly triangular, prominently 3- veined, about 2.5 mm long, narrowly triangular stigma lobes becoming almost cylindrical at top, and broadly lanceolate blades, which are glabrous on the upper and sparsely pubescent on the lower surface. Pfaffia costaricensis has extremely small floral parts and conspicuously finely branched in¬ florescences, representing 3—4 times branched rac¬ emose structures with strictly opposite branches. The tepals are ovate, 1.1—1.3 mm long, and indis¬ tinctly 3-veined. The stigma lobes are very broadly rounded, even shorter and broader than in P. au¬ rata. The anthers have a length of 0.2 mm, in con¬ trast to the other species, which have anthers that are about 0.4 mm long. Burger (1983) described the characteristic elliptic or broadly elliptic leaf blades of P. costaricensis, with up to 12 pairs of major secondary veins. Pfaffia aurata probably ranges throughout the Neotropics. In Ecuador the species is well estab¬ lished in virgin forests as well as rastrojos, accord¬ ing to the Flora of Ecuador (Eliasson, 1987). There the species is erroneously called Pfaffia paniculata (Martius) Kuntze, a name of a different taxon be¬ longing to a group of closely related species that were described as Hebanthe by Martius (1826). Suessenguth (1934) described Pfaffia completa as occurring from Guatemala to Costa Rica. This range is partly a consequence of the misinterpretation of the species’s identity. In herbaria many older spec¬ imens identified as Iresine completa Uline & W. L. Bray belong in fact to Iresine calea (Ibdnez) Stan- dley. Apart from the type locality in Honduras, Pfaffia completa is at the moment proven to occur in Nicaragua and Panama. Two specimens exist from Nicaragua. One has been collected recently on Isla de Ometepe in Lago de Nicaragua, and the other is the type of Iresine wrightii. The single spec¬ imen from Panama has been collected in the Darien and is also cited by Duke (1961). Pfaffia costari¬ censis is endemic to Costa Rica and is known only from the central part of the country (Burger, 1983). A recent collection has been made in a remote area in Puntarenas, where the plant was climbing on secondary woody vegetation (Hammel, pers. comm.). Additional specimens examined. Iresine calea (Iba¬ nez) Standley (specimens incorrectly identified as Iresine completa Uline & W. L. Bray): COSTA RICA. Puntar¬ enas: Santo Domingo de Golfo Dulce, Tonduz 9861 (BM). HONDURAS. Santa Barbara: San Pedro Sula, Thieme s.n., renumbered by Donnell Smith 5447 (BM,US). NIC- Volume 5, Number 3 1995 Borsch New Combinations in Pfaffia 233 ARAGUA. Managua: S of Managua, J. M. & M.T. Green- man 5725 (MO). Pfaffia aurata (Martius) Borsch: NIC¬ ARAGUA. Boaco: San Jos6 de los Remates, Moreno 20337 (MO). Rivas: Isla de Ometepe, al NE del Volc&n Concepcidn, Sandino 4233 (FR, HNMN not seen). Ze- laya: La Pimienta, Pipoly 6291 (FR); along new road from Siuna to Matagalpa, ca. 31.4 km beyond Rfo Uli, near Wani, Stevens 7493 (FR). Pfaffia completa (Uline & W. L. Bray) Borsch: NICARAGUA. Rivas: Isla de Ometepe, Volc£n Madera, Robleto 291 (FR, MO). PANAMA. Dari¬ en: Paca, Williams 706 (US). Pfaffia costaricensis (Standley) Borsch: COSTA RICA. Puntarenas: Canton de Osa, Fila Costena, Fila Cruces, cabezeras del Rfo Piedras Blancas, Hammel 19289 (MO). Acknowledgments. I thank the curators and di¬ rectors of the herbaria MO, US, and WU for loaning specimens, of BM and K for loaning specimens and permitting the removal of pollen for SEM studies, and of M for allowing me to study the herbarium of C. F. P. v. Martius. I am grateful to Alan J. Paton for his kind help during my work on the collections at Kew, to Troels M. Pedersen, Mburucuya, Argen¬ tina, for hinting to me the true identity of Hebanthe paniculata Martius, and to Barry Hammel, Missouri Botanical Garden, for information on Pfaffia cos¬ taricensis. The SEM photographs were taken at the Forschungsinstitut Senckenberg using a CamScan CS 24 Compact scanning electron microscope. I am especially grateful to the research staff of the Mis¬ souri Botanical Garden, who first drew my attention to the Amaranthaceae. Literature Cited Borsch, Th. 1993. Amaranthaceae. Pp. 18-26 + Appen¬ dix I in L. Brako & J. L. Zarucchi, Catalogue of the Flowering Plants and Gymnosperms of Peru/Cat£logo de las Angiospermas y Gimnospermas del Perti. Monogr. Syst. Bot. Missouri Bot. Gard. 45. Burger, W. C. 1983. Amaranthaceae. In: Flora Costari¬ censis. Fieldiana, Bot. 13: 142-180. Cuadrado, G. A. 1988. Granos de polen de Amarantha¬ ceae del nordeste Argentino: 2. G£nero Pfaffia. Bol. Soc. Argent. Bot. 25(3—4): 385-394. Dietrich, D. 1839. Synopsis Plantarum. Vol. 1. Bernhard Friedrich Voigt, Weimar. Duke, J. A. 1961. Amaranthaceae. In: Flora of Panama. Ann. Missouri Bot. Gard. 48: 6-50. Eliasson, U. H. 1987. Amaranthaceae. Pp. 1-138 in G. Harling & L. Andersson, Flora of Ecuador, Vol. 28. Ber- lings, Alov. -. 1988. Floral morphology and taxonomic rela¬ tions among the genera of Amaranthaceae in the New World and the Hawaiian Islands. J. Linn. Soc., Bot. 96: 235-283. Erdtman, G. 1952. Pollen Morphology and Plant Taxon¬ omy. Angiosperms. Almqvist & Wiksell, Stockholm. Martius, C. F. P. von. 1823—1832. Nova Genera et Spe¬ cies Plantarum. 3 Vols. Lindauer et Wolf, Monachii. Nowicke, J. W. 1975. Pollen morphology in the order Centrospermae. Grana 15: 51-77. -& J. J. Skvarla. 1979. Pollen morphology: The potential influence in higher order systematics. Ann Missouri Bot. Gard. 66: 633-700. Schinz, H. 1934. Amaranthaceae. Pp. 7-85 in A. Engler & K. PrantI, Die natiirlichen Pflanzenfamilien, ed. 2, 16c. Leipzig. Stutzer, 0. 1935. Die Gattung Pfaffia. Repert. Spec. Nov. Regni Veg. Beih. LXXXVIII, 1-46. Berlin. Suessenguth, K. 1934. Neue und kritische Amarantaceen aus Siid-und Mittelamerika. Repert. Spec. Nov. Regni Veg. 35: 298-337. -. 1935. Neue und kritische Pflanzen aus Suda- merika, insbesondere Amarantaceen, sowie eine neue Gattung der Podostemonaceae. Repert. Spec. Nov. Reg¬ ni Veg. 39: 1—20. Townsend, C. C. 1993. Amaranthaceae. Pp. 70-91 in K. Kubitzki et al., Families and Genera of Flowering Plants, Vol. 2. Springer, Berlin. Uline, E. B. & W. L. Bray. 1896. Synopsis of North Amer¬ ican Amaranthaceae V. Bot. Gaz. (Crawfordsville) 21: 348-356. Paspalum morichalense (Poaceae: Paniceae), a New Aquatic Species from South America Gerrit Davidse Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A. Fernando 0. Zuloaga Instituto de Botanico Darwinion, Casilla de Correo 22, San Isidro, 1642, Argentina Tarciso S. Filgueiras Reserva Ecologica do IBGE, Caixa Postal 08770, 312-970 Brasilia, DF, Brazil Abstract. Paspalum morichalense is described, illustrated, and compared to related species of the group Dissecta. It occurs in Venezuela, Surinam, Brazil, and Bolivia. Its confusion with Paspalum commutatum Nees and Paspalum lacustre Chase ex Swallen is discussed. We have been aware for some time of the exis¬ tence of a possibly undescribed, aquatic species of Paspalum L. However, because of its confusion with two other species, P. commutatum and P. la¬ custre, it was not possible to describe it until the types of those two names had been studied. Pas¬ palum is a largely American genus with approxi¬ mately 330 species. The group names used in this paper are the ones used by Chase (1929, ined.) in her infrageneric classification of the genus. Paspalum morichalense Davidse, Zuloaga & Fil¬ gueiras, sp. nov. TYPE: Venezuela. Guarico: ca. 32 km SSE of Calabozo on Finca Becerra, in and around morichal with standing water, morichal dominated by Mauritia flexuosa, 65 m, 6 Nov. 1973, Davidse 3770 (holotype, MO; isotypes, AAU, K, LE, MEXU, MG, PRE, SI, US, VEN). Figure 1. Gramen perenne, repens vel fluitans, aquaticum; culmi 13-25 cm alti, ramosi. Folia caulina; ligulae 0.6-2.0 mm longae, glabrae. Racemi 1-2, 3-6 cm longi; rhachis 1.1- 3.5 mm lata, alata, herbacea, scaberula. Spiculae 2.1—2.9 X 0.9-1.1 mm, ellipticae, singulares, non imbricatae, gla¬ brae, acutae. Antherae 3, 0.O—0.8 mm longae, purpureae. Creeping or floating, aquatic perennial. Culms many-noded, decumbent and rooting at the lower nodes, freely branching, the erect portion of the culms 13—25 cm tall; intemodes hollow, glabrous; nodes dark, glabrous or sparsely pilose. Leaves cauline. Sheaths 2-8 cm long, glabrous, slightly Novon 5; 234-237. 1995. keeled; collar dark glabrous. Ligules 0.6—2.0 mm long, membranous, truncate, glabrous. Blades 3.5- 11 cm long, 1.5-3 mm wide, linear to linear-lan¬ ceolate, flat, narrowed at the base, widely spreading to reflexed, glabrous to scabrous on the adaxial sur¬ face and glabrous on the abaxial surface, the mar¬ gins scaberulous. Inflorescence terminal, partially included in the upper leaves; peduncle 2 cm long, cylindric; racemes 1-2, 3-6 cm long; rachis 1.1— 3.5 mm wide, winged, herbaceous, green, incon¬ spicuously veined, glabrous, terminating in a spike- let, the margins minutely scaberulous, usually fold¬ ed and partially embracing the spikelets, the spikelets not crowded, not overlapping on the same side of the rachis; pedicels solitary, 0.3-0.5 mm long, scaberulous. Spikelets 2.1-2.9 mm long, 0.9— 1.1 mm wide, elliptic, concave-convex, broadly acute, whitish green, glabrous. Lower glume absent. Upper glume and lower lemma subequal or the up¬ per glume 0.1-0.2 mm shorter than the upper flo¬ ret, hyaline, 3—5-nerved, with one central nerve and with the submarginal and marginal nerves close together. Lower floret sterile. Lower lemma 5- nerved. Lower palea absent. Upper floret 2.1—2.8 mm long, 0.9—1.1 mm wide, as long as the spikelet, bisexual, elliptic, acute, whitish, indurate, smooth and shining, papillose and with prickle hairs and bicellular microhairs at the apex; lodicules 2, 0.3 mm long, truncate. Stamens 3; anthers 0.5-0.8 mm long, purple. Styles 2, separate; stigmas ca. V4 as long as the styles, plumose, purple. Caryopsis 1.6— 1.8 mm long, 0.9 mm wide, elliptic, brownish; hi- lum ca. % as long as the caryopsis, linear; embryo ca. % as long as the caryopsis. This species is known from Venezuela to Brazil and Bolivia. We have studied it in the field in GuA- rico, Venezuela, and Bahia, Brazil. In GuArico it grows on mud and in shallow water of swamps and Volume 5, Number 3 1995 Davidse et al. Paspalum morichalense 235 Figure I. Pnnicum morichalense Davidse, Zuloaga & Filgueiras (Davidse 3770 MO). —A. Habit. B. Inflorescence. -C. Spikelet, upper glume view. -D. Spikelet. lateral view. E. Spikelet, lower lemma vtew. F. Uppeir floret, lemma view. -G Upper floret, palea view. -H. Detail of apex of the upper lemma. -I. Caryops.s, embryo s.de. - J. Caryopsis, hilum side. ponds at an elevation between 65 and 125 m, es¬ pecially in morichales, palm swamps dominated by Mauritia flexuosa L. f. (Arecaceae). The Bahian gathering was collected at an elevation of approx¬ imately 450 m in a pond ca. 20 m across and ca. 1-2 m deep. Here it grew at the edges and at the center of the pond and was much sought after by cattle and horses. The latter entered into the deeper waters of the pond to reach the soft, floating culms. The specific epithet is derived from the word morichal, which is the Venezuelan designation for a Mauritia swamp. The equivalent Brazilian vege¬ tation type is commonly known as vereda. Because of its few racemes, foliaceous, winged 236 Novon rachis, aquatic habit, creeping or floating culms with many elongated nodes, and glabrous spikelets, Paspalum morichalense belongs to the informal group Dissecta of Chase (1929). Within this group it is most closely related to P. lacustre and P. dis- sectum (L.) L. Paspalum lacustre differs from P. morichalense by its more numerous racemes ('1—6 vs. 1—2), spike- lets sharply acute (vs. broadly acute), the upper floret shorter than the upper glume and lower lem¬ ma (0.4—0.9 mm shorter vs. as long as or only 0.2 mm shorter), longer pedicels (ca. 1.0 mm vs. 0.3- 0.5 mm), and rachis scabrous versus minutely sca- berulous. Paspalum lacustre is known only from the type collection [BRAZIL. Amapa: Mun. Macapa, Ambe, localidade no afluente do Rio Pedreira, em lago fundo. 8 jul. 1951, R. L. Frdes & G. A. Black 27312 (holotype, INPA not seen, fragment US; iso¬ type, US)]. Paspalum dissectum differs from P. morichalense in its more numerous racemes (2-A vs. 1-2), spike- lets generally smaller (1.8-2.2 X 1.2-1.5 vs. 2.1- 2.9 X 0.9-1.1 mm), obovate (vs. elliptic), and more densely arranged (i.e., overlapping in the same row on one side of the rachis vs. not overlapping in one row), veins more strongly developed in the rachis wings, and the rachis wing margins scabrous (vs. minutely scaberulous). Although specimens of Paspalum morichalense have been known at least since the 1864 collection of Warming in Brazil, the species has remained for¬ mally unrecognized until now and has been con¬ fused with two other species. As noted by Chase (ined.), Doell (1877) in Flora Brasiliensis partially misapplied the name Paspalum commutatum. Un¬ der this species he correctly cited a Riedel collec¬ tion, but also included the Warming collection. Paspalum commutatum, as shown by an examina¬ tion of a fragment of the holotype [BRAZIL: pr. Porto Est. locis tempore pluviore inundatis . . . Langsdorff s.n. (holotype, LE not seen, fragment US)], belongs to the group Plicatula of Chase (1929) because it has compressed sheaths, 4-5 racemes, broadly obovate spikelets, and a dark brown, shin¬ ing upper floret. There is no close relationship be¬ tween groups Dissecta and Plicatula. As shown by her annotations and her unpub¬ lished manuscript. Chase (ined.) recognized that P. morichalense was an undescribed species, but she did not differentiate it from P. lacustre, a concept also followed by Judziewicz (1990). On the other hand, Swallen (1967), who effectively published P. lacustre, apparently did distinguish between P. mor¬ ichalense and P. lacustre, since he cited only the type collection, Frdes & Black 27312, under his new species. Unfortunately, he mistakenly classi¬ fied this species as a member of the Plicatula group. In annotations of the US specimens Chase had equated the two collections of Frdes and Black ( 27195 and 27312 ) under the same, unpublished name. We, on the other hand, include Frdes & Black 27195 under P. morichalense, and Frdes & Black 27312 under P. lacustre. As noted earlier, the description of P. lacustre in Judziewicz (1990: 484) is based on a mixture of P. morichalense and P. lacustre, but the illustration (fig. 84A—C) represents only P. morichalense. These species of group Dissecta may be distin¬ guished through the following key: la. Rachis of the raceme extending beyond the spikelets as a naked point. 2a. Spikelets 0.6-0.8 mm wide, in 2 rows; Unit¬ ed States to Argentina. P. repens Bergius 2b. Spikelets 0.8-1.2 mm wide, in 1 row; Mex¬ ico (Nayarit to Chiapas) . . . P. longicuspe Nash lb. Rachis ending in a spikelet. 3a. Upper floret as long as the lower lemma and upper glume or at most 0.1-0.2 mm shorter; Venezuela, Surinam, Brazil, Bolivia . . . P. morichalense Davidse, Zuloaga & Filgueiras 3b. Upper floret at least 0.2 mm shorter than the lower lemma and upper glume. 4a. Spikelets on the same side of the rachis not overlapping; Brazil (Amapa) . . . P. lacustre Chase ex Swallen 4b. Spikelets on the same side of the rachis overlapping, especially toward the base of the rachis. 5a. Spikelets 1.8-2.2 mm long, obtuse; eastern United States, Cuba . . . P. dissectum (L.) L. 5b. Spikelets 3—3.9 mm long, acumi¬ nate. 6a. Racemes 2, rarely 1; blades mostly 3-6 mm wide; Cuba, Jamaica . . . P. serratum Hitchcock & Chase 6b. Racemes 3-5, rarely 2; blades mostly more than 8 mm wide; southern United States to Ar¬ gentina . . . P. acuminatum Raddi Paratypes. VENEZUELA. Bolivar: small swamp in open savanna at base of Serranta de Pijiguao, 160 km SW of Caicara del Orinoco, 6°35'N, 66°45'W, 100—125 m, 12 Sep. 1985, Steyermark, Holst & Manara 131802 (MO, VEN). Guariro: 48 km N of Santa Rita along Hwy. 12 between Cabruta and Chagauramas, shallow pond with standing water in Trachypogon savanna with Curatella and Bowdichia, 110 m, 21 Nov. 1973, Davidse 4287 (COL, K, MO, P, RB, US, VEN). SURINAM. Sipaliwini, savannah, 30 Jan. 1965, Fraser 3 (US). BRAZIL. Amapa: Mun. de Macapa. Rio Macaeoari, Sao Benedito do Bra?o, 4 Jan. 1951, Frdes & Black 27195 (US). Bahia: natante na lagoa Pipiri, Rio Santa Clara, 17 Dec. 1987, Filgueiras 1295 (CEN, IBGE, K, SP). Minas Gerais: Lagoa Santa, in aquis stagnantibus, fluitans, 5 Apr. 1864, Warming s.n. (US). Mato Grosso: just S of the Rio Suia Missu Ferry, Volume 5, Number 3 1995 Davidse et al. Paspalum morichalense 237 ca. 40 km NW of base camp at 12°49'S, 51°46'W, edge of small lake in shallow water, grass with weak procum¬ bent rooting stems with ascending culms, 24 Nov. 1968, Harley 11213 (MO, UB, US). Mato Grosso do Sul: Mun. de Corumbti, Fazenda Santa Delfina, erva prostrada, pouco arraigada ao solo, frequente em va^ante seca, solo aren- oso, 3 Dec. 1978, Allem, Vails, Vieira & Comastri 2557 (CEN, MO); Mun. de Corumb£, Fazenda Sao Bento, erva prostrada, pouco arraigada ao solo, frequente em vazcante seca, 2 Nov. 1978, Allem, Vails, Vieira & Comastri 2529 (CEN, MO). Para: Gavinho, Maraj6 Island, Mar. 1920, Goeldi 284 (US). BOLIVIA. Rfo Guarap6, Santo Antonio de Manoelito, 25 Jun. 1952, Black & Cordeiro 52-15320 (US). Acknowledgments. We thank Vladimiro T. Du- das for the excellent illustration and Robert J. So- reng for checking the identity of the types at LE. Fieldwork by Davidse was made possible through the support of NSF grant GB40630. Filgueiras thanks CNPq (Brazilian Research Council), Brasi¬ lia, for a fellowship. Literature Cited Chase, A. 1929. The North American species of Paspa¬ lum. Contr. U.S. Natl. Herb. 28: 1-310, i-xvii. -. Ined. Paspalum of South America. Unpublished manuscript. Hitchcock and Chase Library, Botany De¬ partment, Smithsonian Institution, Washington, D.C. Doell, J. C. 1877. Paniceae. In: C. F. P. Martius, FI. Bras. 2(2): 33-358. Judziewicz, E. J. and collaborators. 1990. Poaceae (Gra- mineae). FI. Guiananas, Ser. A: Phanerogams 8: 1-727. Swallen, J. R. 1967. New species of Paspalum. Phyto- logia 14: 35B-389. A New Species of Brachynema (Olacaceae) from South America Rodrigo Duno de Stefano Fundacion Institute) Botanico de Venezuela, Apartado 2156, Caracas 1010-A, Venezuela Paul E. Berry Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A. Giovannina Orsini Veldsquez Universidad Central de Venezuela, Facultad de Ciencias, Escuela de Biologfa, Apartado 20513, Caracas 1040, Venezuela Abstract. A new species is described and illus¬ trated in the previously monotypic genus Brachy¬ nema (Olacaceae). Brachynema axillare is known from the base of Sierra de la Neblina in southern Venezuela and northern Brazil, as well as in Am¬ azonian Peru; it differs from B. ramiflorum Ben- tham in its axillary inflorescences, non-striped co¬ rolla tube, glabrous corolla lobes, and mostly 4-merous flowers. Resumen. Se describe una nueva especie en el genero Brachynema (Olacaceae), previamente con- siderado monotfpico. Brachynema axillare se cono- ce de la base de la Sierra de la Neblina en el sur de Venezuela y en el norte de Brasil, as! como en la Amazonia peruana. Difiere de B. ramiflorum en sus inflorescencias axilares, tubo de la corola no rayado, ldbulos de la corola glabros, y las flores mayormente 4-meras. In the most recent revision of neotropical Ola¬ caceae (Sleumer, 1984), Brachynema was consid¬ ered monotypic and endemic to central Amazonian Brazil. In a series of expeditions to the Sierra de la Neblina in southern Venezuela between 1984 and 1987 (Brewer-Carfas, 1988), numerous addi¬ tional collections of the genus were made and sub¬ sequently determined as Brachynema ramiflorum Bentham. However, more detailed study revealed that the Venezuelan specimens are significantly dif¬ ferent from this species and merit recognition as a distinct species, due to their axillary (vs. ramiflo- rous), often tetramerous (vs. strictly pentamerous) flowers, tricolpate instead of triporate pollen (Lob- reau-Callen, 1980), larger fruits, smaller calyx at anthesis, glabrous corolla lobes (vs. internally hairy), and the base of the corolla tube purple and lacking the characteristic “zebra” stripes of B. ram¬ iflorum. Brachynema axillare R. Duno & P. E. Berry, sp. nov. TYPE: Venezuela. Estado Amazonas: Dpto. Rio Negro, 0 to 2 km W of Cerro La Neblina Base Camp, along Rio Mawarinuma, 0°50'N, 66°10'W, 140 m, 7 Feb. 1984, Liesner 15707 (holotype, VEN; isotype, MO). Figure 1. Haec species Brachynemati ramifloro Bentham affinis, sed ab eo infloreseentia axillari non ramiflora, floribus plerumque tetrameris, corollae lobulis glabris etiam tubo basaliter purpureo, polline tricolpato differt. Sparsely branched shrub or small tree 3-8 m tall. Leaves chartaceous, simple, alternate, glabrous, es- tipulate; blade narrowly to broadly elliptic or ovate, 20-40 cm long, 5-20 cm wide, base rounded to cu- neate, apex long-acuminate to caudate, the tip often somewhat curved to one side, penninerved, with 9- 13 subopposite to alternate pairs of parallel second¬ ary veins looping near the margin to form a sub¬ marginal vein, prominent on the lower surface, sunk¬ en on the upper surface, tertiary veins conspicuous and running perpendicular to the secondary veins; margin shallowly to prominently sinuate-dentate, each tooth ending in a gland; petiole (1—)3—25 cm long, terete, noticeably thickened at both the base and the apex. Inflorescence an axillary fascicle or dense corymb of 10-20 flowers on a thickened, gla¬ brous to puberulous axis 5-10 mm long. Flowers actinomorphic, bisexual, 4- or 5-merous; pedicels 1— 2.5 mm long. Calyx cupular, greenish white when fresh, shallowly lobed, ca. 0.5 mm tall and 2 mm wide at anthesis. Corolla tube 16-25 mm long at anthesis, firm-fleshy when fresh, 2-4 mm wide at the base, glabrous to puberulous, uniformly violet-purple at the base, turning white toward the rim, the lobes white, glabrous, 4-6 mm long. Stamens 4 or 5, epi- petalous, included at the base of the tube, filaments ca. 1 mm long; anthers basifixed, ca. 3 mm long, narrowly pointed at the apex, longitudinally dehis- Novon 5: 238-240 1995 Volume 5, Number 3 1995 Duno de Stefano et al. Brachynema axillare 239 cent; pollen suboblate, polar axis 22.5 /am long, equatorial axis 28.7 /am long, circular or subtrian- gular in polar view, tricolpate, the colpi broadened, 14 /am long by 5 /am wide, exine very finely retic¬ ulate or psilate, 3 /am thick, without apertural thick¬ enings. Ovary subglobose, 3—5-locular; ovules 1 per locule, pendulous; stigma sessile, capitate. Drupe globose or depressedly globose with a pointed apical projection, 2—2.5 cm diam., green to purple-black when fresh, the calyx enlarged and persistent, sur¬ rounding the lower part of the fruit. Seed 1. subglob- ular, longitudinally striate, 1.5-2 cm diam. Ecology and distribution. Evergreen lowland forest on slopes and along gravelly and silty river- banks, 100-200 m elevation, known mainly from the base of Sierra de la Neblina in adjacent parts of Estado Amazonas, Brazil, and Estado Amazonas, 240 Novon Venezuela; also known in Amazonian Peru from Departamento Loreto near the confluence of the Rios Maranon and Ucayali. Brachynema is still of uncertain placement in the Olacaceae and is unique in the family in its glan¬ dular-dentate leaves and pulvinar thickenings at both the base and apex of the petiole (Sleumer, 1984). The leaves of B. axillare are very similar to those of B. ramiflorum, but the flowers and fruits are always borne near the ends of young branches in the leaf axils, rather than lower down on the trunk or on branches where the leaves have already dehisced. Neither the distinctive “zebra” striping of the lower part of the corolla of of B. ramiflorum nor the lines of hairs present on the inside of the corolla lobes have been observed in any specimen of B. axillare. Paratypes. VENEZUELA. Amazonas: 0-2 km E of Cerro La Neblina Base Camp, on Rfo Mawarinuma, 0°50'N, 66°10'W, 140 m, 7 Feb. 1984, Liesner 15707 (MO), 20 Feb. 1984, Liesner 16136 (MO, VEN); along Rfo Baria (= Rfo Mawarinuma) just upstream from Base Camp, SW side of Cerro de La Neblina, 0°49'50"N, 66°09'40"W, 140 m, 10 Apr. 1984, Gentry & Stein 46512 (MO), 11 Apr. 1984, Thomas & Plowman 3002 (MO, NY), 16 Apr. 1984, Gentry & Stein 46668 (MO), 4-5 July 1984, Davidse & Miller 27009 (MO), 27 Nov. 1984, Croat 59373 (MO), 4 Dec. 1984, Anderson 13480 (MICH, MO), 17 Feb. 1985, Nee 30915 (MO, VEN), 18 Feb. 1985, Boom & Weitzman 5894 (MO, NY), Stannard 459 (VEN), Stannard 460 (VEN); upper Rfo Barfa, ca. 0°55'N, 66°15'W, Dav¬ idse & Miller 26908 (MO); Cerro de la Neblina, lowland and slope forests, 170 m, 29 Jan. 1954, Maguire , Wurdack & Bunting 37401A (MO, NY). BRAZIL. Amazonas: Ser- ra da Neblina Base Camp on Cano Tucano, Rio Cauaburf, 125 m, 12 Nov. 1965, Maguire, Steyermark & Maguire 60160 (MO, NY). PERU. Loreto: Dtto. Nauta, Rfo Mar¬ anon, Quebrada de Sapira del caserfo Florida, ca. 8 km above Nauta, 13 June 1979, Rimachi 4465 (MO). Acknowledgments. We thank Roy Gereau for revising the Latin diagnosis. Literature Cited Brewer-Carfas, C. (Editor). 1988. Cerro de la Neblina. Resultados de la expedici6n 1983-1987. Fundacidn para el Desarrollo de las Ciencias Ffsicas, Matem4ticas y Naturales, Caracas, Venezuela. Lobreau-Callen, D. 1980. Caractferes compares du pollen des Icacinaceae et des Olacaceae. Adansonia II, 20(1): 29-80. Sleumer, H. O. 1984. Olacaceae. Flora Neotropica 38. New York Botanical Garden, New York. Dos Especies Nuevas del Genero Drosera (Droseraceae) en Venezuela y Otros Comentarios Taxonomicos Rodrigo Duno de Stefano Fundacion Instituto Botanico de Venezuela, Apartado 2156, Caracas 1010-A, Venezuela Alastair Culham Department of Botany, Reading University, Reading, Whiteknights, RO. Box 221, Reading RGG 2AS, United Kingdom Resumen. Se describen dos especies nuevas del genero Drosera provenientes de la Guayana Vene- zolana: Drosera hirticalyx y D. yutajensis. Las es¬ pecies D. colombiana A. Fernandez y D. panamen- sis M. D. Correa & A. S. Taylor son consideradas sin6nimos de D. cayennensis Sagot ex Diels, mien- tras que D. esmeraldae (Steyermark) Maguire & Wurdack es citada de Colombia y D. communis A. Saint-Hilaire es excluida de Venezuela. Abstract. Two new species of Drosera are de¬ scribed from the Venezuelan Guayana: D. hirticalyx and D. yutajensis. Drosera colombiana A. Fernan¬ dez and D. panamensis M. D. Correa & A. S. Taylor are considered nonspecific with D. cayennensis Sa¬ got ex Diels. Drosera esmeraldae (Steyermark) Ma¬ guire & Wurdack is reported for Colombia, and the occurrence of D. communis A. Saint-Hilaire in Ven¬ ezuela is disputed. Se describen dos especies nuevas del genero Drosera (Droseraceae), las cuales presentan hojas simples estipuladas, y gineceo con 3 estilos bipar- tidos desde la base, caracterfsticas dstas tfpicas del subg6nero Drosera secci6n Drosera. Drosera hirticalyx R. Duno & Culham sp. nov. TIPO: Venezuela. Amazonas: Dpto. Atabapo, Cerro Marahuaca-Huha, cumbre altiplanicie no arbolada, m&s o menos plana, con piedras expuestas, rfo abajo, 2580 m, 3°40'N, 65°30'0, 31 ene. 1982, Steyermark, Guariglia, Holmgren, Luteyn & Mori 125982 (holotipo, NY; isdtipo, VEN). Figura 1A-E. Speciei Drosera roraimae affinis, sed foliis brevioribus. Lamina basi gradatim attenuata; petiolus glaber vel subtus pilis raris, unicellularibus, praeditus. Inflorescentia 4-15 cm, (1—)2—6-flora; pedunculus glandulis sessilibus. Sem¬ ina elliptica, faveolata, 100—150 X 50-100 /cm. Planta herbdcea arrosetada. Rafces fibrosas. Tallo corto. Hojas espatuladas, altemas, simples, 5—12 mm de largo; estfpulas intrapeciolares, 2-4 mm de largo, 4-6 partidas, soldadas hasta 1.0-1.5 mm de su base; el peci'olo ensanchdndose hacia la ldmina y poco diferenciado, 4—6 mm de largo, glabro o con escasos pelos simples en el envds; lamina obovada o muy ampliamente obovada, 2.5-5.0 mm de largo y 2.0-4.0 mm de ancho, glabra o con pocos pelos simples en el enves; pelos glandulares (tentdculos) en el centra y margen de la Mmina, hasta 2 mm de largo. Escapo 16 2 por planta, 4—15 cm de largo. Inflorescencia monocdsica, terminal. Flores (1—)2—6; pedunculos 4—13 cm de largo, con pubescencia muy variable a lo largo del mismo: en el extremo proximal con glandulas sdsiles diminutas, pelos glandulares y a veces glabros o pelos simples, mientras que en el extremo terminal con gldndulas sesiles y pelos glan¬ dulares; brdcteas lanceoladas 2 mm de largo y 1 mm de ancho, con pelos glandulares; pedicelos 0.5-1.0 mm de largo, con pelos glandulares. S6palos 5, lan- ceolados, soldados hasta 1.5 mm de su base, 5.0 mm de largo y 2.0-2.2 mm de ancho, acuminados, con pelos glandulares, margen entero. P6talos 5, espa- tulados, 6 mm de largo y 2 mm de ancho. Ovario tricarpelar, 1-locular, rudimentos seminales nume- rosos; estilo 3, bipartido, 2 mm de largo; estigmas como un abanico. Fruto tipo capsula que se abre por 3 valvas. Semillas faveoladas, elfpticas, 100-150 gm de largo y 50-100 ptm de ancho, con abundan- tes granos de cera en la superficie. Drosera hirticalyx estd presente en la Guayana Venezolana (Cerro Marahuaca) y crece en aflora- mientos de arenisca entre 1200 y 2700 m sobre el nivel del mar en lugares humedos. Drosera hirticalyx es afin a D. roraimae (Klotzsch ex Diels) Maguire & J. R. Laundon; esta ultima especie dene las hojas nris laigas (10-20 mm), el pectolo muy pubescente y claramente diferenciado de la lamina, el numero de floras es mayor (4-20), los pelos glandu- losos en el caliz son mds pequeiios (comparar Fig. ID y F). Por uldmo la semilla es mas larga (400-600 pm) Novon 5: 241-245. 1995. 242 Novon Figura 1. A-E. Drosera hirticalyx R. Duno & Culham Steyermark & Holst 130701 (VEN). —A. Habito. —B. Hoja cara superior. —C. Hoja cara inferior. —D. CSliz. —E. Semilla. F, G. Drosera roraimae (Klotzsch ex Diels) Maguire & J. R. Laundon (Liesner 23990, VEN). —F. CAliz. —G. Semilla. y tiene una forma angosto-oblonga con los extremos de la testa inflados (comparar Fig. IE y G). Pardtipos. VENEZUELA. Amazonas: Dpto. Atabapo, open rocky plateau of Cerro Marahuaca above Salto los Monos on tributary of headwaters of rfo Iguapo, 3°37'N, 65°23’W, 2555 m, near stream, flowers white, 26 Feb. 1985, Liesner 17998 (MO); Cerro Marahuaca, summit on SE cor¬ ner, open rocky plateau and ravines, in small pool of water, 3°37'N, 65°21'W, 2700 m, 12 Oct. 1988, Liesner 24685 (MO); Cerro Marahuaca, summit on SE comer, open rocky plateau and ravines, forming cushions on edge of pond 3°37'N, 65°21'W, 2700 m, 12 Oct. 1988, Liesner 24689 (MO); Cerro Marahuaca, summit, SW side of center, open rocky plateau and ravines, on slope, buds pink, 3°39'N, 65°26’W, 2660 m, 24 Oct. 1988, Liesner 25286 (MO); Cerro Marahuaca, summit on undulating plateau with island of shrubs and Marahuacaeae with stream branch leading to Salto de Monos and southeastern edge to Cano Iguapo, S and SE of summit camp, on moist ground, growing with 130700, leaves dense, short, spreading, 337 3. 65°23 W, 2520-2650 m, 26-27 Feb. 1985, Steyermark & Holst 130701 (MO, VEN); Cerro Marahuaca, summit on undu¬ lating plateau with island of shrubs and Marahuacaeae with stream branch leading to Salto de Monos and southeastern edge to Cano Iguapo, S and SE of summit camp, petals white, 3°37'N, 65°23'W, 2520-2650 m, 26-27 Feb. 1985, Steyermark & Holst 130753 (MO, VEN). Drosera yutajensis R. Duno & Culham, sp. nov. TIPO: Venezuela. Amazonas; Dpto. Atures, val¬ ley of rfo Coro-Coro, W of Serrania de \utaje, sandstone plateau W of river valley, along trick¬ ling stream in slightly depressed area, flowing across exposed sandstone, 1100 m, 5 41 N, 66°09’W, Feb. 1987, Holst & Liesner 3277 (holo- tipo, MO; isotipos, RNG, VEN). Figura 2. Volume 5, Number 3 1995 Duno de Stefano & Culham Drosera de Venezuela 243 Figura 2. Drosera yutajensis R. Duno & Culham (Huber 12636, VEN). -A. Hibito. -B. Hoja cara superior. -C. Hoja cara inferior. —D. C4liz. —E. Semilla. Speciebus Drosera villosa atque D. arenicola affinis. fo¬ lia lineari-spathulata 10—20 mm, distincte petiolata, pe- tiolis dense pubescentibus. Inflorescentia 20—25 cm, 16- 30-flora; pedunculus glandulis raris praeditus. Petala 5, spathulata, 4.5-5 X 2.5 mm; semina faveolata, ovoidea, 400-430 X 230-250 pm. Planta herb&cea arrosetada. Rafces fibrosas. Ta- Ho corto. Hojas linear-espatuladas, 10-20 mm de largo; estipulas intrapeciolares, 5-6 mm de largo, 5-6 partidas, soldadas hasta 1 mm de su base; pecfolo ensanch&ndose hacia la lamina pero bien diferenciado, 5-7 mm de largo y 1-2 mm de ancho, pelos simples en el enves; laminas oblanceoladas 244 Novon u obovadas, 6-12 mm de largo y 2-4 mm de ancho, pelos simples en el env6s, pelos glandulares (ten- taculos) en centra y margen de la lamina, hasta 4.0 mm de largo. Escapos 1-3 por planta, 20-25 cm de largo. Inflorescencia monoeasica, terminal. Flo¬ res 16-30; pedunculo 10—12 cm de largo, erecto con pelos glandulares; braeteas lineares, 2 mm de largo y 0.1 mm de ancho, con pelos glandulares escasos; pedicelos 2-3 mm de largo, con pelos glandulares. Sepalos 5, ovados, 3—5 mm de largo y 1 mm de ancho, con pelos glandulares en el env6s, margen entero. Petalos 5, espatulados, 4.5—5.0 mm de largo y 2.5 mm de ancho. Ovario tricarpelar, 1- locular, rudimentos seminales numerosos; estilos 3, bipartidos, 2 mm de largo; estigmas como un aban- ico. Fruto tipo cApsula que se abre por 3 valvas. Semillas faveoladas, ovadas, 400-430 pm de largo y 230-250 pm de ancho. Drosera yutajensis es solo conocida de tres co- lecciones provenientes del cerro Yutaje, en la cuen- ca del rio Manapiare. Crece en afloramientos de arenisca entre 600 y 1800 m sobre el nivel del mar, en lugares humedos. Drosera yutajensis es afm a D. villosa A. Saint- Hilaire (presente solo en Brasil), ambas con hojas linear-espatuladas, pero mAs largas en la especie brasilera (2-5 cm), donde adernas el pecfolo apenas se diferencia de la ldmina; la semilla en este ultimo caso es tambiAn faveolada pero angostamente ob- longa y mucho rods larga (600-700 pm). Tambien es afln a D. arenicola Steyermark, las hojas con la misma forma y pubescencia pero mds pequefias en D. arenicola (8—12 mm); en esta ultima el pedun¬ culo es mAs corto (1-8 cm), el numero de floras menor (1-10) y la semilla es casi elfptica. Pardtipos. VENEZUELA. Amazonas: Dpto. Atures, Rto Coro-Coro, W of Serrania de Yutajg, 8.5 km N of settlement of Yutaj6, small tributary, E of river, semi-ex¬ posed creek running over igneous fault horded by sand¬ stone, vegetation low, in sandy areas; 650 m, 5°42'N, 66°08’W, 25 Feb. 1987, Holst & Liesner 3179 (MO); Cum- bre del cerro Yutaj6, sector oriental de la Serrania Yutaje, herbazal inundable en planicie aluvial, bosquetes de pal- mas al pie de las vertientes, arbustal y herbazal denso sobre las vertientes rocosas, ocasionalmente en sitios abiertos del arbustal y herbazal, 5°45'N, 66°03'0, 1800 m, 21 mar. 1989, Huber 12636 (MYF, PMA, VEN). TambiAn fueron estudiadas las especies D. ca¬ yennensis Sagot ex Diels (1906), D. colombiana A. FemAndez (1965) y D. panamensis M. D. Correa & A. S. Taylor (1976). Estas especies comparten im- portantes atributos como son: forma, tamano y pu¬ bescencia de las hojas; tipo (pelos simples) y den- sidad de la pubescencia en el pedunculo y sepalos; la condici6n de sepalos libras desde la base y el tipo de dehiscencia de la capsula. A pesar de que las semillas encontradas en todas las muestras eran inmaduras, presentaban apariencias muy similares. Todas estas similitudes sugieren que nos encontra- mos en presencia de una sola especie. Previamente Brummer-Dinger (1955) consider6 D. sanariapoana Steyermark coespeclfica a D. cayennensis. Drosera cayennensis Sagot ex Diels, Pflanzenfam. 4(112): 86. 1906. TIPO: French Guiana, Sagot s.n. (P). Drosera sanariapoana Steyermark, Fieldiana Bot. 28(2): 243. 1952. TIPO: Venezuela. Amazonas: collected on wet savanna, vicinity of Sanariapo, near Rfo Sana- riapo, tributary of Rfo Orinoco, 100 m, “petal rose,” 8 Sep. 1944, Steyermark 58472 (hol6tipo, NY). Drosera colombiana A. Fernandez. Caldasia 9(43): 226, fig. 1 1965. Sin. nov. TIPO: Colombia. Meta: Llanos de San Martin, en sabana, “pelos finos rojos sobre fonde verde, floras blancas,” jul. 1948, Dryander 3019 (holntipo. COL; isotipo. US). Drosera panamensis M. D. Correa & A. S. Taylor, Ann. Missouri Bot. Gard. 63: 390. 1976. Sin. nov. TIPO: Panam4. La Yeguada, altos de Baltazar y el Veladero, 650 m, 4 ago. 1976, Correa et al. 2215 (holdtipo, PMA; isotipo-. CHR no visto, COL, DUKE no visto, F no visto, K, MEXU no visto, MO no visto, NSW no visto, U no visto, US no visto). Material adicional estudiado. GUAYANA FRANCESA: Leprieuer 145 (parAtipo de D. cayennensis, M), Spruce 1068 (parAtipo de D. cayennensis, M). VENEZUELA. Amazonas: Guanchez 1083 (MO, VEN). Drosera esmeraldae (Steyermark) Maguire & Wurdack fue conocida anteriormente solo de las tierras bajas de Guayana (Sabana de Esmeralda, Estado Amazonas). Material coleccionado reciente- mente en Colombia (Vaupes y Amazonas) pertenece a este tax6n. Drosera esmeraldae (Steyermark) Maguire & Wur¬ dack, Mem. New York Bot. Gard. 9(3): 335. 1957. Drosera tenella var. esmeraldae Steyer¬ mark, Fieldiana Bot., 28(2): 244. 1952. TIPO: Venezuela. Amazonas: wet places in savanna, between Esmeralda savanna and southeastern base of Cerro Duida, 200 m, 22 Aug. 1944, Steyermark 60920, (holdtipo, F; isdtipo, NY). Material adicional estudiado. COLOMBIA. Vaupes: Schultes 22610 (COL). Amazonas: Idrobo 8970 (COL), Sastre & Reichel 5209 (COL). VENEZUELA. Amazonas: Clark 6475 (NY), Holst & Liesner 3275 (MO), Liesner & Holst 21800 (MO), Huber 2697 (MO), Maguire et al. 29616 (BM, FM, NY), Stergios & Aymard 4277 (PORT). En relacion con Drosera communis A. Saint-Hi- laire, Maguire & Wurdack (1957) indican la pre¬ sencia de esta especie para Venezuela; sin embargo no incluyen lista de exsiccatas. Los pocos espect- Volume 5, Number 3 1995 Duno de Stefano & Culham Drosera de Venezuela 245 raenes recolectados posteriormente en Venezuela e identificados por J. A. Steyermark: (Krai 72009 (VEN), Steyermark 128551 (MO, VEN) y Steyer¬ mark 128978-A (MO, VEN)) difieren claramente del material tipo de esta especie: Saint-Hilaire 695 (holotipo, P); estas muestras fueron identificadas como D. roraimae (Klotzsch ex Diels) Maguire & J. R. Laundon. Por lo tanto consideramos que esta espeeie no esta presente en Venezuela ni Colombia y su Area de distribucion queda limitada al hemis- ferio Sur (Brasil meridional y Uruguay). Drosera communis A. Saint-Hilaire, PI. Usual. Bras. XV plate 15. 1824. TIPO: Brasil. Minas Ger¬ ais: Saint Hilaire 695 (holotipo. P). Drosera parvifolia A. Saint-Hilaire, Hist. PL Remarq. Bre- sil, p. 263, t. 25. 1824. TIPO: Brasil. Minas Gerais: Saint Hilaire D377 (hol6tipo, P). Drosera communis var. pauciflora Eichler en Martius, FI. Bras., 14: 394. 1872. TIPO: B. Material adicional estudiado. BRASIL. Minas Ge¬ rais: Glaziou 10258 (P). Rio de Janeiro: Glaziou 14483 (P). Sao Paulo: Saint Hilaire D 722 (P), Gardner 3573 (K), Luetzelburg 21485 (M), Luetzelburg 21509 (M). PA¬ RAGUAY: Hassler 5271 (K, P), Hassler 5634 (K, P). Agradecimientos. Expresamos nuestro mds sin¬ cere agradecimiento a los curadores de los siguien- tes herbarios: B, BM. COL. K, F. M. MA. MO. MYF, P. PORT. US. Asf como a J. FemAndez-Casas (MA) por sus comentarios y descripciones en latin. tam- bi£n a Otto Huber, Paul Berry y Bruce Holst por la correcion del manuscrito. Literatura Citada Brummer-Dinger, C. H. 1955. Notes on Guiana Drosera- ceae. Acta Bot. Neerl. 4(1): 136-138. Correa, M. D. & A. S. Taylor B. 1976. Droseraceae. Ann. Missouri Bot. Gard. 63: 389—392. Diels, L. 1906. Droseraceae. En: Engler, Das Pflanzen- reich 26 (4): 1-136. Femfindez-PArez, A. 1965. Plantas Insectfvoras, II Dro¬ seraceae de Colombia. Caldasia 11(43): 219-232. Maguire, B. & J. J. Wurdack. 1957. The Botany of the Guayana Highland—Part II. Mem. New York Bot. Gard. 9(3): 331-336. Palisota flagelliflora (Commelinaceae), a New Species from Cameroon with a Unique Habit Robert B. Faden Department of Botany, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. Abstract. Palisota flagelliflora Faden (Comme¬ linaceae), a rosette species, is unique in Palisota in bearing its inflorescences on axillary, flagelliform shoots that lie on the forest floor. These shoots con¬ tinue to grow sympodially and give rise to addi¬ tional inflorescences or viviparously produce a new rosette. Other rare or unique features include: in¬ florescences composed of single cincinni, long ped¬ icels, often vertical flowers, yellow staminode fila¬ ment hairs, bearded filament and connective in the unpaired stamen, and bearded style. The genus Palisota Reichenbach f. ex Endlicher, with ca. 18 species (Faden & Hunt, 1991), is the largest African genus of Commelinaceae occurring predominantly in forests (Faden, 1983). Its center of diversity is Central Africa, especially Cameroon and Gabon. Until recently, all described species were either acaulescent rosette plants or herba¬ ceous shrubs, and all had terminal or terminal and axillary, thyrsiform inflorescences. Recently de¬ scribed species (Brenan, 1984) have habits and in¬ florescences that depart from these types, including a decumbent habit and strictly axillary inflores¬ cences. A unique habit, rosettes producing flagel¬ liform flowering shoots, is described below in the new species P. flagelliflora. Palisota flagelliflora Faden, sp. nov. TYPE: Cam¬ eroon. South Prov. [South-Central Prov.]: Kri- bi-Lolodorf road 3.5 km along the road to Edca that branches off to the N at km 18 (just past Bissiang), 200 m on a track that branches off the Bissiang-Edea road, 3°02'N, 10°02'E, ex¬ ploited forest with much of the structure and understory vegetation remaining, nearby pure stand of Cecropia ( Musanga ) cecropioides along the Bissiang-Edea road, 29 Jan. 1986, Faden & Mbamba 86/50 (holotype, US sheet no. 3264040; isotypes, B, BR, C, COI, G, GC, HBG, IEC, K(3), LISC, MA, MO, P, S, UPS, US(4), WAG, YA(2)). Field-collected inflores¬ cences and fruits, preserved in FAA (US vial B470), are part of the type collection. Figures 1 , 2 . Herbae perennes foliis rosulatis, 14-50 cm longis, 3.5— 10(—11) cm latis, lamina elliptica vel oblongo-elliptica ad oblanceolatam vel obovatam, 10-40 cm longa; inflores- centiis terminalibus et bracteas oppositis in surculis ax- illaribus, flagelliformibus (1.5—)2S—100(—175) cm longis, cincinnis solitariis et contractis, bracteis binatis subtentis compositis; floribus bisexualibus et masculis, stamine im- pari apice filamenti et in connective barbato, sacculo an- therae in flore bisexuali indehiscenti, in flore masculo de- hiscenti, ovulis biseriatis; fructibus triangulari-ellipsoideis ad triangulari-ovoideos, apice acuto ad rostratum, 1.4—2.3 cm longis, 0.9—1.2 cm latis, coccineis vel rubiginosis, dense pilosis; seminibus uni- vel biseriatis. Perennial with a rhizome to 15 cm long, fre¬ quently producing offsets, leaves all or mainly in rosettes, petiolate, 14—50 cm long, 3.5-10(-ll) cm wide, petiole 4—19 cm long, appressed pubescent, lamina elliptic or oblong-elliptic to oblanceolate or obovate, 10-40 cm long, apex abruptly acute or acuminate, base cuneate to attenuate, surfaces dis- colorous, adaxial green, glabrous or appressed whitish sericeous and glabrescent (this pubescence ± persistent in cultivation), abaxial persistently ap¬ pressed whitish sericeous. Inflorescences terminal, usually becoming bract-opposed, pedunculate, borne on axillary, leafless, bracteate, flagelliform shoots (1.5-)25-100(-175) cm long, at first ascend¬ ing, soon becoming prostrate and creeping through the leaf litter, rarely obviously branched, rarely rooting, many-noded, extending sympodially from the axil of the bract subtending the peduncle, the flagelliform shoot in this manner often producing a succession of terminal/bract-opposed inflores¬ cences, often viviparously producing a new rosette distally; intemodes appressed sericeous, nodes cov¬ ered by persistent maroon bracts 1.5-4 cm long, appressed whitish sericeous; peduncle 1-8.5 cm long, usually appressed sericeous, sometimes densely lanate; each inflorescence consisting ol a single contracted, several-flowered cincinnus sub¬ tended by a pair of maroon bracts 1.5-2.5 cm long, similar in shape, color, and pubescence to those of the flagelliform shoot. Flowers bisexual and male, predominantly vertical or with the unpaired (outer) sepal uppermost, occasionally horizontal, ca. lb- 18 mm wide, odorless; pedicels erect in flower, be- Novon 5: 246-251. 1995. Volume 5, Number 3 1995 Faden Palisota flagelliflora 247 Figure 1. Palisota flagelliflora Faden. —A. Habit, showing flagelliform flowering shoot. —B. Inflorescence terminal on flagelliform flowering shoot, the shoot growth continuing sympodially from the axil of the bract at the base of the peduncle. —C. Unpaired stamen, bisexual flower, ventral view. —D. Unpaired stamen, bisexual flower, dorsal view. - E. Paired stamen, dorsal view. —F. Paired staminode. —G. Staminode filament hairs. —H. Gynoecium, lateral view. —I. Fruit, showing contorted pedicel. —j. Seeds, dorsal and ventral views. All from Faden & Mbamba 86/50. 248 Novon f r, re 2 ‘ Pa l“ 0 ^ l fl a Z dl f ora Fade »- —A. Male flower, front view. —B. Male flower, side view. —C. Bisexual flower, front View. —D. Bisexual flower, side view. All from Faden & Mamba 86/50. coming spirally contorted in fruit, 1.5-3.5 cm long, white, sometimes tinged with reddish purple, pi¬ lose; sepals subequal, boat-shaped, lanceolate-ob¬ long to oblong-elliptic, (3-)6-7.5 mm long, 2-3 mm wide, reddish purple outside, the color shining through, densely hirsute with reddish hairs forming an apical tuft, usually intermixed with finer, shorter, white hairs; petals subequal, outer slightly broader than the others, elliptic, (4—)6 mm long, (2—)3—3.5 mm wide, white, inner petals oblong to spathulate- Volume 5, Number 3 1995 Faden Palisota flagelliflora 249 oblong, (4—)6.5—7 mm long, (1.5-)2-2.5 mm wide, usually white, sometimes maroon with white mar¬ gins; unpaired staminode lacking or with a filament 1.5—2 mm long and bearded subapically with ap¬ parently moniliform, white-based yellow hairs; paired staminodes with filaments 2—3 mm long, white, bearded in the distal half with hairs like those of the outer staminode; paired stamens with filaments (2-)3—3.5 mm long, glabrous, white, an¬ thers broadly elliptic to oblong-elliptic or ovate-el¬ liptic, (1—)1.5—1.75 mm long, 0.7—1 mm wide, de¬ hiscence extrorse, connective yellow, glabrous, pollen yellow; unpaired stamen dimorphic, in bi¬ sexual flower slightly shorter to slightly longer than the paired stamens, filament (1.5—)2.5-3.5 mm long, strongly declinate, white, usually densely bearded with minute yellow hairs just below the anther, anther broadly ovoid, ca. (1—)1.8 mm long and wide, yellow, pollen sacs C-shaped, indehis- cent, connective usually densely bearded with min¬ ute yellow hairs; in male flower unpaired stamen distinctly longer than the paired, more central in the flower, usually not declinate, filament ca. 5 mm long, white, bearded at apex, as in bisexual flower, anther similar to that of bisexual flower, including densely bearded connective, but smaller (ca. 1.4 mm long and wide) and pollen sacs dehiscent, in- trorse, with pollen concolorous with that of the paired stamens; gynoecium reduced and apparently abortive in the male flower, in the bisexual: ovary ovoid, deeply 3-lobed, ca. 2-2.5 mm long, densely pubescent with forward-directed, white or reddish hairs, locules 5-7-ovulate, ovules biseriate, style exceeding the stamens, 3.2—4 mm long, straight or declinate, yellow, sparsely bearded basally with ap- pressed hairs like those of the ovary, stigma deeply 3-lobed, yellow with a central brownish area. Fruits triangular-ellipsoid to triangular-ovoid berries, acute to beaked at the apex, 1.4—2.3 cm long, 0.9— 1.2 cm wide, scarlet or reddish brown, densely red¬ dish pilose. Seeds uni- or biseriate, 1—4 per locule, ovate-triangular in outline, ca. 4—5 mm diam. Habitat. Secondary and exploited forest, 10— 520 m. Flowering. January, February, July, September. In cultivation the flowers open at about 0940 and begin to fade at 1230-1330. Distribution. Endemic to Cameroon. Palisota flagelliflora is unique in Palisota be¬ cause of its flagelliform shoots, bract-opposed inflo¬ rescences, and vivipary. It may further be distin¬ guished by its inflorescences composed of a single cincinnus, long pedicels, large, often vertical flow¬ ers, yellow staminode filament hairs, bearded sta¬ men filament and connective in the unpaired sta¬ men, and bearded style. It also has the earliest flower opening time for any Palisota species thus far recorded or observed. In the above description the dimensions of the floral parts were measured from preserved flowers of a cultivated plant of Faden & Mbamba 86/50. The much smaller dimensions for some parts, listed parenthetically, came from a pressed flower on a small plant of Leeuivenberg 8296 (WAG). No field- preserved flowers were available. The larger flowers were the largest observed in Palisota. The staminode filament hairs were described above as “apparently moniliform” because of the beadlike appearance of the cells at low magnifica¬ tion. However, higher magnification (Fig. 1G) re¬ vealed that the cells of the hairs were individually dumbbell-shaped, not spherical or ellipsoid, as in typical moniliform hairs. The nature of the flagelliform shoots requires some comment. Morphologically they could be con¬ sidered one of the following: long, axillary inflores¬ cences; axillary shoots bearing terminal inflores¬ cences; inflorescence-shoots; or stolons bearing inflorescences. Their actual structure is clear, so the problem is more semantic than morphological. Forman (1962) tried to distinguish between an ax¬ illary inflorescence and a reduced axillary shoot bearing a terminal inflorescence, which he termed an inflorescence-shoot. Faden (1991: 11) found the difference between the two, which was based on the absence (inflorescence) or presence (inflorescence- shoot) of a basal prophyll, not very useful because the whole range from leafy lateral shoots with a terminal inflorescence to reduced lateral shoots with only bracts (inflorescence-shoot) could be found within a single plant in some Aneilema spe¬ cies. The axillary inflorescence might have evolved by further reduction, namely loss of the prophyll. Many forest Commelinaceae produce stolons: ax¬ illary, elongate, horizontal shoots with only leaf sheaths basally (Faden, 1988). Distally stolons pro¬ duce new vegetative shoots, not inflorescences. The inflorescence-terminated shoots of Palisota flagel¬ liflora are dissimilar because they bear only bracts and they are primarily for sexual, not vegetative, reproduction. When they viviparously produce a new plant, they are more like the reproductive shoots of the epiphytic Belosynapsis vivipara (Dal- zell) C. E. C. Fischer than like stolons of other terrestrial species. Because they are unique in Pal¬ isota, and perhaps within the family, I have used the term “flagelliform shoots” to describe them. The regularly biseriate ovules were a surprising find in view of the fact that the seeds may be uni- 250 Novon seriate. The alignment of the seeds in the fruit seems to be a function, at least partly, of seed num¬ ber per locule. When three or fewer the seeds ap¬ pear uniseriate, when four or more, biseriate. Only two fruits were available for dissection, so which of these arrangements predominates and whether more seeds are ever present is unknown. One of the two sheets of Leeuwenberg 8296 (WAG) bears two small plants each with a short inflorescence (only 1.5 cm long in one case). Had these been the sole plants collected, treating them as Palisota flagelliflora would have had to have been based largely on the very long pedicels and bearded connective of the unpaired stamen. The second sheet, as well as specimens of this collec¬ tion seen elsewhere, clearly demonstrate that the small plants are merely young P. flagelliflora with short inflorescences. Faden & Mbamba 86/47 was noted in the field as differing from Faden & Mbamba 86/50 in not forming dense stands, in being sterile (it had pro¬ duced new rosettes at the ends of axillary, flagel- liform shoots), less robust, with a reddish petiole and smaller, less discolorous lamina more abruptly narrowed into the petiole, and with smaller bracts on the flagelliform shoots. Although exact chromo¬ some counts were not obtained from either popu¬ lation, 2 n = ca. 40 (approximately diploid) was counted for Faden & Mbamba 86/47 and 2n = ca. 60 for Faden & Mbamba 86/50 (approximately trip- loid). Faden & Mbamba 86/50 was selected as the type, despite its being possibly triploid, because abundant reproductive material was present in the field, including ripe fruits and buds, and that pop¬ ulation also flowered in cultivation, whereas Faden & Mbamba 86/47 did not flower before it died. The inclusion of Bos 3877 in Palisota flagelliflo¬ ra was questionable. Although it came from very close to the type locality, it differed most obviously from the other collections by its broader leaves, more robust flowering shoots that perhaps were not lying in the leaf litter, peduncles completely cov¬ ered by lanate, brownish hairs, and by lacking ster¬ ile bracts between the bracts at the bases of the peduncles. Dissection of flower buds, nearly all partly eaten or bored, revealed that all three stamen filaments and connectives were glabrous, differing from all other collections of P. flagelliflora. In the absence of a larger number of collections, Bos 3877 is provisionally included in P. flagelliflora. Bos 3877 approached Palisota lagopus Mild- braed in robustness and peduncle pubescence, dif¬ fering most obviously by its longer, axillary, sym- podial flowering shoot with several inflorescences and longer, spirally contorted fruiting pedicels. Its somewhat intermediacy between this species and P. flagelliflora suggested a possible hybrid origin, but it could just as readily represent an undescribed species or a form of P. flagelliflora. Palisota flagelliflora is an understory species that has been collected in secondary and exploited primary forest. It should also occur in undisturbed primary forest. In Littoral Province the primary veg¬ etation at the locality for this species would be “At¬ lantic forest of the northwestern type with Caesal- piniaceae rare” (Letouzey, 1985). At the localities east of Kribi the primary forest type would be “At¬ lantic littoral forest with Caesalpiniaceae relatively rare, with Saccoglottis gabonensis ” (Letouzey, 1985). In the field Palisota flagelliflora may form dense stands because of its ability to reproduce vegeta- tively. That tendency, along with the small size of the rosettes, compared to other species, allows it to be spotted from a distance. Closer up, the flagelli¬ form shoots and vivipary are very distinctive, as is the presence of hairs on the connective and fila¬ ment apex of the unpaired stamen. Palisota flagelliflora appears related to the red- fruited species of Palisota because of its fruit color, hairy fruits (glabrous in some species), persistent faded flowers, and long, spirally contorted fruiting pedicels. Most of those species, such as P. barteri Hooker and P. mannii C. B. Clarke, have uniseriate seeds and would thereby belong to section Monos- tichos C. B. Clarke. Palisota flagelliflora, however, with its biseriate ovules and sometimes biseriate seeds, would have to be placed in section Palisota (synonym: section Distichos C. B. Clarke, 1881: 133) with the blue- and black-fruited species such as P. ambigua (P. Beauvois) C. B. Clarke and P. hirsuta (Thunberg) K. Schumann. This apparent anomaly may be explained either by Clarke’s sec¬ tions being unnatural, which well may be the case, or by seed arrangement not always reflecting ovule arrangement, about which we have insufficient data. Palisota flagelliflora is probably most closely re¬ lated to P. satabiei Brenan and P. bogneri Brenan, small, red-fruited species with axillary inflores¬ cences, biseriate ovules, and, like P. flagelliflora, a restricted distribution in the species-rich coastal forests of Cameroon and Gabon. Palisota flagelli- fora uniquely shares with P. satabiei the features of the inflorescence consisting of a single cincinnus, yellow staminode filament hairs, bearded anther connective, and bearded style. Preserved fruits of P. satabiei (Satabid & Letouzey 384, YA) showed partly biseriate seeds in one locule, as in P. fla¬ gelliflora. Volume 5, Number 3 1995 Faden Patisota flagelliflora 251 Palisota flagelliflora, P. satabiei, and P. bogneri extend the range of habit and inflorescence types previously known in Palisota : decumbent habit (P. satabiei), vivipary, strictly axillary inflorescences, inflorescences composed of single cincinni, and fla- gelliform flowering shoots. My research on the Commelinaceae of Cameroon has revealed addi¬ tional habit types in the genus: scandent (to 15 m in length) in P. thollonii Hua, and decumbent with a terminal thyrse in an undescribed species from Cameroon and Gabon. The diversity of habit types now known within Palisota rivals that in the much larger neotropical genus Dichorisandra, which is the ecological counterpart of Palisota in South America. It should be noted that the bisexual flower de¬ picted in Figure 2 has the lower (unpaired) stamen longer and less declinate than in the most extreme cases. Insufficient flowers have been examined to generalize about the “typical” curvature and length of the filament. Paratypes. CAMEROON. Littoral Prov.: Foret du Bukaka, 3 km E of kbcnr. a village on km 11 Nkongsam- ba-Loum road, 4°50’N, 9°54'E, alt. 520 m, 8 Sep. 1971, Leeuwenberg 8296 (US, WAG—2 sheets, YA). South Prov.: ca. 18 km from Kribi, Lolodorf road, 3°00'N, 10°02'E, 5 Feb. 1969, Bos 3877 (WAG, YA); 4 km N of km 20 Kribi-Lolodorf, 3°02'N, 10°03'E, 4 July 1970, Bos 7014 (US, WAG—3 sheets, YA); Kribi-Lolodorf road, km 20, end of track ca. 2 km N of the road, ca. 3°01'N, 10°03'E, alt. 10 m, 29 Jan. 1986, Faden & Mbamba 86/ 47 (P, US, YA). Acknowledgments. I thank B. Satabid for facil¬ itating my fieldwork in Cameroon, Alice Tangerini for the line drawings, and my wife, Audrey J. Fa¬ den, and other volunteers for taking care of my re¬ search plants for many years. Literature Cited Brenan, J. P. M. 1984. Two new species of Palisota (Com¬ melinaceae) from West Africa. Kew Bull. 39: 829-832. Clarke, C. B. 1881. Commelinaceae. Pp. 113—324 in A. & C. de Candolle, Monographiae Phanerogamarum 3. Sumptibus G. Masson, Paris. Faden, R. B. 1983. Phytogeography of African Comme¬ linaceae. Bothalia 14: 553—557. -. 1988. Vegetative and reproductive features of forest and nonforest genera of African Commelinaceae. Pp. 521—526 in: P. Goldblatt & P. P. Lowry II (editors). Modem Systematic Studies in African Botany. Monogr. Syst. Bot. Missouri Bot. Gard. 25. -. 1991. The morphology and taxonomy of Aneile- ma R. Brown (Commelinaceae). Smithsonian Contr. Bot. 76: 1-166. -& D. R. Hunt. 1991. The classification of the Commelinaceae. Taxon 40: 19—31. Forman, L. L. 1962. Aetheolirion, a new genus of Com¬ melinaceae from Thailand, with notes on allied genera. Kew Bull. 16: 209-221. Letouzey, R. 1985. Notice de la carte phytog£ographique du Cameroun: 1:500,000. Institut de la Recherche Agronomique (Herbier National), Yaounde, and Institut de la Carte Internationale de la Vigdtation, Toulouse. Typification of Pinus apulcensis Lindley (Pinaceae), a Misinterpreted Name for a Latin American Pine Alps Farjon Oxford Forestry Institute, Department of Plant Sciences, University of Oxford, South Parks Road, Oxford 0X1 3RB. England, U.K. Abstract. The Mexican taxon Pinus apulcensis Lindley is lectotypified with an ovuliferous cone found among carpological material in the herbar¬ ium of the Natural History Museum in Vienna (W), which was sent to Endlicher by Lindley. This cone confirms treatments of that name in publications on Mexican pines up to 1945—1948, when Marti¬ nez interpreted it in a different sense, using a specimen he had received from the putative type locality. The resulting confusion has been cor¬ rected, and a revised synonymy of Lindley’s taxon is presented. During work being carried out to complete the monographic study of pines for Flora Neotropica started by Brian T. Styles, a problem was encoun¬ tered regarding the current interpretation of one of John Lindley’s species from Mexico, Pinus apulcen¬ sis. This interpretation, commonly accepted since Martinez (1945, 1948), and followed by Stead & Styles (1984) in their revision of the “ pseudostro- bus" group, was due to the lack of a type specimen, and was based on what may be described as the topotype method: the assumption that whatever pine grows today at the type locality must be Lin¬ dley’s taxon. Lindley (1839) published two new species of pine based on collections made by C. T. Hartweg in Mexico in 1838: Pinus pseudostrobus from An- gangueo (“Anganguco,” Michoacdn) and P. apul¬ censis from “ravines near Apulco” (Hidalgo). (The confusion of Apulco with Acapulco seems to have originated with Endlicher (1847: 154).) The origi¬ nal descriptions of both are concise. Apart from short leaves and glaucous shoots, P. apulcensis is said to differ from P. pseudostrobus mainly in its cones, “covered closely with pyramidal elevations, which are sometimes prolonged and contracted in the middle, especially those near the points of the cones.” Lindley’s herbarium is largely at CGE; a few types are also at BM and K (Stafleu & Cowan, 1981). A catalog of conifers (Diimmer, 1913) lists what was in the Lindley herbarium (CGE) early in Novon 5: 9r>9-95f> 1QQ5 the twentieth century. Neither of the two taxa with which we are concerned here is mentioned in that catalog. Searches for original material of Pinus apulcensis in the above-mentioned herbaria were unsuccessful (Stead & Styles, 1984). Loudon (1842: 1014—1015) translated Lindley’s diagnosis and repeated his English description of Pinus apulcensis but stressed even more the “backs of the scales. . .sometimes prolonged into a hook, particularly those nearest the base and the point.” He stated that the species was introduced in 1839, “by cones sent home by Hartweg, from which many plants have been raised.” And finally, he presented figures 1899 and 1900, “from specimens sent home by Hartweg.” Figure 1899 is a natural-size (12 cm long) line drawing of a closed (green?) cone clearly showing what is meant by “prolonged and contract¬ ed pyramidal elevations” (Lindley, 1839) and “scales prolonged into a hook” (Loudon, 1842). There can be little doubt that this drawing repre¬ sents both Hartweg’s original collection and Lin¬ dley’s P. apulcensis (Fig. 1). Shaw (1909) reduced Pinus apulcensis Lindley to P. pseudostrobus var. apulcensis, citing the basio- nym, Loudon’s (1842) figure 1899, Endlicher (1847: 153), and three Mexican collections: Nelson 985 (US) and Nelson 2539 (US), both from Oaxaca, and Pringle 8788 (MEXU, US) from Mexico, as his references. In his plate 12, figures 8, 7, and 6 rep¬ resent cones of these collections respectively. Like Lindley and Loudon, Shaw emphasized the “pe¬ culiar development of the apophyses of the cone, which may attain remarkable prominence.” These three collections present considerable variation in development of the apophysis, with Nelson 2539 closely resembling figure 1899 in Loudon (1842), and Nelson 985 attaining much longer apophysal elongations and very strongly developed umbos. The present study, based on many more collections, demonstrates that Nelson 2539 and Nelson 985 rep¬ resent the two extremes of a cline in these char¬ acters. Pringle 8788 has strongly developed umbos, but the apophyses of the scales are flat and the Volume 5, Number 3 1995 Farjon Pinus apulcensis 253 1914 ARBORETUM ET FRUTICETUM BRITANNICUM. t 48. P. apulce'nsis Lindl. The Apulco Pine. Identification. Lindl. in Bot. Reg. M. Cbron., 1039, No. 100. Synonyme. P. acapulcensis G. Don in Sweet's Hort. Brit. ed. 3. p. 769. Engravings. Our figs. 1899,1900. from specimens sent home by Hartweg- 5 cm (cone) Figure 1. Illustration of Pinus apulcensis Lindley (Fig. 1899) in Loudon’s An Encyclopaedia of Trees and Shrubs; Being the Arboretum et Fruticetum Abridged (1842); drawn “from specimens sent home by Hartweg.” scales rather thin, which places it outside this cline. Martinez (1945: 192; see also 1948: 199) refuted Shaw’s inclusion of the two Nelson cones, in the belief that the specimens from Oaxaca were not conspecific with those from Apulco: “The speci¬ mens from Apulco and surrounding areas lack the characteristic elongations seen in specimens found further south, that is, in Veracruz and Oaxaca” [translated from Spanish]. He observed that Lindley 254 Novon did not mention these elongations (“prolonga- ciones,” meaning the elongated apophyses) but only “a recurved tip,” which sounds like an attempt to bend Lindley’s description into shape with what Martinez saw from Apulco. His subsequent “new” combination Pinas pseudostrobus Lindley var. apul- censis Martinez is superfluous because it repeats Shaw (1909) by including Lindley’s type, the iden¬ tity of which was totally unknown to Martinez. The specimens he referred to (e.g., Martinez 3434 from Apulco, Hidalgo “var. nov.” MEXU; Martinez, 1948: 201, f. 162) indeed lack any pronounced elongation of apophyses and/or umbos. However, they do not represent Lindley’s species, but Mar¬ tinez’s interpretation of it, based on the cited lo¬ cality. Collected around 1939, almost exactly a cen¬ tury after Hartweg made his collections, they are not necessarily from the same location. North of the city of Tulancingo there are two localities with the name Apulco, one at the railwayhead and another ca. 4 km further down a road. To the east of these are mountains (Cerro Viego) and streams (canyons) with pine forests. This constitutes a large area, and it is impossible to know exactly where Hartweg made his collection. Given the depletion of pine forests in many parts of Mexico in the last 100 years, it may well be that the trees Hartweg saw and collected are no longer there. For the collections from Oaxaca that Shaw had cited under Pinus pseudostrobus var. apulcensis (Nelson 985 and Nelson 2359), Martinez (1945: 195; see also 1948: 202) proposed a new combi¬ nation, P. pseudostrobus var. oaxacana, citing in synonymy P pseudostrobus var. apulcensis (Lindley) Shaw. This was an error: he clearly meant to ex¬ clude Lindley s type (Mirov, 1958) and to describe a new variety based on the two Nelson collections. Further, the name was not validly published be¬ cause he failed to provide a Latin diagnosis. Mirov (1958), referring to the specimens cited by Shaw (1909) and the description given by Martinez (1945), and excluding P. apulcensis Lindley, de¬ scribed it as a new species: P. oaxacana Mirov. Harrison (1965) then validated Martinez’s earlier combination under P. pseudostrobus at the rank of variety. Martinez, in the absence of Lindley’s type and with no easy access to European herbaria or nine¬ teenth-century botanical literature (he studied these pines during the Second World War), relied on the topotype method to establish the identity of Pinus apulcensis Lindley. Understandable as this may be in the case of Martinez, given the circumstances under which he worked, it is re¬ markable that later authors (Loock, 1950; Mirov, 1958; Harrison, 1965; Stead, 1983; Stead & Styles, 1984) simply accepted his judgment and never pursued the identity of Lindley’s name by typification. Stead (1983) undertook a numerical study of variation in which one of his collecting sites (No. 14) was the putative type locality of Pinus apul¬ censis Lindley at Apulco, Hidalgo. In his Principal Component Analysis (PCA), the Apulco material (from 25 trees) groups with sites 13 (P. pseudo¬ strobus var. estevezii Martinez) and 15 (P. pseudo¬ strobus var. coatepecensis Martinez) on character states such as “increasing cone size, needle width and stomatal lines,” but without any clear discon¬ tinuity. Cone size also increased for site 8 (rep¬ resenting P. pseudostrobus var. oaxacana (Mirov) Harrison), but no other characters, including its “very distinctive cones” would separate out this taxon. The conclusions outlined in Stead (1983) were adopted by Stead & Styles (1984), who identified the material from sites 13-15 as Pinus pseudostro¬ bus subsp. apulcensis (Lindley) Stead. The two va¬ rieties of Martinez (from sites 13 and 15) were cited as synonyms, as well as P. pseudostrobus var. apul¬ censis “sensu Shaw pro parte” (which, as can be understood from the synonymy under the previous taxon treated in their revision, means as to type, not to specimens cited). The type, however, was cited as “Apulco, Hartweg s.n. (not seen).” (Lindley (1839) did not indicate any lack of a collection number and only mentioned that it was “found by Mr. Hartweg”). It is, of course, the identity of Pinus apulcensis Lindley that needs to be established before adopt¬ ing this epithet. From the above it is obvious that an ovuliferous cone is needed; there is general agreement that foliage characters within Pinus pseudostrobus s.l. show considerable overlap and that leaves alone are inadequate for certain deter¬ mination (Stead & Styles, 1984; Perry. 1991; Car- vajal & McVaugh, 1992). Lindley, when he was Secretary of the Horticultural Society of London, distributed material collected by Hartweg to various botanists in Europe, among whom were Parlatore in Florence and Endlicher in Vienna. On a recent visit to Vienna (W), a number of conifer cones of Hart- weg’s gatherings were found. Among them are two cones of a Mexican pine with Lindley’s labels: P- apulcensis Lindley” and a vial with seeds and scales. The cones are 9.5 X 6 cm (closed) and 10 X 8 cm (open) and strongly resemble figure 1899 in Loudon (1842). They are original material and Volume 5, Number 3 1995 Farjon Pinus apulcensis 255 5 cm Figure 2. Closed ovuliferous cone of Pinus apulcensis Lindley, with a label written by John Lindley; it is part of the lectotype in the carpological collection of the Natural History Museum in Vienna (W). are here designated as the lectotype of Pinus apul¬ censis Lindley (Fig. 2). The synonymy given here pertains only to Pinus apulcensis Lindley; no full synonymy is given for P. pseudostrobus Lindley. Pinus pseudostrobus Lindley var. apulcensis (Lindley) Shaw, Publ. Arnold Arbor. 1: 19, t. 12, f. 6—8. 1909. Pinus apulcensis Lindley, Ed¬ wards’s Bot. Reg. 25: 63. Aug. 1839 [& Allg. Gartenzeitung 7: 325. 1839]. Pinus pseudostro¬ bus subsp. apulcensis (Lindley) Stead, Bot. J. Linn. Soc. 89; 269. 1984. TYPE: Mexico. Hi¬ dalgo: ravines near Apulco, C. T. Hartweg s.n., ex Hort. Soc. London, 2 ovuliferous cones with labels: P. apulcensis Lindley in Lindley’s hand¬ writing, vial with seeds and scales (lectotype, here designated, W). Pinus pseudostrobus var. oaxacana Martfnez, Anales Inst. Biol. Univ. Nac. Mexico 16: 1945, f. 164—168. 1945, nom. inval. Pinus oaxacana Mirov, Madrono 14: 145. 1958. Pinus pseudostrobus var. oaxacana (Mirov) S. G. Harrison, Taxon 14: 247. 1965. TYPE: Mexico. Oaxaca: La Parada, E. W. Nelson 985 (holotype, US). Pinus pseudostrobus Lindley is a widely distrib¬ uted species with polymorphic ovuliferous cones, occurring from Sinaloa and Nuevo Le6n in Mexico to Honduras (Stead & Styles, 1984). Pinus pseu¬ dostrobus var. apulcensis (Lindley) Shaw is a cone variant distinguished by more or less clinal char¬ acters becoming more abundant in southern Mex¬ ico. It is usually sympatric with P. pseudostrobus Lindley var. pseudostrobus but apparently absent from large parts of the range of the species (Car- vajal & McVaugh, 1992). Acceptance at any other rank than variety therefore seems unwarranted for this taxon. Current use of the epithet apulcensis in the sense of Article 57 (formerly 69.4) of the Code (Greuter et al., 1994) is only at infraspecific rank. The name P. oaxacana is occasionally used, but this taxon, too, is treated in most of the literature, especially forestry literature, at infraspecific rank. There is therefore only a weak case for a proposal either to reject P apulcensis Lindley or to conserve it with a conserved type, and I strongly prefer to correct the errors made before I take up the name in the sense of its type in Flora Neotropica. Acknowledgments. Thanks are due to staff at the Herbarium of the Museum of Natural History in Vienna, in particular B. Wallnofer, who alerted me to the existence of old carpological material in W despite its presumed destruction during World War II, and who granted a loan to Oxford (FHO) of the relevant material. Literature Cited Carvajal, S. & R. McVaugh. 1992. Pinus L. Pp. 32-100 in: R. McVaugh, Flora Novo-Galiciana 17. Gymno- sperms and Pteridophytes. Univ. Michigan Herbarium. Ann Arbor. Dfimmer, R. A. 1913. The conifers of the Lindley her¬ barium; Botany School, Cambridge. J. Roy. Hort. Soc. 39 (1): 63-91. Endlicher, S. L. 1847. Synopsis Coniferarum. Scheitlin & Zollikofer, Sangalli (Sankt Gallen). Greuter. W„ F. R. Barrie. H. M. Burdet, W. G. Chaloner, V. Demoulin, D. L. Hawksworth, P. M. Jorgensen. D. H. Nicolson, P. C. Silva, P. Trehane & J. McNeill, Editors. 1994. International Code of Botanical Nomenclature (The Tokyo Code), Adopted by the XVth International Botanical Congress, Yokohama, August-September, 1993. Regnum Veg. 131. Harrison, S. G. 1965. Note on gymnosperm nomencla¬ ture. Taxon 14: 247. Lindley, J. 1839. Miscellaneous notices: Mexican pines. Edwards’s Bot. Reg. 25: 63. Loock, E. E. M. 1950. The pines of Mexico and British Honduras. S. Africa Dept. Agric. and Forest. Bull. 35. Loudon, J. C. 1842. An Encyclopaedia of Trees and Shrubs; Being the Arboretum et Fruticetum abridged. Published by the author, London. 256 Novon Martinez, M. 1945. Las pinaceas mexicanas. Anales Inst. Biol. Univ. Nac. Mexico 16: 1-345. -. 1948. Los Pinos Mexicanos, ed. 2. Edicionas Botas, Mexico City. McVaugh, R. 1972. Botanical exploration in Nueva Ga¬ licia, Mexico, from 1790 to the present time. Contr. Univ. Michigan Herb. 9: 205—357. Mirov, N. T. 1958. Pinus oaxacana, a new species from Mexico. Madrono 14: 145-150. Perry, J. P. 1991. The Pines of Mexico and Central Amer¬ ica. Timber Press, Portland, Oregon. Shaw, G. R. 1909. The pines of Mexico. Publ. Arnold Arbor. 1. Stafleu, F. A. & R. S. Cowan. 1981. Taxonomic Literature 3 [TL-2]. Regnum Veg. 105, Bohn, Scheltema & Hol- kema, Utrecht. Stead, J. W. 1983. Studies of variation in Central Amer¬ ican pines V: A numerical study of variation in the Pseudostrobus group. Silvae Genet. 32: 101-114 [115]. -& B. T. Styles. 1984. Studies of Central Amer¬ ican pines: A revision of the “pseudostrobus” group (Pinaceae). Bot. J. Linn. Soc. 89: 249—275. Aureliana darcyi, a New Species of Solanaceae from Brazil L. d’A. Freire de Carvalho and Massimo G. Bovini Jardim Botanico do Rio de Janeiro, Segao de Botanica Sistematica, Rua Pacheco Leao 915, CEP 22460-030, Rio de Janeiro, RJ, Brazil Abstract. A new species of the genus Aureliana from the coastal vegetation and secondary forest of Paraty Municipality is described. It is distinguished from other species in this genus by the long-acu¬ minate calyx lobes and sparse pubescence. The genus Aureliana Sendtner was recently re¬ vised by Hunziker & Barboza (1991), who recog¬ nized eight taxa, all found in Brazil and concen¬ trated in the central-western, southeastern, and southern regions. During a floristic survey carried out by the Rio de Janeiro Botanical Garden in the Cairugu Envi¬ ronmental Protection Area, Praty Municipality, Rio de Janeiro state, a morphologically distinct species of Aureliana was discovered. Aureliana darcyi Carvalho & Bovini, sp. nov. TYPE: Brazil. Rio de Janeiro: Paraty, Apa- Cairugu, ilha das Almas, na orla da ilha, 15 Mar. 1989, A. L Almeida, B. Kurtz et al. 7 (holotype, RB; isotype, MO). Figures 1—7. Arbor ca. 2—3 m alta; ramis dichotomis puberulis. Folia solitaria, membranacea, glabrescentia, trichomatibus sim- plicibus vestita. Inflorescentiae ex fasciculis 4—12-floris constantes, receptaculo recto; pedicellis filiformibus apice vix dilatato. Flos in alabastra globosus; calyce campanu- lato, lobulis longe acuminatis aequalibus aestivatione lib- eris; corolla viridi; staminum filamentis ca. 2.5 mm longis, antheris ca. 1.5 mm longis. Bacca globosa; placenta ex- pansa protuberationes formante; seminis testa reticulata. Shrub or small tree ca. 2—3 m, branches dichot¬ omous with distinct longitudinal ribs, puberulent, with simple hairs and lenticels. Leaves solitary, membranaceous, dark green above and light green underneath, 3.0-12.0 X 0.8-4.5 cm, lanceolate, acuminate, base slightly decurrent, venation camp- todromous, glabrate to pubescent, hairs simple; pet¬ iole flattened, 1.0-1.5 cm, puberulent. Inflores¬ cence 4—12-fasciculate, receptacles straight, peduncle 0.3—1.2 cm. Flowers green, axillary. Ca¬ lyx whitish green, campanulate, ca. 3 mm, lobes 1.0-2.0 mm, regularly long-acuminate, glabrate ex¬ ternally. Corolla green, when mature white with green macula, 1.7-1.9 cm, rotate-stellate; lobes 5.0-6.0 mm, lanceolate, reticulate-veiny, margins ciliate. Pedicel 0.9—1.0 cm, filiform, slightly en¬ larged at the apex. Anthers light brown, ca. 1.5 mm, basifixed, filaments white, ca. 2.5 mm, filiform. Ovary less than 1.0 mm, globose to ovoid; style ca. 1.0 mm, stigmatic region apical. Fruit green, glo¬ bose, ca. 1.0 cm diam., pericarp thick, placenta with protuberances. Calyx not persistent in fruit, ca. 70 seeds, 0.2-0.3 mm long, oblong, testa reticulate. This plant was growing in the sun near sea level on an island (ilha das Almas) in the Atlantic Ocean. At the time of collection, it appeared to be in full flower with different stages of flowering present. The type specimen was collected in secondary for¬ est, in full sunlight. It is hairier and the leaves are wide-lanceolate. The characters that distinguish A. darcyi from other species of the genus are the long-acuminate calyx lobes and the absence of indumentum on the exterior of the calyx. Aureliana darcyi is closely related to A. wettstein- iana (Witasek) A. T. Hunziker & Barboza, as shown by characteristics of the calyx lobes and flower bud. Overall appearance plus the sparse pubescence place it near A. fasciculata (Vellozo) Sendtner var. fasciculata. The specific epithet honors William G. D’Arcy, eminent botanist from the Missouri Botanical Gar¬ den, for his important work on Solanaceae. Paratypes. BRAZIL. Rio de Janeiro: caminho para a Toca do Ouro, pr6x. 4 estrada Parati-Cunha, 03 Jul. 1992, L. C. Giordano, E. F. Guimaraes et al. 1312 (RB); passando a 18 ponte entrar & direita (sentido RJ/SP) i ca. de 13 km do trevo de Praty, subindo o Rio Corisco, 29 Jun. 1993, R. C. C. Reis, R. Marquete et al. 32 (RB). Novon 5: 257-258. 1995. 258 Novon F'guresl 7 AureUana darcyi Carvalho & Bovini (holotype). —1. Habit. —2. Receptacle of the inflorescence. —3. R U . ' - <- , V '^' s owm 8 e ong-acuminate lobes. —5. Relation of the calyx to the corolla, external face. —6. Relation ol the filaments to the anthers, internal face. —7. Globose fruit. Acknowledgments. The authors thank Angela M. Sturdart Vax of the Brazilian Institute of Geog¬ raphy and Statistics (IBGE) for the Latin descrip¬ tion, and Dorothy Sue Dunn de Araujo of the State Environmental Protection Foundation (FEEMA) for translating the text into English. Literature Cited Hunziker, A. T. & G. E. Barboza. 1991. Estudios sobre Solanaceae XXX. Revisibn de Aureliana. Darwiniana 30(1-4): 95-113. Carex radfordii (Section Laxiflorae: Cyperaceae), a New Species from the Southern Appalachians L. L. Gaddy The Wild Wood Associates, Inc., The Wild Wood Way, Walhalla, South Carolina 29691, U.S.A. Abstract. Carex radfordii Gaddy is a new spe¬ cies of Carex sect. Laxiflorae from the southern Ap¬ palachian Mountains of North Carolina, South Car¬ olina, and Georgia. It is a robust species most closely allied to Carex purpurifera Mackenzie and Carex laxflora Lamarck. Its long awns, glaucous blue-green blades, and brownish white basal sheaths distinguish it from other species of Carex. Carex sect. Laxiflorae has been studied by vari¬ ous investigators in this century. Recently, Bryson (1980) carried out a morphological analysis, and Manhart (1984, 1985, 1986) conducted studies on the morphology, chemotaxonomy, and cytology of the section. Carex laxflora Lamarck, the namesake for sec¬ tion Laxiflorae, ranges across eastern North Amer¬ ica from Maine and southern Quebec south to North Carolina west to Wisconsin and Indiana (Gleason & Cronquist, 1991). It is probably the most variable and complex species in section Laxiflorae and is easily confused with other species in this section. Cariciologists have often wondered whether Carex laxflora was one species or a complex of species. Mackenzie (1935) described C. purpurifera Mac¬ kenzie from specimens that were previously thought to be C. laxflora, and Brysons (1985) description of C. manhartii Bryson indicates that C. purpurifera is, in fact, made up of at least two distinct elements. The following species was first seen in the spring of 1979. Since then, I have observed the plant at numerous sites and mapped its distribution. I now conclude that it is a new taxon. Carex radfordii Gaddy, sp. nov. TYPE: U.S.A. South Carolina: Oconee County, in shallow soil, over calcareous rocks. Station Cove, 25 Apr. 1986, L. L Gaddy s.n. (holotype, CLEMS; isotypes, GA, MO. NCU, NY. US, USCH, WCUH, herbarium of C. T. Bryson). Figure 1. A Carex laxflora aristis squamis pistillatis longis et laminis venetis glaucis differt; a Carex purpurifera vaginis basalis albidis brunneis differt. Plant caespitose perennial, culms 4—6 dm tall. strongly aphyllopodic, triangular, usually 1.5 X longer than blades. Basal sheaths brownish white, 3—9 cm long, cataphylls well developed, 1-7 cm long. Blades blue-green, glaucous on undersides, 2—5.5 cm long, 7-15 mm wide, midrib well devel¬ oped, with small teeth dorsally. Staminate spikes 2-A cm long, 3-6 mm wide, usually solitary, peduncles 4—12 cm long. Pistillate spikes 3-5, pe¬ duncles 4—12 cm long, bracts 4—10 cm long, 1-2 mm wide, apices acuminate, sheaths 1-3 cm long, ligules scarious, 4—6 mm long. Perigynia 3—10 per spike. 3-A.5 mm long, obovoid, light green, beak slightly recurved, to 1.5 mm long; scales acute to aristate, often asymmetrical, rarely retuse, serrulate awns 1—5 mm long, usually overtopping the beak of the perigynia, total scale and awn length 3—9 mm, midrib green, margins scarious and hyaline. Achenes obovoid, filling body of the perigynia, 1— 2.5 mm long, 0.8-1.2 mm wide, styles exserted, stigmas 3. I take pleasure in naming this species after Al¬ bert E. Radford. Professor Emeritus, University of North Carolina (Chapel Hill), and a lifelong student of Carex. In the mid 1970s, Dr. Radford told me that the Brevard Belt of the Carolinas and Georgia was a rich floristic zone that offered many potential discoveries. I honor him with this discovery. Some of the essential characters of C. radfordii, C. purpurifera, C. laxflora, C. striatula Michaux. and C. manhartii are compared in Table 1. As may been seen in the table, C. radfordii shares many characters with C. purpurifera and C. laxflora, and fewer characters with C. striatula and C. manhartii. Furthermore, measurements of the height of the flowering culms, pistillate scale body length, and pistillate scale awn length overlap somewhat in these five species. In height of flowering culm, C. radfordii is a robust species that can only be com¬ pared to C. purpurifera. Its blue-green blades (not obvious in herbarium specimens), which are glau¬ cous on the undersides, are similar to those of C. purpurifera, except for the lack of purplish pigment in the basal sheaths. Its pistillate scale awns are usually much longer than those of C. laxflora, C. Novon 5: 2.59—261. 1995. 260 No von Figure 1. Variation in pistillate scale shape and awn length in Carex radfordii Gaddy: from lower and upper pistillate spikes of the type specimen. 9X. Illustration by Hu Ye. striatula, and C. manhartii. Occasionally, the awns of C. purpurifera' s pistillate scales approach those of C. radfordii in length; however, on a given plant, the mean pistillate scale awn length is always great¬ er in C. radfordii. The shape of the pistillate scale awn in C. radfordii is also distinct from that of other related species. In C. laxiflora, C. striatula, and C. manhartii, the pistillate scales are acute to awned; in C. purpurifera, the scales are more variable, of¬ ten being retuse; in C. radfordii, the scales are long-awned to rarely retuse, with a majority usually asymmetrical. Chromosome number in C. radfordii has been determined to be n = 23, which separates it from all related species (Robert Naczi, unpubli¬ shed data; Manhart, 1986). Carex radfordii can be delimited from related species by several characters, as may be seen in the key below. From Carex manhartii and C. pur¬ purifera, to which it appears to be most closely re¬ lated, it can be separated by its brownish white culm bases. From C. laxiflora and C. striatula, it can be distinguished by its blue-green blades with glaucous undersides and its asymmetrical pistillate scale bodies with extremely long awns. Upon first examination, C. radfordii appears to be an “albino” C. purpurifera. Closer inspection, however, reveals that the staminate spikes of C. radfordii are significantly longer than those of C. purpurifera and that the pistillate scales are more variable in shape in C. radfordii. Other good field characters for C. radfordii are its erect flowering culms (those of C. laxiflora and C. striatula usually lean over) and its strongly aphyllopodic culm bases. Carex radfordii is found in rich, often rocky, cal- Table 1. Carex radfordii and its allies. Character C. radfordii C. purpurifera C. laxiflora C. striatula C. manhartii Height of flow¬ ering culms 4-6.5 dm 3.7-6.7 dm 3.1—5 dm 3-5.5 dm 2.1-3.9 dm Blade color Blue-green, glaucous underside Blue-green, glaucous underside Dark green Whitish blue- green Dark green Basal sheath color Brownish white Purple Brownish white Brownish white Reddish purple Shape of pis¬ tillate scale Long-awned to asymmetrical to retuse Long-awned to retuse Acute to awned Acute to awned Acute to awned Scale length 2-4 mm 1.7—5 mm 2-4.5 mm 1.5-5.7 mm 3.5-4 mm Awn length 1-5 mm 1—5 mm 0.5-2.3 mm 0.8-3.7 mm 0.1-3.5 mm Chromosome number n = 23 n = 17, 18, 19 n = 20 n = 18, 20 n = 14 Volume 5, Number 3 1995 Gaddy Carex radfordii 261 careous woodlands on well-drained soils. The plant is found from about 300 m to just under 650 m in elevation. Of the closely related species discussed in Table 1, Carex radfordii is sympatric only with C. manhartii (see below). Most known sites are lo¬ cated in the Brevard or Chauga Belt, a belt of meta¬ sedimentary rock along a transition zone between the Piedmont and the Blue Ridge of the Carolinas and Georgia. The rocks of this belt have weathered to produce soils with high concentrations of calci¬ um and magnesium (Hatcher, 1969). The soil pH at C. radfordii sites is fairly high for the generally acidic southern Blue Ridge region; at Station Cove, the type locality, the pH is 6.0 (Gaddy, 1990). Al¬ though it occurs in rich coves with C. austro-caro- liniana Bailey, Carex radfordii is usually found at rockier, better-drained microsites than those occu¬ pied by C. austro-caroliniana. Below Whitewater Falls, North Carolina, C. radfordii grows alongside of C. manhartii. When a plant of C. radfordii from the type locality was experimentally grown in low calcium, low pH soils with a plant of C. laxiflora from the Blue Ridge of North Carolina and a plant of C. striatula from the Piedmont of South Carolina, all three species retained their distinguishing mor¬ phological characteristics. Key to C. radfordii and Related Species la. Base of culms brown, brownish white, or white. 2a. Blades whitish green to blue-green, under¬ sides white striolate; pistillate scales sym¬ metrical, awns 0.5-3.7 mm long. 3a. Blades whitish green to blue-green . . . C. striatula 3b. Blades dark green. C. laxiflora 2b. Blades blue-green, undersides glaucous; pistillate scale bodies often asymmetrical, awns 1—5 mm long. C. radfordii lb. Base of culms reddish, reddish purple, or pur¬ plish. 3a. Blades green, undersides not glaucous . . . . C. manhartii 3b. Blades blue-green, undersides glaucous . . .C. purpurifera Paratypes. U.S.A. Georgia: Stephens County, Cedar Creek, Camp Mikell Rd. off Ga. Rt. 184, just W of Camp Mikell, N of Toccoa, elev. ca. 300 m, deep gorge, 19 June 1975, D. E. Boufford 16552 & E. Wood (GA). North Car¬ olina: Macon County, Whitewater Falls, just below falls on lower portion of rich slopes, 15 May 1994, L L Gaddy s.n. (CLEMS). South Carolina: Greenville County, with Carex austro-caroliniana and Cladrastis kentuckea off Bentfield Rd. N side of Saluda River, 17 May 1993, S. R. Hill 25034 (CLEMS); Oconee County, in rich, calcareous woods along Ramsey Creek below Rich Mountain, 20 Apr. 1992, L L Gaddy s.n. (CLEMS); Rich woods, Station Cove, 22 Apr. 1986, L. L Gaddy s.n. (CLEMS); NE side of ridge at Tamassee Creek, Rich cove woodland, Comp. 17, Stnd. 16, 8 May 1993, S. R. Hill 24994 (CLEMS). Pickens County, Rich woods, below “the Narrows” in the Eastatoe Gorge, 23 Apr. 1987, L L Gaddy s.n. with D. A. Rayner (CLEMS); 1.3 mi. S of NC line, N of Rocky Bottom, elev. ca. 2100 ft., diverse mesic forest slope near creek crossing, soil a rich alkaline or neutral loam, oc¬ casional to frequent, conspicuously glaucous perennial, 10 Apr. 1988, S. R. Hill 19102 (CLEMS); shaded rich mesic woodland, 1.3 mi. S of NC line, N of Rocky Bottom, elev. ca. 2100 ft., occasional glaucous herb on steep slope, 9 May 1988, S. R. Hill 19271 with C. Horn (CLEMS). Acknowledgments. I thank individuals who commented on drafts of this paper over the last two years. Special thanks go to A. A. Reznicek, Charles T. Bryson, and Robert Naczi. I also acknowledge the curators of the following herbaria for loans and assistance: CLEMS, GA, MICH, NCU, OKL, UARK. Literature Cited Bryson, C. T. 1980. A Revision of the North American Carex Section Laxiflorae (Cyperaceae). Ph. D. Disser¬ tation, Mississippi State University, Mississippi State. -. 1985. A new species of Carex (section Laxiflo¬ rae) from the southern Appalachians. Castanea 50: 15— 18. Gaddy, L. L. 1990. Glade Fern Ravine, a rich fern site in the Blue Ridge Province of South Carolina. Castanea 55: 282-285. Gleason, H. A. & A. Cronquist. 1991. Manual of the Vascular Plants of Northeastern United States and Ad¬ jacent Canada. Van Nostrand, 2nd ed.. The New York Botanical Garden, Bronx, New York. Hatcher, R. D. 1969. Stratigraphy, petrology, and struc¬ ture of the low rank belt of the Blue Ridge of the north- westernmost South Carolina. Geologic Notes 13: 105- 141. Mackenzie, K. K. 1935. Carex purpurifera and Carex lax¬ iflora. In: North American Flora 18: 253—255. Manhart, J. R. 1984. A biosystematic study of Carex (sec¬ tion Laxiflorae). Ph.D. Dissertation, University of Geor¬ gia, Athens. -. 1985. Foliar flavonoids of the North American members of Carex section Laxiflorae (Cyperaceae). Bio- chem. Syst. and Ecol. 14: 85-90. -. 1986. Cytology of Carex purpurifera Mack. (Cy¬ peraceae). Rhodora 88: 141—147. New Species of Southern African Moraea (Iridaceae—Iridioideae), and the Reduction of Rheome 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 National Botanical Institute, P. Bag X7, Claremont 7735, South Africa Abstract. Two new species of the large African genus Moraea (125 species), M. rivulicola and M. regalis, both members of subgenus Viemseuxia, are described here. Moraea rivulicola has cream to greenish flowers, unusually long capsules, and is a robust plant restricted to the banks and beds of seasonal streams in Namaqualand, in the north¬ western part of South Africa. Moraea regalis is known from one locality, near De Rust in the Little Karoo, South Africa. Both species occur in the semiarid winter rainfall part of southern Africa, a region known for the richness and diversity of its flora. The genus Rheome, now believed to be most closely related to species of Moraea subg. Moraea, is reduced to synonymy, and the three species of Rheome are transferred to Moraea as M. maximi- liani, M. nana, and M. umbellata. Over half of the estimated 1750-1800 species of the petaloid monocot family, Iridaceae, occur in southern Africa (Goldblatt, 1991, 1994). The area is well known botanically, yet despite more than 250 years of botanical exploration, novelties con¬ tinue to be discovered there. Although the family is a significant element of the flora throughout the subcontinent, Iridaceae are most diverse in the southern and western parts of South Africa, a region of winter rainfall and summer drought. Two new species of Moraea are described here, both from this portion of southern Africa, and the genus Rheome is reduced to synonymy in Moraea. Moraea is the largest African genus of subfamily Iridoideae, and the central genus of Irideae-Homeriinae (Gold¬ blatt, 1976, 1986a). At the last revising, Goldblatt (1986a) admitted 119 species to the genus. Since then one species, M. vlokii (Goldblatt, 1992), has been described. With the addition of the two new species described here and three transferred from Rheome, there are now 125 species in the genus. According to the revised taxonomy, some 110 spe¬ cies of Moraea occur in southern Africa, that is, south of the Zambezi—Cunene axis, most of them endemic, and by far the majority restricted to the semiarid and arid southwestern portion of the re¬ gion. New Species 1. Moraea rivulicola Goldblatt & J. C. Manning, sp. nov. TYPE: South Africa. Northern Cape Province: Namaqualand, Springbok District, streambed at the eastern end of Spektakel Pass, 11 Sep. 1993, Goldblatt & Manning 9710 (holotype, NBG; isotypes, K, MO, PRE, WAG). Figure 1A-D. Plantae 50-80 cm altae, cormo globoso ca. 14—18 mm in diametro tunicis reticulatis, folio solitario lineare can- aliculato ca. 10 mm lato, caule ramoso, floribus cremeis vel virescentibus vel pallide brunneis, tepalis exterioribus unguiculatis atropunctatis ad bases limborum 20—24 mm longis, limbis lanceolatis ca. 14 mm longis, unguibus can- aliculatis 7—10 mm longis, tepalis interioribus 13—15 mm longis tricuspidatis, filamentis ca. 8 mm longis in columno ca. 5.5 mm longo connatis, antheris 4—5 mm longis, ramis styli ca. 6 mm longis, cristis 3—5 mm longis erectis, cap- sulis cylindricis 25-32 mm longis. Plants 50-80 cm high. Corm globose, 14—18 mm diam., the tunics of pale, coarse vertical libers con¬ nected by fine cross strands in herringbone pattern. Cataphylls usually dry and brown at flowering time, extending 5-10 mm above the ground. Foliage leaf Figure 1. Morphology of Moraea rivulicola Goldblatt & J. C. Manning ( Goldblatt & Manning 9710 ) and M. unguiculata Kew Gawler ( Goldblatt & Manning 9711). M. rivulicola: —A. Corm and basal part of stem. —B. Upper part of stem. C. Outer and inner tepal. D. Capsule. M. unguiculata: —E. Leaf and upper part of stem. —F. Outer and inner tepal. —G. Capsule. Novon 5: 262-269. 1995. Volume 5, Number 3 1995 Goldblatt & Manning Southern African Moraea 263 264 Novon Figure 2. Distribution ranges of Moraea rivulicola (closed circles), M. regalis (open circle), and M. unguiculata (delimited by heavy lines). solitary, basal, 2-2.5 times as long as the stem, channeled, 6-10 mm wide (when opened flat). Stem erect, consisting of 4(-5) intemodes, straight and erect, the intemodes 10-15 cm long, simple or with 1-3 branches, the branches held close to the axes, bearing an entirely sheathing bractlike leaf at each of the Sf-A) nodes, sheathing leaves 50-65 mm long, the apices attenuate-filiform. Inflorescences rhipidia, these single and terminal on each branch, several-flowered; spathes enclosing the rhipidia un¬ equal, green, firm-textured, with dry brown atten¬ uate apices, the outer ca. 45 mm long, the inner 55-67 mm long. Flowers cream, greenish or beige, with large nectar guides at the bases of the outer tepal limbs, these consisting of dark spots on a dull yellow field, the claws similarly dotted, both guide and outer tepal claws glabrous, inner tepals light to reddish brown, also densely spotted; outer tepals 20—24 X 10—13 mm, the claws ascending, chan¬ neled, 7-10 mm long, each with a large basal nec¬ tary, the nectary opposed by a lobe ca. 2 mm long, the limbs reflexed 45°-60° below horizontal, 14 X 10 mm; inner tepals 13-15 X 4—5 mm, the claws ascending, ca. 8 mm long, channeled, the margins usually partly conduplicate, the limbs trilobed, the median lobe linear, curving inward laxly and asym¬ metrically coiled, lateral lobes broad and weakly defined, obtuse, 1—2 mm long. Filaments ca. 8 mm long, united in a cylindric column ca. 5.5 mm long, free and diverging in the upper 2.5 mm; anthers 4— 5 mm long, each appressed to the opposed style branch, shortly exceeding the stigma lobes, orange. Ovary oblong-trigonous, 11—14 mm long, exserted from the spathes; style dividing at the apex of the filament column, the branches ca. 6 X ca. 2 mm, diverging, each with paired terminal lanceolate crests 3—5 mm long. Capsules cylindric, 25—32 mm long; seeds angular, only known in the immature state. Flowering mid August to late September. Etymology. From the Latin rivulieola, “living in streams,” referring to the specialized habitat of the species. Distribution and habitat. Moraea rivulicola is a fairly rare endemic of northern Namaqualand, Northern Cape Province, South Africa. It extends from the Spektakel Mountains west of Springbok southward to the Kamiesberg (Fig. 2). Plants appear to be restricted to the beds of seasonal streams and along drainage lines, usually with their corms deep¬ ly wedged in crevices in the granite bedrock. Re- Volume 5, Number 3 1995 Goldblatt & Manning Southern African Moraea 265 cords are all from areas of fairly high elevation, and for Namaqualand, relatively high rainfall. Although Moraea rivulicola was first collected in 1883 by Harry Bolus and in the past was thought to be no more than a large-flowered variant of the widespread and fairly common M. unguiculata Ker Gawler (Goldblatt, 1976, 1986a), field observations in Namaqualand in 1993 and 1994 show that this treatment is incorrect. At two sites some 100 km apart, M. rivulicola was found growing along and in small seasonal streams, while fairly typical M. un¬ guiculata grew on adjacent well-drained slopes. Seen at the same time and in quite different habi¬ tats, it became evident that these were different species, not merely ecotypes or local variants of a single polymorphic one. Critical comparison of liv¬ ing plants showed that not only is M. rivulicola more robust vegetatively, and has larger flowers, but that there are several critical qualitative differences between it and M. unguiculata. The flowers of Moraea rivulicola (Fig. 1A—C) are cream with a pale greenish or beige tinge, and the outer tepals are ca. 24 mm long with the limbs typically held at 60° below the horizontal. The nec¬ tar guides, located at the base of the tepal limb, are whitish covered with small dark brown to black¬ ish spots, and both the nectar guide area and the tepal claws are smooth. The inner tepals are also distinctive: the central lobe of the tripartite limb is drawn into a slender obliquely coiled cusp, while the lateral lobes are obtuse and very short. The claws of the inner tepals are channeled and the margins are partly conduplicate, thus forming a nearly closed tube. The ovary is oblong-cylindric and 11—14 mm long, and gives rise to cylindrical capsules 25-32 mm long. Ripe seeds are unknown, but are probably larger than those of M. unguicu¬ lata. Moraea unguiculata, both in Namaqualand and elsewhere across its range, has smaller flowers with the outer tepals seldom exceeding 16 mm (Fig. 1E-G). Larger dimensions for the species given by Goldblatt (1976, 1986a) included plants now re- ferred to M. rivulicola. The tepal limbs in Nama¬ qualand populations of M. unguiculata are 8-10 mm long, and are held slightly below the horizontal, or flexed up to 45°, but in populations in the south¬ ern Cape, the limbs may be fully reflexed. Both the small nectar guide area and the outer tepal claws are densely pubescent. The inner tepals are 7—10 mm long, and also tripartite. The central cusp is long and symmetrically coiled inward, but the outer lobes are relatively longer and more distinctly formed than in M. rivulicola, and may be subacute. The inner tepal claws are more or less terete or slightly flattened on the adaxial surface. Flower col¬ or in M. unguiculata varies, ranging from pure white to cream, brownish, and even blue, and floral form (but not size) is unusually plastic (Goldblatt, 1986a). The outer tepal limbs may extend outward or, in some populations outside Namaqualand, may be completely reflexed. The distinctive pale green leaves of Moraea ri¬ vulicola are relatively long, sometimes up to 2.5 times as long as the stem, thus over 1 m in length, broadly channeled, and 6-10 mm wide. The stems are 4—5 intemodes long and bear 3-4 sheathing leaves 50-65 mm long. Leaves of M. unguiculata are darker gray-green, shorter, seldom more than a third again as long as the stem, and they are nar¬ rowly channeled, 2—3(-4) mm wide. The stems are 3 intemodes long and bear 2 sheathing leaves TO¬ SS mm long. Thus the two species differ both in leaf and stem morphology as well as in floral structure. Differences in the leaf texture and venation of the two species reflect notable anatomical differ¬ ences (Fig. 3). In transverse section the leaf of Mo¬ raea unguiculata (Fig. 3B) is relatively thin. The heavily sclerified sheaths of the major bundles touch those of the opposing minor bundles and there is a large strand of subepidermal marginal sclerenchyma, not apparently associated with a vas¬ cular bundle (although a small central cavity sug¬ gests the presence of minor veins in early ontoge¬ ny). In comparison, the leaf of M. rivulicola is much thicker in transverse section (Fig. 3A), the sheaths of the major bundles are less heavily sclerified and the opposing minor bundles are well separated from them, and most significantly, there is a small mar¬ ginal vein with a small sclerified phloem cap. These differences in leaf anatomy between the two species are of the kind most commonly associated with xer- ic versus more mesic habitats. Habitat differences between Moraea rivulicola and M. unguiculata appear to be absolute. The lat¬ ter grows on open exposed rocky sites, or on stony slopes, in both cases in well-drained and often arid situations. Moraea rivulicola has, however, so far only been found in seasonally wet habitats, water¬ logged or even inundated for part of the growing season. In Namaqualand M. unguiculata may be found either on soils derived from shale, as in the Spektakel Mountains or near Steinkopf (Goldblatt, 1986a), or on granite-derived soils. Moraea rivuli¬ cola has only been found on pockets of granite- derived sand in rocky granite on the banks or in the beds of streams. Both Moraea rivulicola and M. unguiculata be¬ long to subgenus Vieusseuxia, members of which share the following synapomorphies: long-lasting flowers, a solitary foliage leaf, distinctive reticulate 266 Novon corm tunics composed of pale, soft-textured fibers, and an asymmetric karyotype with x = 6. Within the subgenus, M. rivulicola and M. unguiculala ap¬ pear to be allied to those species that have similar trilobed inner tepals, inconspicuous nectar guides, and a long filament column. Other species in the alliance include the southwestern Cape species M. tricuspidata (L. fil.) G. Lewis and M. bellendenii (Sweet) N. E. Brown, and the eastern southern Af¬ rican M. trifida R. Foster, M. pubiflora N. E. Brown, and M. brevistyla (Goldblatt) Goldblatt. Paratypes. SOUTH AFRICA. Northern Cape: 2917 (Springbok) 22 km W of Springbok, eastern slopes of Spektakel Mountains, in streambed (DA), 26 Sep. 1974, Goldblatt 2791 (MO, PRE); streambed at the eastern end of Spektakel Pass, 4 Sep. 1994, Goldblatt & Manning 9971 (MO, NBG); rocky places near Modderfontein (DB), Sep. 1883, H. Bolus 6618 (BOL). 3018 (Kamiesberg) east¬ ern slopes of Rooiberg, ca. 1050 m (AC), 14 Oct. 1976, Goldblatt 4313 (MO, PRE); Kamiesberg Mountains, farm Welkom, streambed near the top of Langkloof, 14 Sep. 1993, Goldblatt & Manning 9770 (MO). 2. Moraea regalis Goldblatt & J. C. Manning, sp. nov. TYPE; South Africa. Western Cape: Little Karoo, De Rust, rocky hillside near old cem¬ etery, 19 Sep. 1994, /. A. Vlok & A. L Schutte 345 (holotype, NBG; isotypes, K, MO, PRE, WAG). Figure 4. Plantae 18-25(—35) cm altae, cormo globoso 12—14 mm in diametro tunicis reticulatis, folio solitario lineare can- aliculato, caule eramoso, floribus violaceis, tepalis exter- ioribus albis notatis unguiculatis (26—)30-A0 mm tongis, limbis lanceolatis 13—20 mm longis, tepalis interioribus 20—24 mm longis erectis saepe unguiculatis et limbis tn- Volume 5, Number 3 1995 Goldblatt & Manning Southern African Moraea 267 cuspidatis, filamentis 12—15 mm longis connatis in col- umno 9—10 mm longo, antheris 6-9 mm longis, ramis styli 10—12 X 2 mm divergentibus, cristis anguste lanceolatis vel linearibus 5-8 mm longis. Plants 18-25(-35) cm high. Corms globose, 12- 14 mm diam., the tunics of medium to coarsely textured reticulate fibers, light brown. Cataphylls dry and brown at flowering time. Foliage leaf sol¬ itary, basal, half again to twice as long as the stem, linear and canaliculate, ca. 3 mm wide (when opened flat), upright below, trailing above. Stem erect, 3 intemodes long, straight and erect, un¬ branched, the lowermost internode shortest and the uppermost longest, bearing an entirely sheathing bractlike leaf at each node, these 25-40 mm long, the apices attenuate. Inflorescence a single, usually 2-flowered terminal rhipidium; enclosing spathes unequal, green, firm-textured, with dry brown at¬ tenuate apices, the outer 25-40 mm long, the inner 45-65 mm long. Flowers violet, the limbs of the outer tepals each marked at the base with a small white triangular nectar guide outlined with a dark purple zone, the base of the outer tepal limb mi¬ nutely pubescent, the claw heavily pubescent, the hairs dark violet, evidently without nectar, scent¬ less; outer tepals (26—)30-40 mm long, the claws ascending, 12-16 mm long, with a smooth deltoid zone at the base, evidently a vestigial perigonal nectary, the limbs 13—20 mm long, directed down¬ ward at right angles to the ground; inner tepals erect and ± linear to slightly wider just below the mid¬ line, thus obscurely clawed, 20-24 mm long, the claw ca. 12 mm long, the limb aristate or occasion¬ ally tricuspidate, then the lateral lobe 1-2 mm long, and central cusp ca. 10 mm long. Filaments 12-15 mm long, united below in a cylindric column 9-10 mm long, free and diverging in the upper 3-5 mm; anthers 6-9 mm long, each appressed to the op¬ posed style branch, just shorter than or just ex¬ ceeding the stigma lobes, the pollen orange. Ovary oblong-trigonous, 8-10 mm long, usually fully ex- serted from the spathes, style dividing just above the apex of the filament column, the branches 10- 12 X 2 mm, diverging, each with paired terminal, narrowly lanceolate to linear crests 5-8 mm long. Capsules ovoid-oblong, 12-16 mm long; seeds an¬ gular and prismatic, ca. 1.5 mm long. Flowering mid August to mid September. Etymology. From the Latin regalis, “royal,” re¬ ferring to the dark blue-purple or violet color of the flowers. Distribution and habitat. Moraea regalis ap¬ pears to have been collected for the first time in 1993, when it was found near the Little Karoo town 268 Novon of De Rust (Fig. 2). So far it is known only from a single locality, south-trending slopes of a rocky hill south of De Rust. The substrate is sandy soil de¬ rived from Enon Conglomerate. Like Moraea rivulicola, described above, M. re- galis is a typical member of subgenus Vieusseuxia. It is distinguished in the genus and subgenus by the striking dark violet flower color, the relatively large flower size with the outer tepals 26-40 mm long, the long tepal claws, and the narrow, either aristate and more or less linear or tricuspidate inner tepals (Fig. 4). Its affinities lie with M. unguiculata and M. algoemis Goldblatt, both of which have smaller flowers and trilobed inner tepals with a large incurved central cusp and small but broadly obtuse lateral lobes. The slender filament column and narrow style branches are also typical of this group of species. Paratype. SOUTH AFRICA. Western Cape: 3322 (Oudtshoom) Little Karoo, De Rust, hill S of the town, stony S-facing slopes (BC), Sep. 1993, J. A. Vlok & A. L Schutte s.n. (MO). Reduction of Rheome Goldblatt The genus Rheome was erected by Goldblatt (1980) to accommodate two species of Homeria, H. umbellata (Thunberg) G. Lewis and H. maximiliani Schlechter. These differed from other species then included in Homeria in their brown, nonreticulate corm tunics, apically diverging outer inflorescence spathes, and apically crowded rhipidia (inflores¬ cence units) on the stem. The remaining species of Homeria have reticulate corm tunics composed of coarse black fibers, sheathing outer rhipidial spathes, and rhipidia that are not crowded apically. The basic chromosome number also differs in the two groups. Species of Homeria have x = 6, and secondarily 2 n = 10, 9, or 8, whereas H. umbellata and H. maximiliani have x = 10, considered to be the plesiomorphic chromosome number for Hom- eriinae (Goldblatt, 1976, 1980; Rudall & Goldblatt, 1993). Moreover, a series of crossing experiments showed that although most of the species of Hom¬ eria could be crossed with one another, none could be crossed with H. maximiliani. The latter, how¬ ever, could be crossed with one species of a second genus Hexaglottis, H. nana L. Bolus, which also has the corm tunics, rhipidial spathes, and arrange¬ ment of the rhipidia characteristic of Rheome. Nei¬ ther Homeria maximiliani nor Hexaglottis nana could be crossed with a selection of species of Hex¬ aglottis. It was evident on the basis of morpholog¬ ical, cytological, and crossing evidence that Hom¬ eria umbellata, H. maximiliani, and Hexaglottis nana were a monophyletic group parallel to Hex¬ aglottis and Homeria and that the close floral sim¬ ilarity to the latter two genera was due to conver¬ gence, most likely through the parallel reduction of the more complex Moraea- type flower, plesio¬ morphic in subtribe Homeriinae in its dimorphic inner and outer tepals and narrow style branches either bearing a reduced crest or lacking them en¬ tirely. What was not evident when Rheome was de¬ scribed was that the distinctive corm tunics, inflo¬ rescence spathes, and arrangement of rhipidia characteristic of that genus have a parallel in Mo¬ raea longiflora Ker Gawler (subg. Moraea sect. Tubiflora). The second species of section Tubiflora, M. coopen Baker, also has brown, nonreticulate corm tunics, but it has entirely sheathing rhipidial spathes and laxly disposed rhipidia. Basic chro¬ mosome number in subgenus Moraea is x — 10, which also accords with the number in Rheome. The discovery of M. linderi Goldblatt (Goldblatt, 1982) provided the missing link between Rheome and Moraea. This species has a flower typical of Moraea, having large outer tepals with conspicuous nectar guides, narrow inner tepals, broad style branches with paired petaloid crests, and a trans¬ verse stigma, but corms, corm tunics, rhipidial spathes and rhipidial arrangement of the type char¬ acteristic of Rheome, and a basic chromosome num¬ ber of x = 10 (Goldblatt, 1986b). Finally, a phylogenetic study of subtribe Hom¬ eriinae (Rudall & Goldblatt, 1993), using several leaf anatomical characters as well as morphology and chromosome cytology, provided evidence that Rheome, Moraea sect. Tubiflora, and two other spe¬ cies of section Moraea, M. linderi and M. margar- etae Goldblatt, comprise a clade. The study did not include enough species to show, with any degree of confidence, the species or section most closely al¬ lied. Nevertheless, the evidence now available makes it clear that the species of Rheome and Hex¬ aglottis nana are nested within a small clade of species of Moraea subg. Moraea. Under these cir¬ cumstances, continued recognition of Rheome, a small genus nested within a section of the large and diverse Moraea, is clearly inconsistent with modem phylogenetic classification (Hennig, 1966; Dono- ghue & Cantino, 1988). We therefore propose to synonymize Rheome in Moraea. As noted above, Rheome, H. nana, M. linderi, and M. margaretae comprise a clade with the following apomorphic characteristics: nonreticulate, brown corm tunics; apically diverging outer rhipidial spathes, and a tendency for the crowding of rhipidia and foliage leaves near the stem apex. Some of these characters Volume 5, Number 3 1995 Goldblatt & Manning Southern African Moraea 269 are also shared by the members of section Tubiflora which, in addition, have the tepals united in a tube, apomorphic in Moraea. The absence of some of the above synapomorphies in M. cooperi is puzzling, and we suspect may be due to character reversal. The taxonomy of species transferred to Moraea is outlined below. 1. Moraea maximiliani (Schlechter) Goldblatt & J. C. Manning, comb. nov. Basionym: Homeria maximiliani Schlechter, Bot. Jahrb. Syst. 27: 94. 1900. Rheome maximiliani (Schlechter) Goldblatt, Bot. Not. 133: 92. 1980. TYPE: South Africa. Western Cape: Hoek, Clanwil- liam District, Schlechter 8700 (lectotype, des¬ ignated by Goldblatt (1980), B; isotypes, BOL, K. MO, S). 2. Moraea nana (L. Bolus) Goldblatt & J. C. Man¬ ning, comb. nov. Basionym: Hexaglottis nana L. Bolus, Kew Bull. Misc. Inf. 1932: 326. 1932 [1933]. TYPE: South Africa. Western Cape: Clanwilliam District, without precise lo¬ cality, Weintraub s.n. (holotype, BOL 19972). 3. Moraea umbellata Thunberg, Dissertatio de Moraea no. 16. 1787. Homeria umbellata (Thunberg) G. Lewis, J. S. African Bot. 14: 86. 1948. Rheome umbellata (Thunberg) Gold¬ blatt, Bot. Not. 133: 92. 1980. TYPE: South Africa. Western Cape: near Piketberg, Thun¬ berg s.n. (holotype. Herb. Thunberg 1230 at UPS). Acknowledgments. We thank Jan Vlok, Western Cape Department of Nature Conservation, for his assistance with fieldwork and for bringing M. re¬ gain to our attention. Literature Cited Donoghue, M. J. & P. D. Cantino. 1988. Paraphyly, an¬ cestors and the goals of taxonomy: A botanical defense of cladism. Bot. Rev. 54: 107—127. Goldblatt, P. 1976. The genus Moraea in the winter rain¬ fall region of southern Africa. Ann. Missouri Bot. Card. 63: 657-786. -. 1980. Redefinition of Homeria and Moraea (Ir- idaceae) in the light of biosystematic data, with Rheome gen. nov. Bot. Not. 133: 85-95. -. 1982. A synopsis of Moraea (Iridaceae) with new taxa, transfers and notes. Ann. Missouri Bot. Gard. 69: 351-369. -. 1986a. Convergent evolution of the Homeria flower type in six new species of Moraea (Iridaceae) in southern Africa. Ann. Missouri Bot. Gard. 73: 102—116. -. 1986b. The Moraeas of Southern Africa. Ann. Kirstenbosch Bot. Gard. 14: 1—224. -. 1991. An overview of the systematics, phylogeny and biology of the southern African Iridaceae. Contr. Bolus Herb. 13: 1—74. -. 1992. New species, chromosome cytology and notes on the southern African Iridaceae—Irideae: Mo¬ raea, Roggeveldia and Homeria. S. African J. Bot. 58: 209-214. -. 1994. Geography of African Iridaceae. Pp. 931— 940 in J. H. Seyani & A. C. Chikuni (editors). Pro¬ ceedings of the XHIth plenary Meeting AETFAT, Ma¬ lawi, Volume 2. National Herbarium and Botanic Gardens of Malawi, Zomba. Hennig, W. 1966. Phylogenetic Systematics. Translated by D. D. Davis & R. Zangerl. Univ. Illinois Press, Ur- bana. Rudall, P. & P. Goldblatt. 1993. Leaf anatomy and sys¬ tematics of Homeriinae (Iridaceae). Bot. J. Linn. Soc. Ill: 379-397. A New Rhynchospora (Cyperaceae) from the Cordillera de Talamanca, Costa Rica Jorge Gdmez-Laurito Escuela de Biologfa, Universidad de Costa Rica, Ciudad Universitaria 2060, San Jose, Costa Rica Abstract. Rhynchospora cabecarae G<5mez-Lauri- to (Cyperaceae), a new species belonging to section Racemosae C. B. Clarke, is described and illustrated. A new species of Rhynchospora Vahl (Cypera¬ ceae) is described as a result of revision of material recently collected during a field trip to the Carib¬ bean slope of the Cordillera de Talamanca in Costa Rica. Rhynchospora cabecarae Gbmez-Laurito, sp. nov. TYPE: Costa Rica. Li rnon: Talamanca, N flank of Cerro Casma, along Ujarras-San Jose Cab^car trail. Cordillera de Talamanca, 9°20'30"N, 83°13'30"W, 2250-2270 m, 17 Mar. 1993, M. Grayum 10354 (holotype, USJ). Figure 1. Ex affinitate R. boliviensis C. B. Clarke atque R. dissi - tiflorae Steudel ex Bockeler, ab utroque characteribus in- florescentiarum et achaeniarum praecipue differt. Perennial, loosely caespitose with short rhizomes and coarse roots. Culms covered by old sheaths, erect, stiff, ca. 60 cm tall, 2.5 mm thick, nodose, leafy with many cauline leaves. Leaves 8-20 cm long, erect, somewhat wiry; blades linear, 2-2.8 mm wide, gradually narrowed apically, the margins ciliate to scabrous, channeled, the midrib adaxially pilose, raised and ciliate abaxially. Sheaths striate, reddish brown, villous with soft, curled hairs. Inflo¬ rescence paniculate, of ca. 5 partial panicles, the panicles not exceeding the leaves, 1.5—2.5 cm long, 1.3—2 cm wide, of 11-23 ascending spikelets; the peduncles 1.3-2 cm long, terete to complanate, hir¬ sute, the axis sparsely hirsute. Bracts leaflike. Bractlets subulate, 4—6 mm long, hirsute. Spikelets 6-7 mm long, 1—1.3 mm wide, acute, reddish brown, 2-fruited. Lower glumes empty, ovate, 2.5- 3 mm long, mucronate, other flower-bearing glumes lanceolate, 4 mm long, 1.5 mm wide, membranous, mucronate at the tip. Achenes ovate-elliptic to el¬ liptic, tumid, 1.2 mm long, 0.7 mm wide, with a conspicuous basal stipe 0.5 mm long, shiny-casta- neous, cancellate with minute isodiametrical cells. Style-base conical, 1 mm high, 0.8 mm broad at base, dorsiventrally compressed, brown. Hypogy- nous bristles 5, upwardly scabrous, rather slender, shorter than the achene body, or often only 2 very short rudimentary ones. Stamens 3. Etymology. This species is named after the Ca- becar people, who inhabit the Talamanca region. Rhynchospora cabecarae is a member of section Racemosae C. B. Clarke because of its short rhi¬ zome, culms covered by old sheaths, many cauline leaves, inflorescence paniculate, number of hypog- ynous bristles shorter than the achene body, often only 2 vestigials, and cancellate achenes. It is ap¬ parently related to R. boliviensis C. B. Clarke and R. dissitiflora Steudel ex Bockeler. As has recently been stated by Thomas (1993), this group is quite variable and rare. Rhynchospora cabecarae is known only from the type collection. It is distin¬ guished from the above-mentioned species by the combination of characters in Table 1. Rhynchospora cabecarae was found in an open scrubland with Sphagnum and shrubs and treelets of several families, including Clusia (Clusiaceae), Table 1. Comparison of three related species of Rhynchospora. R. boliviensis R. dissitiflora R. cabecarae Inflorescences A single open panicle with slender peduncles 1—5 cm long with solitary spikelets Achenes Widely obovate, lenticular to plano-convex; brownish gray to straw color. Stipe none 3-5 racemes, each with 1-4 spikelets with slender and pendant peduncles Widely obovate, lenticular, pale brownish gray. Stipe none 5 partial small panicles, each with 11-23 spikelets; pe¬ duncles short, 1—3 cm long Ovate to ovate-elliptic, tumid, shiny-castaneous, with a distinct stipe 0.5 mm long Novon 5: 270-271. 1995. Volume 5, Number 3 1995 Gomez-Laurito 271 Rhynchospora cabecarae from Costa Rica figure 1. Rhynchospora cabecarae Gdmez-Laurito (Grayum 10354, USJ). a. Habit. b. Partial panicle. Achene. Prunus (Rosaceae), Vaccinium (Ericaceae), Ilex (Aquifoliaceae), Miconia (Melastomataceae), Pali- courea (Rubiaceae), Gaiadendron (Loranthaceae), Monnina (Polygalaceae), Neomirandea (Astera- ceae), and Phyllonoma (Grossulariaceae) (M. Gra¬ yum, in litt. 1994). Acknowledgments. I am grateful to Michael H. Grayum for reviewing the manuscript and to Judy Hoomans for the excellent illustration. Literature Cited Thomas, W. W. 1993. A synopsis of Rhynchospora (Cy- peraceae) in Mesoamerica. Brittonia 44: 1 4-44 . Two New Species in Cuphea (Lythraceae), and a Note on Alzateaceae Shirley A. Graham Department of Biological Sciences, Kent State University, Kent, Ohio 44242 U.S.A. Abstract. Cuphea (sect. Melvilla) alaniana from central Mexico and C. (sect. Diploptychia) armata from Central America are described and illustrated. The monotypic family Alzateaceae, long known from Peru and Bolivia, and more recently discov¬ ered in Costa Rica, Panama, and Colombia, is here reported from Ecuador for the first time. Of the two Ecuadorian collections, one belongs to the north¬ ern, broad-leaved Alzatea verticillata subsp. ampli- folia; the other is most similar to the southern sub¬ species verticillata. Apical leaf glands are a newly recognized feature of the genus. They are inter¬ preted as extra-floral nectaries. In revisionary studies of Cuphea, two new North American species were discovered among collections on loan from a number of North American herbaria. Cuphea alaniana is a large-flowered perennial of section Melvilla, subsection Polyspermum (Koehne, 1903). It is one of four species of the subsection, which is endemic to the western and southern moun¬ tains of Mexico. Cuphea armata, discovered in neighboring areas of Guatemala, El Salvador, and Honduras is a new species of section Diploptychia. Cuphea alaniana S. A. Graham, sp. nov. TYPE: Mexico. Guerrero: Distrito Adama, Temisco, Barranca del Consuelo, 500 m, 15 Nov. 1937, Mexia 8810a (holotype, UC; isotype, GH). Fig¬ ure la-d. Suffmtices vel frutices, probabiliter ad 1.5 m, caulis simplex, erectus, viscosissimus. Petioli 5-10 mm longi, ad summum caulis sessiles; folia 25-100 mm longa, 5-35 mm lata, anguste lanceolata vel linearia. Flores 25-32 mm longi. rubro-purpurei, glanduloso-hispidi; lobus dorsalis calycis ceteris paullo major; appendices prominentes, se- tosae. Petala 6, roseo-purpurea, inaequalia, 2 dorsalia 7- 9 mm longa, manifeste unguiculata, 4 ventralia 1.0-2.5 mm longa. Ovula 40-50. Discus 2-3 mm longus, 2.5 mm latus, 2 mm crassus, basi ovarii late affixus. Suffrutescent or shrubby perennials, probably 0.5-1.5 m tall, stems simple, erect, from a fibrous- rooted base, branched distally in the inflorescence; stems and pedicels highly viscous, the vestiture densely, minutely glandular-puberulous and glan¬ dular-hispid, the hairs thin, erect, colorless or red- Novon 5: 272-277. 1995. dish purple, bulbous-based, minute (< 0.25 mm) or 0.5-1 mm long; intemodes up to half as long as the subtending leaves. Leaves with petioles 5—10 mm long, becoming subsessile or sessile distally in the inflorescence; blades 25-100 mm long, 5-35 mm wide, membranous, crowded on the upper stem, narrowly lanceolate-linear, base acute, apex acute; blade surface minutely scabrous, or scabrous with sparse to abundant white, weak, slightly ap- pressed hairs to 1 mm long, the margins of young leaves densely ciliate, the hairs erect, glandular- viscous, to 1 mm long; upper leaves very gradually diminishing in size to become the lanceolate-linear bracts of the inflorescence. Inflorescences densely flowered, terminal, leafy racemes; flowers solitary at the nodes, interaxillary; pedicels 4—6 mm long; bracteoles 1.5 mm long, lanceolate-linear or re¬ duced to a few fleshy hairs, positioned 1.0-1.5 mm below the apex of the pedicel. Floral tubes 25-32 mm long, including a rounded, horizontal base ex¬ tending 1 mm beyond the pedicel, 4—5 mm wide in lateral view at anthesis, dorsally slightly convex, the neck slightly or not at all narrowed, the mouth flaring, oblique in lateral view, the ventral side ex¬ tended 1 inin or less beyond the dorsal side, the floral tube more distinctly convex and distended in fruit, the neck contracted; outer surface reddish purple, straw-colored ventrally at the mouth, glan¬ dular-puberulous and glandular-hispid with abun¬ dant, erect, thin, colorless or reddish purple hairs to 1 mm long; inner surface pale, neither bialate nor vesiculate, glabrous or lightly villous above the stamens, glabrous below; calyx lobes subequal, the dorsal lobe deltoid, 2X3 mm, the others 1X2 mm, margins not ciliated; appendages at the sinus¬ es of the lobes prominent, 1 mm long, equaling the calyx lobes, thick, green or reddish purple, the apex bulbous, crowned by 5-8 setae 1—2 mm long. Petals 6, rose-purple, the 2 dorsal 7—9 mm long including a slender claw 1-2 mm long, narrowly obovate, the 4 ventral 1.0-2.5 mm long, linear-spa- thulate, caducous. Stamens 11, all inserted at % the length of the floral tube, the 2 dorsalmost shortest, included, the 5 antesepalous exserted, the 4 ante- petalous included to subexserted; filaments gla- Volume 5, Number 3 1995 Graham Cuphea 273 Figure 1. a-d, Cuphea alaniana S. A. Graham, —a. Habit. —b. Lateral view of flower. —c. Nectariferous disc at base of the ovary within the floral tube. —d. Floral tube opened dorsally, ovary removed to show position and number of stamens. —e. Cuphea armata S. A. Graham, lateral view of flower. Drawn from the holotypes. brous to lightly villous, reddish; anthers purple. Pollen triangular in polar view; tricolporate, syn- colpate, pores not protruding; exine uniformly fine¬ ly striate, the striae extending ca. Vi the distance to the poles, partially anastomosing; diameter 24- 28 gm in lactic acid. Style exserted at maturity, glabrous; stigma punctiform; ovary gibbous dorsal¬ ly, the gibbosity pronounced in flower; ovules 40- 50. Seeds numerous, 2 mm long, 1.5 mm wide. Disc massive, 2-3 mm long, 2.5 mm wide, 2 mm thick, horizontal, oblong-rectangular in dorsal view, broadly attached at the base of the ovary. Phenology. Collected in flower and fruit in No¬ vember. 274 Novon Figure 2. Distribution of Cuphea alaniana and C. armata in Mexico and Central America. Distribution and habitat (Fig. 2). In southern Mexico state and adjacent Guerrero; low spiny, de¬ ciduous woods, dry slopes, and sandy streamsides; 500-700 m. Etymology. The species is named after Alan Graham, who has generously and ably assisted on field excursions many times since our first trip to Mexico for Cuphea in 1961. The recognition of Cuphea alaniana adds a fourth species to subsection Polyspermum of section Melvilla. Species of the subsection are endemic to the western and southern mountains of Mexico. The subsection is characterized by spurless, deeply col¬ ored, thick-bodied floral tubes 20-32 mm long and ovules numbering 40-120. In C. alaniana, the red¬ dish purple floral tube with two large erect, clawed dorsal petals, four small ventral petals, and a mas¬ sive horizontal nectary termed the “disc” serve best to distinguish it from C. micropetala var. hirtella Koehne, to which it is surely most closely related. The latter differs in having six minute petals of equal size and a small disc. Both species share an unusual, dorsally gibbous ovary in addition to sim¬ ilar vegetative, trichome, and pollen morphology. Within the subsection, C. alaniana also resembles the poorly known C. heteropetala Koehne from Oa¬ xaca by its slightly enlarged dorsal lobe and large petals, but C. heteropetala lacks a gibbous ovary and has a small, erect disc. Attempts to relocate C. alaniana in the states of Mexico and Guerrero were unsuccessful. The type area, along the river near Temisco, Guerrero, is now occupied by dwellings and cultivated fields. Exploration further upstream in less disturbed sites also failed to locate the spe¬ cies. The paratype locality, directly north of Tem¬ isco in Mexico, was not accessible by road in 1989, and was not visited by us. Paratypes. MEXICO. Mexico: Cerro de San Antonio, Tlatlaya, Matuda 32110 (GH, MEXU). Cuphea armata S. A. Graham, sp. nov. TYPE: El Salvador. Chalatenango: 4 km SSE of La Pal¬ ma, pine forest, 950 m, 11 June 1970, G. Dav- idse & R. Pohl 2073 (holotype, F; isotype, MO). Figure le. Suffrutices, 0.5-1 m, caules erecti vel suberecti, pilis minutis albidis antrorsis adpressisque. Folia sessilia vel subsessilia, 20-50 mm longa, 5-12 mm lata ad medium caul is, ovata, oblonga, vel sublinearia, basi obtusa vel cor- data, utrinque scabrella. Inflorescentia sat distincta, ra- cemosa, interdum subcomposita; pedicelli gracili, 2-5 mm longi, persistentes; bracteolae 0.5 mm longae, lineari-lan- Volume 5, Number 3 1995 Graham Cuphea 275 ceolatae. Flores 12—15 mm longi, pallidi purpureo-rosei, basi puberuli saepe prominente bulboso-setosi, calcari 4 mm longo, distincte ascendente etiam conico; intus infra stamina bialatus, alae glabrae. Petala 6, 2 dorsalia intense violacea, majora, 7—9 mm longa; 4 ventralia minora 3-5 mm longa, pallide violacea. Stamina 11, 9 longiora exser- ta. Stylus demum exsertus; ovula 8—10. Semina 2 X 1.5 mm, oblonga, margine obtusa. Discus crassus, deflexus. Semi-shrubby perennials, 0.5—1 m, erect or weakly erect and sprawling, irregularly branched from above the base, branches shorter than to lon¬ ger than the main stem, bearing minute white, op¬ pressed, antrorse, sharp-pointed hairs and a lon¬ gitudinal row of minute, falcate, antrorse hairs; intemodes mostly equal to or shorter than the sub¬ tending leaves. Leaves sessile or subsessile, petioles 0-1 mm; blades at midstem 20-50 mm long, 5—12 mm wide, decreasing in size above and below the midstem, membranous, ovate, oblong, or sublinear, base rounded to cordate, apex acute to acuminate; surfaces finely scabrous, appearing glabrous, the hairs minute, appressed, the margins of the upper leaves and bracts ciliolate, bearing sparse bulbous- based setae, upper surface darker green than the lower; upper leaves gradually diminishing in size to become the bracts of the inflorescence. Inflores¬ cences terminal bracteolate racemes or loose, short- branched thyrses; one flower interaxillary at a node, 1—3 additional flowers sometimes present on slen¬ der, alternate, axillary branchlets ca. 1 cm long; pedicels 2—5 mm, slender, persistent; bracteoles 0.5 mm, linear-lanceolate, persistent at the apex of the pedicel. Floral tubes 12—15 mm long, including an ascending, conical spur 4 mm long, the apex sometimes slightly descending, floral tubes dorsally concave, 1-2 mm wide at anthesis, in lateral view the base wider than the midportion, the neck not strongly contracted at anthesis, the mouth oblique, the ventral side extended 1 mm or less beyond the dorsal, the dorsal and adjacent lateral ribs diverg¬ ing approximately 1 mm; outer surface pale rose- purple, darkest dorsally, puberulent with sparse but prominent red-purple, bulbous-based setae present on the dorsal and lateral ribs, especially toward the base of the tube, rarely individual flowers lacking setae, setae 0.5 mm long; inner surface lightly vil¬ lous to glabrous above the stamens in a shallow ventral pouch, glabrous below, the alae glabrous; calyx lobes subequal, broadly triangular, 1.0 X 1.5 mm, margins of the dorsal lobe setulose-ciliolate, rarely glabrous, the others glabrous; appendages 0.5 mm long, terminating in a short bristle. Petals 6, unequal, the 2 dorsal largest, 7—9 mm long, ob¬ long to nearly orbicular, not clawed, deep purple, held erect or reflexed slightly back over the dorsal side of the tube, subtended at the base by a tri¬ angular to comiculate, thickened scale free at the tip for 0.2 mm, the 4 ventral petals 3-5 mm, oblong to obovate, pale purple, held horizontally. Stamens 11, the 2 dorsalmost stamens shortest, included, 9 inserted at ca. 2-3 mm below the rim of the floral tube, exserted; filaments probably pale purple, gla¬ brous; anthers purple. Pollen oval-triangular to nearly spherical, pores very slightly protruding; tri- colporate, syncolpate; exine distinctly and uniform¬ ly striate, striae covering the surface of the grain, and possibly extending to the poles without disag¬ gregation; diameter 18-20 gm in lactic acid. Style ultimately well-exserted; stigma punctiform; ovules 8-10. Seeds 2 X 1.5 mm, oblong, brown, margin thick, rounded. Disc 1 mm long, 0.5 mm wide at base of ovary, thick, triangular in outline, deflexed into the spur. Phenology. Collected in flower and fruit in June, September, and December. Distribution and habitat (Fig. 2). Southeastern Guatemala, Honduras, and El Salvador; pine for¬ ests; 900—1640 m. Etymology. The specific epithet refers to the red-purple, bulbous setae that arm the floral tube and upper bracts of the inflorescence. The species is placed in section Diploptychia by the presence of the key character of the section, the alae, two wings of tissue that extend from the base of the dorsalmost stamens in the floral tube to the base of the tube (Koehne, 1903). Cuphea ar- mata bears close resemblance to Cuphea pinetorum Bentham and C. avigera Robinson & Seaton but differs from both by its unequal, bicolored petals. The prominent reddish, bulbous-based setae and pale purple floral tube further distinguish it from C. pinetorum, which has a white to rose, hirsute and pubescent floral tube. Cuphea armata resembles C. avigera especially by its conical, ascending spur, but lacks the glandular-viscous patches on the stems that distinguish C. avigera. Neither C. pine¬ torum nor C. avigera has been collected within the range of C. armata. Because agamospermy with ac¬ companying infertile pollen is present in several species of the section, pollen of C. armata was test¬ ed for pollen viability. In cotton blue-lactic acid after 24 hours, 99% of all pollen stained deeply, indicating viable protoplasm. The pollen exine seems to lack the meshwork of striae at the poles that is typical of the pollen group to which C. avi¬ gera and C. pinetorum pollen types belong (Gra¬ ham, in prep.), but this needs to be confirmed by SEM observations when more abundant material becomes available. Paratypes. EL SALVADOR. Chalatenango: carretera 276 Novon Figure 3. Distribution and variation in leaf shape, size, and petiole length in Alzatea verticillata. Examples 1-5 represent subspecies amplifolia, characterized by large, broad, oblong, sessile or subsessile leaves. Examples 6-8 represent subspecies verticillata, characterized by small, elliptical, oblong or obovate, distinctly petiolate leaves. Petioles on plants represented by examples 5 and 6 are approximately intermediate between the two subspecies. The major distinction remaining between the subspecies is leaf size, especially leaf width. a La Palma, Montalvo 3682 (MO); Rfo Nunuapa near La Palma, R. Seiler 574 (F). GUATEMALA. Chiquimula: pine forest between Guatemala-Honduras border and Atu- lapa, Molina R. * Molina 25260 (F, MO). HONDURAS. Intibuca: 12.7 mi. N of Marcala along the road from Mar- eala to La Esperanza, Davidse et al. 35191 (MO). Note on Alzateaceae The New World tropical family Alzateaceae is represented by the single species Alzatea verticil¬ lata Ruiz & PavtSn. The species had been placed in eight families in five orders, including the Lyth- raceae, before its segregation to the monotypic fam¬ ily Alzateaceae (Graham, 1984). Alzatea verticillata was described first from the eastern side of the Pe¬ ruvian Andes in 1798 (Ruiz & Pavdn, Syst. Veg. FI. Peruv. Chil. 1: 72). It was considered restricted to Bolivia and Peru until 1936, when it was col¬ lected in the low montane forests of Costa Rica. In 1978, it was discovered in the elfin cloud forest of Veraguas Province, Panama, and subsequently, in 1986, was found in Colombia on the Cerro del Tor- rd, Departamento del Choco (Silverstone-Sopkin & Graham, 1986). The distribution of the species is now further enlarged by collections made by the late Alwyn Gentry on his last field expedition to Ecuador. Two collections. Gentry 80008 and 80334 (MO), from the Cordillera del Condor, near the dis¬ puted Ecuador-Peru border in Morona-Santiago Province, represent the first record of the family in Ecuador, and tenuously establish its continuous distribution in mid- to low-montane forests along the eastern slopes of the mountains from Bolivia to Costa Rica (Fig. 3). The species is one of an esti¬ mated 26% of Ecuadorian species restricted to mid-elevations (900-3000 m) with widespread dis¬ tribution in the Neotropics (Balslev, 1988). The two Ecuadorian collections differ in leaf shape and size, and represent two subspecies. Gen¬ try 80334, collected from an 18-m-tall tree at 2100 m, is typical of the elliptical to oblong or obovate, Volume 5, Number 3 1995 Graham Cuphea 277 petiolate, small-leaved subspecies verticillata from Peru and Bolivia (Fig. 3). Gentry 80008, collected from a 5-m-tall tree at 1350 m, has the broad, ob¬ long, large leaves of subspecies amplifolia, until now known from Colombia, Panama, and Costa Rica. Both collections have short-petiolate leaves and are intermediate for this character between the distinctly petiolate-leaved subspecies verticillata and the sessile-leaved subspecies amplifolia. The distinctions between the subspecies, which were described before the species was known from Co¬ lombia or Ecuador (Graham, 1984), are diminished by the partial intermediacy of the Gentry collec¬ tions. If future collections of Alzatea from northern Peru and Ecuador sample a fuller range of leaf shape and size, it should be easy to decide whether or not to continue recognition of these intraspecific taxa. Leaves of Gentry 80334, among all collections of Alzatea, are the first to display apical leaf glands. The glands are similar to the hydathodes and extra¬ floral nectaries described for several other genera of the Myrtales (Ross & Suessenguth, 1926; Rao & Chakraborti, 1982; Turner & Lersten, 1983; Gra¬ ham, 1984). The shiny, hardened secretion at the Alzatea leaf apex suggests the secretory product is a sugar solution and the gland is probably an extra¬ floral nectary. Specimens examined. ECUADOR. Morona-Santia- go: Gualaquiza Cantdn, Cordillera del Cdndor, ridge top above Banderas, primary forest, 03°28'S, 78°22'W, pri¬ mary forest, 17 July 1993, A. Gentry 80008 (MO); Cam- pamento Achupalla, 15 km E of Gualaquiza, 03 ; 27'S. 78°22'W, dense tangled scrubby forest, 22 July 1993, A. Gentry 80334 (MO). Acknowledgments. I thank Ishmael Calzada for his energetic assistance in the field during our un¬ successful attempts to relocate C. alaniana. I am also grateful to Ronald Liesner, who first deter¬ mined the Alzatea collections, and whose intimate knowledge of plant families has resulted in spe¬ cialists everywhere receiving many interesting and important specimens that otherwise might have gone undetermined or unnoticed among exsiccatae. Wendy Mahon-Hils prepared the illustrations in Figure 1. This research was supported by NSF grant DEB 8806523. Literature Cited Balslev, H. 1988. Distribution patterns of Ecuadorean plant species. Taxon 37: 567—577. Graham, S. A. 1984. Alzateaceae, a new family of Myr¬ tales in the American tropics. Ann. Missouri Bot. Gard. 71: 757-779. Koehne, E. 1903. Lythraceae. Pp. 1-326 in A. Engler (editor). Das Pflanzenreich, IV. 216. Heft 17. H. R. En- gelmann, Leipzig. Rao, T. A. & S. Chakraborti. 1982. A little looked at attribute of the leaves of Sonneratia caseolaris (L.). Curr. Sci. 51: 303-305. Ross, H. & K. Suessenguth. 1926. Das Apikalorgan der Blatter von Lafoensia. Flora 120: 1—18. Silverstone-Sopkin, P. A. & S. A. Graham. 1986. Alza¬ teaceae, a plant family new to Colombia. Brittonia 38: 340-343. Turner, G. W. & N. R. Lersten. 1983. Apical foliar nec¬ tary of pomegranate (Punica granatum: Punicaceae). Amer. J. Bot. 70: 475-480. Muhlenbergia aguascalientensis (Poaceae: Chloridoideae: Eragrostideae), a New Species from Mexico Yolanda Herrera-Arrieta CIIDIR Unidad Durango, COFAA-Instituto Politecnico Nacional, Apartado Postal 738, Durango, Dgo., 34000 Mexico Margarita De la Cerda-Lemus Centro Basico, Universidad Autonoma de Aguascalientes Avenida Universidad s/n, Aguascalientes, Ags., 20100, Mexico Abstract. Muhlenbergia aguascalientensis, from Sierra Frfa, Aguascalientes, Mexico, is described and illustrated. A tabular comparison with its re¬ lated species, M. durangensis Herrera and M. mich- isensis Herrera & Peterson, is given. The genus Muhlenbergia Schreber (Poaceae, Chloridoideae) is comprised of over 160 species (Clayton & Renvoize, 1986), most of which are dis¬ tributed in North and South America. In the south¬ western United States and northern Mexico, species of Muhlenbergia make up about 25% of the grass¬ land flora and provide a considerable amount of forage for livestock (Cronquist et al., 1977). The majority of species are xeromorphic, and a large percentage of the Mexican species are endemic (44% of the M. montana (Nuttall) Hitchcock com¬ plex; Herrera-Arrieta & Grant, 1993). A distinct taxon belonging to this complex was found during an examination of herbarium specimens at HUAA to complete the monographic descriptions of the Muhlenbergia montana complex, and to make a Ho¬ listic study of the grasses from Aguascalientes. Ad¬ ditional field collections were made in the Sierra Fria, Aguascalientes, Mexico, where the species was first collected. The new species is named after this region. Muhlenbergia aguascalientensis Herrera & De la Cerda, sp. nov. TYPE: Mexico. Aguasca¬ lientes: Mpio. de San Jos6 de Gracia, NW of Aguascalientes, 12 km NW from La Congoja, Playa Mariquitas, Monte Grande de la Sierra Frfa, 2850 m, 22°15'6.7"N, 102°37'24.9"W, 8 Nov. 1986, G. Garcia 2801 (holotype, HUAA; isotypes, CIIDIR, ENCB. IEB). Figure 1. A Muhlenbergia michisensis culmis 30-58 cm altis, lig- ulis 0.5-0.7(-l) mm longis, auriculis 1—2(—6) mm longis; laminis basalis 5-10(-15) cm longis; paniculis 5-8(-15) cm longis; spiculis 5-6.5 mm longis; aristis 2—3.5 cm lon¬ gis; antheris 2.4—3 mm longis differt. Caespitose perennial, rhizomatous, 30-58 cm tall. Sheaths up to 1.5 cm long, glabrous to sca- berulous, striate, shorter than the intemodes, with hyaline margins, elongated toward the blade form¬ ing auricles l-2(-6) mm long. Ligules 0.5-0.7(-l) mm long, membranous, hyaline, fimbriate, rounded. Basal blades 5—10(—15) cm long, 1-2.5 mm wide, scaberulous, flat to folded, the margins and central rib sclerosed, scabrous. Panicles 5—8(—15) cm long, 2-4 cm wide, open, loosely flowered, pyramidal, largely exserted, olive, the ascending branches ap- pressed to spreading at maturity, 20—40° from the culm axis, 2—5 cm long; pedicels 1-2.5 mm long, scabrous; inflorescence branches 2—5 cm long; cen¬ tral axis 8-ribbed at the base of the panicle, sca¬ brous. Spikelets 5—6.5 mm long, erect, 1-flowered. Glumes 5-6.5 mm long, oblanceolate, entire, longer than the lemma, usually equal in length, the apex sharply apiculate to mucronate, the mucron 0.5-1 mm long, 1-nerved, olivaceous; second glume with the 2 lateral nerves inconspicuous, scabrous along the entire length. Lemma 4.5-5 mm long, oblong- lanceolate, hyaline to yellowish, awned, glabrous to shortly pilose on the lateral nerves at base; apex acute, minutely bifid, the lobes to 0.2 mm long; awn 2-3.5 cm long, flexuous, yellowish. Palea 4.2-4.8 mm long, oblanceolate, entire, glabrous to pilose on the nerves at the base. Anthers 2.4-3 mm long, purplish. Caryopsis not seen. Chromosome number 2n = 20. Distribution and habitat. Muhlenbergia aguas¬ calientensis is known from northwestern Aguasca¬ lientes, northwest of La Congoja, San Jose de Gra¬ cia, at 2850 m. It occurs in grassland in open forest of Juniperus deppeana Steudel and Quercus side- roxyla Humboldt & Bonpland. Muhlenbergia aguascalientensis resembles M. Volume 5, Number 3 1995 Herrera-Arrieta & De la Cerda-Lemus Muhlenbergia aguascalientensis 279 Figure 1. Muhlenbergia aguascalientensis Herrera & De la Cerda ( G. Garcia 2801). A. Habit. B. Ligule. C. Inflorescence branch. —D. Spikelet. —E. Glumes. —F. Lower glume, dorsal view. —G. Floret. —H. Lemma, lateral view. —I. Palea, dorsal view. 280 Novon Table 1. Characters distinguishing Muhlenbergia aguascalientensis, M. durangensis, and M. michisensis. Characters M. durangensis M. aguascalientensis M. michisensis Rhizomes Present Present Absent Ligule apex Truncate-fimbriate Truncate-fimbriate Acuminate Ligule length 0.5—0.7(-l) mm 0.5—0.7(-l) mm 4—7 mm Leaves Caulinar Basal Caulinar Panicle length 10—25 cm 5—8(—15) cm 12-17 cm Spikelet length 6-7(-7.5) mm 5-6.5 mm 4.5-5 mm Glume length (5—)6—7(—7.5) mm 5-6.5 mm 4.5—5 mm Lemma length 5-6.7(-7) mm 4.5-5 mm 3.5—4 mm Lemma apex Entire Minutely bifid Minutely bifid Lemma lobes length 0.0 mm 0.2 mm 0.2 mm Lemma awn length 15—20 mm 20-35 mm 12-13 mm Palea length 5-6 mm 4.2-4.8 mm 2.8-3.5 mm Palea apex Entire-apiculate Entire Bilobed Anther length (2.5—)3—3.5 mm 2.4—3 mm 2—2.4 mm michisensis Herrera & Peterson and M. durangensis Herrera; however, these two species differ in many morphological characteristics (Table 1). Muhlenber- gia aguascalientensis is similar to M. flaviseta Scribner, a species of restricted distribution in the Sierra Madre Occidental, sharing with it macro- morphological features such as basal leaves and a long-exserted panicle, but differing in most of the spikelet features. The flavonoid chemistry of M. aguascalientensis is very similar to M. eriophylla Swallen and M. michisensis Herrera & Peterson. They share: quer¬ cetin 3-0 rhamnosilxyloside, luteolin 6 galactoside, luteolin 6 glucoside, luteolin 8 glucoside, apigenin 7-0 arabinoside, apigenin 7-0 diarabinoside, api¬ genin 7-0 glucoside, tricin 5-0 glucoside, tricin 7- 0 glucuronide, and 4'hydroxyflavone 7-0 glucoside (Herrera & Bain, 1991). Muhlenbergia aguascalien¬ tensis also contains luteolin 6 xylosilglucoside, which was previously thought to be a unique com¬ pound of M. durangensis. Paratypes. MEXICO. Aguascalientes: Playa Mari- quitas. Sierra Frfa, 12 km NW from La Congoja, San Jos6 de Gracia, 22°15'6.7"N, 102°37'24.9"W, 2850 m, 25 Nov. 1993, Herrera & De la Cerda 1185 (CIIDIR, HUAA, IEB, ENCB, MEXU, US). Acknowledgments. This study was partially supported by the Banco de Mexico, COFAA-IPN and UAA. Literature Cited Clayton, W. D. & S. A. Renvoize. 1986. Genera Gramin- um. Grasses of the World. Kew Bull., Addit. Ser. XIII: 1—389. Her Majesty’s Stationery Office, London. Cronquist, A., A. H. Holmgren, N. H. Holmgren, J. L. Reveal & P. K. Holmgren. 1977. Intermountain Flora. Vol. 6, The Monocotyledons. Columbia Univ. Press, New York. Herrera A., Y. & J. F. Bain. 1991. Flavonoid profiles in the Muhlenbergia montana complex (Poaceae). Bio- chem. Syst. Ecol. 19: 665-672. Herrera-Arrieta, Y. & W. F. Grant. 1993. Correlation be¬ tween generated morphological character data and fla¬ vonoid content of species in the Muhlenbergia montana complex. Canad. J. Bot. 71: 816-826. New Taxa and Combinations in the Tribe Juanulloeae (Solanaceae) Sandra Knapp Department of Botany, The Natural History Museum, Cromwell Road, London SW75BD, United Kingdom ABSTRACT. A new species of Trianaea from east¬ ern Ecuador, T. naeka, is described and illustrated. Several new combinations in the genera Trianaea, Schultesianthus, and Juanulloa are made in prep¬ aration of a larger work on the phylogeny of the tribe Juanulloeae. A recent phylogenetic study of the tribe Juan¬ ulloeae (Persson et al., 1994) has necessitated sev¬ eral new combinations in the genera Trianaea, Schultesianthus, and Juanulloa. These nomencla- tural changes are made here in preparation for a conspectus of the tribe and a more ample discus¬ sion of relationships within the groups. In addition to the nomenclatural changes identified, several new taxa were discovered, one of which is de¬ scribed here. Other new taxa will be described as more material becomes available. Trianaea naeka S. Knapp, sp. nov. TYPE: Ecuador. Morona Santiago: Colinas, above the Mision Sa- lesiana-Shuar, 800-1000 m, 3°25'S, 78°35'W, 6 Nov. 1986, Neill & Cerdn 7449 (holotype, MO; isotypes, NY, QCNE). Figure 1. Species Trianaeae nobili Planchon & Linden similis, sed foliis linearis pubescentibus, inflorescentiis uniflori- bus differt. Epiphytic shrub or liana at forest edge; stems densely hirsute, the trichomes 4—5 mm long, con¬ sisting of a simple, 2-4-celled, uniseriate tip aris¬ ing from a multicellular, multiseriate, pustular base, the apex minutely glandular, golden brown, the uniseriate tips of the trichomes often deciduous, leaving the pustulate base; bark of older stems pale tan, loose and somewhat exfoliating. Leaves 0.4—1 X 5-10 cm, linear, coriaceous, drying dark olive green, the venation obscure, ca. 8—10 primary veins, densely hirsute with simple, uniseriate, mi¬ nutely glandular trichomes like those of the stems, but lacking the pustular bases, arising directly from the epidermis, the base attenuate, the margins rev¬ olute, the apex acuminate; petiole 5—8 mm, some¬ what winged from the attenuate leaf base, drying dark brown. Inflorescence terminal, consisting of a single flower. Buds not known. Pedicel at anthesis 18-20 cm long and flexuous, 2 mm diam. at the base, ca. 10 mm diam. at the apex, densely pubes¬ cent with simple uniseriate 2—4-ee]|ed trichomes, the trichomes 4—5 mm, less rigid than those of the stems, lacking pustulate bases. Flowers with the calyx tube 1.5-2 cm, conical, the lobes 4—5 cm, triangular, pubescent with simple uniseriate tri¬ chomes like those of the pedicel, sparsely pubes¬ cent within; corolla campanulate, greenish brown or purplish (“clear lila” fide van der Wetff & Gudifio 11246), the tube flaring, ca. 5 cm long, the lobes rounded, 1.5—2 cm long, reflexed at anthesis, sparsely pubescent with trichomes like the calyx, the trichomes weak and crumpling; stamens with the filaments ca. 3.5 cm, inserted in the basal Vs of the corolla tube, the anthers ca. 7X3 mm, ven- trifixed; ovary glabrous, the style 5.5-6 cm long, glabrous, the stigma globose, ca. 5 mm diam., pa¬ pillose. Fruit and seeds not known. Distribution. Known only from wet premontane rainforest in eastern Ecuador, from 800 to 1000 m elevation. Trianaea naeka is a striking plant and cannot be confused with any other species of Trianaea, or with any other species of epiphytic Solanaceae. The nar¬ rowly linear, densely pubescent leaves are found nowhere else in the tribe. Pollen morphology is of the Trianaea nobilis type (Persson et al., 1994), where the exine ornamentation is reticulate at the equator and at the poles but the muri are not joined in the mesocolpium to form a tectum. The specific epithet, naeka, is the Shuar name (given by Antonio Wampash, Shuar informant) for this plant. Paratype. ECUADOR. Morona Santiago: along new road Mendez-Morona, km 30—35, 800 m, 19 Aug. 1989, van der Werjf A' Gudifio 11246 (QCNE). Trianaea brevipes (Cuatrecasas) S. Knapp, comb, et stat. nov. Basionym: Trianaea spectabilis Cuatrecasas var. brevipes, J. Wash. Acad. Sci. 49: 271. 1959. TYPE: Colombia. Cundina- marca: Monterredondo between Guaybetal and Limoncito, Cordillera Oriental, forests, 1800— 2100 m, 6 July 1955, Vogel 6 (holotype, US; isotype, M). 282 Novon MISSOURI BOTANICAL GARDEN HERBARIUM N? 4355395 Figure 1. Holotype of Trianaea naeka S. Knapp (Neill & Cerdn 7449, MO). Volume 5, Number 3 1995 Knapp Tribe Juanulloeae (Solanaceae) 283 Schultesianthus venosus (Standley & C. V. Mor¬ ton) S. Knapp, comb. nov. Basionym: Markea verwsa Standley & C. V. Morton, Publ. Field Mus. Nat. Hist., Bot. 18: 1065. 1938. TYPE: Costa Rica. Alajuela: Vara Blanca de Sarapi- quf, 1770 m, Oct. 1937, Skutch 3355 (holo- type, US 1643419). Schultesianthus crosbianus (D’Arcy) S. Knapp, comb. nov. Basionym: Markea crosbiana D’Arcy, Ann. Missouri Bot. Gard. 60: 654. 1973 [1974], TYPE: Panama. Chiriquf: Cerro Horqueta NW of Boquete, 5000-5800 ft., Dwyer et al. 430 (holotype, MO). Schultesianthus uniflorus (Lundell) S. Knapp, comb. nov. Basionym: Markea uniflora Lun¬ dell, Phytologia 1: 340. 1939. TYPE: Mexico. Chiapas: Volcan Tacana, 2800 m, 27 Mar. 1939, Matuda 2816 (holotype, MICH; iso¬ types, MEXU, MO). Juanulloa wardiana (D’Arcy) S. Knapp, comb, nov. Basionym: Rahowardiana wardiana D’Arcy, Ann. Missouri Bot. Gard. 60: 671. 1973 [1974]. TYPE: Panama. Cocld beyond La Mesa towards El Llano and the Panama/ Code border N of El Valle de Anton, 800-860 m, Luteyn 3150 (holotype, DUKE, isotype, MO). Juanulloa globifera (S. Knapp & D’Arcy) S. Knapp, comb. nov. Basionym: Rahowardiana globifera S. Knapp & D’Arcy, Novon 3: 429. 1993. TYPE: Colombia. Antioquia: Parque Nacional Natural “Las Orquideas,” Sector Venados, 890 m. 6°33'N, 76°19'W, 1 Apr. 1988, Cogollo et al. 2895 (holotype, JAUM- 018167; isotypes, COL, FMB, JAUM, MO). Acknowledgments. I thank Viveca Persson and Steve Blackmore, my coworkers on pollen mor¬ phology of Solanaceae; Roy Vickery and the cura¬ tors of the herbaria mentioned in the text for help with the loan of specimens; Tamara Nunez and Da¬ vid Neill for asistance at QCNE; and the Photo¬ graphic Unit of the Natural History Museum for the photographic plate. Literature Cited Persson, V., S. Knapp & S. Blackmore. 1994. Pollen mor¬ phology and systematics of tribe Juanulloeae A. T. Hun- ziker (Solanaceae). Rev. Palaeobot. Palynol. 83: 1-30. A New Species of Sicydium Schlechtendal (Cucurbitaceae: Zanonioideae, Zanonieae, Sicydiinae) for the Flora Mesoamericana Rafael Lira Herbario Nacional de Mexico, Departamento de Botanica, Instituto de Biologfa, UNAM, Apdo. Postal 70-367, Delegacion Coyoacdn, 04510, Mexico, D.F. Abstract. Sicydium daviliae, from the region of Ocozocuautla, Chiapas, Mexico, is described as new. It can be distinguished from the other species in the genus by the presence of a soft and dense pubescence composed of long, multicellular, non- glandular hairs covering stems, blade surfaces, pet¬ ioles and tendrils, by filaments 2—3 times longer than anthers, by petal margins entire, and by sta- minate pedicels articulate at the middle or further up. Resumen. Se describe como nueva especie a Si¬ cydium daviliae, de la region de Ocozocuautla, Chiapas, Mexico. Esta nueva especie se distingue de las otras especies del genero por la pubescencia densa y suave, compuesta de tricomas largos, mul- ticelulares y no glandulares, la cual cubre los tal- los, ambas superficies de la lamina, los peciolos y los zarcillos, ast como tambi£n por presentar fila- mentos 2-3 veces mas largos que las anteras, pe¬ tal os con los mdrgenes enteros y pedicelos estam- inados articulados en la mitad o poco mas arriba. As a result of examination of herbarium speci¬ mens for the treatment of Sicydium for Flora Me¬ soamericana, one specimen was found whose sta¬ men and vestiture features set it apart as a new species, described below. Sicydium daviliae Lira, sp. nov. TYPE: Mexico. Chiapas: Mpio. Oeozocuauda, Camino Horizon¬ te a Santa Laura, 12 June 1983, F. Vdzquez B. 1024 (male plant) (holotype, XAL). Figure 1. Sicydio tamnifolio (HBK) Cogniaux similis, sed differt staminum filamentis quam antheris 2—3 plo longioribus, foliorum, caulium, petiolorumque indumento densiore, tri- chomatibus longis apice non glandularibus constato, pe- talorum marginibus integris, florum masculorum pedicellis in parte media vel altiore articulatis. Herbaceous vine, perennial, dioecious, drying yellowish green; stems, petioles, upper and lower surfaces, and tendrils densely pubescent to tomen- tose, with long (1—2 mm), soft multicellular, non- glandular hairs. Stems angulate-sulcate. Petioles angulate-sulcate, l-1.5(-2) cm long; blades subcoriaceous, ovate-cordate, 3.5—5.5 cm long, 3— 5 cm broad, entire, apex acute to shortly acuminate, margins entire to slightly undulate-dentate, basal sinus cordate. Tendrils simple. Flowers small, ar¬ ranged in loose panicles. Staminate panicles 4-6 cm long, the branches commonly with a reduced leaf in the axils; pedicels filiform, 1—1.3 mm long, articulate at the middle or a little above the middle, subtended by a lanceolate reduced leaf; receptacle disk-shaped, less than 0.2 mm long, 1-1.2 mm wide, puberulent; sepals 5, triangular, ca. 0.6 mm long, 0.3 mm wide, puberulent; petals 5, ovate-lan¬ ceolate, spreading, greenish cream or greenish white, 1—1.3 mm long, 0.5-0.6 mm wide, puberu¬ lent, the margins entire; stamens 3, totally free, two with bithecial anthers, one with a monothecial an¬ ther; filaments inserted in the base of the recepta¬ cle, 0.4-0.6 mm long, slightly bent inward, forked toward the apex in the stamens with bithecial an¬ thers; anthers oblong-elliptic, 0.2—0.3 mm long, opening by a longitudinal suture. Plants with pis¬ tillate flowers and fruits unknown. Sicydium daviliae is known only from the type, collected in the Municipio Ocozocuautla, Chiapas, Mexico, growing in disturbed tropical rainforest at 900 m, on sandy, clayish, and rocky soils. The approximately eight species of Sicydium are distributed in the Neotropics. They have been tra¬ ditionally distinguished mainly on the basis of sta¬ men structure, and presence/absence and distri¬ bution of vestiture (Cogniaux, 1881, 1916; Dieterle, 1976; Jeffrey & Trujillo, 1992; Nee, 1993; Schle¬ chtendal, 1832; Wunderlin, 1978). So far, three species were found in Mexico—Mesoamerica: S. tamnifolium (HBK) Cogniaux, a species ranging from central Mexico to South America, and S. tuerckheimii J. Donell-Smith and S. schiedeanum Schlechtendal (= S. glabrum Standley & Steyer- mark), both ranging at least from southern Mexico to Central America. Volume 5, Number 3 1995 Lira Sicydium daviliae 285 0.5 mm Figure 1. Sicydium daviliae Lira. —A. Habit. —B. Staminate flower. —C. Comparison among the tnchomes present on the leaves of S. daviliae (1), S. tuerckheimii (2), and S. tamnifolium (3). Based on type specimen F. Vdzquez B. 1024 (XAL). 286 Novon Sicydium daviliae can readily be distinguished from the above-mentioned species by its unique combination of stamens with filaments 2-3 times longer than the anthers; petals with entire margins; a dense, soft indument made up of long, multicel¬ lular, nonglandular hairs; and staminate pedicels that are articulate at or above the middle. In con¬ trast, Sicydium tamnifolium includes populations with leaf blades, stems, and petioles densely and softly pubescent, but the pubescence is less dense and the hairs are much shorter and gland-tipped. In addition, the stamens of S. tamnifolium have very short or obsolete filaments, the petal margins are somewhat fimbriate or laciniate, and the pedi¬ cels of the staminate flowers are articulate near the base. A species with a similar vestiture to that of Si¬ cydium daviliae is S. araguense Steyermark & Tru¬ jillo from Venezuela (Jeffrey & Trujillo, 1992). However, in S. araguense the stems are only pu- berulent with age, in adult leaves the pubescence is not as dense or soft, and the leaf blades are nar¬ rowly lanceolate and longer. The new species is named to honor Patricia Davila, Mexican agrostol- ogist. A Key to the Mexican/Mesoamerican Species of Sicydium la. Plants essentially glabrous or only slightly pu- berulent; leaves always unlobed, usually lustrous in both surfaces; filaments 2-2.5 times longer than anthers.5. schiedeanum Schlechtendal lb. Plants conspicuously soft-puberulent, pubescent, or tomentose at least on the lower surface of the leaves; leaves unlobed to angulate or slightly lo- bate, not lustrous; filaments obsolete to 3 times longer than anthers. 2a. Leaves pubemlent and rough-scabrous on the upper surface, densely and softly pubes¬ cent on the lower surface, the pubescence commonly dark reddish brown when dry; fil¬ aments 2-3 times longer than anthers . . . .S. tuerckheimii J. Donell-Smith 2b. Leaves pubescent, puberulent to tomentose on both surfaces, the pubescence yellowish or whitish in dry specimens; filaments ob¬ solete to 2—3 times longer than anthers. 3a. Plants puberulent to villous, the vesti¬ ture mainly of glandular hairs; filaments obsolete; margins of the petals slightly fimbriate or laciniate; pedicels of the staminate flowers articulate near the base . ... S. tamnifolium (HBK) Cogniaux 3b. Plants densely and softly pubescent to tomentose, the vestiture mainly of non¬ glandular hairs; filaments 2—3 times longer than anthers; margins of the pet¬ als entire; pedicels of the staminate flowers articulate in the middle or slightly above. S. daviliae Lira Acknowledgments. I thank the curators of BM, BR, CHAPA, ENCB, F, HUH, K, LL, MICH, MO, NY, P. TEX, US, and XAL for allowing me to review collections of Sicydium, especially Sergio Avendano (XAL) for making available the specimen on which the description of the new species is based. Eloisa Duarte prepared the excellent drawings, and David Sutton (BM), Fernando Chiang, and two anonymous reviewers commented critically on the manuscript; Fernando Chiang also prepared the English version of the paper, as well as the Latin diagnosis. Literature Cited Cogniaux, A. 1881. Cucurbitacfes. Pp. 325-951 in A. de Candolle & C. de Candolle (editors), Monographiae Phanerogamarum. G. Masson, Paris. -. 1916. Cucurbitaceae—Fevilleae et Melothrieae. Pp. 1—277 in A. Engler (editor). Das Pflanzenreich. Regni Vegetabilis Conspectus. Verlag von Wilhelm En- gelmann, Leipzig. Dieterle, J. V. A. 1976. Cucurbitaceae. In: Flora of Gua¬ temala. Part XL Fieldiana Bot. 24: 306-395. Jeffrey, C. & B. Trujillo. 1992. Cucurbitaceae. Pp. 11- 201 in G. Morillo (editor). Flora de Venezuela. Fondo Editorial Acta Cientffica Venezolana, Caracas, Venezue¬ la. Nee, M. 1993. Cucurbitaceae. In: V. Sosa (editor). Flora de Veracruz. Fasctculo 74. Instituto de Ecologfa A.C./ University of Riverside, Xalapa, Veracruz. Schlechtendal. D. F. L. 1832. De Plantis Mexicanis. Lin- naea 7: 380-400. Wunderlin, R. P. 1978. Cucurbitaceae. In: R. E. Wood- son, Jr. & R. W. Schery (editors). Flora of Panama. Part IX. Ann. Missouri Bot. Card. 65: 285—366. Conceveiba parvifolia (Euphorbiaceae), a New Species from Panama and Colombia Gordon D. McPherson Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, U.S.A. ABSTRACT. Conceveiba parvifolia from Darien, Panama, and Antioquia, Colombia, is described and illustrated. Two wind-thrown trees encountered on the for¬ ested slopes above the abandoned Cana gold mine in Darien are apparently the first known represen¬ tatives of the genus Conceveiba Aublet from Pana¬ ma, although the genus was previously known from both neighboring Costa Rica and Colombia. Spec¬ imens from these two trees key readily to the sub¬ tribe Conceveibinae of the Alchomeae (Acalyphoi- deae) in Webster’s synopsis (1994). Within that subtribe, however, the Panamanian collections combine characters currently used to distinguish Conceveiba and Gavarretia Baillon, i.e., on the one hand the sepals of the pistillate flower are separate (usually), the style branches blunt, and the anthers muticous (as in Conceveiba), while on the other hand the sepals of the pistillate flower are eglan- dular, and the ovary is 2-locular (as in Gavarretia, although some species of Conceveiba are similarly eglandular). The generic boundaries were not clear in this subtribe even before this intermediate taxon further blurred them, and I have chosen to name the new species in Conceveiba since I agree with Webster (1994) that Mueller’s inclusion (1874) of Gavarretia as a section of Conceveiba may be jus¬ tified. Conceveiba parvifolia McPherson, sp. nov. TYPE: Panama. Darien: near Cana mine S of El Real (Cerro Pirre massif), 7°45'N, 77°40'W, ca. 900 m, 26 Aug. 1987. McPherson 11607 (holotype, PMA; isotypes, BM, COL. CR, DAV. EAP, F, G, GH. HUA. K, MEXU, MICH, MO, NY, P, US). Figure 1. Species foliis minoribus (2.7—16.5 cm X 0.8-5.5 cm), infiorescentiis masculis brevibus (10 cm) fere eglandula- tis, floribus masculis noil fasciculatis, sepalis floris femi- nei 4(—5) eglandulatis, ovario biloculari, stylis 2 semicon- natis a congeneribus diversa. Tree (6-) 15-20 m; colored exudate absent; youn¬ gest branches stellate-puberulent, without lenticels, terete or somewhat ridged, the older ones glabrate and bearing elongated lenticels. Leaf blade (2.7—) 3.5- 8.0(-16.5) cm long, (0.8-)1.8-4.0(-5.5) cm wide, elliptical-oblong, obtuse at the base, acumi¬ nate at the apex, the margin crenate (sometimes obscurely so) with 5—8 mucronate teeth, the upper surface sparsely stellate-puberulent before maturi¬ ty, rapidly glabrate, the lower surface densely stel¬ late-puberulent before maturity, eventually gla¬ brate, often with 1—several embedded glands, secondary veins 6—10 on each side of the midrib, the reticulum slightly raised on the upper surface and more strongly raised on the lower; petiole 1.3— 4.0(—4.8) cm long, 0.5-1.0 mm diam., semicircular in cross section, sparsely stellate-puberulent, a dis¬ tal pulvinus usually discolored and thereby distin¬ guishable, 2-3 mm long, sometimes slightly swol¬ len, a smaller proximal pulvinus often present; stipules subulate, 2—4 mm long, up to 0.5 mm wide, stellate-puberulent, quickly caducous. Plants di¬ oecious; inflorescences terminal or from the axils of the most distal leaves. Staminate inflorescences paniculate or less often racemose, up to 10 cm long, the axes stellate-puberulent, bracts triangular, up to 2 mm long, stellate-puberulent, each bract often accompanied by two smaller bracteoles, both bracts and bracteoles sometimes bearing weakly differ¬ entiated glabrous glandular patches; pedicels 1—1.5 mm, not fasciculate, glabrous; buds ca. 2 mm diam., apiculate; sepals valvate, 2 or 3 in number, 2.5- 3.5 mm long, up to 3 mm wide, concave, eglan¬ dular, green, glabrous without, somewhat granular- puberulent within at base of filaments; petals and disk absent; stamens 35-42, filaments up to 3 mm long, free or the central ones connate basally, gla¬ brous, anthers ca. 0.8 mm long, rounded, somewhat puberulent; pistillode absent. Pistillate inflores¬ cences racemose, up to 5 cm long (occasionally the most proximal flowers subtended by only slightly modified leaves), the axes stellate-puberulent, eglandular; bracts not seen, rapidly caducous; ped¬ icels 4—13(—18) mm long, somewhat inflated dis- tally, stellate-puberulent, usually bearing two bracteoles ca. 2 mm long, these caducous, usually attached near the pedicel base but sometimes at¬ tached subapically and then mistakable for sepals; Nnvniv 5- 9R7-98Q 1QQ5 288 Novon Figure 1. Conceveiba parvifolia McPherson. —A. Twig bearing staminate inflorescences. —B. Portion of stammate inflorescence. —C. Staminate flower. —D. Pistillate inflorescence. —E. Pistillate flower with 4 sepals and a bracteole. (A—C based on McPherson 11621; D, E based on McPherson 11607.) Volume 5, Number 3 1995 McPherson Conceveiba parvifolia 289 sepals 4(-5) in number, 2-5 mm long, subulate, free or occasionally two of them partially connate, eglandular, pale green, stellate-puberulent; petals, disk, and staminodes absent; pistil 6-9 mm long, stellate-puberulent, ovary ca. 3 mm long, 2-locular; styles 2 in number, fused basally into a column 1.5—3 mm long, free and bifid distally, these free portions stigmatic adaxially and ca. 3 mm long in total, the ultimate branches ca. 2 mm long, blunt- tipped; ovules 1 per locule. Fruit unknown. From the seven or eight other members of its genus and from the monotypic Gavarretia, Concev¬ eiba parvifolia can be separated by its smaller leaves, shorter and only obscurely glandular sta- minate inflorescences, non-fasciculate staminate flowers, and pistillate flower with 4(—5) eglandular sepals, a bilocular ovary, and 2 styles connate half their length. At the suggestion of Grady L. Webster, I exam¬ ined two additional specimens that proved to be¬ long to the new species. These collections, made by R. Callejas, F. J. Roldan, and J. Betancur of HUA, extend the distribution of Conceveiba parvi¬ folia from easternmost Central America well into the northwestern corner of South America. Paratypes. PANAMA. Darien: near Cana mine S of El Real (Cerro Pine massif), 7°45'N, 77°40'W, ca. 900 m, 26 Aug. 1987, McPherson 11621 (BM, COL, CR, DAV, F, HUA, K, MEXU, MICH, MO, NY, PMA, US). COLOM¬ BIA. Antioquia: Municipio Anon, via Dos Bocas-Provi- dencia, 7°26'N, 74°56'W, 100-120 m, 9 jul. 1987, Cal¬ lejas et al. 4477 (MO); Municipio Remedies, 10-12 km NW de Remedios en la via a Zaragoza, 7°05'N, 74°15’W, 300 m, 16 sep. 1987, Callejas et al. 5261 (MO). Acknowledgments. I thank David Olson, who accompanied me to Cana to collect ants, and John Myers, who drew the illustration. Literature Cited Mueller, J. 1874. Euphorbiaceae [part 2], Pp. 293-750 in Martius, Flora Brasiliensis, Vol. 11. Cramer, Wein- heim. Webster, G. L. 1994. Synopsis of the genera and supra- generic taxa of Euphorbiaceae. Ann. Missouri Bot. Card. 81: 33-144. A New Votomita (Melastomataceae) from Panama Thomas Morley Department of Plant Biology, University of Minnesota, St. Paul, Minnesota 55108, U.S.A. Frank Almeda Department of Botany, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118-4599, U.S.A. Abstract. A new species of Votomita, V. cupuli¬ formis, has been found on the north side of western Panama, the first member of the genus known to occur in Central America. The new species is most similar to V. roraimensis Morley of central Roraima, Brazil. Distinguishing features are discussed, and a key to all nine species of the genus is provided. Votomita cupuliformis Morley & Almeda, sp. nov. TYPE: Panama. Bocas del Toro: Laguna de Chiriqui, rincon Sur-Este Nuri, campamen- to cerca pequena poblacion Guaymi, 15 km Oeste de Punta Cricamola, entrando Ensenada de Catavela, y subiendo Quebrada Nuri, 8°55'N, 81°49'W, unos 5 km al Sur del cam- pamento, 10-25 m, 18 mar. 1993, Robin Fos¬ ter, A. Herre, E. Kalko <£ C. Handley 14563 (holotype, CAS; isotypes, MO, PMA, SCZ, US). Figures 1, 2. Arbor glabra usque 10 m alta: cryptae stomatophorae Typi 11. parvae, simplices; epidermis supera folii stratiis 1 cellularum; limbus calycis cupuliformis lobis apiculatis; stamina monadelpha; ovarium 2-loculare ovuliis 16 axil- laribus. Glabrous tree 10 m high; young twigs rounded. Petioles 4—6 mm long; blades 6.5-11.9 cm long, 2- 4-4.5 cm wide, elliptic to elliptic-ovate or seldom slightly obovate, abruptly acuminate at apex, acute to abruptly short-attenuate on the petiole at base; midrib grooved above, narrowly 2-winged below; lateral nerves obscurely visible to invisible above and below when dry. Midrib xylem tubular; sto- matal crypts Type II or borderline I—II (see Morley, 1976); upper epidermis one cell thick, lacking mu¬ cilaginous walls; hypodermis none; foliar sclereids all terminal, variable in shape, mostly columnar with large branches at each end, sometimes slant¬ ing, sometimes with a horizontal central body at veinlet level bearing branches upward and down¬ ward. Peduncles terminal, axillary, and at the up¬ permost leafless nodes, 1 per side, of 1 intemode 3- 7 mm long, 1-flowered; bracts narrowly Novon 5; 290-293. 1995. triangular, acute, entire, 1.2—1.5 mm long, decid¬ uous at anthesis. True pedicels 8-11 mm long; flowers known only from buds, the buds yellow; flowers 4-merous; calyx including inferior ovary 4.5—5 mm long, obconic to slightly campanulate, 1.8-2.2 mm long, the surface flecked with scurfy patches at least when moist; calyx limb abruptly expanded and shallowly cup-shaped, 2.5 mm long, ca. 3.8 mm diam.; calyx lobes 0.7 mm long, 2.5 mm wide, broadly rounded-triangular and promi¬ nently apiculate. Immature petals ovate, acute, ses¬ sile, adherent to each other in bud, 8 mm long, ca. 3.5 mm wide. Stamens monadelphous, 4.24.5 mm long, 0.9-1.0 mm wide, adherent on the back to the petals in bud, this condition at anthesis un¬ known; filaments ca. 0.4 mm long; anthers 3.74.2 mm long; thecae 2.4—2.7 mm long; gland adaxial at the apex, shallowly concave, 0.7 mm long, 0.4— 0.5 mm wide. Ovary 2-locular with axile placen- tation, each locule with ca. 8 ovules crowded at different levels on the placenta, the ovules 16 in all. Calyx prominent on the immature fruit; ripe fruit and seed unknown. Distribution. Known only from the type locality near the southeast comer of the Laguna de Chiriqui on the north side of western Panama. “Terra firme, bosque alto, humedo sobre filos y pendientes.” The new species is the first member of Votomita known to occur in Central America. Its leaves are overall the simplest in the genus in their average size and shape, lack of hairs, undivided upper epi¬ dermis, strictly terminal foliar stone cells with only a moderate columnar tendency, and small simple stomatal crypts. The flowers, however, are of an intermediate level of reduction from the pre¬ sumed ancestral type: in the nine species of Vo¬ tomita there is a general trend from large-flowered plants with 4-locular ovaries and numerous ovules to small-flowered ones with 1 locule and 5—10 ovules. Votomita cupuliformis has a 2-locular ova¬ ry with 16 ovules; in this feature the plant is most similar to V. roraimensis of central Roraima, Bra- Volume 5, Number 3 1995 Morley & Almeda Votomita cupuliformis from Panama 291 Figure 1. Votomita cupuliformis Morley & Almeda, habit drawing (by Ellen del Valle). zil, which has 4 locules and 15-16 ovules. The new species is easily distinguished from V. rorai- mensis, however, by the former’s longer petioles, simpler leaf anatomy, larger petals and stamens, and 2-locular ovary. A key is provided to all nine species to illus¬ trate how the three named since the latest mono¬ graph (V. cupuliformis, V. pubescens, V. roraimensis) compare to the six included in the monograph (Morley, 1976): la. Calyx including inferior ovary 7.5-10.5 mm long; ovules 20-48. 2a. Leaf blades pubescent beneath, 10.5—15.5 cm long. V- pubescens Morley 2b. Leaf blades (excluding the midrib) glabrous beneath, 7—12 cm long. за. Petioles 3—5 mm long, placentation axile. V. plerocarpa (Morley) Morley 3b. Petioles 7-12 mm long; placentation basal in each locule . . V. monadelpha (Ducke) Morley lb. Calyx including inferior ovary 2.2—5.6 mm long; ovules 5—16 and 36. 4a. Calyx including inferior ovary 5-5.6 mm long; stamens 5.“T-6.1 mm long; ovules ca. зб, in 4 whorls. V. orbinaxia Morley 292 Novon F igure 2. Votomita cupuliformis. —A. Leaf shapes (traced). B—I (camera lucida): —B. Cleared portion of leaf blade showing veins and terminal sclereids. —C. Cross section of leaf blade showing sclereids, stomatal crypts, and upper epidermis. D. Cross section of leaf midrib. —E. Large flower bud. —F. Longisection of flower before anthesis. G. Petal. —H. Anthers, face and side views. —1. Immature fruit. (All by TM.) Volume 5, Number 3 1995 Morley & Almeda Votomita cupuliformis from Panama 293 4b. Calyx 2.2-5 mm long; stamens 2.9—5.1 mm long; ovules 5-16. 5a. Calyx including inferior ovary 4.5—5 mm long; ovules 15-16. 6a. Petioles 1.5-2.5 mm long; petals ca. 4.7-5 mm long; anthers 2.6-2.8 mm long; thecae 1.2-1.3 mm long; ovary 4-locular. . . V. roraimensis Morley 6b. Petioles 4—6 mm long; petals ca. 8 mm long; anthers 3.7—4.2 mm long; thecae 2.4-2.7 mm long; ovary 2- locular . . . . V. cupuliformis Morley & Almeda 5b. Calyx 2.2^1 mm long; ovules 5-10. 7a. Apex of blade abruptly acuminate; petioles 4—9 mm long; blades (7—) 9—12 cm long . . . V guianensis Aublet 7b. Apex of blade rounded to acute and often mucronulate; petioles 1.5-4 mm long; blades 2.5-7.5 cm long. 8a. Blades 22—42 times as long as the petiole; pedicels 1—1.3 mm diam. 2 mm below the calyx teeth when dry . . V. orinocensis Morley 8b. Blades 10-20 times as long as the petiole; pedicels 1.5—2.3 mm diam. 2 mm below the ca¬ lyx teeth when dry . . . . .V. monantha (Urban) Morley Literature Cited Morley, T. 1976. Memecyleae (Melastomataceae). FI. Neotrop. 15: 1—295. New Names in Papaver Section Meconella (Papaveraceae) David F. Murray Herbarium (ALA), University of Alaska Museum, Fairbanks, Alaska 99775-6960, U.S.A. Abstract. Two combinations ( Papaver macounii subsp. discolor and P. radicatum subsp. kluanensis) and a subspecies ( P. nudicaule subsp. americanum) are published. To complete a treatment of the scapose poppies (section Meconella of Papaver) for Flora of North America, it is necessary to publish one new com¬ bination and to validate one combination and a subspecies that have been previously proposed, but not in compliance with the International Code of Botanical Nomenclature (Greuter et al., 1994). Papaver macounii subsp. discolor (Hulten) Ran- del ex D. F. Murray, stat. nov. Basionym: Pa¬ paver macounii var. discolor Hulten in Hulten, FI. Alaska Yukon 5: 803. 1945. TYPE: U.S.A. Alaska: Nome, Anderson 3250 (holotype, S). This combination was made by Randel (1977: 425) without a full and direct reference to the bas- ionym’s author and place of valid publication as required by Article 32.2 (Greuter et al., 1994), which I have now supplied. Papaver nudicaule subsp. americanum Randel ex D. F. Murray, subsp. nov. TYPE: [Canada.] Klotassin area Yukon T[erritory], SW of Yukon River, between Selkirk and White River, Aug. 1916, D. D. Cairnes 91890 (holotype, CAN). Randel (1977: 437) determined that the speci¬ mens from Alaska and Yukon represented a taxon distinct from the Asiatic material she had studied. She proposed Papaver nudicaule subsp. american¬ um, for which she supplied a diagnosis but failed to select a type as required by Article 37.1 (Greuter et al., 1994). As holotype I have chosen a specimen complete with leaves, flowers, and fruits, which Randel had seen and annotated as subspecies americanum. Papaver radicatum subsp. kluanensis (D. Love) D. F. Murray, comb, et stat. nov. Basionym: Papaver kluanensis D. Love in Love & Freed¬ man, Bot. Not. 109: 178-180. 1956. TYPE: [Canada], Yukon [Territory]: N of Quill Creek camp, 20 mi. W of Burwash landing, alt. ca. 5000 ft., 19 June 1953, Freedman s.n. (holo¬ type, MAN). Doris Love described Papaver kluanensis based on specimens from southwestern Yukon, which she later (1969) showed to be more widely distributed southward along the Rocky Mountains. Whereas she maintained this taxon as distinct from the Utah populations in this complex, I am, at least for the time being, including them within P. radicatum as subspecies kluanensis. Specimens of subspecies kluanensis from the type locality and elsewhere in the very northern part of its range are taller, with larger flowers and ellipsoid-subglobose capsules. Plants from the southernmost part of the range (Col¬ orado, Utah, and New Mexico) are consistently shorter, with smaller flowers and oblong-obconic capsules. Taken alone, the Utah specimens are uni¬ form and distinct; nevertheless, I agree with Randel (1975) that the northern and southern populations are linked in Colorado and Wyoming by interme¬ diate forms. Welsh (1986) has applied the name Papaver rad¬ icatum var. pygmaeum (Rydberg) Welsh to the Utah specimens that I have determined as P. radicatum subsp. kluanensis. I view P. pygmaeum as morpho¬ logically distinct from the P. radicatum complex, being more closely related to the European P. al- pinum group and different from the Utah plants in several respects. Papaver pygmaeum has leaves sparsely strigose to commonly glabrous, leaf lobes short and orbicular, and capsules with whitish (ivo¬ ry) trichomes, whereas P. radicatum subsp. klu¬ anensis has leaves coarsely strigose, lobes obovate, and capsules with light (straw colored) to dark brown trichomes. Papaver pygmaeum is a diploid (2 n = 14; Pack¬ er, 1968). Faberge (1942, 1944) applied the name P. pygmaeum to a specimen from which he obtained a chromosome count of n = 21. The origin for this material was given by Faberge as “Pike’s Peak Utah.” Clearly, this count was actually derived from a specimen of P. radicatum subsp. kluanensis from Pike’s Peak, Colorado, sent to Faberge by [W. P] Cottam from his home base in Utah. Papaver rad¬ icatum subsp. kluanensis has a chromosome num¬ ber of 2 n = 42 (Love, 1969, based on Colorado material) or 2 n = 56 (fide Moss & Packer, 1983, based on material from Alberta). Volume 5, Number 3 1995 Murray Papaver Section Meconella 295 Acknowledgments. My thanks to Robert Kiger and John McNeill for reviewing the manuscript and for suggesting improvements; and to the curators of CAN, MAN, NY, and S for loans or their many courtesies during visits to view types and other specimens. Literature Cited Faberge, A. C. 1942. Genetics of the Scapiflora section of Papaver 1. The garden Iceland poppy. J. Genetics 44: 169-193. -. 1944. Genetics of the Scapiflora section of Pa¬ paver 3. Interspecific hybrids and genetic homology. J. Genetics 46: 125-149. Greuter, W., F. R. Barrie, H. M. Burdet, W. G. Chaloner, V. Demoulin, D. L. Hawksworth, P. M. J0rgensen, D. H. Nicolson, P. C. Silva, P. Trehane & J. McNeill, Editors. 1994. International Code of Botanical Nomenclature (The Tokyo Code), Adopted by the XVth International Botanical Congress, Yokohama, August-September, 1993. Regnum Veg. 131. Love, D. 1969. Papaver at high altitudes in the Rocky Mountains. Brittonia 21: 1-10. Moss, E. H. & J. G. Packer. 1983. Flora of Alberta, ed. 2. Univ. Toronto Press, Toronto. Packer, J. G. 1968. IOPB chromosome number reports XVII. Taxon 17: 285-288. Randel, U. 1975. Die Beziehungen von Papaver pyg - maeum Rydb. aus den Rocky Mountains zum norda- merikanischen P. kluanense D. Love sowie zu einigen nordost-asiatischen Vertretern der Sektion Scapiflora Reichenb. im Vergleich mit P. alpinum L. (Papavera- ceae). Feddes Repert. 86(1-2): 19—37. Randel, U. 1977. Ober Sippen des subarktisch-arktisch- en Nordamerikas, des Beringia-Gebietes und Nordost- Asiens der Sektion Lasiolrachyphylla Bernh. (Papaver- aceae) und deren Beziehungen zueinander und zu Sippen anderer Arealteile der Sektion. Feddes Repert. 88(7-8): 421^50. Welsh, S. L. 1986. New taxa and combinations in the Utah Flora. Great Basin Naturalist 46: 254-260. Validation of a New Species of Diospyros (Ebenaceae) from Yunnan, China Wu Zhengyi (C. Y. Wu) and Zhu Hua (H. Chu) Kunming Institute of Botany, Academia Sinica, Kunming 650204, Yunnan, China ABSTRACT. Diospyros xishuangbannaensis, previ¬ ously invalidly described, is validated. Diospyros xishuangbannaensis was described in 1990 (Zhu & Wu, 1990), but two types, represent¬ ing flowering and fruiting states, were designated, thus contravening Article 37 of the International Code of Botanical Nomenclature (Greuter et al., 1994) and making the name invalid. The species is herein validated by designating the fruiting ma¬ terial as the holotype. Diospyros xishuangbannaensis C. Y. Wu & H. Chu, sp. nov. TYPE: China. Yunnan: Xish- uangbanna, Mengla, Bubeng, 20 July 1988 (fr), Chu Hua & Wang Hong 2451 (holotype, KUN). Proxima Diospyroti martabanicae Clarke, sed ab ea flore masculo solitario, corolla breviore, staminibus 8, endos- permio laevigato differt; a D. pilosula (A. de Candolle) Hiem foliis subtus persaepe dense pubescentibus, nervis lateralibus utrinsecus 10-15, tertiariis utrinque conspi- cuis, flore femineo 5-mero, flore masculo solitario, antheris attenuato-ovatis apice acutis differt. Trees ca. 20 m tall; branchlets densely brown pubescent. Leaves oblong or oblong-lanceolate, 8- 18 cm long, 3-4.5 cm wide, stiffly chartaceous to subcoriaceous, somewhat pubescent adaxially only on midvein when young, soon completely glabrous, abaxially with pubescence along midvein like that of young branchlets, base rounded or subcordate, apex acuminate; midvein sunken adaxially, elevat¬ ed abaxially; lateral veins 10-15 on each side, flat adaxially, elevated abaxially, tertiary veins conspic¬ uous on both surfaces; petiole 2—3 mm long, pu¬ bescent as in branchlets. Flowers solitary; stami- nate flowers 4-merous, on pubescent pedicels 8—10 mm long; calyx 6-7 mm long, divided nearly to base into linear-lanceolate lobes, appressed seri¬ ceous outside, glabrous inside; corolla salverform, sericeous outside as in calyx, glabrous inside, tube ca. 7 mm long, lobes ca. 9 mm long; stamens 8, unequal, glabrous; anthers attenuate-ovate, acute at apex; rudimentary ovary small villous; pistillate flowers 5-merous, on pubescent pedicels 10-16 mm long; calyx and corolla as in staminate flowers; ovary ovoid, pubescent, 4-loculed. Fruit globose, 2—2.5 cm diam., densely sericeous; fruiting pedicel 1—1.6 cm; fruiting calyx reflexed outward; seeds 4; endosperm smooth. Paratype. CHINA. Yunnan: Xishuangbanna, Mengla, Manzhuan, 7 Mar. 1983, Zhou Shochin & Wang Hong 2452 (KUN). Acknowledgments. Michael G. Gilbert is thanked for pointing out the problem and Ihsan A. Al-Shehbaz for helping with the manuscript. Literature Cited Greuter, W., F. R. Barrie, H. M. Burdet, W. G. Chaloner, V. Demoulin, D. L. Hawksworth, P. M. Jorgensen, D. H. Nicolson, P. C. Silva, P. Trehane & J. McNeill, Editors. 1994. International Code of Botanical Nomenclature (The Tokyo Code), Adopted by the XVth International Botanical Congress, Yokohama, August—September, 1993. Regnum Veg. 131. Zhu Hua & Wu Zhengyi. 1990. Contributions to the Flora of Xishuangbanna (1). Acta Bot. Yunnan. 12: 375—380. Lectotypification of Kengyilia mutica (Keng ex Keng & S. L. Chen) J. L. Yang, Yen & Baum (Poaceae), and Change of Circumscription Chi Yen and Jun-Liang Yang Triticeae Research Institute, Sichuan Agricultural University, Dujiangyan City 611830, Sichuan, China Bernard R. Baum Centre for Land and Biological Resources Research, Agriculture and Agri-Food Canada, Central Experimental Farm, K.W. Neatby Building, Ottawa, Ontario, K1A 0C6, Canada Abstract. A lectotype of Kengyilia mutica (Keng ex Keng & S. L. Chen) J. L. Yang, Yen & Baum is here designated based on an illustration of the ho- lotype; the latter has been lost. A more complete description than the original, which was based on a single and incomplete specimen, is provided. Professor Y. L. Keng identified a single specimen obtained from the Province of Qinghai, China, as Roegneria mutica Keng, nomen nudum. During 1944, Y. L. Keng collected grasses in Qinghai with his son, P. C. Keng. At that time Guide County was occupied by a military lord, Ma Bu-Fong and his herds of horses. Keng did not obtain permission to collect in this region. A friendly person who worked on the horse farm brought Keng a specimen. Keng examined the plant, decided that it differed from related species, and called it Roegneria mutica (P. C. Keng, pers. comm. 1992). He subsequently pub¬ lished Roegneria mutica Keng as a nomen nudum (Keng et al., 1957) and later provided a description (Keng et al., 1959), but in Chinese only, which re¬ sulted therefore in an illegitimate name. The Latin description was finally published in 1963 (Keng & Chen, 1963), giving it invalid status. The single specimen on which this species was based may have been destroyed during the “Cul¬ tural Revolution” (Song Gui-Qing, pers. comm. 1992), or may not have been returned to the her¬ barium when it was lent for drawing the figure in Keng et al. (1959, and P. C. Keng, pers. comm.). This specimen is “No. 4, Military Horse Farm, Guide, Qinghai”—the precise locality is unknown; however, it was collected in an area near the two subsequent known locations (Fig. 1) and was con¬ served in the herbarium of Nanjing University (N). The only remains of the type collection are thus the illustration, figure 337 in Keng et al. (1959), be¬ cause the protologue (Keng & Chen, 1963) referred to this figure. This interpretation is in accord with the definition of what constitutes type material in the Code (Greuter et al., 1994; Article 9.7 Foot¬ notes). Thus, figure 337 in Keng et al. (1959) is here designated as the lectotype (Fig. 2). Kengyilia mutica (Keng ex Keng & S. L. Chen) J. L. Yang, Yen & Baum, Hereditas 116: 27. 1992. Basionym: Roegneria mutica Keng ex Keng & S. L. Chen, Acta Nanjing Univ. 1: 87. 1963. TYPE: China. Qinghai Province: Guide County, Military horse farm, 4 [“No. 4, Military Horse Farm, Guide, Qinghai”] (holotype, N now lost). Lectotype, here designated: figure 337 in Keng et al. (1959). Roegneria mutica Keng in Keng et al., Clav. Gen. & Sp. Gram. Sin. 75, 188. 1957. Nomen nudum. Roegneria mutica Keng in Keng et al., FI. lllustr. Plant. Prim. Sin. Gram. 408, figure 337. 1959. Nomen inval., based only on Chinese description. An expedition to Guide County, Qinghai Prov¬ ince, was conducted in 1993 in order to study the morphological variation at the type locality. There, we realized that what was described as an awnless species was in fact an unusual type and that most of the individuals of Kengyilia mutica have short lemma awns. Although we collected awnless spec¬ imens in 1986 on the Lajishan mountains, at 3200 m, we found in the 1993 collections only a few specimens with awnless florets intermixed with short-awned florets on the same spike. Most indi¬ viduals of this species, based on study of extensive collections, are short-awned. Totally awnless plants have yet to be found. We provide here a revised description of Kengyilia mutica and a drawing (Fig. 3). Perennial with fibrous roots. Culms 40-70 cm Novon 5: 297-300. 1995. 298 Novon Distribution of KengyiNa mutica Figure 1. Distribution of Kengyilia mutica (Keng ex Keng & S. L. Chen) J. L. Yang, Yen & Baum. tall, puberulent below the spikes; 3-noded, genic¬ ulate at the lowest node (Fig. 3A). Leaf sheath smooth, glabrous or puberulent; ligules scabrous, ca. 1 mm long; leaf blades flat or involute, glabrous or puberulent, 4—6 mm wide; tillers with narrow leaf blades, usually up to 19 cm long. Spikes erect or slightly curved, 5-9 cm long, 8 mm wide (Fig. 3A); rachis intemodes puberulent, the upper ones 3-4 mm long, the lower ones 8-9 mm long. Spike- lets yellow-green, 12 mm long, with 4—6 florets (Fig. 31), the uppermost not well developed; rachilla intemodes puberulent, ca. 1.5 mm long; glumes ob¬ long ovate, 5-6 mm long, smoothly glabrous, acute, usually 3-nerved with the midnerve prominent (Fig. 3F, G); lemmas 5-nerved, densely hirsute (Fig. 3H), with rounded back, tips acute, mucronate (Fig. 3B) or ending in an awn 1.5-5 mm long (Fig. 3H); 1st lemma ca. 9.5 mm long; palea equal to or shorter than lemma, with an obtuse to slightly emarginate apex, ciliate toward tip on the two keels (Fig. 3C); anthers yellow or black, ca. 3 mm long. Caryopsis tip hirsute (Fig. 3E). Distribution in China. Qinghai Province, Guide County (Fig. 1), endemic. Specimens examined. All collected in 1993: CHINA. Qinghai Province: Guide County, I^ajishan Mountain, alt. 3200 m, in Stipa steppe, J. L Yang & R. von Bothmer 860020, 860021, 930002, 930007 (all SAUTI, DAO). Figure 337 Roegneria mutica Keng sp. nov. 1. Adult plant; 2. Spikelet; 3. Florelet; 4. Dorsal view of palea; 5. Caryopsis; 6. Lodicules, anthers with filament ovary with stigmata. (No. 4, Guide Military Ranch, Qinghai, Holotype) Figure 2. Illustration of the lectotype of Kengyilia mu¬ tica (Keng ex Keng & S. L. Chen) J. L. Yang, Yen & Baum. 1 Volume 5, Number 3 1995 Yen et al. Kengyilia mutica 299 H Figure 3. Detailed drawing of Kengyilia mutica (Keng ex Keng & S. L. Chen) J. L. Aang. Aen & Baum. A. Mature plant. —B. Dorsal view of floret, showing an awnless lemma. —C. Ventral view of floret. —D. Ventral view of palea. —E. Caryopsis. —F. Lower glume. —G. Upper glume. —H. Dorsal view of an awned lemma. —I. Spikelet. 300 Novon Literature Cited Greuter, W., F. R. Barrie, H. M. Burdet, W. G. Chaloner, V. Demoulin, D. L. Hawksworth, P. M. j0rgensen, D. H. Nicolson, P. C. Silva, P. Trehane & J. McNeill, Editors. 1994. International Code of Botanical Nomenclature (The Tokyo Code), Adopted by the XVth International Botanical Congress, Yokohama, August-September, 1993. Regnum Veg. 131. Keng, Y. L. & S. L. Chen. 1963. A revision of the genus Roegneria C. Koch of China. Acta Nanjing University (Biology) 1: 1-92. [In Chinese, with Latin descriptions of new species, and with an English summary.] -(editor), P. C. Keng, Y. C. Tang, B. Z. Gao, S. L. Chen & L. Liu. 1957. Claves Generum et Specierum Graminearum Primarum Sinicarum, Appendice Nomen- clatione Systematica. Science Press, Beijing [in Chi¬ nese]. * -& S. L. Chen, J. R. Feng, Z. Y. Feng, B. Z. Gao, L. Z. Jia, X. Q. Jiang, P. C. Keng, L. Liu, W. Q. Shi, Y. C. Tang & S. Q. Zhong. 1959. Flora Illustrata Plantar- urn Primarum Sinicarum, Gramineae. 403, f. 337. Sci¬ ence Press, Beijing [in Chinese]. Dracontium croatii (Araceae), a New Species from the Western Slopes of the Andes in Ecuador Guanghua Zhu Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A. and Department of Biology, University of Missouri-St. Louis, 8001 Natural Bridge Road, St. Louis, Missouri 63121, U.S.A. Abstract. Dracontium croatii G. Zhu, a new spe¬ cies from the western slopes of the Andes in Ec¬ uador, is described and illustrated. Morphological¬ ly, this species does not appear to be particularly close to any member of the genus and is readily distinguished by its showy greenish spathe, large and globose berries, rounded and strongly inter¬ rupted irregular warty seeds, and matte upper blade surface. Resumen. Dracontium croatii G. Zhu, una espe- cie nueva de las laderas occidentales de los Andes de Ecuador se describe e ilustra. A1 parecer, la nueva especie no esta cercanamente relacionada con otros taxa en el g^nero. Es fdcilmente recono- cible por la siguiente combinacion de caracteres: la espata es vistosa y verdosa, la superficie adaxial de las hojas es opaca y las semillas y los frutos son redondeados. Dracontium L. (Araceae), with 22 species and 2 varieties, is known exclusively from the New World tropics. One of the most poorly known genera of the Araceae, it is represented in Ecuador by at least 3 species, 1 of which is new to science. This new species was detected in the course of a taxonomic revision of the genus. Dracontium croatii G. Zhu, sp. nov. TYPE: U.S.A. Missouri: St. Louis, cultivated plant in the Missouri Botanical Garden, Zhu 1493 (ho- lotype, MO); originally collected along unfin¬ ished road between Lita and San Lorenzo, 15.5 km W of Lita, 705 m, 0°55'N, 78°28'W, San Lorenzo Canton, Esmeraldas, Ecuador, Croat 72368. Figures 1, 2. Herba terrestris, magna, tuberosa, unifoliata, usque plus quam 3 m alia; tuber hemisphaericum, usque 15 cm diam., 7 cm crassum, apice planum interdum tuberculis; folium bene ante vel post anthesin prodiens; petiolus 1 1 3 m longus, cylindricus, camosus, atro aut purpureo-vir- idibus maculis, guttatus pallide viridis aut sordide albus; lamina tripartita usque 20 cm longis, 9 cm latis, apicibus acuminatis, interdum bifidis; pedunculus 1-1.7 m longus, 1.3—2 cm diam., cylindricus; spatha 23—50 cm longa, 8— 20 cm lata, cucullata basi, obovata aspectu frontali, extus impolita, atroflava-viridis suffusa purpurea, intus velutina, flava-viridis apice et secus margines exteriores, cum ner- vis atropurpureis, viridi-alba basi usque 3 cm prope mar¬ gines et usque 7 cm in medio; spadix 3—5 cm longus, 1.3 cm diam.; tepala 7-8, subcucullata apice; baccae globo- sae, usque 1—2 cm diam.; semina rubra, suffusa aurantia- ca, plus minusve rotunda, usque 0.8-1 cm diam., 0.5 cm crassa. Terrestrial herb, perennial, to more than 3 m tall. Tuber hemispherical, to 15 cm diam., 7 cm thick, 20-30 cm underneath the ground, flat at apex, with or without a few tubercles among numerous roots, underneath smooth, convex, without tubercles and roots. Leaf 1, arising from tuber 4—30 cm below the ground level; petiole 1.4—3 m long, 3.5-6 cm diam. at base, 1.5-2.5 cm diam. at apex, cylindric, fleshy, dark or purplish green, mottled pale green or dirty white forming a reptilian pattern, distal half smooth, proximal half with many protuberances; major divisions of blade 3, each to 1 m long, matte, sometimes fenestrate, upper surface green, lower surface slightly paler; ultimate segments to 20 cm long, 9 cm wide, oblanceolate, apex gradually acu¬ minate or caudate, or bilobed, lobes gradually acu¬ minate; smaller leaflets triangular, or ± oblanceo¬ late, 3.5-15 cm long, 2.5-6 cm wide; midrib and major veins convex, sunken adaxially, round-raised abaxially, secondary veins impressed on upper sur¬ face and raised on lower surface; rachises pale green, smooth, faintly mottled as petiole, rachis (from petiole apex to the first major subdivision) of the middle division up to twice as long as that of lateral divisions. Inflorescence arising from apex of tuber before or after the leaf development; cata- phylls to 20 cm long, unribbed, drying light brown toward base, darker brown toward apex; peduncle 1-1.7 m long, 1.3-2 cm diam. at mid point, cylin¬ dric, fleshy, dark blackish purple, sparsely and weakly streaked, violet or dark yellow-green, be¬ coming somewhat more brownish with larger swatches of yellow-green above middle, these be¬ coming obliquely arranged, larger and paler toward apex, beset with irregular protuberances through- 302 Novon Figure 1. Dracontium croatii G. Zhu (Croat 72368). —A. Leaf. —B. Peduncle (left) and petiole (right). out; spathe 23-50 cm long, 8—20 cm wide when open, conspicuously hooded, convolute at base, ob- ovate in front face view, outside matte, green tinged purple, apex and the lower Ps of spathe densely purplish speckled, midrib broadly convex, pale yel¬ lowish green in lower half of spathe, purplish on upper half of spathe, with 11-12 dark purplish ma¬ jor veins on either side, these veins interconnected by oblique cross veins, inner surface velvety, tri¬ color: apex and along outer margins yellow-green, with dark purple veins, area 4-6 cm from the outer margins dark purple and becoming more velvety, area 3 cm high near the margins and 7 cm high medially from the base solid greenish white, the border between the white and purple portions abrupt but with smoothly feathered margin, apex probably rounded or weakly cuspidate (apparently somewhat damaged and split upon opening), outer margins thin and weakly curled back, moderately undulate toward base; spadix 3-5 cm long, 1.3 cm diam. excluding protruding styles, purplish, axis of spadix pinkish within, aerenchymatous. Flowers ir¬ regularly rounded, 2.5-3 mm diam.; tepals 7-8 per flower, 1.8—2.2 mm long, 1-1.5 mm wide, somewhat cucullate at apex, inner margin broadly rounded, purplish with paler, weakly raised projections, pal¬ er and somewhat scalloped along the edges; styles 1-1.2 mm long, 0.8-1 mm diam., spindle-shaped, purplish, covered with weakly raised projections as tepals; stigmas somewhat cushion-shaped, 2- or rarely 3-lobed, 0.4 mm high, 0.8 mm diam., cov¬ ered with brushlike translucent bristles, becoming purplish after anthesis; stamens not emerging above tepals; filaments ca. 1.5 mm long; anthers brown¬ ish, clearly visible between tepals and style prior to anthesis. Berries globose, 1—2 cm diam., olive- green with purplish spots, becoming deep red; seeds 2 per berry, red tinged orange, with thick, strongly interrupted, irregular warts, round, 0.8-1 cm diam., 0.5 cm thick. Dracontium croatii is apparently restricted to the western slopes of the Ecuadorian Andes between 450 and 1330 m. In addition to the type locality, collections have also been made from Carchf Prov¬ ince. The species may also be expected in Los Rios Province in Ecuador, as well as adjacent areas of Colombia. Morphologically, Dracontium croatii does not ap¬ pear to be particularly close to any member of the genus. It is readily distinguished by its showy spathe with a unique coloration: outer surface greenish with raised, dark purple veins, inner sur¬ face velvety, dark purple with 4—6 cm wide, yellow- green outer margins and a solid greenish white area 3-7 cm high at the base. The globose berries of the species, at 1-2 cm diam., are the largest in Dra¬ contium. Other species, such as D. spruceanum, may also have globose berries, but they never ex¬ ceed 1 cm diam. Seeds of D. croatii are definitely the largest known in the genus; the distinctive seed morphology is illustrated on one of the cover pages of Aroideana (Madison, 1980). Dracontium croatii is also often marked by a matte upper blade sur¬ face, which is glossy or semiglossy in other species. Inflorescences of Dracontium croatii have been collected in November, fruits in July. Under culti¬ vation, D. croatii flowers in September and from November to January. It produces inflorescences before or immediately after new leaf development. Volume 5, Number 3 1995 Zhu Dracontium croatii 303 Figure 2. Dracontium croatii G. Zhu. —A. Infructescence (Croat 72368). —B, C. Inflorescence (Zhu 1493, type). — D. Inflorescences opened to show the spadix and the creamy-white area at the base of the inner spathe surface (Zhu 1493, type). 304 No von Dracontium croatii is named in honor of Thomas B. Croat, from whose collection of living Araceae in the Missouri Botanical Garden I recognized this new species and prepared the holotype. Two other live collections are known: one from the Marie Sel¬ by Botanical Gardens (75-491), the other from the Missouri Botanical Garden (received from the Selby Gardens, 78-2173); both were collected by Calaway Dodson in Pichincha Province, Ecuador, in 1975 and 1978. Other plants of this species have been distributed by the Selby Gardens to various other places under the name D. ornatum. One of these plants is cultivated by Dewey Fisk in Florida. Paratypes. ECUADOR. Carchf: Forestry Reserve, vi¬ cinity of encampment in Aw4 Ethnic, 0°58'N, 78°16’W, 1330 m. Hoover 3279 (QCA); trail from San Marcos to Tobar Donoso, 660 m, 1°07'N, 78°20'W, Barfod 41597 (AAU, MO, QCA). Esmeraldas: along unfinished road between Lita and San Lorenzo, 15.5 km W of Lita, 705 m, 0°55'N, 78°28'W, Croat 72368 (MO). Pichincha: bridge over Rfo Chiguilpe near junction with Rfo Baba, 7 km from junction of entrance road at 7 km from Santo Domingo on highway to Quevedo, 450 m, Dodson 5951 (SEL); La Centinela, at crest of Montanas de Ila on road 12 km from Patricia Pilar to 24 de Mayo, 600 m, Dodson 7288 (MO, SEL), Dodson & Fallen 7762 (MO, QCNE, SEL); La Centinela, 0.2 km past Escuela Mixta La Cen¬ tinela, along trail to left of road, exactly 13 km E from main Santo Domingo-Quevedo Highway in Patricia Pilar, 1000 m, 0°32'S, 79°11'W, Croat 73040 (MO). Cultivated plants: Marie Selby Botanical Gardens (75-461, vouch- ered by Dodson 5951), Christenson 1576 (SEL), Croat 71798 (MO), Madison 4140 (SEL); Marie Selby Botanical Gardens (78-2173, vouchered by Dodson 7288), Christen¬ son 1154 (SEL), Plowman 10926 (F); Missouri Botanical Garden (vouchered by Croat 72368), Zhu 1518 (MO); Missouri Botanical Garden (received from the Marie Selby Botanical Gardens, 78-2173, vouchered by Dodson 7288), Zhu 1453 (MO). Acknowledgments. This paper is part of my doctoral dissertation to be submitted to the Uni¬ versity of Missouri-St. Louis Department of Biology. I thank my major advisor, Thomas B. Croat, for discussions and critical suggestions on this subject. I also thank Mike Grayum, Sue Thompson, German Camevali, Eleanor Sauer, P. Mick Richardson, Da¬ vid Boufford, Bruce Bartholomew, Petra Malesev- ich, Donna Atwood, Kathy Upton, and Hong Song for comments on the manuscript and other assis¬ tance. I extend my appreciation to Dewey Fisk, Ju¬ lius Boos, and curators of the Marie Selby Botanical Gardens and Herbarium for providing specimens and photos. The research was conducted at the Mis¬ souri Botanical Garden and was partially supported by NSF doctoral dissertation research grant DEB- 9310171 to T. B. Croat and G. Zhu, and by a grant from the Andrew W. Mellon Foundation to G. Zhu. Literature Cited Madison, M. 1980. Cover photo. Aroideana 3(1): 1-2. Volume 5, Number 3, pp. 209-304 of NOVON was published on 05 September 1995. Volume 5 Number 4 1995 NOVON MISSOURI BOTANIC"! DEC 2 7 1995 GARDEN LlbRrtRt' A New Blakea (Melastomataceae: Blakeeae) from Tapantf National Park, Costa Rica Frank Almeda Department of Botany, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118-4599, U.S.A. Gina Umaha Dodero Herbario Nacional de Costa Rica, Museo Nacional, Apartado 749-1000, San Jos6, Costa Rica Abstract. Blakea tapantiana, a new Costa Rican endemic, is described, illustrated, and compared with B. grandiflora, an allopatric Costa Rican en¬ demic that occurs at higher elevations and flowers sporadically throughout the year. The new species is distinguished by its short-pedicellate flowers, caudate-acuminate outer floral bracts, white petals, and unappendaged anther thecae. Blakea, a neotropical genus with over 100 de¬ scribed species, continues to yield new Mesoamer- ican taxa of local distribution (Umana & Almeda, 1991). Unlike many species of Blakea, which tend to flower sporadically throughout the year, the spe¬ cies proposed here appears to flower for a short period during the rainy season followed by a rather lengthy period of fruit maturation. This probably accounts, in part, for the paucity of flowering and mature fruiting specimens in herbaria and empha¬ sizes the need for repeated visits to mid-elevation montane forests in Costa Rica and elsewhere in tropical America. Blakea tapantiana Umana & Almeda, sp. nov. TYPE: Costa Rica. Cartago: Parque Nacional Tapantf, Sendero Oropdndola, 9°48'18"N, 83°57'12"W, 1200 m, 3 Sep. 1992 (fl), Umana et al. 497 (holotype, CR; isotypes, CAS, COL, F, K, MEXU, MO, US, USJ, WIS). Figure 1. Frutex epiphyticus vel terrestris. Ramuli primum sub- quadrangulati demum teretes sicut foliorum subtus venae primariae et secundariae pedicelli bracteaeque primum densiuseule furfuracei, demum glabrescentes. Lamina el- liptiea vel oblongo-elliptiea, apice caudato-acuminato. Flores 6-meri in quoque nodo superiore 1—3, pedicellis 0.6-1 cm longis: bracteae exteriores ca. 3-6 mm coalitae apice hebeti-acuminato. Antherae 4.5-5 X 1—1.5 mm in¬ ter se non cohaerentes; connectivum dorsaliter paulo ele- vatum ecalcaratum. Terrestrial or epiphytic shrubs 2-6 m tall. Upper branchlets rounded-quadrate to somewhat sulcate, becoming rounded and glabrous with age. Vegeta¬ tive buds, young branchlets, young petioles, pri¬ mary and secondary abaxial leaf veins, and imma¬ ture inner and outer floral bracts densely to moderately covered with dendritic and scurfy hairs. Mature leaves of a pair equal to somewhat unequal in size, essentially glabrous adaxially or sparingly beset with a mixture of paleaceous, dendritic, and/ or conic hairs along the elevated veins; petioles 1.5-3.5 cm long; blades coriaceous, 8.3-17 cm long and 4.1-7.6 cm wide, elliptic to oblong-ellip¬ tic, rarely elliptic-obovate, apex acuminate to cau¬ date-acuminate, base acute varying to obtuse, mar¬ gin denticulate to subentire, 5-nerved or 5-plinerved with the innermost pair of primary veins diverging from the median vein 3—6 mm above the blade base, the striolate transverse sec¬ ondary veins spaced 1.5—3 mm apart at the widest Novon 5: 305-308. 1995. 306 Novon igure 1. Blakea tapantiana Umana & Almeda. —A. Habit. —B. Representative leaf (abaxial surface). —C. Enlarge¬ ment of a primary leaf vein (abaxial surface). —D. Immature fruit enveloped by outer and inner floral bracts. —E. \oung fruit (top view) showing ovary summit, torus, and calyx lobes. —E Fused inner floral bracts. —G. Petal. —H. Stamens, lateral view (left) and ventral view (right). (A—C from Almeda et al. 6860 ; D—H from Umana et al. 497.) Volume 5, Number 4 1995 Almeda & Umana Blakea from Costa Rica 307 portion of the blade. Flowers erect, 1-3 in each leaf axil of uppermost branches; pedicels 0.6-1 cin long, thick but somewhat compressed and 2-edged, glabrous and lenticellate. Floral bracts sessile, en¬ tire; outer bracts 1.8-2.5 X 0.8-1.1 cm, fused ba- sally for 3-6 mm, ovate, apex acuminate to caudate-acuminate, margin entire, 3-veined with only the median vein usually prominulous, glabrous throughout; inner bracts 1—1.2 cm long, connate basally for nearly half their length to form a 2-lobed cuplike collar that envelops the hypanthium but is somewhat concealed by the outer bracts, the free portions of the inner bracts rounded-triangular and glabrous, 0.4-0.7 X 0.7-0.8 cm. Hypanthium (in young fruit) campanulate, 0.8-1 cm long to the to¬ rus and 0.6—0.8 cm diam., glabrous. Calyx tube 2.5-3 mm long; calyx lobes 4.5-6 mm long and 5- 6 mm wide basally, rounded-triangular, margin ± entire and conspicuously callose-thickened, gla¬ brous on both surfaces. Petals 6, glabrous, 17—21 X 7-10 mm, white, narrowly obovate, apically rounded (sometimes obliquely so) and often abrupt¬ ly and bluntly acute, essentially entire and cilio- late. Stamens 12, isomorphic; filaments 5-7 mm long, complanate, glabrous and declined to one side of the flower opposing the style; anthers 4.5-5 mm long, 1—1.5 mm wide, yellow, oblong, laterally com¬ pressed with two well-separated pores at the trun¬ cate apex; connective slightly dilated dorso-basally ca. 0.5 mm above the filament insertion into a low blunt thickening. Ovary wholly inferior, 6-celled, glabrous at the slightly elevated bluntly conic apex but lacking a collar. Style somewhat declinate, gla¬ brous, 10-11 mm long; the expanded stigma sub¬ truncate to narrowly conic. Mature berry and seeds not seen. Distribution. This little-collected cloud forest species is known only from Cartago Province, Costa Rica, in a region extending from the recently des¬ ignated (1993) Tapantf National Park (formerly Re¬ fugio Nacional de Vida Silvestre) to Valle Escon¬ dido on the Atlantic slope of the Cordillera de Talamanea at 1150-1450 m. Blakea tapantiana is notable for its caudate-acu¬ minate leaf apices, short-pedicellate flowers, acu¬ minate to caudate-acuminate outer floral bracts, white petals, and unappendaged anther thecae. Among the few white-flowered Costa Rican spe¬ cies of Blakea that exhibit both an epiphytic and free-living shrubby habit, B. tapantiana is likely to be confused only with B. grandiflora Hemsley. The latter species is also endemic to Costa Rica, but it is most commonly encountered in the wet wooded slopes and valleys of the Cordillera Central and Cordillera TilarAn. Many of the diagnostic features that can be used to separate Blakea tapantiana from B. grandiflora are quantitative. The leaf blades of B. grandiflora are consistently larger (13-25.5 X 7.7-14.7 cm), as are the pedicels (2.5-4.5 cm), outer floral bracts (1.3-2 cm wide), and fully expanded petals (25-35 X 25-32 cm). In B. grandiflora the outer floral bracts are fused basally for 8-15 mm, but they are fused for only 3-6 mm in B. tapantiana. Flower color also appears to provide a diagnostic field character for distinguishing the two species. The apex of floral buds in B. tapantiana is pale pink, but the petals are completely white when fully ex¬ panded. In B. grandiflora, however, the buds are deep pink, and the expanded petals are flushed pink apically on one or both surfaces. Other more subtle but consistent differences can be used to distinguish these two species. In overall aspect the vegetative and floral organs of Blakea grandiflora are coarser and thicker, but these are difficult to quantify. The anthers of B. grandiflora have a distinctive blunt appendage borne dorsally on the connective about 1 mm above the base of the thecae. The anther connective in B. tapantiana is merely thickened and not elaborated into a knob¬ by appendage. Blakea grandiflora, like II. tapan¬ tiana, has a sparse to dense indument on upper¬ most branchlets, vegetative and young floral buds, and petioles that consists of coarse dendritic hairs that vary toward or are intermixed with paleaceous hairs that have somewhat lacerate margins. In B. tapantiana the dendritic hairs are not as coarse or as well developed and vary toward a furfuraceous indument on abaxial leaf surfaces. Taken together these consistent differences, cou¬ pled with the elevational and geographical sepa¬ ration between B. tapantiana and B. grandiflora, provide a compelling rationale for recognizing two species. This taxonomic interpretation is consistent with a pattern of sharply discontinuous quantitative differences seen in other species pairs such as B. austin-smithii Standley and B. chlorantha Almeda and in B. calycosa Gleason and B. tuberculata Don¬ nell Smith. Paratypes. COSTA RICA. Cartago: Refugio Nacional de Fauna Silvestre Tapantf, Sendero de la Orop^ndola bor¬ dering Rfo Macho, 22 Mar. 1986 (fr), Almeda et al. 5699 (CAS, CR, MO, NY); Refugio Nacional de Vida Silvestre, Sendero de la (iropcndola about 1.2—2.7 km from the re¬ serve entrance, 1 Mar. 1991 (fr), Almeda et al. 6860 (CAS, CR, F, US); Refugio Nacional de Vida Silvestre, Sendero de la I iropcndola. 13 Feb. 1992 (fr), Almeda et al. 6971 (CAS, CR, MO, US); bank of Rfo Grande de Orosf, 4 km S of Tapantf, 15 Sep. 1967 (fl). Lent 1280 (F); Valle Es- 308 Novon condido, 15 Apr. 1966 (fr), Schnell 689 (F); Parque Na- cional Tapantf. Sendero (Iropdndola, 9°48'18''N, 83°57'12"W, 1200 m, 20 Aug. 1992 (fl), Umana et al. 496 (CR), 29 Sep. 1992 (fl), Umana et al. 498a (CR), 24 Sep. 1993 (fl), Umana et al. 498b (CR). Acknowledgments. We thank Colleen Sudekum for preparing the line drawings, Barrett Anderson, Bruce Bartholomew, Thomas F. Daniel, Leticia Pa¬ checo, and Nelson Zamora for assistance in the field, the staff of the Herbario Nacional de Costa Rica for logistical support, and the Servicios de Parques Nacionales de Costa Rica for permission to collect in protected areas of Costa Rica. Literature Cited Umana D., G. & F. Almeda. 1991. Blakea costaricensis (Melastomataceae): A remarkable new epiphyte from southeastern Costa Rica. Selbyana 12: 1-4. Placement of Arabidopsis parvula in Thellungiella (Brassicaceae) Ihsan A. Al-Shehbaz and Steve L. O’Kane, Jr. Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. Abstract. The genus Thellungiella is recog¬ nized, and the characters separating its two species from Arabidopsis are given. Arabidopsis parvula (Schrenk) O. E. Schulz is recognized in Thellun¬ giella. The limits of Arabidopsis have been the subject of considerable controversy, and as many as 50 spe¬ cies were previously placed in the genus (Al-Sheh- baz, 1988; Price et al., 1994). Detailed morpholog¬ ical and molecular studies by the authors reveal that Arabidopsis includes less than ten species, most of which were previously assigned to other genera, and that the remaining species are more appropriately placed in five or six genera. The re¬ sults of these on-going studies will be published elsewhere. The present paper deals with two species here recognized as members of Thellungiella O. E. Schulz. Thellungiella parvula has generally been accepted in Arabidopsis in all recent floras (e.g.. Ball, 1964; Busch, 1939; Hedge, 1965, 1968; Pav¬ lov, 1961) that cover the areas of its range. The generic placement of T. salsuginea (Pallas) 0. E. Schulz has been more controversial; although sev¬ eral authors followed Schulz (1924) in treating it in Thellungiella, Al-Shehbaz (1988) and Rollins (1993) retained the species in Arabidopsis. Schulz (1924, 1936) placed Arabidops is and Thellungiella in different subtribes separated solely on the basis of presence versus absence of seed-coat mucilage. As shown by Vaughan and Whitehouse (1971), the presence or absence of mucilage in wetted seeds can be observed in various species of the same ge¬ nus and, therefore, is not a reliable taxonomic char¬ acter. Although the seeds of T. parvula produce mu¬ cilage upon wetting and those of T. salsuginea do not, it is abundantly clear that this and other dif¬ ferences given in the key below do not justify the placement of the two species in different genera. In addition to Thellungiella salsuginea, Schulz (1924), Busch (1939), and Pavlov (1961) recog¬ nized another species, T. halophila (C. A. Meyer) D. E. Schulz, which they separated from T. salsug¬ inea primarily by having coarsely dentate to divid¬ ed instead of entire to repand basal leaves. It is highly unlikely that these differences alone justify the recognition of T. halophila as distinct from T. salsuginea. Because the types of these two species were not available for this study, we refrain from formally reducing T. halophila to the synonymy of T. salsuginea. As here delimited, Thellungiella consists of two (or perhaps three) species that are centered pri¬ marily in Kazakhstan and neighboring Russia. Thellungiella parvula is distributed in Kazakhstan, southern Russia, Turkmenistan, and Turkey, where¬ as T. salsuginea is widespread in Kazakhstan, southern Russia, Mongolia, northern China, Cana¬ da, and the United States. Both species are restrict¬ ed to strongly saline soils, and such edaphic ad¬ aptation to salinity, which is rare elsewhere in the Brassicaceae, is not exhibited by any species of Arabidopsis. In addition, both species are glabrous throughout, are glaucous on stems and leaves, and do not produce well-defined basal rosettes. All spe¬ cies of Arabidopsis produce basal rosettes that often persist well after fruit maturity, and none has glau¬ cous stems and leaves. The characteristic indu¬ mentum of mixed simple and stalked furcate tri- chomes should readily distinguish species of Arabidopsis from members of Thellungiella. The two species of Thellungiella are easily separated as fol¬ lows: la. Cauline leaves oblong to ovate or cordate, not fleshy, auriculate to rarely amplexicaul at base; infructescence axis straight; petals much longer than sepals; seeds not mucilaginous when wetted . T. salsuginea lb. Cauline leaves linear to linear-oblong, fleshy, not auriculate at base; infructescence axis flexuous; petals subequaling sepals or absent; seeds mu¬ cilaginous when wetted . T. parvula Thellungiella parvula (Schrenk) Al-Shehbaz & O’Kane, comb. nov. Basionym: Diplotaxis? parvula Schrenk in Fischer & C. A. Meyer, Bull. Cl. Phys.-Math. Acad. Imp. Sci. Saint lY'tcrsbourg ser. 2, 2: 199. 1844. TYPE: [Ka¬ zakhstan]. “In desertis salsuginosus ad Ters- akkan,” A. Schrenk s.n. (holotype, LE; iso¬ type?, G). Acknowledgments. This research was supported by the National Science Foundation grant DEB- Novon 5: 309-310. 1995. 310 Novon 9208433 and the National Geographic Society grant 5068-93, both of which were awarded to Al- Shehbaz. We thank Barbara Schaal (Washington University) for her support in conducting the mo¬ lecular studies leading to these results. Literature Cited Al-Shehbaz, I. A. 1988. The genera of Sisymbrieae (Cru- ciferae; Brassicaceae) in the southeastern United States. J. Arnold Arbor. 69: 213-237. Ball, P. W. 1964. Arabidopsis. In: T. G. Tutin et al. (edi¬ tors), FI. Europaea 1: 267-268. Busch, N. A. 1939. Arabidopsis. In: V. L. Komarov (edi¬ tor), FI. URSS 8: 76-80. Hedge, 1. C. 1965. Arabidopsis. In: P. H. Davis (editor), FI. Turkey 1: 489-490. -. 1968. Sisymbrieae. In: K. H. Rechinger (editor), FI. Iranica 57: 309-342. Pavlov, N. V. 1961. Flora Kazakhstana 4: 1—545. Price, R. A., J. D. Palmer & I. A. Al-Shehbaz. 1994. Systematic relationships of Arabidopsis: A molecular and morphological perspective. In: E. M. Meyerowitz & C. R. Sommerville (editors), Arabidopsis. Cold Springs Harbor Laboratory Press, New York. Rollins, R. C. 1993. The Cruciferae of Continental North America. Stanford Univ. Press, California. Schulz, O. E. 1924. Cruciferae-Sisymbrieae. In: A. En- gler (editor), Pflanzenreich IV. 105(Heft 86): 1-388. -. 1936. Cruciferae. In: A. Engler & K. Prantl (ed¬ itors), Natiirlichen Pflanzenfamilien ed. 2. 17B: 227— 658. Vaughan, J. G. & J. M. Whitehouse. 1971. Seed structure and the taxonomy of the Cruciferae. Bot. J. Linn. Soc. 64: 383^09. New Species of Cestrum (Solanaceae) and Synonymy under Two Widespread Species Carmen Benitez de Rojas Facultad Agronomfa, Universidad Central de Venezuela, Maracay, Apartado Correos 4597, Aragua, Venezuela William G. D’Arcy Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A. Abstract. Three new species of Cestrum, C. til- lettii, C. jaramillanum, and C. ruizteranianum, are newly described from Venezuela. Cestrum tomen- tosum L. f. and C. strigilatum Ruiz & Pav6n have been identified as the correct names for two spe¬ cies for which many names have been proposed, and synonymies for these two species are pre¬ sented. Three new species are described from different parts of Venezuela: Sierra de Perija in northeastern Zulia State, the western, Andean region of Merida State, and the more or less central part of the coast¬ al range in the Federal District. The last species is also found in Ecuador. Two other widespread species have been known under many different names, and because con¬ cepts of their ranges have been fragmented with so many names in use, we present the synonymy here, rather than waiting until a revision of the genus can be published for some major region. One of these, C. tomentosum L. f., ranges from Mexico to Peru and Venezuela; the other, C. stri¬ gilatum Ruiz & Pavdn, ranges from Costa Rica to Venezuela, Peru, Brazil, and Argentina. Cestrum embraces many species with local or regional dis¬ tributions, but these two add to the short list of species that are widespread in both Central and South America: C. alternifolium (Jacquin) O. E. Schulz, C. latifolium Lamarck, C. megalophyllum Dunal, C. microcalyx Francey, C. rejlexum Sendt- ner, C. racemosum Ruiz & Pav6n, and C. scandens Vahl. Cestrum jaramillanum Benttez & D’Arcy, sp. nov. TYPE: Ecuador. Pichincha: Centenela, Montafias de Ila, 12 km E of Patricia Pilar, virgin rain forest, 550-650 m, 79°19'W, 0°34'S, 10 July 1979 (fl). Lojtnant & Molau 15835 (holotype, AAU; isotype, CB). Figure 1. Arbor frutexve 2—3 m, foliis ovatis, basi rotundatis, truncatis, subcordatisve, membranaceis, glabrescentis, fo¬ liis minoris ovatis. Inflorescentia axillaria, subumbellata, axibus 6-7 mm longis, tomentulosis glabratisve. Bracteae sessiles foliaceae, lanceolatae, ad maturitatam fructus per¬ sisted, lateraliter calycem involvens. Shrub or small tree 2-3 m, branched, the branches slender, at first compressed, sometimes drying grooved, later terete, tomentulose with mostly reduced, crinkled ascending hairs, the in- temodes 3-4.5 cm long. Leaves ovate, 10—14 X 4-7.5 cm, basally rounded, truncate, or subcor- date, apically short-acuminate, membranous or chartaceous, glabrous above on emerging, the bas¬ al half beneath tomentulose on emerging, glabres- cent, the veins inconspicuous above, the costa and major veins slightly reddish, somewhat elevated and finely puberulent beneath, the lateral veins 7- 8 on each side, ascending, distally arcuate or looping and forming a submarginal vein 0.5—1 cm from the margin, the margin plane; petiole 5—10 mm long, finely pubescent; minor leaves ovate, 7 X 1.5 mm, sessile, glabrate. Inflorescences axil¬ lary, the peduncle ca. 1 mm long, longer in fruit, subumbellate with 3 axes 6-7 mm long, tomen¬ tulose to glabrate, in fruit 15 mm long. Flowers sessile; bracts foliaceous, narrowly lanceolate, 10 X 3.5 mm, persistent in fruit and partially envel¬ oping the calyx laterally, sessile, acuminate, gla¬ brate or pubescent on both sides; calyx tubular, 4 X 2.5 mm, drying stramineous, basally rugulose, glabrate, 5-lobed, the lobes deltoid to broadly acu¬ minate, faintly 5-costate; corolla pale green, 29- 30 mm, the tube 26 mm long, gradually expanded upward, the mouth not contracted, 2.5—3 mm wide, 5-lobed, the lobes 6-6.5 X 1 mm, oblong, the apex acute, the pleated margin short-pilose; stamens 22-23 mm long, the filaments straight, smooth, glabrous, basally adnate for 21 mm, free 1 mm; anthers orbicular, 1 mm across; ovary 0.7 mm diam., slightly rugose, glabrous; ovules 5-6; Novon 5: 311-317. 1995. 312 Novon Figure 1. Oestrum jaramillanum Benitez & I)'A re v. -—A. Flowering branch. —B. Flowers. —C. Flower opened lon¬ gitudinally showing corolla, anthers, and pistil. —D. Fruit. (A, B, C after iMtnant & Molau 15835; D after Steyermark 92147.) style 23 mm long; stigma slightly bilobed, includ¬ ed. Fruit ovoid, 8-10 X 6-7 mm, the pericarp thin, about equaling and loosely enclosed in the bract; seeds 3—4, dark brown, 5-6.5 mm long. Cestrum jaramillanum is found in moist forests on the western slopes of the Ecuadorean Andes and on slopes of the Venezuelan coastal cordillera at 550-1300 m elevation. Volume 5, Number 4 1995 Benitez & D’Arcy New Species and Synonymy in Oestrum 313 This species is named in honor of Jaime Lucio Jaramillo Asanza, professor and active plant col¬ lector at the Pontificia Universidad Catolica del Ec¬ uador in Quito, who has greatly assisted many bot¬ anists visiting his country. This new species is suggestive of Oestrum reflex- um Sendtner and C. amictum Schlechtendal, both described from Brazil and having conspicuous bracts, but in C. reflexum the corolla tube is abrupt¬ ly expanded at the throat, and the plant is scandent; in C. amictum the leaves are basally acute or acu¬ minate and decurrent on the petiole, and the minor leaves are reniform. This new species is apparently a plant of lower or middle elevations, the two col¬ lections being made at 550-1300 m. Paratype. VENEZUELA. Distrito Federal: Cerro Naiguata, laderas pendientes del Norte, arriba del pueblo de Naiguatd, bosque humedo denso, Lomas de Las Deli- cias entre Quebrada de Basenilla y Quebrada Guayoyo, 9—12 km suroeste de Hacienda Cocuizal, 1000—1300 m, 15-19 Nov. (fl, fr), Steyermarh 92147 (NY, US, VEN). Cestrum ruizteranianum Benitez & D’Arcy, sp. nov. TYPE: Venezuela. Merida: Distrito Ran¬ gel, trail from La Negrita downstream towards Puente de La Escalera, montane cloud forest, 2550-2950 m, 1-2 Nov. 1978 (fl), Luteyn et al. 6171 (holotype, NY). Figure 2. Flores copiosi, calyce atrobrunneo, extus glanduloso, pilos sparsos ferens, corolla 20-26 mm longa, dilute viridi ex purpurea. Filamenta 7—9 mm ad corollam coalita, ad basim parte adnata pubescenti, in loco conjunetionis tum- idis, geniculatis, parte libera 7.5—9.5 mm. Fructus haud suppetunt. Shrub 1-4 m tall, erect, branched, young stems angular, terete when mature, scurfy pubescent with dark, crinkled, perhaps branched hairs. Leaves narrowly elliptical to elliptical, 4—6 X 1.5-3 cm, basally obtuse, apically acute or obtuse, firmly membranous, glabrous on both sides except for some scurfy hairs on minor veins, the major veins impressed above, elevated beneath, 8-10 on each side, ascending, looping and uniting near the mar¬ gins, the margins slightly revolute; petioles 6-8 mm long, slender, inrolled, tomentulose above; minor leaves sometimes present, ovate, 10—13 X 5—7 mm, with petioles 0.5-1 mm long. Inflorescences axil¬ lary racemes, sometimes appearing as terminal panicles, 2.5—10 cm long; peduncles 0.7—5 cm long; bracts 1.5-2.5 mm, linear. Flowers with ped¬ icels obsolete; calyx drying dark brown, tubular, 6— 8 X 3—3.5 mm, faintly striate, thick, glandular and with sparse hairs outside, pubescent within and with glandular hairs halfway down, 5-lobed, the lobes 2X2 mm; corolla pale green, purplish out¬ side, pale yellowish green inside, 20—26 mm long, exserted ca. 19 mm from the calyx, the tube 17— 20 mm long, 3-3.5 mm at its widest, contracted around the ovary and then gradually expanded up¬ ward, the throat not constricted, 5 mm wide at the mouth, 5-lobed, the lobes 3-5 X 1.5-2 mm, nar¬ rowly triangular-acuminate, sometimes sparingly pubescent, the pleated margin indexed and tomen- tose; stamens 15.5-19 mm long, adnate 7-9 mm, the adnate portion pubescent for the basal 3—1 mm, free part 7.5—9.5 mm, the insertion geniculate-tu¬ mid, 1.5-2.5 mm long; anthers orbicular, 1 mm across; ovary lobed, 0.7 diam., glabrous, papillose 4-5 mm below the stigma, exceeding the stamens by 1.5 mm; style 15-19 mm, stigma subcapitate, slightly bilobed, included; ovules 16-18. Fruit not seen. Cestrum ruizteranianum is found in cloud forest ravines in the Venezuelan Andes at elevations of 2500-2900 m. This species is named in honor of Luis Rufz- Teran, a botanist who for many years collected in¬ teresting plants in Andean northwestern Venezuela. As a herbarium specimen, this species resembles showy yellow-flowered species from upland Central America, e.g., C. laxum Bentham, but the flowers are reported as pale green and purple. It differs also in its glandular calyx. Although three collections are cited, they were made the same day by the same collecting party, perhaps at the same place. Paratypes. VENEZUELA. Merida: Distrito Rangel, Gavidia, orillas del camino entre el Hato de La Escalera y el puente sobre la quebrada La Escazes, 2550—2950 m, 1 Nov. 1978 (fl), RuCz-Terdn et al. 16171 (MERF, MY), 16154 (MY). Cestrum strigilatum is one of the most wide¬ spread species of Cestrum. It ranges from coastal Brazil and northern Argentina to Costa Rica, oc¬ curring from sea level to about 1600 m. In his monograph of the genus, Francey (1935) used the later name C. calycinum for this species. His ra¬ tionale for use of names of Willdenow in Roemer and Schultes (1819) instead of those of Kunth (1818) was stated on p. 142 of the monograph, which cites Kuntze’s Revisio Generum and Sendtner in Flora Brasiliensis. Some of the following synon¬ ymy was expressed by D’Arcy (1972) and Cabrera (1983), but not by Smith and Downs (1966). We have chosen not to lectotypify the species at this time. Francey included Cestrum lundianum Dunal (in DC., Prodr. 13(1): 658) in synonymy under this spe¬ cies. Photographs of the type (Brazil, Sao Paulo, 314 Novon figure 2. Cestrum ruizteranianum Benftez & D’Arcy. —A. Flowering branch. —B. Flower. —C. Flower opened longitudinally showing corolla, anthers, and pistil. (After Luteyn 6191.) 2cm Volume 5, Number 4 1995 Benitez & D’Arcy New Species and Synonymy in Oestrum 315 Lund 134 (G-DC, -IDC microfiche, =F photo 006898 (MO))) did not permit verification, but its identity with C. strigilatum seems unlikely. Cestrum strigilatum Ruiz & Pav6n, FI. Peruv. 2: 29, t. 156. 1799. TYPE: Peru. Nemoribus ver¬ sus Pozuzo, Chinchao et Cuchero vicos, Ruiz & Pavdn s.n. (holotype, M; isotypes, B, HAL, fide Francey, neither seen, F photo of element once at B, 018394). Cestrum calycinum Kunth, Nov. Gen. Sp. 3: 58. 1818. TYPE: Peru. Prope Gonzanama, 1180 hex, Bonpland (B-W 4461 not seen, microfiche). Synonymized by D’Arcy (1974); Cabrera (1983: 471). Cestrum viridiflorum Hooker, Bot. Mag. t. 4022. 1843. TYPE: Brazil. Porto Alegre, Tweedie (K). Synony¬ mized by Francey (1935: 137); Cabrera (1983: 471). Cestrum unibracteatum Dunal in DC., Prodr. 13(1): 656. 1852. SYNTYPES: Peru. Sylvis Cochero, Dombey s.n. (G-DC, MPU, neither seen), Poeppig s.n. (sub “96 C. longium, Ruiz & Pav6n 1246”) (G-DC not seen, =IDC microfiche, F photo 006899 MO). Syn¬ onymized by Francey (1935: 142). Cestrum unibracteatum var. [p] brachystachys Dunal in DC., Prodr. 13(1): 657. 1852. TYPE: Brazil. Circa Cujaba uhique, da Silva Manso 34 (G-DC not seen, = IDC microfiche, =F photo 023178 MO). Synony¬ mized by Francey (1935: 142). Cestrum cancellatum Dunal in DC., Prodr. 13(1): 657. 1852. TYPE: Peru. Poeppig 3080 (G-DC not seen, = IDC microfiche, =F photo of B, not extant 002969 MO). Cestrum strigilatum (strigillatum) var. [a] laxiflorum Kun- tze. Rev. Gen. PI. 3: 220. 1893. SYNTYPES: Argen¬ tina. Salta: Or£n, Lorenz & Hieronymus s.n. (NY); Bolivia. Cochabamba: Yungas “Juntas,” Kuntze s.n. (NY). Synonymized by Francey (1935: 142); Cabrera (1983: 471). Cestrum calycinum Kunth, Nov. Gen. Sp. 3: 58. 1818. Cestrum strigilatum (strigillatum ) var. [/3] calycinum (Kunth) Kuntze, Rev. Gen. PI. 3: 220. 1893. SYN¬ TYPES: Bolivia. Matto Grosso: West-Velasco bis Tunarigebirge, 200-2400 m, collector not named. Cestrum impressum Rusby, Bull. New York Bot. Gard. 4: 425. 1905. TYPE: Bolivia. La Paz: Bang 2516 (NY not seen). Synonymized by Francey (1935: 137); Smith & Downs (1966: 230). Sessea rugosa Rusby, Bull. New York Bot. Gard. 8: 119. 1912. TYPE: Bolivia. Apolo, 4800 ft., R. S. Williams 2449 (NY not seen). Synonymized by Francey (1935: 142). Cestrum calycinum var. tenuijlorum F rancey, Candollea 6: 142. 1935. Syn. nov. SYNTYPES: Paraguay. Villa Fmcamaci6n, Bettfreund 131; Argentina. Yaguara- Zapa, Niederlein 268b (G not seen, F photo 002998 MO). Cestrum strigilatum var. tenuijlorum Francey, Candollea 6: 144. 1935. Syn. nov. SYNTYPES: Ecuador. Balao, Eggers 14274 (lectotype, designated here, W; isolec- totype, US fragment). Guayaquil, Jameson 57 (B, M, W, none seen). Cestrum aristeguietae Steyermark, Acta Bot. Venez. 6(1- 4): 86. 1971. TYPE: Venezuela. Carabobo: wooded slopes along Rfo San Gian, 2 km below Planta Elec¬ trics, S of Borburata, alt. 350 m, J. & C. Steyermark 95463 (holotype, VEN; isotypes, NY, P, US). Cestrum longifolium Ruiz & Pavdn ex Dunal in DC., Prodr. 13(1): 657. 1852. Invalid name, cited in syn¬ onymy of C. unibracteatum. Cestrum tiilettii Benftez & D’Arcy, sp. nov. TYPE: Venezuela. Zulia: Distrito Perijfi, headwaters of Rfo Guasare, Sierra de PerijA, Serranfa de Va- lledupar, environs of Campamento Frontera V, along international boundary, 2700-3300 m, 10°23'07.8"N, 72°52'42.5"W, 10-19 July 1974 (fl), lillett 747-1021 (holotype, MY; is¬ otypes, AAU, MO, MYF, VEN). Figure 3. Arbor 2—3 m alta, foliis glabris coriaceis, marginibus revolutis, foliis minoribus carentibus. Inflorescentia axil¬ laris, congesta, tomentosa. Flores fragrantes diurni casta- nei, sessiles, calyce viride apice purpureo tubuloso, co¬ rolla 15—18.5 mm longa, staminibus 10-13 mm longis, geniculatis tumidisque ad insertionem; ovula 5. Fructus ovoideus 9X6 mm. Tree 2-3 m tall; stems brown, glabrous, striate and scarred. Leaves firm, subcoriaceous, narrowly elliptical, basally narrowly cuneate, slightly acute near the top, the tip obtuse, 4-6 X 1.3-1.5 cm, dark green, shiny above, dull green beneath, the major veins sunken above, salient beneath, 6—8 on each side, the minor veins impressed beneath, the margin revolute, glabrous on both sides; pet¬ iole canaliculate, 3—7 mm long, glabrous. Minor leaves wanting. Inflorescences axillary, congested, the main axes tomentose, bright matte green, 1.5— 3 cm long, the peduncles 5—7 mm long, pubes¬ cent, thickened, with circular scars from fallen flowers, flowers few per node. Flowers sessile (pedicels obsolete) with faint, sweet fragrance dur¬ ing the day; calyx yellow-green, flushed distally with dark purple, tubular, thick, costate, the cos¬ tas especially conspicuous distally, slightly pu¬ bescent outside with hairs slightly thickened near the base, 4.5-5.5 mm long, 2-2.5 mm thick, 5- lobate, the lobes 1.5-2.5 mm long, narrowly tri¬ angular, the apex pubescent; corolla 15-18.5 mm long, the tube 13—16 mm long, slightly expanded toward the apex, the throat not noticeably con¬ tracted, the mouth ca. 2 mm wide, 5-lobate, the lobes 2-2.5 mm long, the folded margins puber- ulent; stamens 10-13 mm long; filaments white, adnate 6-9 mm, free 4.5 mm, the point of inser¬ tion geniculate-tumid, slightly pilose, 1 mm long; anthers brown, spherical, 0.5 mm diam.; ovary ovoid, glabrous, smooth, 1 mm diam., 5-ovulate; style 12-13.5 mm long, papillose 1-2 mm below the stigma, exceeding the stamens by 1.5 mm; stigma green. Fruit ovoid, 9X6 mm, pericarp thick, opaque; seeds 6—7, 3—3.5 mm long. 316 Novon Figure 3. Cestrum tillettii Benftez & D’Arcy. —A. Flowering branch. —B. Flower. —C. Flower opened longitudinally. —D. Fruits. —E. Leaf, abaxial side. (A, B, C after Tillet 747-1021; E after Tillet 747-929.) Cestrum tillettii occurs in cloud forests at the headwaters of rivers at elevations of 2700-3650 m. It has been found on the eastern side of the Sierra de PerijA, a mountain range shared by Venezuela and Colombia. This species is named in honor of Stephen S. Tillett, professor in the Faculty of Pharmacy at the Universidad Central de Venezuela, Caracas. He was the leader of the expedition that collected the only known specimens of this species. 1 1cm Volume 5, Number 4 1995 Benitez & D’Arcy New Species and Synonymy in Oestrum 317 Paratypes. VENEZUELA. Zulia: Distrito Perij4, headwaters of Rfo Ap6n, Sierra de Perij£, Serranfa Valle- dupar. Hacienda Buena Vista to N side of Cerro Lami- nado, ca. 5 km N of Buena Vista, along international boundary, 3300-3650 m, 10°20'23"N, 72°54'14"W, 9-10 July (fr), Tillett & Honig 747-929 (MY); Dist. Perijd, headwaters of Rfo Guasare along international boundary, “Campamento Frontera V,” 2700-3300 m, 10°23'7.8"N, 72°52'45.5"W, 20-23 July (fl, fr). Wood & Berry 88 (MO, VEN). Oestrum tomentosum is one of the most wide¬ spread species of Oestrum , and in many places it is common, especially as a successional element, occurring from about 1000 to 2600 m elevation. The species has been collected from as far north as Sonora, Mexico, and as far south as extreme southern Peru. It is also found in northwestern Ven¬ ezuela but is generally absent from the Amazon re¬ gion of South America. It is a shrub or small tree, usually erect, with pubescent leaves, calyces, and corolla tubes, and it has a shiny, purple-black, fleshy fruit. The identity of many of the names cited here was suggested by Nee (1986: 49), who also provided a good description (as C. lanatum). Cestrum tomentosum L. f., Suppl. PI. 150. 1781. TYPE: Colombia. Mutis 94 (holotype, LINN 258.6 not seen, =IDC microfiche). Oestrum hirsutum Jacquin, PL Hort. Schoenbr. 3: 41, t. 324. 1798. Syn. nov. TYPE: Indias Occidentales: cult. hort. Schoenbrunn (W, B-W 4449, not seen, =IDC microfiche, =F photo 033032 MO). Cestrum lanuginosum Ruiz & Pav6n, Fl. Peruv. 2: 30, tab. 157. 1799. Syn. nov. TYPE: Peru. Prov. Arequipa: Camana, collibus arenosis, Ruiz s.n. (lectotype, des¬ ignated here, G not seen, =F photo 002984; isolec- totype, F). Cestrum lanatum Martens & Galeotti, Bull. Acad. Roy. Sci. Bruxelles 12(2): 146. 1845. TYPE: Mexico. Ve¬ racruz: dans les bois et haies de la colonie de Mir- ador, 3000 pieds, Galeotti 1208 (holotype, BR not seen; isotypes, G, K, NY, US, W). Cestrum moritzii Dunal in DC., Prodr. 13 (1): 619. 1852. Syn. nov. TYPE: Venezuela. Caracas, Moritz 309 (ho¬ lotype, G-DC not seen, =IDC microfiche, =F photo 002988; isotypes, BM, MO). Cestrum miersianum Weddell, Chlor. And. 2: 97. 1857. TYPE: Colombia. Sierra Nevada de Santa Marta, 3,300 m, Linden 1615 (holotype, G-DC not seen, = IDC microfiche). Cestrum miersianum Pittier, J. Wash. Acad. Sci. 22(2): 37. 1932, not Weddell (1857). Syn. nov. TYPE: Vene¬ zuela. Merida: San Rafael de Mucuchfes, 3150 m, A. Jahn 767 (holotype, VEN; isotype, MO). Cestrum neomiersianum Benftez, Rev. Fac. Agron. (Maracay) 7: 90. 1974. New name. Cestrum diasae Pittier, J. Wash. Acad. Sci. 22 (2): 29. 1932. Syn. nov. TYPE: Venezuela. M6rida: Misintd, arriba de Mucuchfes, 3500 m, Pittier 12919 (holo¬ type, VEN; isotype, US). Cestrum meridanum Pittier, J. Wash. Acad. Sci. 22(2): 36. 1932. Syn. nov. TYPE: Venezuela. Vecinidades de Merida, 1700 m, Pittier 12858 (holotype, VEN; iso¬ type, US). Cestrum densiflorum Francey, Candollea 6: 195. 1935. Syn. nov. SYNTYPES: Venezuela. Merida: entre Chachopo y Timotes, Pittier 13294 (lectotype, des¬ ignated here, NY; isolectotypes, F, MO, US, VEN); Pittier 13323 (NY, VEN). Cestrum densiflorum Francey var. puberulum Francey, Candollea 6: 196. 1935. Syn. nov. TYPE: Venezuela. Trujillo. San Pablo de Mendoza, Pittier 13323 (ho¬ lotype, VEN; isotype, US). Cestrum verbascifolium Zuccarini ex Francey, Candollea 6: 191. 1935. Syn. nov. TYPE: cultivated in Berlin and Munich (B-W 4449 not seen, =IDC microfiche). Cestrum ambatense Francey, Candollea 6: 169. 1935. Syn. nov. TYPE: Ecuador. Prov. Tungurahua: vicinity of Ambato, Pachano 75 (holotype, NY; isotype, US fragment). Acknowledgments. The authors acknowledge grants from the Consejo Nacional de Investiga- ciones Cientfficas y Tecnoldgicos, CONICIT (PI- 056) and the National Science Foundation, NSF (INT-9116039). Michael Nee, The New York Botanical Garden, reviewed the manuscript and augmented parts of it. Literature Cited Cabrera, A. L. 1983. Flora de la Provincia de Jujuy, Re- publica Argentina. Colecci6n Cientffica del INTA, Bue¬ nos Aires. D’Arcy, W. G. 1972 [1973]. Solanaceae. In: R. E. Wood- son et al., Flora of Panama. Ann. Missouri Bot. Gard. 60: 573-870. Francey, P. 1935-1936. Monographic du genre Cestrum L. Candollea 6: 46-398; 7: 1-132. Kunth, C. S. 1818. Solaneae. In: Nova Genera et Species Plantarum, Vol. 3. Paris. Nee, M. 1986. Solanaceae. In: Flora de Veracruz. 49: 1- 191. Inst. Nac. Invest. Recursos Bi6t., Xalapa. Roemer, J. J. & J. A. Schultes. 1819. Caroli a Linn6 equitis Systema vegetabilium secundum classes ordines genera species, Vol. 4. Stuttgart. Smith, L. B. & R. J. Downs. 1966. Solan£ceas. Parte I. In: P. R. Reitz (editor). Flora Illustrada Catarinense. Herb£rio “Barbosa Rodrigues,” Itajai, Santa Catarina, Brazil. Two New Species of Fuchsia Section Fuchsia (Onagraceae) from Southern Ecuador Paul E. Berry Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A. Abstract. Two new species of Fuchsia sect. Fuchsia (Onagraceae) from the Andes of southern Ecuador are described and illustrated. Fuchsia campii P. E. Berry is known from mid-elevation cloud forest of Azuay and Loja Provinces, and Fuchsia summa P. E. Berry is a high-elevation spe¬ cies currently known only from Loja Province. Both occur in Podocarpus National Park, together with several other rare species of Fuchsia. Studies and collections of plants from southern Ecuador have increased dramatically in the past de¬ cade, especially in areas such as Podocarpus Na¬ tional Park in Loja Province. Many novel plants have been located, and this paper describes two new spe¬ cies of Fuchsia that occur in cloud forest areas on the slopes of Cerro Toledo, one of them also known further north in Azuay Province. Three other species previously thought to be extremely rare in southern Ecuador are also known from Cerro Toledo: F. andrei I. M. Johnston, F. scherffiana Andre, and F. steyer- markii P. E. Berry. All these taxa belong to the larg¬ est section of the genus, section Fuchsia, which now approaches 65 species. They are also examples of the high degree of local endemism in cloud forests of southern Ecuador, where moist montane habitats are often separated by dry valleys and show a wide range of precipitation regimes. Fuchsia campii P. E. Berry, sp. nov. TYPE: Ec¬ uador. Azuay: Quebradas leading into the Rfo Collay, 3—8 km N of Sevilla de Oro, 7000— 8300 ft., 27 Aug. 1945, W. H. Camp E-4994 (holotype, RSA herbarium # 71952; isotypes, G-DEL, MO, NY, UC, US). Figure 1. Frutex 1.5-4 m altus, foliis 3-vel 4-verticillatis, inter- dum oppositis, 3—9 X 1—3 cm, margine subintegro vel 7— 10-dentato, floribus axillaribus apicem versus, pedicellis 18-30 mm longis, tubo florali roseo 29-45 mm longo, se- palis lanceolatis 12-15 mm longis, petalis auriantiacis 8- 11 mm longis, 4—6 mm latis. Erect to recumbent shrub 1.5-4 m tall, well branched with mosdy ascending branches; young growth pubescent, the trichomes 0.1-0.5 mm long, appressed to erect; branchlets terete to slighdy an- Novon 5: 318-322. 1995. gled, older branches with purplish, exfoliating bark. Leaves mostly in whorls of 3 or 4, sometimes opposite or subopposite; blade membranous, narrowly elliptic, 3—9 cm long, 1-3 cm wide, acute to acuminate at the apex, acute at the base, subglabrous to sparsely stri- gose when young on both surfaces and margin, tri¬ chomes persisting mainly along the midvein and sec¬ ondary nerves on lower surface when fully expanded, upper surface smooth, shiny, and light to dark green, lower surface paler green; margin nearly entire, usu¬ ally with 7-10 glandular teeth projecting slightly from the edge especially in the upper Vi to % of the blade; secondary nerves (5-)8-10 on either side of the mid¬ vein; petiole strigose, (4—)6-25 mm long; stipules nar¬ rowly lanceolate when young, 1—1.5 mm long, ca. 0.2 mm wide, the tip quickly abscising and the thicker lower third persisting or abscising later. Flowers ax¬ illary in the upper leaf axils, 2-several per node, ped¬ icel 18-30 mm long; floral tube narrowly funnelform, (29-)32-45 mm long, 2.5-3.5 mm wide at the base, narrowed to 2—2.5 mm wide in the lower V4, then gradually widened until 5-8 mm wide at the rim, sparsely strigose outside, densely pubescent inside in the lower narrowed portion; nectary a doughnut¬ shaped ring 1—2 mm high and 2—3 mm wide at the base of the tube surrounding the style; ovary 4-angled, 5-6 mm long, 2-3 mm thick; sepals narrowly trian¬ gular or lanceolate, 12-15 mm long, 4—5 mm wide at the base; petals elliptic to obovate, 8-11 mm long, 4— 6 mm wide, broadly acute at the apex, sometimes with a small mucro; tube and sepals bright pink except for the green sepal tips, petals orange and darker than the tube; style pink, 40-64 mm long, densely pubes¬ cent in lower third, stigma pink, capitate, ca. 2 mm high, ca. 2.5 mm wide; filaments 10-12 mm and 7— 9 mm long, anthers oblong, 2-3 mm long, 1-1.5 mm wide. Fruit oblong-ellipsoid before maturity, subro¬ tund when ripe, 9-15 mm long, 5-9 mm thick; seeds flattened, obtriangular in outline, 1.8-2.2 mm long, 1.2—1.6 mm wide, ±200 per fruit. Gametic chromo¬ some number n = 11. Ecology and distribution. Occurring in thick¬ ets, along streams, and along roadsides in pasture zones on moist mountain slopes, 2300—2700 (—3350) m, in Azuay and Loja Provinces, Ecuador. Volume 5, Number 4 1995 Berry Fuchsia from Southern Ecuador 319 Figure 1. Fuchsia campii P. E. Berry. —A. Habit. -B. Detail of stipules. -C. Flower at anthesis. —D. Flower in longitudinal cross section. —E. Petal. —F. Anther. Drawn from Berry & Brako 4646. Etymology. Named for Wendell H. Camp (1904—1963), a botanist who made extensive plant collections in southern Ecuador for the New York Botanical Garden in the mid 1940s. The type specimen of Fuchsia campii was treated by Munz (1974) under Fuchsia ayavacensis HBK, but that species has wider and shorter petals, larger and more pubescent leaves, and occurs in drier habitats in southern Ecuador and in northern Peru. One of the paratypes cited here (Camp E-4257) was listed by Munz under F. canescens Bentham, but that species is confined to southern Colombia and 320 Novon has thick, firm flowers, short pedicels, trowel¬ shaped petals, tetragonous sepals in bud, and most¬ ly 4-whorled leaves. Fuchsia campii might be con¬ fused with F. vulcanica Andre (sensu lato) but differs in its narrower, orange petals contrasting with the pink floral tube and sepals (the sepals also turn green toward the apex) and in the narrower leaves with margins entire or else with small teeth. It is apparently diploid, as chromosome counts of n = 11 and 2n = 22 were obtained from Escobar 1543, and three collections identified as F. vulcan¬ ica from Azuay Province were tetraploid (Berry, 1982). Paratypes. ECUADOR. Azuay: E Cordillera, 1—8 km N of the village of Sevilla de Oro, 8000-9000 ft., 27 July 1945, Camp E-4257 (NY, RSA); quebradas leading into the Rfo Collay, 3—8 km N of Sevilla de Oro, 7000-8000 ft., 27 Aug. 1945, Camp E-4994 (MO); Partidero Llantera- Chiquintad-Saucay-Guandum, 10,000 ft., 27 May 1979, Jaramillo 1068 (AAU, QCA). Loja: road from Yangana to Cerro Toledo, 2360 m, 13 Nov. 1988, Berry & Brako 4646 (MO, QCA); between Yangana and Toledo, 7700-8000 ft., 26 July 1979, Escobar 1543 (MO, TEX), 1544 (MO, TEX), 1545 (MO, TEX). Fuchsia summa P. E. Berry, sp. nov. TYPE: Ec¬ uador. Loja: shrub patches in open pArarno above tree line, road from Yangana to Cerro Toledo, 3150 m, 13 Nov. 1988. P. E. Berry & L Brako 4644 (holotype, MO; isotypes, QCA, US). Figure 2. Frutex 30—100 cm altus, ramulis erectis foliis apicem versus congestis 3-vel 4-verticillatis, 15-35 mm longis, petiolis 5-8 mm longis. Flores axillares, 4-angulati, ped- icellis 4—7(—10) mm longis, tubo florali 18-26(-32) mm longo, sepalis 10—14 mm longis, petalis suborbicularibus 8-12 X 7-9 mm. Erect shrub 30—100 cm tall; branches erect, mostly bare except for the leafy tips, with short intemodes mostly 6-12(-25) mm long and pro¬ truding leaf base scars, branchlets appressed-pu- bescent with whitish trichomes. Leaves mostly in whorls of 3 or less often 4, densely clustered at the stem apices; blade subcoriaceous, broadly el¬ liptic, 15—35 mm long, 12—18 mm wide, broadly to narrowly acute or rounded at the apex and base, usually with scattered appressed trichomes along the midvein on both surfaces, upper surface dark green, lower surface paler green; margin gland- denticulate with trichomes between the teeth; sec¬ ondary nerves 4—6 on either side of the midvein; petiole 5-8 mm long; stipules narrowly lanceolate, 1—2 mm long, 0.3—0.5 mm wide. Flowers axillary and pendent in the upper leaf axils, 2-several per node, pedicel 4—7 mm long in flower, to 10 mm long in fruit; floral tube narrowly funnelform, 18— 26(-32) mm long, 2-2.5 mm wide at the base, 4— 6 mm wide at the rim, 4-angled in cross section, appressed-pubescent to nearly glabrous in the lower half outside, densely villous in lower half inside; nectary a doughnut-shaped ring ca. 1.5 mm high and 2 mm wide at the base of the tube surrounding the style; ovary 4-angled, 5—6 mm long, 2-3 mm thick; sepals broadly elliptic, acute to subacuminate at the apex, 10—14 mm long, 5— 6 mm wide at the base, quadrangular in cross sec¬ tion when joined before opening; petals suborbi- cular, slightly spreading, 8-12 mm long, 7—9 mm wide, broadly acute to mucronate at the apex, broadly acute to spathulate at the base; tube pale red, sepals pale green in bud but becoming flushed with red by anthesis, petals slightly darker red than the tube; style red, 28—35 mm long, vil¬ lous in lower half, stigma pink, capitate, ca. 2 mm high by 2 mm wide; filaments 5—7 mm and 4-5 mm long, anthers broadly oblong, ca. 2 mm long and 1.5 mm wide. Fruit 4-angled before fully ma¬ ture, 9-12 mm long, ca. 5 mm thick; seeds flat¬ tened, obtriangular in outline, 1.9—2.1 mm long, ca. 1.5 mm wide. Ecology and distribution. Occurring in low shrub patches among open grassy areas at and above tree line, on very humid upper slopes of Cer¬ ro Toledo east of Yangana in Loja Province, Ecua¬ dor, 3100-3450 m elevation. Etymology. From the Latin summus, meaning highest or uppermost, in reference to this species occurring at tree line and at higher elevations than any other species of Fuchsia on the same mountain. This species forms small, dense thickets in ex¬ posed areas close to tree line. Its leaves are small, with few secondary veins, and are tightly congest¬ ed toward the stem apices. The flowers are dis¬ tinctive in their short pedicels that angle sharply downward alongside the stems and in the strongly quadrangular ovary, floral tube, and sepals in bud. Fuchsia summa appears to be morphologically most similar to F. loxensis HBK, a broadly circum¬ scribed species in Berry (1982). Besides the char¬ acters listed above, F. summa differs in the green¬ ish coloration of the sepals and generally more pubescent flowers. Paratypes. ECUADOR. Loja: road from Yangana to Cerro Toledo, 3140 m, 12 Nov. 1988, Berry & Brako 4643 (MO, QCA); Cerro Toledo, Parque Nacional Podocarpus, 3000-3400 m, 27 Nov. 1988, Madsen 75617 (AAU, MO), 1 Dec. 1988, Madsen et al. 75640 (AAU), 21 July 1989, Madsen et al. 86119 (AAU, MO), 30 Oct. 1989, Madsen 86320 (AAU), 30 Oct. 1989, Madsen 86343 (AAU); Cerro Volume 5, Number 4 Berry 1995 Fuchsia from Southern Ecuador e E io I-1 321 Figure 2. Fuchsia summa P. E. Berry. A. Habit. B. Detail of stipules. —C. Leaf and detail of margin. —D. Flower at anthesis. —E. Flower in longitudinal cross section with details of the stigma and nectary; at lower left a cross section of the ovary. —F. Detail of a sepal, petal, and two stamens. Drawn from Berry & Brako 4644. 322 Novon Toledo E of Yangana, Parque Nacional Podocarpus, 3400- 3450 m, 26 Feb. 1985, 0llgaard et al. 58186 (AAU). Acknowledgments. This work was supported by grant 3827-88 of the National Geographic Society. The illustrations were prepared by Bobbi Angell, and Roy Gereau kindly reviewed the Latin diag- Literature Cited Berry, P. E. 1982. The systematics and evolution of Fuch¬ sia sect. Fuchsia (Onagraceae). Ann. Missouri Bot. Gard. 69: 1—198. Munz, P. A. 1974. Flora of Ecuador. 141. Onagraceae. Opera Bot. Ser. B no. 3, 3—46. New Combinations in Chamaesyce (Euphorbiaceae) from Queensland, Australia Paul 1. Forster and Rodney J. F. Henderson Queensland Herbarium, Queensland Department of Environment & Heritage, Meiers Road, Indooroopilly, Queensland 4068, Australia Abstract. The new combinations Chamaesyce carissoides (F. M. Bailey) D. C. Hassall ex P. I. For¬ ster & R. J. F. Henderson, C. centralis (B. G. Thompson) P. I. Forster & R. J. F. Henderson, C. coghlanii (F. M. Bailey) D. C. Hassall ex P. I. For¬ ster & R. J. F. Henderson, C. petala (Ewart & L. R. Kerr) P. I. Forster & R. J. F. Henderson, and C. psammogeton (P. S. Green) P. I. Forster & R. J. F. Henderson are made. Recognition of Chamaesyce S. F. Gray as a genus distinct from Euphorbia L. has received consider¬ able support (e.g., Hassall, 1976; Koutnik, 1984, 1987; Smith, 1981; Webster, 1994) or rejection (e.g., Radcliffe-Smith, 1980; McPherson & Tirel, 1987; Oudejans, 1990) among taxonomists. Revi¬ sions of Chamaesyce and Euphorbia s. str. in Aus¬ tralia were undertaken by Hassall (1977a). These were unfortunately never published in full; only a revision of the latter genus (Hassall, 1977b), valid transfers of some names into Chamaesyce (Hassall, 1976), and discussion of arid zone speciation pat¬ terns (Hassall, 1982) appeared in print. Recognition of Chamaesyce in Australia subse¬ quent to Hassall’s work has not been widespread, with most authors opting for a broadly circum¬ scribed Euphorbia (e.g., Forster, 1993, 1994; OGreen, 1993; Forster & Henderson, 1994; Thomp¬ son, 1992; Weber, 1986; Wheeler, 1987) as op¬ posed to the narrower view of James and Harden (1990). Despite this lack of local support, we are now of the opinion that the arguments proposed by Hassall (1976), Koutnik (1984, 1987), and Webster (1994) for recognizing Chamaesyce as a genus in its own right are convincing and that their views should be supported. As a new edition of “Queens¬ land Vascular Plants Names and Distribution” the Queensland Herbarium (BR1) census of Queens¬ land Plants in which the authors intend to recog¬ nize Chamaesyce as a distinct genus is soon to be prepared, new names are required for certain taxa. While valid combinations exist in Chamaesyce for the majority of taxa that occur in Queensland, no combinations exist for five species. Therefore, these combinations are made here. Two of these combi¬ nations were listed in the unpublished thesis of Hassall (1977a) but never published. 1. Chamaesyce carissoides (F. M. Bailey) D. C. Hassall ex P. I. Forster & R. J. F. Henderson, comb. nov. Basionym: Euphorbia carissoides F. M. Bailey, Queensland Agric. J. 16: 449. 1906. TYPE: Australia. Queensland: Cook District, Herberton, Ringrose [AQ342536] (ho- lotype, BRI). 2. Chamaesyce centralis (B. G. Thompson) P. I. Forster & R. J. F. Henderson, comb. nov. Bas¬ ionym: Euphorbia centralis B. G. Thompson, Nuytsia 8: 353. 1992. TYPE: Australia. North¬ ern Territory: 3 km SW of Alice Springs, 18 Jan. 1990, Thompson 3408 (holotype, DNA not seen; isotypes, BRI; AD & CANB not seen). 3. Chamaesyce coghlanii (F. M. Bailey) D. C. Hassall ex P. I. Forster & R. J. F. Henderson, comb. nov. Basionym: Euphorbia coghlanii F. M. Bailey, Bot. Bull. 13: 12. 1896. TYPE: Australia. Queensland: Gregory North District, Roxborough, Georgina River, Dec. 1895, Bai¬ ley [AQ342538] (holotype, BRI). 4. Chamaesyce petala (Ewart & L. R. Kerr) P. I. Forster & R. J. F. Henderson, comb. nov. Bas¬ ionym: Euphorbia petala Ewart & L. R. Kerr, Proc. Roy. Soc. Victoria, New Series 39(1): 1. 1926. TYPE: Australia. Northern Territory: Wycliffe Well, June 1924, Ewart (holotype, MEL not seen). 5. Chamaesyce psammogeton (P. S. Green) P. I. Forster & R. J. F. Henderson, comb. nov. Basionym: Euphorbia psammogeton P. S. Green, Kew Bull. 48: 314. 1993. TYPE: Aus¬ tralia. Lord Howe Island: Blinky Beach, 13 Nov. 1963, Green 1625 (holotype, K not seen; isotype, A not seen). Literature Cited Forster, P. I. 1993. Rediscovery of Euphorbia carissoides (Euphorbiaceae). Austral. Syst. Bot. 6: 269—272. Novon 5; 323-324. 1995. 324 Novon -. 1994. Tropical beaches and Euphorbia atoto in Australia and Papuasia. Cact. Succ. J. (U.S.) 66: 7-10. -& R. J. F. Henderson. 1994. Euphorbiaceae. Pp. 107—117 in: R. J. F. Henderson (editor), Queensland Vascular Plants: Names and Distribution. Queensland Department of Environment & Heritage, Brisbane. Green, P. S. 1993. Notes relating to the floras of Norfolk & Lord Howe Island, IV. Kew Bull. 48: 307-325. Hassall, D. C. 1976. Numerical and cytotaxonomic evi¬ dence for generic delimitation in Australian Euphor- bieae. Austral. J. Bot. 24: 633-640. -. 1977a. Systematic studies in Australian Eu- phorbieae (Euphorbiaceae). Doctor of Philosophy The¬ sis, University of Queensland. St Lucia, Brisbane. -. 1977b. The genus Euphorbia in Australia. Aus¬ tral. J. Bot. 25: 429-453. -. 1982. Distribution and evolution of Euphorbia and Chamaesyce (Euphorbiaceae) in the arid zone of Australia. Pp. 323-328 in W. R. Barker & P. J. M. Greenslade (editors), Evolution of the Flora and Fauna of Arid Australia. Peacock Publications, Frewville. James, T. A. & G. J. Harden. 1990. Euphorbiaceae. In: G. J. Harden (editor), FI. New South Wales 1: 389—430. Univ. New South Wales Press, Sydney. Koutnik, D. L. 1984. Chamaesyce (Euphorbiaceae) — A newly recognised genus in southern Africa. S. African J. Bot. 3: 262-264. -. 1987. A taxonomic revision of the Hawaiian species of the genus Chamaesyce (Euphorbiaceae). Al- lertonia 4: 331-388. McPherson, G. & C. Tirel. 1987. Flore de la Nouvelle- Cal6donie et Dependences. 14. Euphorbiaceae I. Mu¬ seum National D’Histoire Naturelle, Paris. Oudejans, R. C. H. M. 1990. World Catalogue of Species Names Published in the Tribe Euphorbieae (Euphorbi¬ aceae), With Their Geographical Distribution. R. C. H. M. Oudejans, Utrecht. Radcliffe-Smith, A. 1980. Euphorbia. In: A. K. Airy Shaw, The Euphorbiaceae of New Guinea. Her Majesty’s Stationery Office, London. Smith, A. C. 1981. Euphorbiaceae. In: FI. Vit. Nova 2: 439-575. Pacific Tropical Botanical Gardens, Lawai. Thompson, B. G. 1992. Three new species of Euphorbia L. subgenus Chamaesyce Rafinesque (Euphorbiaceae) from central and northern Australia. Nuytsia 8: 351— 360. Weber, J. Z. 1986. Euphorbiaceae. In: J. P. Jessop & H. R. Toelken (editors), FI. South Australia 2: 735-768. Government Printer, Adelaide. Webster, G. L. 1994. Synopsis of the genera and supra- generic taxa of Euphorbiaceae. Ann. Missouri Bot. Gard. 81: 33-144. Wheeler, J. R. 1987. Euphorbiaceae. In: N.G. Marchant et al., FI. Perth Region, 444-455. Western Australian Herbarium, South Perth. Gladiolus somalensis (Iridaceae), a New Species from Northeastern Somalia Peter Goldblatt B. A. Krukoff Curator of African Botany, Missouri Botanical Garden, P. 0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. Mats Thulin Department of Systematic Botany, University of Uppsala, Villavagen 6, S-752 36 Uppsala, Sweden Abstract. Gladiolus somalensis is a new species restricted to the limestone escarpments of the Sa- naag and Bari Regions of northeastern Somalia. Al¬ though Gladiolus is the largest genus of Iridaceae subfamily Ixioideae and has some 83 species in tropical Africa, only 4 species occur in Somalia and only G. somalensis is endemic there. The affinities of this new species are uncertain, but the relatively small flowers with a short perianth tube appear to place the species in subgenus Gladiolus, a taxon that is most diverse and speciose in southern Af¬ rica. The Old World genus Gladiolus, a member of Iridaceae subfamily Ixioideae and one of the larger, if not the largest genus of the family, consists of some 250 species (Goldblatt, in press; Goldblatt & Manning, ms.). The center of Gladiolus in terms of species numbers and taxonomic diversity is tem¬ perate southern Africa, more particularly the winter rainfall region of southern Africa (Lewis et al., 1972; Goldblatt, 1991). The genus is, however, well represented in tropical Africa, where some 83 spe¬ cies occur south of the Sahara and north of the borders of Namibia, Botswana, and South Africa (Goldblatt, in press), 75 of them endemic. At least 150 species occur in southern Africa (Goldblatt & Manning, in prep.), another 8 in Madagascar (Gold¬ blatt, 1989), and perhaps 10 more in Europe, North Africa, and the Middle East. Here we describe a new species of Gladiolus restricted to northeastern Somalia, one of only four species of the genus re¬ corded from that country, and the only one endemic there (Goldblatt, 1995). Although first collected in 1956, Gladiolus so¬ malensis remained until now too poorly known to be described or even assigned with confidence to genus. The type collection made in January 1995, however, is well preserved, and is accompanied by photographs and spirit material. This has made it possible to draw up a description, formally name the species, and have an illustration made. Gladiolus somalensis Goldblatt & Thulin, sp. nov. TYPE: Somalia. Sanaag Region, escarpment S of Laasqoray, near Ragad, 11°00'N, 48°29'E, evergreen bush on limestone, 16 Jan. 1995, Thulin, Dahir & Hassan 9079 (holotype, UPS). Figure 1. Plantae (7—)12—30 cm altae, eormo ca. 12 mm in dia- metro, foliis 4—5 linearibus (1-12—1 mm latis, spica 2—10 florum, bracteis extemis viridibus (7—)12—17(-23) mm longis internis minoribus, flos aurantiacus, tepalis later- alibus infemis infra flavis (raro tepalo inferiore infra flavo), tubo perianthii infundibuliformi 6-8 mm longo, tepalis lanceolatis tribus supemis majoribus 16-18 X ca. 8 mm, infemis 12—15 X 5—5.5 mm, filamentis 8-10 mm longis, antheris 3—5 mm longis, ramis styli filiformibus ca. 2.5 mm longis. Plants (7-)12-30 cm high. Corm obconic, ca. 12 mm diam., the tunics of softly textured layers, these decaying with age into fine netted fibers. Leaves four or five, the lower three basal and longest, reaching at least to the base of the spike and one or more often slightly exceeding the spike, the blades ± linear, (l-)2-4 mm wide, the upper one or two leaves inserted on the lower half of the stem, smaller than the basal leaves. Stem erect, simple or with one or two branches, ca. 1.2 mm diam. be¬ low the base of the spike. Spike lightly flexuose, 2- 10-flowered; bracts green and soft-textured, the out¬ er (7-)12-17(-23) mm long, the inner about two-thirds as long as the outer. Flowers zygomor- phic, orange, the lower lateral tepals (rarely the lower median tepal) bright yellow in the lower half; perianth tube funnel-shaped, 6-8 mm long; tepals unequal, lanceolate, the upper three larger than the lower, the dorsal inclined over the stamens, 16-18 X 8 mm, the upper laterals about as long, the lower tepals ± parallel to the ground, the lower lateral Novon 5: 325-328. 1995. 326 Novon Figure 1. Gladiolus somalensis Goldblatt & THulin size. (Drawn by John C. Manning.) (from Thulin, Dahir & Hassan 9079). Scale: approximately life Volume 5, Number 4 1995 Goldblatt & Thulin Gladiolus somalensls 327 tepals ca. 15 X 5.5 mm, the margins raised below and the surface channeled in the lower half, the lower median ca. 12 X 5 mm, not normally chan¬ neled. Filaments 8—10 mm long, exserted 1—6 mm from the tube; anthers 3—5 mm long, yellow. Ovary ovoid, 2—3 mm long; style arching over the stamens, dividing ca. 1.5 mm beyond the anther apices, the branches ca. 2.5 mm long, filiform, evidently not expanded apically. Capsules and seeds unknown. Flowering January and February. Distribution and Habitat Still poorly known, Gladiolus somalensis appears to be restricted to the limestone escarpments of the Sanaag and Bari Regions facing the Gulf of Aden in northeastern Somalia. In the type locality, in the eastern part of the Cal Madow Range in Sanaag, only a single plant was seen despite extensive searching. This was growing in evergreen bushland dominated by Buxus hildebrandtii Baillon (Buxa- ceae), a native boxwood, and Cadia purpurea Forss- kal (Fabaceae) at 1350 m elevation. Gladiolus schweinfurthii Baker, a species of moderate eleva¬ tions in Eritrea, Ethiopia, and Kenya, is a fairly common and conspicuous plant in the area. Cal Madow is a major center of endemism in Somalia, described in detail by Thulin (1994a). Two early collections are referred to Gladiolus somalensis, both from the Cal Miskat Range in Bari Region, some 150 km to the east of the type local¬ ity. Nothing is known about the habitat of the spe¬ cies in this area, one that is generally drier than Cal Madow, and from where neither Buxus or Cadia is known. Diagnosis, Variation, and Relationships Gladiolus somalensis is readily distinguished from all other tropical African species of the genus by its small, bright orange flowers with a conspic¬ uous yellow nectar guide located on the lower lat¬ eral tepals (or sometimes, and presumably abnor¬ mally, only on the lower median tepal) and fairly short perianth tube, 6—8 mm long (Fig. 1). The flow¬ ers have a superficial resemblance to species of Tritonia, a genus in which most species have bright orange flowers with yellow markings on the lower tepals. A characteristic feature of most orange-flow¬ ered species of Tritonia is the presence of a large, tooth-like callus in the lower center of each of the lower tepals, and this feature is lacking in G. so¬ malensis. Gladiolus and Tritonia also differ funda¬ mentally in their seeds and in the nature of the floral bracts. The seeds of Tritonia are prismatic to globose, smooth, and have the vascular trace ex¬ cluded (Goldblatt & Manning, 1995), and the bracts of the genus are typically fairly short, more or less membranous to scarious, and have bifurcate or trifurcate apices. The capsules and seeds of G. somalensis are unknown and cannot be used to as¬ sist in generic placement. The bracts, however, are relatively large, soft-textured, and green, hence quite typical of Gladiolus. The style branches are linear and do not appear to be apically expanded as is the case with most species of Gladiolus, but this minor difference in style branch structure does not seem particularly significant in relation to ge¬ neric placement. The two collections from Cal Miskat both consist of small plants up to 12 cm high with leaves 1-2 mm wide, whereas the type specimen is 30 cm high and has leaves up to 4 mm wide. Sacco s.n. is ster¬ ile, but the inflorescence of Azzaroli 6 is unbran¬ ched and has only two flowers. The type, from Cal Madow, has a branched inflorescence with 10 flow¬ ers on the main axis. Also, the flowers are some¬ what smaller in Azzaroli 6 than in the type. To prop¬ erly evaluate these differences further field studies are needed. For the present we think these three collections are best regarded as representing a sin¬ gle variable species. The relationships of Gladiolus somalensis within the genus are uncertain. It bears a fair resemblance to the Ethiopian G. calcicola Goldblatt and to the Eritrean G. mensensis Baker (both subgenus Ophio- lyza), largely because both these species have rel¬ atively small flowers and are fairly small plants with several well-developed foliage leaves (Goldblatt, in press). It is more likely that the affinities of G. so¬ malensis lie with southern African species of sub¬ genus Gladiolus. Members of that subgenus usually have short-tubed flowers of a size comparable to those of G. somalensis. Most southern African spe¬ cies of subgenus Gladiolus have ellipsoid, apically acute capsules, unlike the ovoid to oblong, apically three-lobed capsules of G. calcicola and G. men¬ sensis, more characteristic of subgenus Ophiolyza. Until more is known about G. somalensis, especially the nature of its capsules and seeds, its affinities remain speculative at best. Although a relationship with southern African species seems less likely because of the distance involved, there are phytogeographical connections between the Somalian region and arid parts of southern Africa (de Winter, 1974; Thulin, 1994b). In Iridaceae the distribution of Babiana is a strik¬ ing example. One species of this largely South Af¬ rican genus occurs on Socotra, an island off the coast of Somalia (Lewis, 1959). Another example 328 Novon involving a petaloid monocot concerns a new spe¬ cies of Trachyandra (Asphodelaceae) that grows in the same area as Gladiolus somalensis, but in a different habitat, crevices in shady limestone rocks (Thulin, 1995). Trachyandra is otherwise a mainly southern African genus with a few species extend¬ ing into tropical Africa. Paratypes. SOMALIA. Bari Region: Wadi Hantara, Candala. 6 Feb. 1956, Azzaroli 6 (FT); Azienda Uar Ma¬ han, Jan.-Feb. 1959, Sacco s.n. (FT). Acknowledgments. Mats Thulin’s fieldwork in Somalia was supported by SAREC (Swedish Agen¬ cy for Research Cooperation with Developing Countries). We thank John Manning for the illus¬ tration reproduced here. Literature Cited Goldblatt, P. 1989. Systematics of Gladiolus (Iridaceae) in Madagascar. Bull. Mus. Natl. Hist. Nat., S6r. 4, Sect. B, Adansonia 11: 235-255. -. 1991. An overview of the systematics, phylogeny and biology of the southern African Iridaceae. Contrib. Bolus Herb. 13: 1-74. -. 1995. Iridaceae. Pp. 62-67 in M. Thulin (edi¬ tor), Flora of Somalia, volume 4. Royal Botanic Gar¬ dens, Kew. In press. -. In press (1996). Gladiolus in Tropical Africa. Timber Press, Portland, Oregon. -& J. C. Manning. 1995. Phylogeny of the African genera Anomatheca and Freesia (Iridaceae—Ixioideae), and a new genus Xenoscapa. Syst. Bot. 20: 161-178. Lewis, G. J. 1959. The genus Babiana. J. S. African Bot., Suppl. 3. -. A. A. Obermeyer & T. T. Barnard. 1972. A re¬ vision of the South African species of Gladiolus. J. S. African Bot., Suppl. 10: 1-316. Thulin, M. 1994a. Cal Madow, Somalia. Pp. 194—195 in S. D. Davis, V. H. Heywood & A. C. Hamilton (editors). Centre of Plant Diversity, a Guide and Strategy for their Conservation, Volume 1. Information Press, Oxford, UK. -. 1994b. Aspects of disjunct distributions and en¬ demism in the arid parts of the Horn of Africa, partic¬ ularly Somalia. Pp. 1105-1119 in J. H. Seyani & A. C. Chikuni (editors). Proceedings of the Xlllth plenary Meeting AETFAT, Malawi, Volume 2. National Herbar¬ ium and Botanic Gardens of Malawi, Zomba. -. 1995. Trachyandra triquetra (Asphodelaceae), a new species from the Horn of Africa. Nordic J. Bot. 16. In press. Winter, B. de. 1974. Floristic relationships between the northern and southern arid areas of Africa. Mitt. Bot. Staatssamml. Miinchen 10: 424-^137. Some Taxonomic Changes in Syringa L. (Oleaceae), Including a Revision of Series Pubescentes P. S. Green Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, United Kingdom M. C. Chang Shanghai Museum of Natural History, 260 Yanan Road E., Shanghai, People’s Republic of China Abstract. A number of species of Syringa are reviewed, and series Pubescentes is revised. At in¬ fraspecific rank, the following new combinations in Syringa from Asia are proposed: S. komarowii subsp. reflexa, S. oblata subsp. dilatata, S. reticu¬ lata subsp. amurensis, S. reticulata subsp. pekinen- sis, and S. pubescens subsp. microphylla var. potan- inii. Identification keys are presented. The preparation of an account of Syringa for the forthcoming English version of the Flora of China , combined, for the senior author, with the prepara¬ tion of an account for the European Garden Flora, has led to a review of the genus and a reassessment of the appropriate rank for some of the previously recognized species or varieties. The following are the taxonomic and nomenclatural consequences. A. Syringa komarowii Syringa komarowii Schneider subsp. reflexa (Schneider) P. S. Green & M. C. Chang, stat. nov. Basionym: Syringa reflexa Schneider, Re- pert. Spec. Nov. Regni Veg. 9: 80. 1910. SYN- TYPES: “C.-China. Hupeh: 8-9000'”, Henry 6819 (K); “W. Hupeh,” Wilson 2078 (K). Although McKelvey (1928: 77) stated, “The re¬ lationship of Syringa komarowi to S. reflexa Schnei¬ der is exceedingly close, and it is possible that at some future time S. komarowi may be classified as an extreme form of the Hupeh [Hubei] plant,” it was not until 1990 that the two were formally unit¬ ed at the varietal rank (Chang & Chen, 1990: 35). However, while bearing the morphological differ¬ ences in mind, because S. komarowii and S. reflexa occupy distinct geographical areas, the former in southern Gansu, southern Shaanxi, Sichuan, and northern Yunnan, the latter in western Hubei and northeastern Sichuan, we believe that subspecies is the appropriate infraspecific catagory. They may be distinguished as follows: la. Corolla lobes somewhat erect; inflorescence usu¬ ally ± compact .subsp. komarowii lb. Corolla lobes spreading; inflorescence somewhat pyramidal, often interrupted.subsp. reflexa B. Syringa oblata Syringa oblata Lindley subsp. dilatata (Nakai) P. S. Green & M. C. Chang, stat. nov. Basionym: Syringa dilatata Nakai, Bot. Mag. (Tokyo) 32: 128. 1918. SYNTYPES: Korea. “Pon-san,” Kamibayashi s.n., “Pan-syu-non,” Uchiyama s.n., “in summo montis Nonensan,” Nakai 2209, “Kosui,” Nakai 2602, “Su-hun,” Nakai 2610, and “Chokoku,” Kin-o-sho 268 (synty- pes, TI not seen). As well as exhibiting morphological distinctness, Syringa dilatata and S. oblata have different dis¬ tributions, the former in Korea and northeastern China, and the latter in northern China. The rank of subspecies seems therefore to be the more ap¬ propriate one. They may be distinguished as fol¬ lows: la. Leaves usually slightly broader than long, 2.5-7 X 3—8 cm, base truncate to usually slightly cor¬ date; corolla tube 6-11 mm long, corolla lobes 4—6 mm long.subsp. oblata lb. Leaves usually slightly narrower than long, 3—7 X 2.5-6 cm, base truncate; corolla tube 11—14 mm long, corolla lobes 5—8 mm long. .subsp. dilatata C. Syringa reticulata Syringa reticulata (Blume) Hara subsp. amuren¬ sis (Ruprecht) P. S. Green & M. C. Chang, stat. nov. Basionym: Syringa amurensis Ruprecht, Bull. Cl. Phys.-Math. Imp. Acad. Sci. St. P6- tersb. 15: 371. 1857. TYPE: China/Russia. Ussuri River area, Maack s.n. (syntypes, ?LE not seen). Novon 5: 329-333. 1995. 330 Novon Syringa reticulata (Blume) Hara subsp. pekinen- sis (Ruprecht) P. S. Green & M. C. Chang, stat. nov. Basionym: Syringa pekinensis Ruprecht, Bull. Cl. Phys.-Math. Imp. Acad. Sci. St. Pe- tersb. 15: 371. 1857. TYPE: China. Hebei: ?Kirilov s.n. (holotype, ?LE not seen). Once again, and for similar reasons, it is be¬ lieved that subspecific rank is the appropriate one for these taxa. Syringa reticulata subsp. reticulata is native to Japan, subspecies amurensis to north- easternmost China, adjacent Russia, and Korea, and subspecies pekinensis to northern China and Mongolia. Although in most of the relevant litera¬ ture, the last has been treated at specific rank, it is noteworthy that Maximowicz (1859: 194), only two years after the plant was first described as a species, reduced it to varietal rank. Furthermore, Hemsley (in Forbes & Hemsley, 1889: 82) sank S. pekinensis under S. amurensis and commented, “The differences between the Mandshurian, Japa¬ nese, and Chinese specimens are slight.” They may be keyed out as follows: la. Leaves mostly longer than 7 cm, hairy, especially on the midrib and main veins on the under sur¬ face; capsules blunt.subsp. reticulata lb. Leaves mostly less than 7 cm long, glabrous above. 2a. Petioles stoutish, 1—2 cm long; veinlets slightly sunk; capsule apex blunt. .subsp. amurensis 2b. Petioles slender, 1.5-3 cm long; veinlets not sunk on the upper surface of the leaf; cap¬ sule apex acute .subsp. pekinensis D. A Revision of Series Pubescentes Chang and Chen (1990; see also Chang & Qui, 1992: 63-71) have proposed changes in rank for some of the taxa in series Pubescentes (C. K. Schneider) Lingelsheim, which, following the clas¬ sic monograph of McKelvey (1928), have previously been treated as species by botanists and gardeners alike. The taxa in this section are closely related, and, for some, classification at the rank of species does not seem justified. However, examination of numerous collections of these plants has led to cer¬ tain modifications to the classification proposed by Chang and Chen. These are set out below, following an identification key to the revised series. The nu¬ merous synonyms cited by McKelvey (1928) are not repeated here. Key to Syringa Series Pubescentes la. Leaves with lateral veins ± pinnate, the 2 lowest pairs of veins not closely adjacent. 2a. Leaves usually manifestly longer than broad, sometimes almost as broad as long; corolla lobes 2—4 mm long. 3a. Upper (adaxial) surface of leaves glabrous or glabrescent, except sometimes for a slight pubescence on midrib and main veins on the under surface, rarely pilose. 4a. Leaves usually 1.5-3.5(—7) cm long, I 2.51—1) cm broad.1. S. pubescens 5a. Young stems and inflorescence axes ± 4-angled; corolla tube 10-15 mm long; north-central China .la, subsp. pubescens 5b. Young stems and inflorescence axes — terete; corolla tube 8—10 mm long; northwestern and west-central China.Id. subsp. microphylla var. microphylla 4b. Leaves 5—11 (rarely less) cm long, 2.5-6 (rarely less) cm broad .lb. subsp. patula 3b. Upper surface of leaves finely, often scattered, pilose. 6a. Leaves (2.5—)3—5(—7) cm long, (1.5—)2—2.5(—3.5) cm broad, densely pilose to villous below. 7a. Young stems and inflorescence axes puberulent to densely pilose. 8a. Corolla tube 7—8 mm long; petioles 2—12 mm long .lc. subsp. julianae 8b. Corolla tube 8—10 mm long; petioles 2—5 mm long . . Id. subsp. microphylla var. polaninii 7b. Young stems and inflorescence axes ± densely villous.2. S. mairei 6b. Leaves 1.5-3 cm long, 0.6—1.8 cm broad, glabrous except for a pilose midrib and primary veins below.3. 5. pinetorum 2b. Leaves as broad as long, 1.2—2.2 cm broad, or slightly broader than long, almost circular; corolla lobes 4-5 mm long.4 5. wa rdii lb. Leaves with 2 pairs of veins ± palmately arranged at the lamina base, blade usually 1.5—3(—4) cm long, 1— 2.7(—3) cm broad.5 5. meyeri 1. Syringa pubescens Turczaninow, Bull. Soc. Nat. Moscou 13: 73. 1840. TYPE: China. He¬ bei: 1831, Kirilov s.n.? (holotype, LE, photo¬ graph, K). Erect or spreading shrubs to 5 m tall, young stems glabrous or pubescent. Leaves narrowly to broadly ovate, elliptic or broadly elliptic, 1.5-9(— 11) cm long, l-5(-6) cm broad, glabrous to densely villous, 3 primary veins not closely adjacent at the lamina base. Inflorescence dense or open, glabrous to pilose, 3—15 cm long. Corolla tube 4—15 mm Volume 5, Number 4 1995 Green & Chang Syringa 331 long, lobes 2—4 mm long. Capsule slender or stou- tish, 0.7—1.7 cm long, smooth or with a few lenti- cels. la. Syringa pubescens subsp. pubescens Erect shrubs to 5 m tall, young stems glabrous. Leaves ovate to usually broadly ovate, sometimes elliptic, (1.5-)2.5-3.5(-7) cm long, (l-)1.7-2.5(-4) cm broad, apex acute to obtuse, very slightly acu¬ minate, glabrous above, glabrous to pilose below, especially on the midrib and primary veins toward their bases. Inflorescence ± dense, 4—8(-10) cm long, axes glabrous to pilose. Corolla tube 10-15 mm long, lobes 2-4 mm long, purplish lilac, paler within. Capsule stoutish, 9-10 mm long, smooth with a few lenticels. This subspecies is recorded (Chang in Chang & Qui, 1992) from the following provinces of China, all in the northern part of the country: Hebei, Hen¬ an, eastern Shaanxi, western Shandong, and Shanxi. lb. Syringa pubescens subsp. patula (Palibin) M. C. Chang & X. L. Chen, Invest. Stud. Nat. 10: 34. 1990. Ligustrum patulum Palibin, Acta Hurt. Petrop. 18: 146 (Conspec. FI. Korea 2: 10), 1900. Syringa patula (Palibin) Nakai, Bot. Mag. (Tokyo) 40: 148. 1926. TYPE: Korea, Sontag s.n. (holotype, ?LE not seen). Syringa velutina Komarov, Acta Hort. Petrop. 18: 428. 1901. TYPE: Korea, Komarov s.n. (syntype, ?LE not seen). Syringa palibiniana Nakai, Bot. Mag. (Tokyo) 27: 32. 1913. TYPE: Korea, Faurie s.n. (holotype, TI not seen). Erect shrubs to 3 m tall, young stems slightly pubescent to glabrous. Leaves ovate or broadly ovate to usually elliptic or broadly elliptic, (3—)5— 9(—11) cm long, (2-)2.5—5(-6) cm broad, apex slightly acuminate, glabrous above, rarely scat¬ tered short pilose, pilose to usually glabrous below except for the midrib and primary veins toward their bases. Inflorescence ± dense, 5-9(—15) cm long, pilose. Corolla tube 7-8(-10) mm long, lobes 2(—3) mm long, vinaceous lilac, white within. Cap¬ sules slender, curved, 1—1.5 cm long, with a few lenticels. Recorded from northeastemmost China (Jilin and Liaoning) and Korea. This is a hardy plant that has proved its worth as a garden plant under rigorous climatic conditions. In the West it is per¬ haps best known in cultivation in the form of cv. ‘Miss Kim.’ lc. Syringa pubescens subsp. julianae (C. K. Schneider) M. C. Chang & X. L. Chen, Invest. Stud. Nat. 10: 34. 1990. Syringa julianae C. K. Schneider, Ill. Handb. Laubholzk. 2: 777. 1912 & Bull. Misc. Inform. Kew 1912: 37. 1912. TYPE: cultivated, seed from China, Hubei, Wilson 1220A (holotype, ?W not seen; isotype, K). Spreading shrubs to 2 m tall, young stems pu- berulent. Leaves narrowly ovate to elliptic, (2.5—)4— 5(-7) cm long, (1.5—)2.3—2.5(-3) cm broad, apex acute, slightly long-acuminate, scattered pilose above, pilose below, densely so on midrib and main veins; petioles 2-12 mm long. Inflorescence ± open, (3-)4—6(—10) cm long, densely pilose. Corolla tube 7—8 mm long, lobes 2-3 mm long, violet-pur¬ ple, paler within. Capsules slender, 0.7—1 cm long, smooth. This subspecies has been recorded only from the province of Hubei. l d. Syringa pubescens subsp. niicrophylla (Diels) M. C. Chang & X. L. Chen, Invest. Stud. Nat. 10: 34. 1990. Syringa microphylla Diels, Bot. Jahrb. Syst. 29: 531. 1900. SYN- TYPES: China. Shaanxi: Giraldi 1644 & 1645 (B destroyed). i. Syringa pubescens [subsp. niicrophylla] var. microphylla Spreading shrubs to 2 m tall, young stems finely puberulent to rarely glabrous. Leaves ovate to el¬ liptic. narrowly to broadly so, apex obtuse to acute, very slightly acuminate, glabrous above, rarely scattered pilose, glabrous below, except the midrib and primary veins toward their bases, rarely pilose. Inflorescence ± dense, 4—10f—12) cm long, pilose. Corolla tube 8-10 mm long, lobes 2-3 mm long, pinkish lilac, slightly paler within. Capsules ± slender, 1.2-1.5 cm long, smooth with a few lenti¬ cels. Recorded (see Chang in Chang & Qui, 1992: 67) from the Chinese provinces of Gansu, Hebei, Hen¬ an, Hubei, Ningxia, Qinghai, Shaanxi, Shanxi, and Sichuan. Variety flavoanthera (X. L. Chen) M. C. Chang has been recognized, but the taxonomic val¬ ue of anther color is uncertain, and the recording of it can be suspect, seeing that the pollen of pur¬ plish anthers can make them appear yellow after dehiscence. 332 Novon ii. Syringa pubescens [subsp. microphylla] var. potaninii (C. K. Schneider) P. S. Green & M. C. Chang, comb, et stat. nov. Basionym: Syrin¬ ga potaninii C. K. Schneider, Repert. Spec. Nov. Regni Veg. 9: 80. 1910. TYPE: “Nord- China, O-kansu, am Flusse Tschi lo ku im Ge- birge,” 1885, Potanin s.n. (holotype, ?LE not seen; isotype, K). More or less erect shrubs to 4 m tall, young stems finely puberulent. Leaves ovate to elliptic, (2.5—)3—5(—6) cm long, (1.5—)2—2.5(—3.5) cm broad, apex acute, somewhat acuminate, puberu¬ lent to glabrous above, rarely scattered pilose, scattered to densely pilose below, especially on the midrib and main veins toward the base; peti¬ oles 2-5 mm long. Inflorescence ± dense, 6-10 cm long, finely puberulent. Corolla tube 8-10 mm long, lobes 2—3.5 mm long, pinkish lilac, paler within. Capsules slender, 1.5—1.7 cm long, smooth with a few lenticels. Known only from the Chinese province of Gansu. 2. Syringa mairei (H. Lcveille) Rehder, J. Arnold Arbor. 15: 302. 1934. Ligustrum mairei H. L6veill6, Cat. PI. Yun-Nan 181. 1916. TYPE: China. Yunnan: Maire s.n. (hclotype, E, pho¬ tograph, K). Syringa rugulosa McKelvey, J. Arnold Arbor. 6: 152. 1925. TYPE: China. Yunnan: Maire “169/1914” (ho¬ lotype, E, photograph, K). Shrubs to 4 m tall, young stems densely villous. Leaves ovate to elliptic, somewhat broadly so, (2—) 4-^L5(-6) cm long, (1.5—)2—2.5(—3) cm broad, apex acuminate, pilose above, densely pilose below. In¬ florescence dense, 7—12 cm long, villous. Corolla tube 7—9 mm long, lobes 2—3 mm long, “rose-vio¬ let.” Capsule stoutish, 1 cm long, smooth with a few lenticels. This is a little known plant from Yunnan. The two names, Syringa mairei and S. rugulosa, were almost certainly based on duplicates of the same collection made by E. E. Maire. 3. Syringa pinetorum W. W. Smith, Notes Roy. Bot. Card. Edinburgh 9: 132. 1916. TYPE: China. Yunnan: Forrest 12472 (holotype, E, photograph, K). Slender shrubs to 2.5 m tall, young stems pi¬ lose. Leaves narrowly ovate to elliptic, 1.5-3 cm long, 0.6-1.8 cm broad, apex acute, acuminate, scattered pilose above, glabrous below except for the midrib and primary veins toward their bases. Inflorescence ± dense, 8-12 cm long, pilose. Co¬ rolla tube 9—10 mm long, lobes 3 mm long, “pale lavender-rose.” Capsule with inconspicuous len¬ ticels. A little known plant from Tibet and the province of Yunnan. Plants that have been cultivated under this name have been misidentified. They have fre¬ quently been found to be S. yunnanensis Franchet. 4. Syringa wardii W. W. Smith, Notes Roy. Bot. Card. Edinburgh 9: 132. 1916. TYPE: China. Yunnan: Kingdon Ward 312 (holotype, E, pho¬ tograph, K). Shrubs to 5 m tall, young stems pilose. Leaves ± circular, about 1.2-2.2 cm long and broad, apex rounded, extremely shortly acuminate, glabrous above and below, 3 primary veins on each side, not adjacent at the base of the lamina. Inflorescence somewhat dense, 6-9 cm long, pubescent. Corolla tube 13—14 mm long, lobes 4—5 mm long, pale pinkish lilac? Capsule with inconspicuous lenti¬ cels. This species is only known from the type collec¬ tion and was previously treated as a synonym of Syringa pinetorum by Chang (in Chang & Qui, 1992: 71). However, the small, more or less circular leaves and long corolla tube mark it out as distinct. 5. Syringa meyeri C. K. Schneider in Sargent, PI. Wilson. 1: 301. 1912. TYPE: cultivated at the Arnold Arboretum, origin from cultivation in China, Hebei, Meyer, under Dept, of Agri¬ culture number 23032 (holotype. A). Compact shrub to 1.5 m tall, young stems finely puberulent. Leaves ovate to very broadly ovate-el¬ liptic, (1.5-)2-3(-4) cm long, (l-)1.8-2.7(-3) cm broad, apex obtuse, rarely somewhat acute, very slightly and shortly acuminate, glabrous above, gla¬ brous below except the midrib and the primary veins toward their bases, 2 (or 3) primary veins on each side, subpalmate, arising within 2 mm of the base of the lamina. Inflorescence dense, 5—11 cm long, puberulent. Corolla tube 9-12 mm long, lobes 2-3 mm long, purplish lilac, paler within. Capsules slender, 1-1.2 cm long, lenticular. This compact, relatively dwarf species was first described from cultivated material. It is widely grown in northern China, and in the West under the name ‘Palibin’ (see Green, 1979). It was known only as a cultivated plant until discovered in the wild by Chang as recently as 1989 in the province of Liaoning. This plant has been de¬ scribed as: Volume 5, Number 4 1995 Green & Chang Syringa 333 Syringa meyeri var. spontanea M. C. Chang, Invest. Stud. Nat. 10: 33. 1990. TYPE: China. Liaon¬ ing: M. C. Chang & X. K. Qin 2872 (holotype, SHM; isotype, K). The variety is distinguished by its looser inflo¬ rescence, somewhat shorter corolla tube, and small¬ er leaf. Literature Cited Chang, M. C. & X. L. Chen. 1990. Studies on Chinese Syringa (1). Invest. Stud. Nat. 10: 32—U). [Chinese with English summary.] -& L. Q. Qui (Editors). 1992. Oleaceae. FI. Rei- publ. Popularis Sin. 61: 2—222. Forbes, F. B. & W. B. Hemsley. 1889. Index florae si¬ nensis 2. J. Linn. Soc., Bot. 26: 1—561. Green, P. S. 1979. Syringa meyeri Palibin. Bot. Mag. 182: t. NS. 778. McKelvey, S. D. 1928. The Lilac, a Monograph. London. Maximowicz, C. J. 1859. Primitiae florae amurensis. Mdm. Acad. Imp. Sci. St. Petersburg Divers Savants 9. Homalia pennatula (Musci: Neckeraceae), a New Combination from Southeast Asia, with a Key to the Species of Homalia Si He Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. Johannes Enroth Botanical Museum, P.0. Box 47, FIN-00014 University of Helsinki, Finland Abstract. Symphyodon pennatulus (Dixon) Dix¬ on from Southeast Asia is transferred to the genus Homalia of the Neckeraceae (Musci). A key to the five species and one variety of Homalia is provided. Homalia (Bridel) Schimper is primarily distrib¬ uted in the northern temperate regions and nearby subtropical borders, where it grows on boulders, rock outcrops, or bases of trees. It is characterized by strongly complanate plants with leaves arranged in distinct ranks; by oblong-ovate to spatulate leaves with broad apices and smooth leaf cells; and by smooth capsules with a “hypnoid” peristome. Symphyodon Montagne has 14 species worldwide and is primarily distributed in the tropical and sub¬ tropical regions of Southeast Asia, with one species in Hawaii, South Africa, and Central and South America, respectively. In Southeast Asia, it grows exclusively on tree trunks and tree branches. Be¬ sides its habitat distinction, taxonomic characters of generic importance for Symphyodon are found in the capsule ornamentation and peristomes. It has echinate capsules and an “isobryoid” peristome. Other secondary gametophyte features include: leaves often appressed and imbricate or spreading; leaf apices narrowed; and leaf cells prorate. Symphyodon pennatulus (Dixon) Dixon was ex¬ cluded from the genus Symphyodon, because it is epilithic and has strongly complanate to sub-disti¬ chous leaves (He & Snider, 1992). The type spec¬ imen of Symphyodon pennatulus is meager and con¬ tains no sporophytes. Since no other specimens of this species were available. He & Snider (1992) were uncertain of its generic placement. Recently, numerous collections of this moss have come from Thailand, and though they consistently lack spo¬ rophytes, we now have enough material to make detailed observations on the gametophytes. Sym¬ phyodon pennatulus exhibits many aspects of Hom¬ alia, such as a strongly complanate habit, a hom- alioid-type leaf insertion showing an eight-ranked phyllotaxy (see He, 1992), oblong-ovate to spatulate Novon 5: 334-335. 1995. leaves with broad and rounded apices, and a vari¬ able costal morphology. It is similar to the widely distributed Homalia trichomanoides (Bridel) Schimper in overall similarities of leaf shape and the absence of pseudoparaphyllia, but it differs in having distinct double costae and linear, projecting median leaf cells. On the basis of the double leaf costae and elongate median leaf cells, Symphyodon pennatulus appears most closely related to Homalia glabella (Hedwig) Schimper from Central and South America, except that it lacks pseudoparaphyllia. Although the narrowly linear and projecting median leaf cells of Symphyodon pennatulus and its pres¬ ence in southern Asia differ from those of previ¬ ously recognized Homalia species, the overall ga¬ metophyte features of this moss support its placement in Homalia. Hence, we are making the following new combination. A key to the five spe¬ cies and one variety of Homalia now recognized is provided. A description and an illustration of the species treated here will be published in the forth¬ coming revision of Homalia by the first author. Homalia pennatula (Dixon) He & Enroth, comb, nov. Basionym: Stereodon pennatulus Mitten ex Dixon, Rec. Bot. Surv. India 6: 66. 1914. Sym¬ phyodon pennatulus (Dixon) Dixon in Broth- erus, Nat. Pflanzenfam., ed. 2, 11: 267. 1925. TYPE: Myanmar (Burma). Moulmein, Parish s.n. (holotype, NY; isotype, BM). In his book entitled Burma, Its People and Pro¬ ductions, Mason (1883) published a list of plants, including mosses. Homalia erosa Hampe and Hom¬ alia pennatula Mitten were listed in this publica¬ tion. Both names are apparently nomina nuda, since no descriptions or basionyms were given. The former is clearly a herbarium name, because we have seen several herbarium specimens annotated with the name Homalia erosa Hampe. However, we have seen no specimens bearing the name Homalia pennatula Mitten, nor has it ever been published Volume 5, Number 4 1995 He & Enroth Homalia pennatula from Southeast Asia 335 validly. Although it could be based on Mitten’s her¬ barium name Stereodon pennatulus, Homalia pen¬ natula Mitten has no nomenclatural bearing in our paper. In recognizing that Homalia trichomanoides var. japonica (Besch.) He, stat. nov., cited in He (1992) has not been validly published, we make the fol¬ lowing new combination: Homalia trichomanoides var. japonica (Bes- cherelle) He, comb. nov. Basionym: Homalia japonica Bescherelle, J. Bot. (Morot) 13: 39. 1899. TYPE: Japan. Sengantoge, Faurie 14984 (holotype, PC; isotype, KYO). Additional selected specimens examined. MYANMAR. Moulmein, Kurz 2930 (BM). INDIA. Kerala, Norhett 13377 (BM). THAILAND. Changburi: Larsen et al. 1857 (AAU, MO). Chiangmai: Mt. Suthep, 98°55'E, 18°50'N, Touw 8395 (BM, L, MO, NY). Krabi: Khao Phanom Ben- cha Park, Schafer-Verwimp & Verwimp 16230 (H, MO). Nakhon Nayok: Larsen et al. 3380 (AAU, MO). Nakhon Sawan: 98°55'E, 16°40'N, Touw 8319 (BM, L, MO, NY). Prachinburi: 101°30'E, 14°30'N, Touw 12215 (BM, L, MO). Trat: 102°49'E, 11°59'N, Charoenphol et al. 4982 (AAU, BM, H, MO, NY); Ko Kut, 101°31'E, 12°35'N, Charoenphol et al. 5129 (AAU, BM, H, MO, NY). Key to the Species and Varieties of Homalia 1. Costa single, extending Vi—Y 5 the leaf length ... 2 1. Costa double or forked, less than Vs the leaf length or indistinct. 4 2(1). Pseudoparaphyllia present; costa stout, extend¬ ing 3 A~y s the leaf length; leaf apices dentate, with large teeth; western Europe, northern Af¬ rica, Canary Islands ... 1. H. lusitanica Schimper 2. Pseudoparaphyllia absent; costa slender, ex¬ tending Vi-V a the leaf length; leaf apices mod¬ erately serrate, with small teeth. 3 3(2). Autoicous; secondary stems often blunt near the apex; flagelliform branches uncommon; eastern Asia, Russia, Europe, North America, Mexico. .2a. H. trichomanoides var. trichomanoides (Hedwig) Schimper 3. Dioicous; secondary stems often attenuate near the apex; flagelliform branches commonly pres¬ ent; Japan, Korea, Taiwan . .2b. H. trichomanoides var. japonica (Bescherelle) He 4(1). Median leaf cells rhombic, 8.5-17.5 pm long; Azores, Maderia, Canary Islands, Caucasus . . .3. H. webbiana (Montagne) Schimper 4. Median leaf cells oblong-rhomboidal to narrow¬ ly linear, 35-100 pm long. 5 5(4). Leaves 1.25-3.25 mm X 0.75-1.25 mm; pseu¬ doparaphyllia present; Mexico, West Indies, Central America, Brazil, Venezuela . .4. H. glabella (Hedwig) Schimper 5. Leaves 0.8-1.2 mm X 0.5-0.65 mm; pseudo¬ paraphyllia absent; India, Myanmar, Thailand .5. H. pennatula (Dixon) He & Enroth Acknowledgments. We thank Bruce Allen (MO) and Marshall R. Crosby (MO) for constructive com¬ ments on the manuscript. Literature Cited He, Si. 1992. A worldwide taxonomic revision of the gen¬ era Homalia, Pendulothecium (Musci: Neckeraceae) and Symphyodon (Musci: Symphyodontaceae). Doctoral Dissertation, University Microfilms, Ann Arbor. -& J. A. Snider. 1992. A preliminary survey of the moss genus Symphyodon (Symphyodontaceae, Mus¬ ci). Bryobrothera 1: 283—287. Mason, F. 1883. Burma, Its People and Productions. Vol. II (Botany). Stephen Austin & Sons, Hertford. Polygonum hickmanii (Polygonaceae), a New Species from California Harold Hinds Biology Department, University of New Brunswick, Fredericton, New Brunswick, E3B 6E1 Canada Randall Morgan 3500 N. Main Street, Soquel, California 95073, U.S.A. Abstract. A new endemic species of Polygonum is described from Scotts Valley, California. Differ¬ entiation from closely related taxa P. parryi and P. heterosepnlum is presented. Polygonum (sect. Duravia) hickmanii H. R. Hinds & R. Morgan, sp. nov. TYPE: U.S.A. California: Santa Cruz County, Scotts Valley, grassland N of Navarra Drive, W of Carbonero Creek, 1 July 1993, R. Morgan 2119 (holo- type, CAS; isotypes, MO, RSA, UC, UNB). Hickman’s knotweed. Herba annua. Folia sessilia, 0.5-3.5 cm X 1—1.5 mm, revoluta. Ocrea divisa ad basem in longas rectas libras. Flores singuli in axillis omnium paene bracteorum folior- um. Achenia laevia, (2—)2.3 X 1.6 mm, duobus lateribus concavis et uno latere latiore planioreque. Erect taprooted annual, 2-5 cm, simple in de¬ pauperate plants to profusely branched from near the base. Leaves sessile, linear, acuminate, mucro- nate, 0.5—3.5 cm X 1—1.5 mm, revolute and wing- nerved below, closely investing and with the ocreae ± concealing the stems. Ocrea 6 mm long, silvery, divided nearly to base into long straight filaments, adnate to the base of the leaves and appearing al¬ most as long as the apical leaves. Flowers single in axils of almost all bracteal leaves, tepals white, with two outer tepals cucullate and mucronate, 3 mm long, three inner tepals rounded to obtuse 2 mm long, anthers conspicuous, orange-pink 0.4 mm. Achenes smooth, (2-)2.3 X 1.6 mm, light to dark olive brown, lighter on the angles, with two concave sides and one broader ± flat side; not exserted. Styles 0.3 mm, persistent. Flowering from late May to August (October). Polygonum hickmanii is known from two sites about one mile apart, at the northern end of Scotts Valley, Santa Cruz County, California, at about 700—800 feet elevation and occurs only in very re¬ stricted microhabitats within an isolated relictual grassland. The sites are gently sloping to nearly level, with fine-textured shallow soil over outcrops of Santa Cruz mudstone and Purisima sandstone, sparsely vegetated with small annual herbs. This habitat is shared with a more abundant but equally narrow endemic, Chorizanthe robusta C. Parry var. hartwegii (Bentham) J. Reveal & R. Morgan. The population of Polygonum hickmanii fluctuated from about 200 to 400 plants in 1990-1992 to 1000 or more in 1993 following a relatively wet winter. Polygonum hickmanii is sympatric with no other members of the section Duravia. The nearest lo¬ cation of a closely related species, P. parryi Greene (itself disjunct), is at Mt. Hamilton about 50 miles inland. Polygonum hickmanii differs from P. parryi in its larger white flowers, longer leaves, larger anthers and achenes, and longer, straight ocreae. It differs from P. heterosepalum M. E. Peck & Ownbey in achene shape (more rounded at base as opposed to more tapering), size (1.5-2 mm), and flower struc¬ ture (perianth segments very unequal). Polygonum hickmanii is named for James C. Hickman (1941-1993), distinguished botanist, who confirmed the uniqueness of this taxon and whose advice and inspiration is here acknowledged. Acknowledgments. The authors thank George Flanders, Irene Johnston, and John Geyssen for their considerable assistance with manuscript prep¬ aration. Novon 5: 336. 1995. Una Nueva Especie de Marsdenia (Asclepiadaceae) de Mexico Veronica Judrez-Jaimes Herbario National, Instituto de Biologfa, U.N.A.M., Apartado Postal 70-233, Delegation Coyoacan 04510, Mexico D.F., Mexico W. Douglas Stevens Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. Resumen. Marsdenia rzedouiskiana es descrita como nueva. Esta especie ha sido confundida con M. macrophylla (Humboldt & Bonpland ex Schultes) E. Fournier pero esta mds relacionada con M. prin- glei S. Watson. La distribution de M. rzedowskiana esta restringida al noreste de Puebla y partes ad- yacentes de Veracruz, hacia el sur de la distribu- ci6n conocida de M. pringlei y en altitudes super- iores a las de M. macrophylla. Abstract. Marsdenia rzedowskiana is described. This new species has been confused with M. ma¬ crophylla, but is most closely related to M. pringlei. Marsdenia rzedowskiana seems to be restricted to northeastern Puebla and adjacent Veracruz, to the south of the known distribution of M. pringlei and at higher elevations than M. macrophylla. El genero Marsdenia R. Brown se encuentra bien representado en Mexico por 24—26 especies que se distribuyen en todo el pais. Durante la revision para el tratamiento del genero para Mexico y Cen- troamerica. se encontro material que inicialmente se incluyo en la section Ruehssia, subsection Ma- crophyllae (Rothe, 1915); sin embargo, pertenecia mas bien a la section Rostrata, subsection Edules, y adernas correspondfa a una especie no descrita. Marsdenia rzedowskiana Judrez-Jaimes & W. D. Stevens, sp. nov. TIPO: Mexico. Puebla: Mun- icipio Teziutlan, Agua de Obispo, 30 mar. 1973 (fl), F. Ventura 8102 (holdtipo, MEXU; isotipos, ENCB, MO). Figura 1. Marsdeniae pringlei S. Watson similis sed floribus mi- noribus, apice styli perbreviore et integro recedit. Plantas trepadoras, con ldtex bianco. Tallos gla- bros, gruesos, 5—7 mm de ancho. Pecfolos 1—3 cm, glabrescentes; ldminas 5.0-15.5 X 2.6-10.5 cm, elfpticas, con 7—8 nervios secundarios, base redon- deada a obtusa, apice acuminado a cortamente ac- uminado, coridceas, glabras. Inflorescencias 3—6 X 2-4 cm, con 10-22 flores, extra-axilares, laxamente ramificadas; brdcteas elfpticas y acuminadas, 2 mm de largo, pubescentes; pedunculos 1.0-2.5 cm de largo, ligeramente pubescentes; pedicelos 2—4 mm de largo, pubescentes. Ldbulos del cdliz 2.6-3.0 X 2.2-2.8 mm, orbiculares, dpice algo agudo, ligera¬ mente pubescentes por fuera, glabros por dentro, mdrgenes ciliados, los senos con 1—3 gldndulas por dentro; corola campanulada, 5.0-6.5 X 4.8-6.6 mm, glabra por fuera, ldbulos 3.0-3.2 X 1.8-2.3 mm, oblongos, erectos en la antesis, la superficie interna pilosa en el centra, apices ligeramente emarginados, tubo 2.5-3.0 mm de largo, piloso en la garganta, los pelos largos y blancos; ldbulos de la corona 1.5—1.7 X 1.0-1.5 mm, camosos, redon- damente deltoides, con los mdrgenes laterales sal- ientes, la punta libre y terminando al mismo nivel de la parte apical de las anteras, ginostegio 2—3 mm de largo, membranas de las anteras 0.9-1.2 mm, ampliamente ovadas, anteras 0.4—0.5 mm de largo, elipsoides; corptisculos 0.3-0.4 mm de largo, elfpticos, polinios 0.3-0.4 mm de largo, casi glo- bosos, apice del estilo 2.0-2.5 mm de largo, cdni- co-alargado, con una pequena Ifnea de bifurcacion en la parte apical; carpelos glabros. Folfculos des- conocidos. Marsdenia rzedowskiana estd muy relacionada con M. pringlei S. Watson, puesto que ambas tienen hojas ampliamente elfpticas, cimas ramificadas, flores amarillas, garganta de la corola con pelos blancos, ldbulos de la corona deltoides, dpiee del estilo alargado y polinios casi globosos. Sin embar¬ go, M. rzedowskiana se diferencia por poseer flores mds pequenas, ldbulos de la corola mds o menos del mismo tamano que el tubo, tubo de la corola no urceolado, apice del estilo 3 veces mds corto y presentando apenas una Ifnea de bifurcacidn en la parte apical. Por otra parte, M. rzedowskiana solo es conocida de la Sierra Norte de Puebla y la regidn adyacente a Jalapa, Veracruz, en lo que Rzedowski (1978) considera bosques mesofilos de montana, mientras que M. pringlei se distribuye en un drea mds extensa de la Sierra Madre Oriental. Novon 5: 337-339. 1995. 338 Novon El material de Marsdenia rzedmvskiana tradi- cionalmente ha sido determinado como M. macro- phylla principalmente por su aspecto general, pero la presencia en M. rzedowskiana de una inAores- cencia muy ramificada de Acres amarillas, lobules de la corona deltoides y dpice del estilo cdnico- alargado la separa definitivamente de esta; ademas, M. rzedowskiana crece en bosques superiores a los Volume 5, Number 4 1995 Juarez-Jaimes & Stevens Marsdenia rzedowskiana 339 1300 m, mientras que M. macrophylla es de tierras bajas en Mexico. Marsdenia rzedowskiana Judrez-Jaimes & W. D. Stevens se dedica a Jerzy Rzedowski por sus im- portantes aportes al estudio de la vegetacion y la botariica de Mexico, trabajos por los cuales los au- tores tienen una gran admiracion. Pardtipos. MEXICO. Veracruz: Mpio. Aratlan. El Ce- rro, 3 jun. 1977 (fl), F. Ventura 14030 (XAL); Mpio. Ye- cuatla, along winding road from Naolinco to Misantla, just above Santa Rita and 3 km by road from Paz de Enriquez, 1550 m (fr), M. Nee et al. 26425 (MO). Puebla: Mpio. Hueytamalco, El Popual, 1500 m, F. Ventura 1038 (US). Agradecimientos. Agradecemos a Hector Her¬ nandez, Francisco Lorea, Fernando Chiang y Agus- tln Rudas la revision crftica del manuscrito. Henk van der Werff escrihio la diagnosis en latln; John Myers realizo la ilustracion de la planta. Literatura Citada Rothe, W. 1915. Uber die Gattung Marsdenia R. Br. und die Stammpflanze der Condurangorinde. Bot. Jahrb. Syst. 52: 354-434. Rzedowski, 1. 1978. Vegetaci6n de Mexico. Limusa, Max- New Taxa of Corispermum L. (Chenopodiaceae), with Preliminary Comments on the Taxonomy of the Genus in North America Sergei L. Mosyakin N. G. Kholodny Institute of Botany, 2 Tereshchenkivska Str., Kiev, 252601 Ukraine ABSTRACT. Five new species and three varieties of Corispermum L. (Chenopodiaceae) are described from North America. Corispermum pacificum Mo¬ syakin and C. pallidum Mosyakin are restricted to the Pacific Northwest; C. welshii Mosyakin is known from Utah and adjacent parts of Wyoming; C. navicula Mosyakin occurs in Colorado and prob¬ ably adjacent states; C. hookeri Mosyakin is known from Canada. Corispermum pacificum is closely re¬ lated to the Asian taxa C. crassifolium Turczaninov and C. maynense Ignatov; C. pallidum is possibly related to C. macrocarpum Bunge ex Maximowicz; the three remaining new taxa and all other native North American taxa are allied to the C. pallasii Steven aggregate. The nomenclature of C. pallasii [— C. leptopterum (Ascherson) Iljin = C. sibiricum Iljin subsp. baicalense Iljin = C. bjelorussicum Klo¬ kov & Krasnova], C. villosum Rydberg, and C. americanum (Nuttall) Nuttall is discussed. Three new varieties are described: C. americanum var. rydbergii Mosyakin, C. hookeri var. pseudodeclina- tum Mosyakin, and C. ochotense var. alaskanum Mosyakin. Corispermum ochotense Ignatov, a spe¬ cies recently described from the northeastern part of the Russian Far East, is reported from Alaska and northwestern Canada, where it is represented by two varieties. Corispermum ochotense var. alas¬ kanum differs from the typical variety in having larger fruits. Preliminary results of a study of the genus in North America are discussed. Corispermum has the reputation of being a prob¬ lem for taxonomists and may be regarded as one of the most taxonomically complicated groups within Chenopodiaceae. As an example, the name Coris¬ permum hyssopifolium L., which refers to a species restricted mostly to a small area in southeastern Europe and adjacent parts of western Asia, has been misapplied to almost all other species of the genus. For a long time the representatives of this genus were believed to have been introduced to North America from Eurasia. In the treatment by Maihle and Blackwell (1978) only three species were reported for North America: C. hyssopifolium L., C. nitidum Kitaibel ex Schultes, and C. orientale Lamarck (all Eurasian). However, it is now clear from the fossil evidence that Corispermum was present in North America at least 38,000 years be¬ fore the present (for details see Betancourt et ah, 1984). Native American species of Corispermum are evidently closely related to eastern Asian (es¬ pecially Siberian) ones (see discussion below). I strongly believe that even less than 10,000 years was sufficient for Asian ancestors of North American taxa to evolve into several distinct races (species or subspecies) because the high rate of microevolutionary processes is crucial for a suc¬ cessful life strategy of such annual psammophytes of pioneer and marginal habitats. The important mi¬ croevolutionary patterns within the genus include both allogamy (performed by wind pollination) and autogamy (especially under unfavorable environ¬ mental conditions), high degree of morphological variability (including so-called “cryptic variabil¬ ity”), and high possibility of hybridization (es¬ pecially between closely related species), which is typical for many other chenopods, including Che- nopodium L. However, polyploids, which are fairly common in Chenopodium and many other genera of the family, seem to be absent, or at least extremely rare, in Corispermum: all taxa studied until present were diploids with 2 n = 18 (for additional discus¬ sion, see Adamkiewicz, 1970; Strazdin’sh, 1985; Mosyakin, 1988a). In the course of preparation of a taxonomic treat¬ ment of Corispermum for the Flora of North Amer¬ ica, it has become evident that several new species and varieties should be described. Before describ¬ ing these taxa I will briefly discuss the taxonomy of other Corispermum species from North America. However, I should mention that the taxonomic treatment of the genus in North America is not yet complete, and more efforts, especially study of vari¬ ability in the natural habitats, are needed. At pres¬ ent my experience in North American Corispermum is based exclusively on herbarium studies, plus some experience with Eurasian species, which was especially crucial for this work, and four weeks of fieldwork in the southwestern United States (Utah and Nevada). Because of this, some of the following Novon 5; 340-353. 1995. Volume 5, Number 4 1995 Mosyakin Corispermum from North America 341 considerations (especially concerning the C. pal- lasii Steven— C. americaniirn (Nuttall) Nuttall aggre¬ gate) should be regarded as preliminary, a kind of brief introduction to the taxonomic problems in North American Corispermum. A history of taxo¬ nomic study of the genus in North America, as well as literature citations, may be found in Maihle and Blackwell (1978); thus, some of the dates and ref¬ erences are not mentioned below in order to avoid repetition. Discussion The first taxon described from North America was C. hyssopifolium L. var. americanum Nuttall [= C. americanum (Nuttall) Nuttall], Unfortunately, its authentic description is insufficient, and the lec- totype specimen (Nuttall s.n., PH; labeled “C. hys¬ sopifolium var. pubescens ”) is immature. In my opin¬ ion, this specimen cannot be assigned with certainty to any native or introduced species. Be¬ cause of this, in the early stage of my work on North American Corispermum I regarded the name C. americanum as a nomen dubium. Fortunately, there is another authentic specimen, which evidently was annotated by Nuttall. This specimen is deposited in the Besser memorial collection at KW. It is la¬ beled “Corispermum hyssopifolium * americanum m. Nuttall”; as in other similar cases, an asterisk indicates the taxon described by Nuttall himself. The herbarium sheet contains an immature plant, which is almost identical to the PH specimen, and a branch from another plant, which is more mature. Corispermum americanum was cultivated by Nuttall (see Nuttall, 1818), and probably the more mature specimen represents a fragment collected from a cultivated plant. Because of this find the long-ne¬ glected name C. americanum can be restored. Judging from the description of Corispermum pi- losum Rafinesque (1836) (since no authentic her¬ barium material has yet been traced), this taxon does not belong to Corispermum at all. It was de¬ scribed from Florida, where no species of Corisper- mum are (or have been) known. The name C. pi- losum probably refers to the hispid form of Salsola kali L. aggregate (immature specimens of Corisper¬ mum and Salsola L. are often confused in herbaria). Moreover, the name is a later homonym of C. pi- losum Pallas. In my opinion, Corispermum villosum Rydberg (1897) should be regarded as a distinct native spe¬ cies, which is closely related to the North American c. americanum and Asian-North American C. pal- lasii Steven (see below), and possibly also to C. chinganieum Iljin (more distantly). Intermediate forms between C. americanum and C. villosum oc¬ casionally occur in regions where these species are sympatric. The characteristic features of C. villosum s. str. are small (usually 1.8-3.0 X 1.5-2.0 mm) fruits, almost wingless, or with a narrow wing up to 0.1(-0.15) mm broad; both wingless and narrow¬ winged forms are sympatric, but wingless plants seem to be much more common in the Rockies. In the taxonomy of Corispermum, as well as of many other Chenopodiaceae, the most reliable characters are those of the fruits. The degree of pubescence is an unstable character in most representatives of the genus: it largely depends on the age of the plant (plants often become glabrous at maturity). Because of this I can find no sound justification for main¬ taining C. villosum and C. emarginatum Rydberg (1904) as separate species. The lectotype of the first species (Montana: Manhattan, 1895, Rydberg 2623, US; isolectotype at NY) is younger and, conse¬ quently, more pubescent; the type of C. emargina¬ tum (Wyoming: Laramie, 15 Sep. 1897, Aven Nelson 4282, RM; isotypes at GH, MO, NY) is completely mature and almost glabrous. However, the fruit characters are essentially the same in both species, and, in my opinion, these taxa should be united under the name C. villosum, which has priority. Corispermum imbricatum A. Nelson (1909) and C. simplicissimum Lunell (1910), despite their dif¬ ferences in habit, both have small narrow-winged fruits of similar shape and dimensions. The type of C. imbricatum (Wyoming: Laramie Hills, 29 Aug. 1902, Elias Nelson 733, RM) represents a morpho- type, with stems branched from the base, and in¬ florescences distinctly clavate and dense at the apex. This morphotype is somewhat transitional to¬ ward C. villosum. The authentic specimens of C. simplicissimum (type: North Dakota: Pierce Co., Barton, on a lake shore, 26 Aug. 1890, Lunell 395, MIN) are branched above the base, and their inflo¬ rescences are more lax and narrow: judging from available herbarium specimens, this morphotype seems to be common in the Great Plains region. It differs from typical C. americanum only in branch¬ ing habit. In both C. imbricatum and C. simplicis¬ simum, branches of inflorescences are simple or nearly so. These taxa probably represent extremes of the variability of C. americanum s.l. Corispermum americanum has the largest area of distribution among the native species of Corisper¬ mum, centered mostly in the Rocky Mountains and adjacent portions of the Great Plains, and it is very variable, especially in its vegetative characters. Plants occurring in the Great Plains region usually have winged fruits and often show some degree of 342 Novon morphological transition toward C. pallasii (see be¬ low). Specimens related to the Corispermum pallasii- C. americanum aggregate with unusually large (up to 4.5-5.0 mm long) wingless or narrow-winged fruits are known from the southwestern United States (mostly Texas. Oklahoma, Colorado, New Mexico), and from western central Canada (mostly Alberta and Saskatchewan). The Canadian plants are evidently more closely related to C. pallasii s. str. and C. ochotense Ignatov var. alaskanum Mo- syakin described below, but they are very different in habit: inflorescences are usually short-clavate, dense and thick, with strongly imbricate broad bracts. The large-fruited Canadian plants are prob¬ ably identical to “C. hyssopifolium ” var. robustius Hooker (see Hooker, 1840). However, I have not seen the type of Hooker’s variety. Because of this I am not convinced that it is the same as the large- fruited Canadian plants, and prefer to describe the latter as C. hookeri Mosyakin (see below). The large-fruited plants from the southwestern United States are more closely related to Corisper¬ mum americanum s. str. They differ from northern large-fruited specimens in having fruits more elon¬ gate and obovate (more distinctly broadest above the middle than in the Canadian specimens), often with white warts (formed by loose portions of peri¬ carp) and reddish brown spots (a character that oc¬ curs in several species, including C. pallasii), usu¬ ally with a more prominent triangular “beak” (formed by the style bases in mature fruits) at the apex. Some of these forms remain enigmatic, and more information is needed to determine their tax¬ onomic status. The most striking and recognizable morphotype is described below as C. navicula Mo¬ syakin. Another complicated case is Corispermum mar¬ ginal Rydberg. In the protologue Rydberg (1903) cited one collection from New Mexico and three localities from Colorado. His description matches the characters of some plants occurring in the southwestern United States and northern Mexico. These plants differ from C. americanum s. str. and C. villosum s. str. in having fruits larger, up to 4.0(- 4.5) mm long, and more prominently winged (wing usually ca. 0.2-0.4 mm broad), and usually a more interrupted inflorescence. Thus, Rydberg’s concept of C. marginale as a species distinct from C. vil¬ losum was correct, and it is unfortunate that he cited as the holotype the New Mexican specimen (Albuquerque, 1884, C. L. Herrick s.n., NY) with small narrow-winged fruits. This specimen differs from C. americanum s. str. only in having slightly more lax spikes, which is not sufficient for regard¬ ing it as specifically distinct. As is evident from his comments, Rydberg (1897) believed that C. hysso¬ pifolium s. str. was a species with large fruits (up to 5 mm long); this erroneous concept probably af¬ fected his decision, since he apparently tried to stress the difference between his C. marginale and C. hyssopifolium (sensu Rydberg). The name must stand or fall together with the type, and because of this the epithet “marginale” should be regarded as a synonym of C. americanum s. str. However, I be¬ lieve that other specimens cited by Rydberg (para- types) represent a distinctive variety (or possibly subspecies), and it is validated below as C. amer¬ icanum var. rydbergii Mosyakin. This variety, as well as some other forms of C. americanum, usually has lax spikes, and due to this was constantly mis- identified by North American authors as C. nitid- um. For a better understanding of these taxa, it is necessary to comment briefly on Corispermum ni- tidum Kitaibel ex Schultes (or “so-called C. nitid- um”). I have seen several herbarium specimens of the true C. nitidum collected in North America re¬ cently. Judging from the fruit morphology, almost all American representatives of the “C. nitidum” morphotype belong to the C. pallasii—C. american¬ um aggregate. Because of this I strongly believe that most of the records of this species from North America are referable to C. americanum. The first collections of the “C. nitidum ” morphotype were made as early as 1847 in New Mexico (A. Fendler 702, GH, MO); 1852 in Chihuahua, Mexico (G. Thurber 811, GH); 1873 (71 S. Brandegee 841, MO); 1878 (Marcus A. Jones 655, MO); 1881-1882 in Colorado (“Common along Pueblo,” R. W. Wood¬ ward s.n., GH); 1879 in Texas (E. Palmer 1165, GH); these are only examples based on GH and MO collections. This list may be greatly extended by inclusion of references to specimens from other herbaria. In the earliest collections most of the mentioned localities were on the edge of civiliza¬ tion, and I cannot believe that European C. nitidum had already been widespread in the relative “wil¬ derness” of the North American West. In the mid¬ dle of the nineteenth century this species was poor¬ ly known even in Europe. Its natural area of distribution originally covered only the Pannonian (southeastern Central Europe, mostly along the Danube River and some of its tributaries in Hun¬ gary, Austria, Romania, and Bulgaria) and Pontic (southern Ukraine and southwestern Russia) flo- ristic regions, with a small extension into the Tu¬ ranian area (northwestern Kazakhstan). Besides, C. nitidum does not seem to be a markedly invasive species. It is definitely introduced in some locali- Volume 5, Number 4 1995 Mosyakin Corispermum from North America 343 ties in Europe (e.g., in Poland, Byelorussia, central Russia, etc.), but it cannot be regarded as wide¬ spread in the Old World. At present I know only two cases of successful and comparatively wide ter¬ ritorial expansion of Corispermum species: C. pal- lasii (= C. leptopterum) in central and western Eu¬ rope, and C. declinatum Stephan ex Iljin in eastern Europe and southwestern Siberia. Amazing morphologic parallelism is common in Corispermum. For example, typical specimens of C. heptapotamicum Iljin (central Asia) may be distin¬ guished from C. nitidum by their broader, usually linear-oblanceolate lower leaves, and fruits some¬ what similar to those of C. lehmannianum Bunge; but in many cases herbarium specimens are almost indistinguishable. Such species as C. filifolium C. A. Meyer ex A. Becker, C. aralocaspicum Iljin, and C. declinatum also can be confused with C. nitid¬ um. Fortunately, in all these cases the fruit char¬ acters help to distinguish the taxa. Another exam¬ ple of this striking parallelism between some New World and Old World taxa of Chenopodiaceae is Chenopodium novopokrovskyanum (Aellen) Uotila (see Uotila, 1993), a species of the C. strictum Roth aggregate restricted to arid regions of southeastern Europe and the Irano-Turanian portion of Asia, which is similar in habit (but not closely related) to native North American taxa of the C. leptophyl- lum Nuttall ex S. Watson aggregate. The species that seems to be common in the Great Lakes region is conspecific with Corispermum leptopterum (Ascherson) Iljin, a taxon described from central Europe. However, in Europe this spe¬ cies is evidently introduced; the possible means of its introduction will be discussed in the forthcom¬ ing review of the European species of the genus. Voss (1985) was the first to suggest that the Great Lakes plants may be conspecific with C. leptopter¬ um. Forms identical with Corispermum leptopterum are known from Siberia (especially southern and eastern parts, plus adjacent portions of Mongolia and China), where they are commonly referred to as C. sibiricum subsp. baicalense Iljin (see Straz- din’sh, 1985). At present I cannot find evident dif¬ ferences between most of the Siberian, European, and North American plants of this aggregate, except for usually narrower leaves, more lax and narrow spikes, and slightly smaller fruits in average North American specimens. I believe that this species may also be native in North America, at least in part. Deciding which name has priority for Siberian “Corispermum leptopterum ,” one must consider at least two older names: C. patens Fisher ex Roemer & Schultes and C. pallasii Steven (both described from Siberia; see Steven, 1817). I have not yet traced any mature and unquestionably authentic specimens of the first species, and it still remains a nomen dubium. The type of C. pallasii (Dahuria, Helm s.n., H) belongs to the group that includes plants previously referred to mostly as C. sibiricum s.l. The holotype specimen is not completely ma¬ ture, and because of this I previously hesitated to assign it to a particular race of this group. More¬ over, some presumably authentic specimens of C. pallasii in Besser and Turczaninov memorial col¬ lections at KW belong to C. redowskii Fischer ex Steven, a distinctive Siberian species with pubes¬ cent undulate-winged fruits. However, Steven knew C. redowskii well (see Steven, 1817), and it is im¬ probable that he could confuse it with another spe¬ cies. He mentioned only one collection of his C. pallasii (“Semina in specimine juniore a D. Helm mihi dato, videntur emarginata”), which should be consequently regarded as the holotype. Study of several young fruits of the type of C. pallasii, and their comparison with immature fruits of European C. leptopterum and Siberian C. sibiricum subsp. baicalense, revealed no evident difference between them. Thus, the earliest available name for this spe¬ cies is C. pallasii. Corispermum pallasii may be distinguished from other species occurring in the United States and Canada by its comparatively large (usually 3.0-4.5 X 2.0-3.5 mm), obovate or obovate-elliptic fruits (normally distinctly broadest above the middle); thick wing (translucent only along margins) (0.2-) 0.3-0.4 mm broad, which is usually long-adnate to the style bases, forming a broadly triangular or rounded-triangular apex; style bases often separat¬ ed to below the edge of the wing; and inflorescences usually clavate and rather dense, or occasionally clavate-linear (the last character is very variable). Judging from the morphology of the fruits, this spe¬ cies is closely related to C. americanum. Some European forms of the Corispermum pal¬ lasii aggregate, known mostly as C. membranaceum Iljin or C. leptopterum var. membranaceum (Iljin) Aellen, have smaller and less winged fruits, mor¬ phologically approaching C. americanum. The Asian relatives of C. pallasii are: C. stauntonii Mo- quin-Tandon, C. elongatum Bunge ex Maximowicz s.l. (including C. confertum Bunge ex Maximowicz), C. candelabrum Iljin, C. bardunovii M. Popov ex M. Lomonosova, and some other species. Corisper¬ mum sibiricum Iljin s. str. (subsp. sibiricum = subsp. jenissejense Iljin) is also related to this ag¬ gregate, but more distantly. Corispermum sibiricum subsp. amurense Iljin is conspecific with C. elon- 344 Novon gatum. For additional information about C. pallasii (= C. leptopterum) see Iljin (1929), Aellen (1961, 1964), Strazdin'sh (1985), Kock (1986), and Mo- syakin (1988b). In the Atlas Florae Europaeae (Jalas & Suomi- nen, 1980) and the second edition of Flora Euro- paea (Aellen & Akeroyd, 1993), Corispermum pal¬ lasii (= C. leptopterum) and another distinct species, C. gallicum Iljin, were erroneously sub¬ merged in C. intermedium Schweigger. The latter is a distinctive endemic of the southeastern coast of the Baltic Sea, and its differences from C. pallasii are numerous and evident: constant absence of perianth segments, broadly ovate or almost orbic- ulate fruits with broader wing rounded at the apex, etc. In spite of some similarity between the habits of C. intermedium and the C. canescens Kitaibel ex Schultes-C. marschallii Steven aggregate, their his¬ tory and development were independent, repre¬ senting two hypothetical migration routes in the westward European expansion of the genus: north¬ ern route, affected mostly by glacial factors of the Pleistocene, especially Riss-Wiirm/Mikulino inter¬ glacial; and southern route, affected mostly by the pre-Pleistocene and Pleistocene transgressions and regressions of inland seas. The first migration path may be traced from east to west by the present distribution of C. algidum Iljin (northeastemmost European Russia), C. intermedium (Baltic Sea coasts), and C. gallicum (southern France). Addi¬ tional evidence supporting this geohistorical recon¬ struction may be found in the discovery of fossil fruits of Corispermum in the Late Weichselian de¬ posits in England (ca. 42,000-45,000 years BP) and Belgium (see Godwin, 1975; Bell, 1969). The genus was not known from the British Isles in his¬ torical times until the very recent discovery of in¬ troduced C. pallasii reported by Gibson (1992) as C. leptopterum. The end of the second (southern) migration path of Corispermum in Europe is marked by the westernmost area of natural distribution of C. canescens s. str. and C. nitidum in the Pannonian floristic region. Because of some parallels between arid and nu¬ tritionally deficient habitats (see Small, 1973), the xeromorphic species of Corispermum are well adapted to both hot deserts of central Asia and southwestern North America, and cool river shores of subarctic Siberia and Alaska. Usually they are most abundant in pioneer, marginal, and naturally and/or artificially disturbed habitats. In the Qua¬ ternary the most widescale pre-anthropogeic dis¬ turbance was caused by Pleistocene glaciations (see, e.g., Pielou, 1991); the glacial events affected both distribution and evolution of many species with “weedy” life strategies (see Harlan & de Wet, 1965). An excellent example of a response of “weedy” species to the glacial and post-glacial events of the late Pleistocene and Holocene is Am¬ brosia L. (see Bassett & Terasmae, 1962). I believe that the Pleistocene glaciations and early Holocene post-glacial events played an im¬ portant role in the history of Corispermum. Expan¬ sions and retreats of glaciers, as well as eustatic and isostatic sea level changes, created many pio¬ neer, marginal habitats, especially fluvioglacial, al¬ luvial, littoral deposits in coastal and limnetic ar¬ eas, river valleys, etc. Abundance of such habitats provided excellent migration routes for ancestors of modem species of Corispermum. Since some evi¬ dence exists that paleo-Americans collected and consumed Corispermum fruits (see Betancourt et al., 1984), the possible role of prehistoric people in the local dispersal of seeds should also be taken into consideration. More detailed reconstruction of the history of distribution and migration of Corispermum during the Pleistocene and Holocene will be published lat¬ er. However, it seems to be useful to mention some basic considerations concerning the history of the genus in North America. Corispermum was present in North America at least 38,000 years BP (before present) in Alaska and the Yukon, 11,000-14,000 years BP in Arizona and Utah, ca. 4,000 years BP in New Mexico (for details see Betancourt et al., 1984; Matthews, 1982; Rosendahl, 1948; Young, 1982). In my opinion, ancestors of the modem native North American taxa of this genus migrated to non- glaciated parts of Alaska and the Yukon through the Bering Land Bridge after the Pelukian (Kazan- tsevo/Sangamon) sea transgression, most possibly after Zyryanka/Early Wisconsin glaciation, during the Kargin (Zyryanka-Sartan) interstadial. Their further southward expansion became possible after the retreat of Late Wisconsin glaciers and separa¬ tion of Laurentide and Cordilleran ice sheets. There is also the possibility of their migration to more southern parts of North America during the Middle Wisconsin interstadial; this helps to explain the most ancient records of fossil Corispermum fruits in the southwestern United States. Judging from the present distribution of Coris¬ permum species and available fossil records of the genus, there were at least three main paleo-migra- tion routes of ancestors of modem taxa in North America. All these routes started in non-glaeiated parts of Beringia. The Pacific route followed the coastal line of the Pacific Ocean and main river valleys of northwestern North America. This route Volume 5, Number 4 1995 Mosyakin Corispermum from North America 345 seems to be the main migration path for ancestors of C. pacificum and C. pallidum. The two other mi¬ gration corridors are marked by the present distri¬ bution of species of the C. pallasii—C. americanum aggregate. One of these reconstructed routes went southward and partly southwestward along the Rocky Mountains and adjacent plains, and another crossed Canada in a southeastern direction toward the Great Lakes. During their expansion, species of Corispermum probably followed shorelines of Pleis¬ tocene glacial and post-glacial paleo-lakes (Agas¬ siz, McConnell, precursors of the Great Lakes, etc.) and ancient river valleys, as well as fluvioglacial deposits exposed after the retreat of glaciers. Validation of New Taxa As stated above, Corispermum is represented in North America by several native species belonging to at least three species aggregates (species groups). Five of these species are described below. Corispermum pacificum Mosyakin, sp. nov. TYPE: U.S.A. Washington: Wawawai, appar¬ ently introduced, Oct. 1893, C. V. Piper 1770 (Native ! 1900 C. V. P.) (holotype, GH; iso¬ types, F, GH, MIN, MO, NY, UC). Figure 1A. Planta annua, ca. 15-40 cm alta, a basi ramosa, glabra vel sparsim stellato-pilosa (saepissime in partibus juven- alibus). Folia 2—5(—7) cm longa, 0.2-0.6 mm lata, anguste lanceolata, lineari-lanceolata vel linearia, plana, apice brevissime mucronulata. Inflorescentia plerumque com- pacta, densa vel rarius plus minusve remotiflora, apice solum congesta: ovoidea, elongato-ovoidea, elongato-cla- vata, rarius late linearia. Bracteae saepissime (1.0—)1.5— 2.5 cm longae, 0.3-0.7(-0.9) cm latae; ovato-lanceolatae, anguste ovato-lanceolatae, ovatae, lanceolatae, apice ac- uminatae; saepissime late membranaceo-marginatae, im- bricatim dispositae. Perianthium monophyllum, apice denticulatum. Fructus maturi 3.0-4.0 X 2.7—3.8 mm, sub- rotundati vel rotundato-obovati, per lineam mediam latis- simi vel rarius leviter supra medium, in fronte paulo con- vexi vel rarius subplani, a tergo plerumque plani vel leviter concavi, nitiduli, alati; ala plerumque (0.2—)0.3— 0.6 mm lata, membranacea, subpellucida, margine undu- lala vel irregulariter minutissime denticulata (rarius inte- gerrima), apice rotundata vel rarius subemarginata; nucle¬ us maturus saepissime niger, rarius atro-olivaceus. Annual plant ca. 15-40 cm tall (or occasionally more?), usually branched from the base; glabrous, or sparsely covered with branched and almost stel¬ late hairs (especially when young). Leaves usually 2—5(—7) cm long, 0.2-0.6 mm broad, narrowly lan¬ ceolate, linear-lanceolate, or linear; flat or nearly so; short-mucronulate at apex. Inflorescences usu¬ ally compact, rather dense, or sometimes ± lax, condensed only near the apex; ovate, ovate-elon¬ gate, obovate, elongate-clavate, or occasionally broadly linear. Bracts (1.0-)1.5-2.5 cm long, 0.3— 0.7(—0.9) cm broad; ovate-lanceolate, narrowly ovate-lanceolate, almost ovate, or lanceolate, acu¬ minate at apex; usually with broad membranous margins; overlapping. Perianth segment 1, with erose or denticulate apex. Mature fruits usually 3.0-4.0 X 2.7-3.8 mm, almost orbicular or orbic- ular-obovate in outline, broadest near the middle (or rarely slightly above the middle); slightly convex (rarely almost flat) at the abaxial side, flat or slight¬ ly concave at the adaxial side; shiny; winged. Wing usually (0.2—)0.3-0.6 mm broad, thin, translucent; slightly undulate or indistinctly erose-denticulate at margins; rounded or rarely indistinctly emarginate (notched) at apex. Mature fruit body (nucula, “ker¬ nel”) in most cases black (rarely deep olive green), sharply contrasting with greenish semi-transparent wings. Corispermum pacificum is not closely related to other native or introduced North American taxa. Its closest allies are Siberian C. crassifolium Turczan- inov and C. maynense Ignatov. The latter species occurs in the northeastern Russian Far East and may be expected to occur in Alaska and/or north¬ western Canada. Corispermum pacificum differs from C. maynense by its usually more robust habit, and its wing rounded (rarely rounded-truncate or indistinctly emarginate, but not triangular) at apex. From C. crassifolium it may be distinguished by the constant presence of perianth segments, and more flattened black mature fruits. At present C. pacifi¬ cum is known to me only from cited localities in Washington, Oregon, and northwestern Idaho, but it probably also occurs in adjacent regions of Brit¬ ish Columbia. The specimens of this species were misidentilied by Maihle and Blackwell (1978) partly as Corisper¬ mum hyssopifolium, and partly as C. nitidum. I do not agree with these determinations, nor with rea¬ sons for splitting C. pacificum into two species. Paratypes. U.S.A. Idaho: Nez Perce Co., sandy island in Clearwater River, opposite Lewiston, 12 Oct. 1924, H. St. John 6792 (UC). Oregon: Howell’s Pacific Coast Plants, Josephine Co., Grant’s Pass, Sep. 1887, Thomas Houiell s.n. (OSC); Multnomah Co., sandy bar on the Co¬ lumbia River just E of the “Interstate Bridge.” 23 Sep. 1915, M. A. Flinn s.n. (OSC); Plants of Oregon. Sand bar on Hayden Island opposite Vancouver, Washington, 11 Oct. 1919, J. C. Nelson 2955 (GH); Whitman Co., Wa- wawaii, Snake River, dry river sands, 13 Oct. 1923, F. A. Warren 168 (NY); Flora of Northwest America. Sandy ground, Hayden Island, Oregon, 8 Oct. 1927, J W. Thompson 3915 (MO, PH); Hood River Co., sandy muddy bank of the Columbia River, with Riccia and Cyperus. about 15-20 mi. W of the Dalles, Oregon. 22 Oct. 1961, Georgia Mason 4201 (OSC). Washington: Flora of Wash¬ ington Territory. Northern Transcontinental Survey, Yaki- 346 Novon E E Figure 1. Fruits of Corispermum : comparison between North American and Asian species. —A. Corispermumpacificum (North America). —B. Corispermumpallidum (North America). —C. Corispermum platypterum (Asia). —D. Corispermum maerocarpum (Asia). —E. Corispermum welshii (North America). Volume 5, Number 4 1995 Mosyakin Corispermum from North America 347 ma Region, sands of the Columbia, Oct. 1882, Frank Tweedy s.n. (NY); W. Klickitat Co., sandy banks of the Columbia River, 11 Sep., 29 Oct. 1893, W. N. Suksdorf 1385 (F, GH, MIN, MO, NY, UC) [A mixed collection; see note below—S.M.]; Klickitat Co., Spedio, sandy shores and sandy slopes, very common, 19 Sep. 1925, M. W. Gorman s.n. (OSC); Whitman Co., sandy bank of the Snake River at Wawawai, 15 Oct. 1939, Marion Ownbey & Ruth P. Ownbey s.n. (CAN, GH, OSC, PH, RM, RSA, TEX, UC); Whitman Co., Wawawai—Lewiston road along Snake River, on sandy beach at edge of river, 15 Sep. 1956, R. T. Ogilvie s.n. (DAO, UBC). Some degree of reproductive isolation between the sympatric species of Corispermum exists be¬ cause of different phenology, especially different flowering period. However, occasional hybrids probably occur between partly sympatric C. paci- ficum and C. villosum. These hybrids are similar in habit to C. pacificum: they have usually broad leaves and dense inflorescences. However, many of their fruits (up to 50—70%) are abortive, almost lacking a perisperm. Occasional well-developed fruits are polymorphic and show transitional char¬ acters: they are usually narrow-winged or almost wingless, strongly convex at the abaxial side (char¬ acters of C. villosum), black or nearly so, broadly ovate to orbiculate in outline (characters of C. pa¬ cificum). The high percentage of abortive fruits in¬ dicates that C. pacificum and C. villosum are tax- onomically rather distant species, because hybrids between closely related taxa of Corispermum (i.e., belonging to the same species aggregate) usually show no evident loss of fertility. The studied spec¬ imens of C. pacificum X C. villosum are cited be¬ low: U.S.A. Oregon: Umatilla Co., mouth of Umatilla River, Umatilla, 11 July 1915, M. E. Peck 6667 (OSC); along Columbia, opposite Dalles, Oregon, 1915, J. Evans s.n. (OSC); Baker Co., 28 Nov. 1952, LeRoy Wright s.n. (OSC); Hood River Co., drifting sand of the Columbia River, 10 Sep. (fl), 1 Oct. 1923 (fr), L F. Henderson 344 (of 1924) (MO). Washington: Wallula, sandy soil, alt. 220 m, 17 Sep. 1894, J. B. Leiberg 912 (OSC). Corispermum pallidum Mosyakin, sp. nov. TYPE: U.S.A. Washington: Douglas Co., in drifting sand, Wilson Creeks, 25 July 1893. J. II. Sandberg & J. B. Leiberg 309 (holotype, MO; isotypes, CAN, F, OSC, PH. UC, US). Fig¬ ure IB. Planta annua, ca. 5—25 cm alta, a basi ramosa, sub¬ glabra vel in partibus juvenalibus stellato-pilosa et min- utissime papillosa. Folia 2—5 cm longa, 0.1—0.2(-0.3) cm lata, linearia, anguste linearia vel filiformia, plana vel par- tim convoluta, apice brevissime mucronulata. Inflorescen- tia plerumque laxiuscula, rarius densiuscula; linearia vel elongato-linearia. Bracteae (0.5—)1.0—1.5(—2.0) cm longae, 0.2—0.5 cm latae; plerumque anguste ovato-lanceolatae vel lanceolatae, apice acuminatae; saepissime membran- aceo-marginatae, imbricatim dispositae vel plus minusve remotae. Perianthium monophyllum, apice denticulatum. Fructus maturi saepissime 2.8-3.5(-3.8) X 2.4—2.8(-3.3) mm, rotundato-obovati vel obovati, plerumque supra me¬ dium vel rarius per lineam mediam latissimi, in fronte subplani vel subplano-convexi, a tergo plerumque plani vel leviter concavi, subnitidi vel opaci, alati; ala pler¬ umque 0.7—1.0 mm lata, membranacea, subpellucida, margine erosa vel irregulariter eroso-denticulata (rarius subintegra), apice stylorum basi longe adnata, emarginata vel rotundata; incisura inter basin stylorum margine ala¬ rum profundiore; nucleus maturus pallidus, stramineus vel brunneus, saepe maculatus. Annual plant ca. 5-25 cm tall (or more?), branched from the base, almost glabrous, or cov¬ ered with branched and almost stellate hairs and minute papillae (especially on veins of young bracts and upper leaves). Leaves 2-5 cm long, 0.1-0.2(- 0.3) cm broad, linear, narrowly linear, or filiform; flat or occasionally convolute (especially at matu¬ rity); acuminate or short-mucronulate at apex. In¬ florescences usually lax, rarely rather dense; linear or elongate-linear. Bracts usually (0.5—)1.0-1.5(- 2.0) cm long, 0.2-0.5 cm broad, usually narrowly ovate-lanceolate or lanceolate, acuminate at apex; in most cases with distinctly membranous margins; overlapping or slightly remote. Mature fruits 2.8- 3.5(—3.8) X 2.4—2.8(—3.3) mm, rounded-obovate or obovate in outline, usually broadest above the mid¬ dle or occasionally near the middle; almost flat or slightly convex at abaxial side, plane or slightly concave at adaxial side; slightly shiny or dull; winged. Wing normally 0.7—1.0 mm broad, thin, translucent, with erose or irregularly erose-dentic- ulate (rarely almost entire) margins; long-adnate to the style bases, emarginate (notched) or rounded at apex. Style bases unusually long (ca. 0.7-1.0 mm, including their parts adnate to wing), in most cases divided to below the edge of the wing. Mature fruit body (“kernel,” nucula) flattened, pale, usually straw-colored or yellowish brown, occasionally with reddish brown spots. Corispermum pallidum seems to be a well-de¬ fined species. The combination of its characters is very distinctive: pale, flattened, and small fruit body; very broad (especially as compared to the fruit dimensions), thin translucent wing with erose margins; and long style bases distinctly divided in their upper parts. Young bracts and upper leaves of C. pallidum are often papillose on margins and veins, in combination with typical branched tri- chomes. This character (papillosity) is quite rare in Corispermum, since most of its species are normally covered only with branched or almost stellate hairs. The same combination of branched hairs and min- 348 Novon ute papillae occasionally occurs in some Asian spe¬ cies. It is hard to believe, but C. pallidum seems to be most closely related to the eastern Asian spe¬ cies C. macrocarpum Bunge ex Maximowicz, which occurs at present in the southern Russian Far East and northeastern China, mostly along the Amur River and some of its tributaries. Two Manchurian species, C. sterwlepis Kitagawa and C. platypterum Kitagawa, also belong to the same aggregate (for additional information about these Asian taxa, see Kitagawa, 1935; Baranov, 1969; Tsien Cho-po & Ma Cheng-gung, 1979). Fruits of C. pallidum are very similar to those of C. macrocarpum in their morphological features, but not in their size; the latter species has much larger fruits 5.0-6.0(-6.5) X 3.8-5.5 mm (Fig. 1B-D). Maihle and Blackwell (1978) placed rather uni¬ form specimens of Corispermum pallidum into two taxa, C. hyssopifolium sensu Maihle & Blackwell (Sandberg & Leiberg 309, Thompson 6764) and C. nitidum sensu Maihle & Blackwell (St. John, Court¬ ney & Parker 4948). Paratypes. U.S.A. Washington: Flora of Eastern Wash¬ ington. Wilson Creek, July 1893, J. H. Sandberg & J. B. Leiberg s.n. (MIN); Grant Co., dry sand, SW of Moses Lake, alt. 1000-1150 ft., 25 June 1921, Harold St. John, W. D. Courtney & Charles S. Parker 4948 (GH, UC); Grant Co., sandy sagebrush plains near Quincy, 15 June 1931, J. William Thompson 6764 (GH, MO). Corispermum welshii Mosyakin, sp. nov. TYPE: U.S.A. Utah: Kane Co., sand dunes, with pon- derosa pine, willow, and gambel oak, ca. 1900 m. Coral Pink Dunes, T43S, R8W, S8, ca. 8 mi. due W of Kanab, 25 Sep. 1992, S. L. Welsh & K. H. Thorne 25170 (holotype, MO; iso¬ types, BRY, NY). Figure IE. Planta annua, ca. 10-35 cm alta, a basi ramosa, dense vel sparsim stellato-pilosa. Folia 2-6 cm longa, 0.2-0.4(- 0.5) cm lata, lineari-lanceolata vel linearia, fere plana vel rarissime partim subconvoluta, apice brevissime mucron- ulata. Inflorescentia plerumque compacta, densa vel rarius plus minusve laxiuscula, apice congesta; ovoidea, elon- gato-obovata vel elongato-clavata. Bracteae 1.0-3.0 cm longae, 0.3-0.8 cm latae; ovatae vel ovato-lanceolatae, rarius anguste ovato-lanceolatae, apice acuminatae; late membranaceo-marginatae, imbricatim dispositae. Perian- thium monophyllum, apice denticulatum. Fructus maturi saepissime (3.3—)3.7-4.6 X (2.7—)3.0—3.6 mm, obovati vel rotundato-obovati, plerumque supra medium vel rarius per lineam mediam latissimi, in fronte convexi, a tergo pler¬ umque leviter concavi, subnitidi vel opaci, alati; ala pler¬ umque (0.3—)0.4—0.6 mm lata, membranacea, subpellu- cida (rarius valdiuscula, margine solum subpellucida), margine integra vel irregulariter eroso-denticulata, apice rotundata, truncata vel subemarginata; nucleus maturus stramineus, fulvus vel brunneus, saepissime maculatus et albido-verrucosus. Annual plant ca. 10—35 cm tall (or more?), branched from the base or nearly so, densely or sparsely covered with branched or stellate hairs. Leaves 2-6 cm long, 0.2-0.4(-0.5) cm broad, lin¬ ear-lanceolate or linear, usually plane, or rarely slightly convolute or folded (especially in dried plants), short-mucronulate at apex. Inflorescences usually rather compact and dense, rarely ± lax, condensed only at apex; ovate, oblong-obovate, or oblong-clavate in outline. Bracts 1.0—3.0 cm long, 0.3-0.8 cm broad, ovate or ovate-lanceolate (rarely narrowly ovate-lanceolate), acuminate at apex, with broad membranous margins. Perianth segment 1, minutely dentate at apex. Mature fruits normally (3.3-)3.7—4.6 X (2.7-)3.0-3.6 mm, broadly obovate or obovate-orbicular in outline, in most cases broadest above the middle (rarely near the middle); convex at abaxial side, usually concave at adaxial side; slightly shiny or dull; winged. Wing usually (0.3—)0.4-0.6 mm broad, thin, translucent (or oc¬ casionally more thick, translucent only at margin), with entire or irregularly erose-denticulate margins; rounded, truncate, or indistinctly emarginate at apex. Mature fruit body (nucula, “kernel”) straw- colored (yellowish brown), light brown or brown, usually with reddish brown spots and whitish warts. This species is probably closely related to the broad-winged and large-fruited plants of the south¬ western U.S.A. ( Corispermum americanum var. ryd- bergii Mosyakin), which Rydberg included in his C. marginale (excluding the type of C. marginale). However, C. welshii differs from the mentioned va¬ riety, as well as from C. americanum sensu proprio, in having shorter, broader, and denser spikes, dis¬ tinctly broader leaves and bracts, more rounded (occasionally almost orbicular), and often more prominently winged fruits. Its difference from C. villosum is also evident, since it has larger fruits approximately twice as long and broad as the latter. From C. pallasii it may be distinguished by its more orbicular fruits having membranaceous wings, which are rounded, rounded-truncate, or indistinct¬ ly emarginate at apex. In general appearance C. welshii resembles some eastern Asian taxa, for ex¬ ample, C. elongatum s.l. It is also somewhat similar in habit to European C. marschallii but is not close¬ ly related to it. At present C. welshii is known to me only from Utah and Wyoming, but similar forms (which are in need of additional study) occur also in other southwestern states, particularly in Colo¬ rado and western Texas. For example, the following collection from western Texas probably also belongs to C. welshii (it differs from typical plants in having more lax spikes and prostrate habit, but the fruits Volume 5, Number 4 1995 Mosyakin Corispermum from North America 349 are almost identical to those of the type): Flora of Texas. Gypsum, Hudspeth Co., from 13 mi. E of Dell City on nearly barren gypsum dunes, 14 Oct. 1984, R. D. Worthington 12752 (TEX). Paratypes. U.S.A. Utah: Millard Co., sand dunes, alt. 4500 ft., 18 Sep. 1926, collected by Cottam, determined by Garrett 1010 (BRY); Millard Co., juniper association, sand dunes, 18 Sep. 1926, coll. A. 0. Garrett 1010 (RM); Salt Lake Co., South Salt Lake City, just S of intersection of 7400 South and 20th East in a sand dune area, alt. 4650 ft., T. 25, R. 1 E„ SE ii of Sec. 28, 29 Oct. 1967, L. Arnou 805 (GH), 805a (COLO); Kane Co., Halls Creek Bay, Lake Powell, sandy benchland, 4 June 1972, S. L Welsh & G. Moore 11797 (BRY) [this plant evidently was buried by sand, and because of this it has developed atyp- ically long branches—S.M.]; Kane Co., Coral Pink Sand- dunes State Park S of Mt. Carmel Junction, elev. 5500 ft., 11 Oct. 1972, B. Albee 1215 (UT); Uintah Co., T3S, R22E, SW Sec. 4, Red Fleet Reservoir, ca. 6000 ft. elev., opens in juniper community, sandy soil, 3 Oct. 1980, E. Neese & M. Chatterley 9894 (NY, RM) [these specimens have fruits slightly smaller than in the type—S.M.]; Kane Co., Navajo Sandstone, Ponderosa, Oakbrush community, T43S R7W S30, Moquith Mountain, ca. 7 mi. W of Kanab, 2142 m elev., 25 Sep. 1992, K. Thorne & S. L. Welsh 15192 (BRY) [also a specimen with slightly smaller fruits— S.M.]. Wyoming: Sweetwater Co., sand dunes, near Steamboat Mt., 1 Sep. 1936, Marion Ownbey & Robert L. Lang 1122 (COLO); Niobrara Co., T34N R61W Sec 36 SE V *, sandy roadside, associates: Cenehrus , Salsola ; elev. 4700 ft., 26 Sep. 1986, R. Dorn 4549 (NY) [inflorescences of this specimen are more lax than in the type, showing a transition toward C. americanum —S.M.]. Corispermum navicula Mosyakin, sp. nov. TYPE: U.S.A. Colorado: Jackson Co., North Park, North sand dunes due E of Dowell Ranch, T10N, R78W, Sec. 6, between Govern¬ ment Creek and North Sand Creek, Eagle Hill Quadr., 29 Sep. 1976, F. Martin Brown s.n. (holotype, COLO). Planta annua, ca. 5—15(—25?) cm alta, a basi ramosa, sparsim stellato-pilosa vel glabriuscula. Folia (1.5—)2—4 cm longa, 0.1-0.5 cm lata, anguste lanceolata vel iineari- lanceolata, plana, rarissime partim convoluta. Inflorescen- tia plerumque compacta, densa, ovoidea vel elongato-ovo- idea. Bracteae 0.5-2.0 cm longae, 0.2-0.6 cm latae; imbricatim dispositae. Perianthium monophyllum. Fructus maturi (4.2-)4.5-5.0(-5.2) X 2.5-3.0 mm, elongato-obov- ati, supra medium latissimi; in fronte valde convexi. a tergo plerumque distinctissime concavi; apteri vel anguste alati. Ala plerumque 0.1—0.2(—0.3) mm lata, valdiuscula. non pellucida vel semipellucida; margine integerrima vel minutissime eroso-denticulata, saepe involuta; apice ros- trata, triangularia, stylorum basi longe adnata. Nucleus maturus atro-brunneus vel atro-viridis, plerumque macu- latus et/vel albido-verrucosus. Annual plant ca. 5—15(—25?) cm tall, branched from the base, sparsely covered with branched and almost stellate hairs, or almost glabrous. Leaves (1.5-)2-4 cm long, 0.1-0.5 cm broad, narrowly lan¬ ceolate or linear-lanceolate, flat or rarely convolute. Inflorescences usually compact and dense, ovate or oblong-ovate in outline. Bracts 0.5-2.0 cm long, 0.2-0.6 cm broad, strongly imbricate. Perianth seg¬ ment 1. Mature fruits (4.2-)4.5-5.0(-5.2) X 2.5- 3.0 mm, elongate-obovate or narrowly obovate-el- liptic, broadest above the middle; strongly convex at abaxial side, usually prominently concave at ad- axial side; narrowly winged or almost wingless. Wing usually 0.1-0.2(—0.3) mm broad, rather thick, not translucent, or semi-translucent; with entire or indistinctly erose margins, often involute toward the adaxial side; rostrate-triangular and long-adnate to style bases at apex. Mature fruit body (nucula, “kernel”) usually dark brown or dark green, usually with dark spots and whitish warts. This species is amazingly similar in its fruit mor¬ phology to the Siberian species Corispermum bar- durwvii M. Popov ex M. Lomonosova (see M. N. Lomonosova, 1992). Corispermum navicula and C. bardunovii probably represent results of parallel evolution (or parallel variability?) within North American and Asian representatives of the C. pal- lasii aggregate, respectively. It is also worth men¬ tioning that the most distinctive characters of C. navicula and C. bardunovii, such as elongated fruit body with almost parallel margins in the middle portion and distinctly triangular apex, show a tran¬ sition (or a parallel evolutionary trend?) toward the representatives of Corispermum sect. Declinata Mo¬ syakin (1994). Additional study of C. navicula is desirable for clarification of its relationships with other species. Some specimens from Oklahoma (e.g., Stillwater, Oklahoma, in sand on bank of Ci¬ marron, 6 Nov. 1899, E. E. Bogue s.n., MIN) prob¬ ably also belong to C. navicula, or at least approach this species most closely. Paratype. U.S.A. Colorado: Jackson Co., North Sand Hills, E of Cowdrey, T10N, R78W, S6 NE W, 2575 m, 1 Oct. 1982, D. H. Wilken 13924 (NY, RM). Corispermum hookeri Mosyakin, sp. nov. TYPE: Canada. Saskatchewan: District de Moose Jaw, palier sablonneux au pied du Coteau du Mis¬ souri, 5-6 mi. 4 Test de Mortlach, dune ev- entrce, 11 Sep. 1960, B. Boivin & G. F. Led- ingham 14079 (holotype, DAO; isotypes, NY, TRT, UBC). Planta annua, ca. 10-40 cm alta, a basi ramosa, spar¬ sim stellato-pilosa vel glabriuscula. Folia 2—5 cm longa, (0.1-)0.2-0.5(-0.6) cm lata, anguste lanceolata vel li- neari-lanceolata, plana. Inflorescentia plerumque densa ovoidea, ovato-clavata vel ovato-cylindrica, rarissime in- terrupta. Bracteae 0.5-1.5(-2.0) cm longae, 0.3-1.0 cm latae, ovatae vel ovato-lanceolatae, imbricatim dispositae. 350 Novon Perianthium monophyllum. Fructus maturi (3.2—)3.5- 4.5(—5.0) X 2.2—3.3(-3.5) mm, elongato-obovati vel obov- ati, per lineam mediam latissimi vel leviter supra mediam latiores; in fronte valde convexi, a tergo concavi vel sub- plani; apteri vel angustissime alati. Ala ad 0.2 mm lata, semipellucida; margine integerrima, apice rotundata vel leviter triangularia. Nucleus maturus atro-olivaceus, brun- neus vel rarissime niger, plerumque non maculatus et non verrucosus, rarius maculatus. Annual plant ca. 10-40 cm tall (or occasionally more?), branched from the base, sparsely covered with branched and almost stellate hairs. Leaves usually 2-5 cm long, (0.1-)0.2-0.5(—0.6) cm broad, narrowly lanceolate or linear-lanceolate, plane. In¬ florescences usually dense, ovate, ovate-clavate, or ovate-cylindric, rarely interrupted (especially near the base). Bracts 0.5-1.5(—2.0) cm long, 0.3-1.0 cm broad, ovate or ovate-lanceolate, imbricate (usually strongly overlapping). Perianth segment 1. Mature fruits (3.2-)3.5-4.5(—5.0) X 2.2-3.3(-3.5) mm, oblong-obovate or obovate, broadest near the middle or slightly above the middle; strongly con¬ vex at abaxial side, prominently concave to sub¬ plane at adaxial side; wingless or narrowly winged. Wing (when present) up to 0.2 mm broad, semi¬ transparent; margins entire, apex rounded or indis¬ tinctly triangular. Mature fruit body (nucula, “ker¬ nel”) usually deep olive green, brown, or rarely almost black, normally without spots and/or warts, or occasionally spotted. This species seems to be most closely related to Corispermum pallasii s. str. and C. ochotense var. alaskanum (see discussion above and comments on C. ochotense below). It is also similar to the small- fruited species, C. villosum, and to the large-fruited C. navicula described above. Some specimens of C. hookeri from British Columbia have unusually elon¬ gated fruits somewhat similar to those of Asian spe¬ cies of Corispermum sect. Declinata. These plants are described below as a new variety. Paratypes. CANADA. Alberta: Lethbridge, 27 Aug. 1911, M. 0 . Malt s.n. (CAN); Battle River, S of Wain- wright, sandy soil, near river bank, 8 Sep. 1951, E. H. Moss 9725 (DAO) [the voucher specimen: 2n = 18, de¬ termined by G. A. Mulligan]; SW of Orion, sand dunes, grazed by livestock in the area, 28 Aug. 1952, E. H. Moss 10110 (CAN). British Columbia: very common on Dryas Island in Fraser River at Hope, 19 July 1914, T. L Thack¬ er s.n. (UBC); Lytton, 26 Sep. 1922, W. B. Anderson 6678 (V); in sand landfill near Alex Fraser Bridge, Annacis Island, many plants in sand, 7 Sep. 1988, Frank Lomer s.n. (UBC); sandy ground under the Pattullo Bridge, Sur¬ rey, a single plant, 7 Sep. 1991, Frank Lomer s.n. (UBC); Steveston, Richmond, Lulu Island, 49°07'N, 123°10'W, 7 Sep. 1992, Huber Moore & Terry Taylor s.n. (UBC); sand landfill dredged from Fraser River, near Fraser Surrey Docks, Surrey, many plants in sand, 27 Aug. 1993, Frank Lomer s.n. (UBC) [this plant is transitional toward C. hook¬ eri var. pseudodeclinatum Mosyakin described below— S.M.]. Manitoba: sand hill 35 mi. SE of Brandon, 23 Sep. 1960, G. A. Stevenson 2255 (CAN, DAO). Northwest Territories: Mackenzie District, in dry sand near shore¬ line, sand hills on S shore of Mackenzie River about 16 mi. downstream from Fort Simpson, 61°58'N, 121°45'W, 25 July 1955, W. J. Cody & J. M. Matte 8944 (DAO) [the specimen is immature; fortunately, authors also collected some fragments of dry plants remaining from the previous year—S.M.]; Mackenzie District, occasional in disturbed sand along roadside on mainland about 1 mi. S of Fort Simpson island, 61°52'N, 121°22'W, 3 Aug. 1955, W. J. Cody & J. M. Matte 9130 (DAO) [the specimen is im¬ mature, but very similar to the previous one—S.M.]. On¬ tario: Sarnia, Oct. 1936, W. A. Dent s.n. (DAO); Norfolk Co., Long Point, 22 Sep. 1946, F. A. Clarkson 6684 (DAO) [a specimen with the same collection number at TRT ap¬ proaches C. pallasii —S.M.]; Long Point Village, sand on open shore, 24 Sep. 1966, R. Beschel 15722 (CAN); Nor¬ folk Co., Long Point, South Walsingham Tp., 12 Oct. 1969, P. M. Catling & J. E. Cruise s.n. (TRT); Norfolk Co., Long Point, along S shore E of Courtright Ridge, Lake Erie, blow-out on sand dune, 23 Sep. 1976, K. M. Lindsay & 1. D. MacDonald 1092 (CAN); Bruce Co., boundary between Chiefs’ pt. I.R. and Amabel Twp., on sand, beside road at edge of pine plantation, 29 Sep. 1977, Joseph W. Johnson (TRT); Norfolk Co., Long Point, Squires Ridge to Little Creek Ridges, 12 x /i to 14 Vi mi. from base of Point, on S beach, open, active sand dunes along beach, nu¬ merous, with Ammophila breviligulata and Andropogon scoparius, 20 Sep. 1979, A. A. Reznicek & P. M. Catling 5255 (DAO). Saskatchewan: Beaver Creek, sandy hill¬ side, 9 Sep. 1937, W. J. White s.n. (TRT); Saskatoon, on sands by river, Sep. 1931, A. W. Anderson s.n. (CAN); Saskatoon, 20 mi. SW at Pike Lake, partially covered sand dunes, 10 Sep. 1951, G. F. Ledingham 1219 (DAO); Moose Jaw Region, Caron, SW Va 16-17-29 W. 2nd, shift¬ ing sand in and around blow-out spots, long-abandoned land, 24 Sep. 1955, T. H. Hudson 1768 (DAO); District de Rosetown-Biggar, coulee Agassiz, Canton Bernard, 12- 15 mi. au sud-est de Demaine, Platiferes de la Saskatch- ewan-Sud, dune en mouvement, 18 Sep. 1960, B. Boivin 14156 (DAO); Saskatoon District, Saskatoon, sandflat be¬ tween natural levee and cutbank of S Sask. River, local, with Elymus canadensis, Psoralea lanceolata, 13 Sep. 1964, J. H. Hudson 2214 (DAO); Lake Atabasca, E of William Point, Salix, Myrica on dry, eroding sand beach on shore, 6 Aug. 1975, G. W. Argus & D. J. White 9750 (CAN) [inflorescence interrupted—S.M.]. Corispermum hookeri Mosyakin var. pseudo¬ declinatum Mosyakin, var. nov. TYPE: Can¬ ada. British Columbia: beside Burnaby Lake, sand dune, 3 Oct. 1965, /. Bayly 83 (holotype, UBC). Fructus elongati vel oblongo-obovati, longitudine lati- tudinem duplo vel subduplo superanti; in fronte valde convexi, a tergo concavi. Inflorescentia elongata, interrup- ta, apice solum congesta. Fruits elongate, oblong-obovate, ca. 2 times as long as broad; prominently convex at abaxial side, concave at adaxial side. Inflorescences elongate, interrupted, condensed only at apex. Volume 5, Number 4 1995 Mosyakin Corispermum from North America 351 Paratype. CANADA. British Columbia: Okanagan, Quilchena, 30 July 1906, E. Wilson 776 (UBC). Corispermum americanum (Nuttall) Nuttall var. rydbergii Mosyakin, var. nov. TYPE: Mexico. Chihuahua: sand dunes, 10-19 Oct. 1935, Harde LeSueur 285 (MO). Fructus maturi (3.2 -)3.3 4.0(- 4.3) X 2.3-3.2 mm, sae- pissime late alati, supra medium latissimi. Ala 0.3-0.4(— 0.5) mm lata. Inflorescentia plerumque angusta, laxius- eula, non denst, plus minusve remotiflora. Ceterum typo respondet. Mature fruits (3.2-)3.5-4.0(-4.3) X 2.3-3.2 mm, usually broadly winged, broadest above the middle. Wing 0.3-0.4(—0.5) mm broad. Inflorescences nor¬ mally narrow, lax; flowers/fruits ± remote. Other characters as in C. americanum var. americanum. Paratype. U.S.A. Colorado: Huerfano Valley, near Gardner, sandy creek bottom, alt. 7000 ft., 10 Sep. 1900, F. K. Vreeland 657 (NY, CAN) [this specimen is also a paratype of C. marginale Rydberg; see discussion above— S.M.], Because the Corispermum americanum group is currently under revision, I refrain now from citing additional paratypes of this variety. However, it seems to occur in several localities in Arizona, Col¬ orado, New Mexico, Texas, and Utah. New collec¬ tions of mature specimens from this region would be extremely helpful for detailing its distribution. Due to its characteristic narrow and interrupted in¬ florescence, this variety, as well as many specimens of C. americanum s. str., was constantly misiden- tified as C. nitidum. Corispermum ochotense Ignatov var. alaskan- um Mosyakin, var. nov. TYPE: U.S.A. Alaska: Onion Portage, Upper Kobuk River, 67°06'N, 158°15'W, in sand dunes, open site, 10 Aug. 1967, Charles Schuieger 196 (holotype, ALA). Fructus maturus 3.2 -1.0 mm longus, 2.2—2.7 mm latus. Ceterum ut in var. ochotense. Mature fruit 3.2-4.0 mm long, 2.2-2.7 mm wide. Other characters as in variety ochotense. Paratypes. U.S.A. Alaska: Yukon River between Rampart and Tanana, 9 Aug. 1932, L J. Palmer 13 (CAN); Ester Siding Field, July 1936, G. IF Gasser s.n. (ALA); Kobuk River valley, forming dense “stands’ on sandy river bar at Point Goldie on Kobuk River, 2 Aug. 1974, C. H. Racine 732 (ALA); Kobuk River valley, low, abundant plant covering sand surface of dunes on S side of Kobuk River opposite mouth of Hunt River, 12 Aug. 1974, C. H. Racine 731 (ALA). Corispermum ochotense, a comparatively small (often dwarf) arctic and subarctic plant, has been described recently from the northeastern Russian Far East (Ignatov, 1986). It is normally floriferous almost to the base, often developing fruits even in the axils of lower leaves and branches. The lower and middle bracts in the inflorescence are usually linear or linear-oblanceolate (often broadest toward the apex, as well as some leaves), rather long, leaf¬ like, almost not different from cauline leaves in their shape and size. In addition, plants often turn deep beet red at maturity. The illustration of “C. hyssopifolium” in the Flora of Alaska (Hulten, 1968) gives an accurate impression of the habit of C. ochotense. The type of C. ochotense (“regio Ma¬ gadan, distr. Ola, pagus Talon, syrtis arenosa ripar- ia, 28—30 Aug. 1971, A. P. Khokhrjakov”), which I have studied at MHA, is eonspecific with the Alaskan and northwestern Canadian plants. How¬ ever, some of the Alaskan specimens differ from their trans-Beringian relatives in having larger fruits. The typical variety of C. ochotense also oc¬ curs in North America. The species is known to me from the following additional localities in Canada (as many specimens were not completely mature, the varieties are not indicated; however, most of the specimens seem to belong to var. ochotense): CANADA. Northwest Territories: District of Mac¬ kenzie, Aklavik River near Aklavik, on the section of slit beach, 6 Sep. 1948, Margaret E. Oldenburg 48—1241 (GH); Mackenzie District, Norman Wells, forming thick mats on eroding heavy soil on slope by airstrip, 22 July 1953, W. J. Cody & R. L Gutteridge 7466 (COLO, DAO, GH, MICH, MO, OSC, TRT, UBC); Mackenzie District, rare on gravel beach of Mackenzie River, Royal Canadian Corps Signals Transmitters, 5 mi. upstream from Norman Wells, 27 July 1953, W. ]. Cody & R. L Gutteridge 7603 (DAO, F); Mackenzie District, 13.0 km WSW of Norman Wells, N.W.T. Site No. 8-10-13, beach ridge, sandy, with Populus tremuloides , 10 Aug. 1974, N. J. Walker 2303 (DAO); Mackenzie District, E bank of MacKenzie River, Norman Wells, N.W.T. Site No. 8-18-1, riverbank, many disturbance species, rare, 18 Aug. 1974, N. J. Walker 2386 (DAO). The following collection from northern Saskatch¬ ewan probably also belongs here (although the plants were not completely mature): CANADA. Saskatchewan: S shore of Lake Atabasca at Beaver Point, lower sandy beach line at, and just above, shoreline, scattered, but frequent, 8 Aug. 1979, V. L Harms & R. A. Wright 27215 (DAO, GH, RM). As outlined above, the native North American species of Corispermum are comparable to most of the widely recognized Eurasian taxa of the genus, regarding the species concept applied to them and morphological differences between species. They are also comparable to some other readily recog¬ nized taxa of Chenopodiaceae, e.g., numerous seg- 352 Novon regate species of such North American aggregates as Chenopodium leptophyllum s.l., or C. fremontii S. Watson s.l. However, especially for followers of a broader species concept, I would like to outline the species aggregates (groups) of closely related taxa of North American Corispermum. Because North American representatives of this genus were commonly misidentified and misunderstood, this explanation would be helpful in preventing possible further mistakes. The closest Asian allies of American Corisper¬ mum are C. pallasii (for almost all native taxa), C. macrocarpum (for C. pallidum ), and C. crassifolium (for C. pacificum). In my opinion, in the future C. pacificum and C. pallidum could not be lumped together with other North American taxa. However, using a very broad species concept, ore may regard C. pacificum as a subspecies of C. crassifolium. Corispermum pallasii is most closely related to C. americanum, C. navicula, and C. welshii; under a broad species concept the three latter species could fit at the subspecies level as well. Corispermum ochotense, C. hookeri, and C. villosum form another unit, which is, however, connected to the previous one by occasional forms transitional between C. americanum and C. villosum, as well as by transi¬ tions between C. pallasii and C. hookeri. Using the ultra-broad species concept, one could unite the mentioned taxa under the priority name C. pallasii. However, I believe that at the present stage of our knowledge, such an approach could only obscure the true relationships within this group. At present I prefer to place the native North American representatives of Corispermum into three species aggregates (groups): the Corispermum pallasii aggregate: C. pallasii s. str., C. americanum, C. navicula, C. welshii, C. hookeri, C. villosum, C. ochotense; also the Asian C. sibiricum, C. stauntonii, C. elongatum, C. bar- dunovii, and probably some others. the Corispermum crassifolium aggregate: C. pa¬ cificum plus the Asian C. crassifolium s. str. and C. maynense. the Corispermum macrocarpum aggregate: C. pal¬ lidum and the Asian C. macrocarpum s. str., C. platypterum, and C. stenolepis. A key to the species and species aggregates of Corispermum occurring in North America will be published later, in a treatment of the genus for the fourth volume of the Flora of North America. Nat¬ ural arrangement of some Eurasian species and species aggregates will be discussed in an article submitted to the Ukranian Botanical Journal (Ukrajins’kyj botanicnyj zurnal). New, more complete (and preferably serial) her¬ barium material of North American Corispermum, as well as field and experimental studies, would be helpful for further progress in understanding the complicated patterns of taxonomy and distribution of the genus in North America. Acknowledgments. The present study of North American Corispermum was possible due to the support provided by the Missouri Botanical Garden (MO). I thank all members of the staff of the Flora of North America Project for their extremely kind help and advice. The kind help of the staff of many American, Canadian, Russian, Ukrainian, Finnish, Austrian, Bulgarian, Chinese, and Japanese her¬ baria (listed alphabetically: ALA, BKL, BRY, CAN, COLO, DAO, F, GH, H, KW, KWHA, LE, MHA, MICH, MIN, MO, MU, MW, NY, OSC, PE, PH, RM, RSA, SOM, TEX, TI, TRT, UBC, UC, US, UT, UTC, V, W, WU) is especially greatly appreciated. I thank Charlie Jarvis (BM) and Konstantin Dobolyi (BP) for sending me excellent photographs of type spec¬ imens and fruits of Corispermum; thanks are also due to Amy Scheuler McPherson, Diana Gunter, and Helen Jeude (MO) for their skillful comments on the style of the manuscript. The fieldwork in the southwestern United States (Utah and Nevada) dur¬ ing August and September 1994 was made possible by support from the National Science Foundation and kind help from Noel Holmgren (NY), Stanley L. Welsh (BRY), and Steven E. Clemants (BKL). Special thanks are due to Nancy R. Morin (MO), Leila M. Shultz (GH), Clifford W. Crompton (DAO), and Ihsan Al-Shehbaz (MO) for productive discus¬ sion, and to Yevonn Wilson-Ramsey (MO) and Anne Keats Smith (MO) for their kind assistance in producing illustrations and obtaining necessary literature. Literature Cited Adamkiewicz, E. 1970. Studia cyto-taksonomiczne i em- briologiczne nad czterema gatunkami rodzaju Corisper¬ mum L. (Cytotaxonomic and embryological studies on four species of the genus Corispermum L.). Acta Soc. Bot. Poloniae 39(4): 626-652. Aellen, P. 1961. Corispermum L. Pp. 716-723 in: G. Hegi, Illustrierte Flora von Mitteleuropa. Aufl. 2. Bd.3. T.2. Munchen. -. 1964. Corispermum L. Pp. 99—100 in: T. G. Tu- tin et al.. Flora Europaea. Vol. 1. Cambridge Univ. Press, Cambridge. -(revised by J. R. Akeroyd). 1993. Corispermum L. Pp. 119-120 in: T. G. Tutin et al.. Flora Europaea. Ed. 2. Vol. 1. Cambridge Univ. Press, Cambridge. Baranov, A. I. 1969. The species of Corispermum (Che- nopodiaceae) in Northeastern China. J. Jap. Bot. 44: 161-169; 195-206. Bassett, I. J. & J. Terasmae. 1962. Ragweeds, Ambrosia species, in Canada and their history in postglacial time. Canad. J. Bot. 40: 141-150. Volume 5, Number 4 1995 Mosyakin Corispermum from North America 353 Bell, F. G. 1969. The occurrence of southern, steppe and halophyte elements in Weichselian (last-glacial) floras from southern Britain. New Phytol. 68: 913-922. Betancourt, J. L., A. Long, D. J. Donahue, A. J. T. Jull & T. H. Zabel. 1984. Pre-Columbian age for North Amer¬ ican Corispermum L. (Chenopodiaceae) confirmed by accelerator radiocarbon dating. Nature 311: 653-655. Gibson, Ch. 1992. Corispermum leptopterum (Aschers.) Iljin in North Essex. B.S.B.I. News, No. 62: 41—42. Godwin, H. 1975. The History of the British Flora: A Factual Basis for Phytogeography. Ed. 2. Cambridge Univ. Press, Cambridge. Harlan, J. R. & J. M. J. de Wet. 1965. Some thoughts about weeds. Econ. Bot. 19: 16-24. Hooker, W. J. 1840. Flora Boreali-Americana, or the Bot¬ any of the Northern Parts of British America. 2 vols. London. Hult6n, E. 1968. Flora of Alaska and Neighboring Ter¬ ritories. A Manual of the Vascular Plants. Stanford Univ. Press, Stanford, California. Ignatov, M. S. 1986. Chetyre novykh vida s Dal’nego Vos- toka (Four new species from the Soviet Far East). Byull. Moskovsk. Obshch. Isp. Prir., Otd. Biol. 91 (3): 111- 114. Iljin, M. M. 1929. Novye vidy roda Corispermum L. (Cor- ispermi generis species novae). Izv. Glavn. Bot. Sada SSSR 28: 638-654. Jalas, J. & J. Suominen. Editors. 1980. Atlas Florae Eu- ropaeae. Distribution of vascular plants in Europe. Vol. 5. Helsinki. Kitagawa, M. 1935. Corispermum in Manshuria & Korea. Pp. 99—105 in: Report of the First Scientific Expedition to Manchoukuo. Sect. 4, part 2. Tokyo. Kock, U.-V. 1986. Verbreitung, Ausbreitungsgeschichte, Soziologie und Okologie von Corispermum leptopterum (Aschers.) Iljin in der DDR. I. Verbreitung und Aus¬ breitungsgeschichte. Gleditschia 14: 305-325. Lomonosova, M. N. 1992. Corispermum L. Pp. 166-171 in: I. M. Krasnoborov & L. I. Malyschev (editors). Flora Sibiri (Flora Sibiriae). Vol. 5 (Salicaceae-Amarantha- ceae). Novosibirsk. Lunell, J. 1910. New plants from North Dakota. Amer. Midi. Naturalist 1(8): 204-208. Maihle, N. J. & W. H. Blackwell. 1978. A synopsis of North American Corispermum (Chenopodiaceae). Sida 7(4): 382-391. Matthews, J. V. 1982. East Beringia during Late Wiscon¬ sin time: A review of the biotic evidence. Pp. 127—150 in D. M. Hopkins et al., Paleoecology of Beringia. Ac¬ ademic Press, London. Mosyakin, S. L. 1988a. Pryrodno-vydova dyfferentsiatsiya rodu Corispermum L. na terytoriyi Ukrayiny. (The nat¬ ural-specific differentiation of the genus Corispermum L. in Ukraine.) Ukrayins’k. Bot. Zhum. 45(3): 37-40. -. 1988b. Krytychnyy pereglyad vydiv rodu Cor¬ ispermum L. flory Ukrayiny. (A critical revision of Cor¬ ispermum species in Ukraine.) Ukrayins’k. Bot. Zhurn. 45(5): 19-23. --. 1994. New infrageneric taxa of Corispermum L. (Chenopodiaceae). Novon 4: 153-154. Nelson, A. 1909. Coulter’s New Manual of Botany of the Central Rocky Mountains. Revised ed. American Book, Chicago. Nuttall, T. 1818. The Genera of North American Plants, and a Catalogue of the Species to the Year 1817. Vol. 2. Philadelphia. -. 1834. Collections toward a flora of Arkansas. Amer. Philos. Soc. Trans. 5: 139—203. Pielou, E. C. 1991. After the Ice Age: The Return of Life to Glaciated North America. Univ. Chicago Press, Chi¬ cago & London. Rafinesque, C. S. 1836. New Flora and Botany of North America. Part 4. Philadelphia. Rosendahl, C. 0. 1948. A contribution to the knowledge of the Pleistocene flora of Minnesota. Ecology 29: 284— 315. Rydberg, P. A. 1897. Rarities from Montana. I. Bull. Tor- rey Bot. Club. 24(4): 188-192. -. 1903. Studies on the Rocky Mountain flora. X. Bull. Torrey Bot. Club. 30(4): 247-262. -. 1904. Studies of the Rocky Mountain flora—XI. Bull. Torrey Bot. Club 31: 399-410. Small, E. 1973. Xeromorphy in plants as a possible basis for migration between arid and nutritionally-deficient environments. Bot. Not. (Lund) 126: 534—539. Steven, C. 1817. Observationes in Plantas Rossicas et descriptiones specie mm novarum. M£m. Soc. Imp. Na- turalistes Moscou 5: 334—338. Strazdin’sh, Yu. G. 1985. Rod Corispermum L. Pp. 154— 166 in: L. V. Tabaka & al. Flora i rastitel’nost’ Latviys- koy SSR. Vostochno-Latviyskiy geobotanocheskiy ray¬ on. (Flora and vegetation of the Latvian SSR. The Eastern Latvian geobotanic region). Zinatne, Riga. Tsien Cho-po & Ma Cheng-gung. 1979. Corispermum L. Pp. 50-75 in: Flora Reipublicae Popularis Sinicae. To- mus 25(2). Beijing. Uotila, P. 1993. Taxonomic and nomenclatural notes on Ckenopodium in the Flora Iranica area. Ann. Bot. Fenn. 30(3): 189-194. Voss, E. 1985. Michigan Flora. Part 2. Dicots (Saurura- ceae-Comaceae). Ann Arbor, Michigan. Young, S. B. 1982. The vegetation of Land-Bridge Ber¬ ingia. Pp. 179-191 in: D. M. Hopkins et al., Paleo¬ ecology of Beringia. Academic Press, London. Note added in proof. Since this article was submitted for publication, some additional isotypes and paratypes have been found in the herbaria at US and LE: Corispermum pacificum: C. V. Piper 1770 (isotype, US); W. N. Suksdorf 1385 (paratype, US); C. pallidum: J. H. Sandberg & J. B. Leiberg 309 (isotype, LE); C. hookeri: B. Boivin & G. F. Ledingham 14079 (iso¬ type, LE); C. ochotense var. alaskanum: L J. Palmer 13 (paratype, US). Corispermum welshii is now known from two additional states of the U.S.A.: Arizona: Coconino Co., Teetso Spring, sandy soil, 13 Nov. 1971, N. D. Atwood & D. Kaneko 3420 (US); New Mexico: Thornton, 19 Oct. 1903, D. Griffiths 6209 (US). A New Species of Geonoma (Arecaceae) from Panama Gregory C. de Nevers California Academy of Sciences, Golden Gate Park, San Francisco, California 94118, U.S.A. Michael H. Grayum Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. Abstract. Geonoma mooreana is described from wet forests on the Atlantic slope of western Pana¬ ma. Review of herbarium material for floristic and monographic projects involving Central American palms (Arecaceae) has revealed the following new Panamanian species of Geonoma (Arecoideae: Geonomeae). Geonoma mooreana de Nevers & Grayum, sp. nov. TYPE: Panama. Veraguas: vicinity Santa Fe, along road between Santa Fe and Calove- bora, 1.7 mi. past Alto Piedra School, 1.5 mi. beyond Quebrada Cosilla (previously referred to as Rio Primero Braso), 8°33'N, 81°08'W. 570 m, 13 July 1994 (fruit), Croat & Zhu 76826 (holotype, MO; isotypes, CAS, PMA). Species cum G. tenuissima H. E. Moore optime con- gruens, sed differt foliis regulariter pinnatis pinnis angus- tis, inflorescentiis majoribus, labio infero fovearum flor- alium bifidoque. Stems caespitose, with to at least 4 stems per plant, 2-3.5 m high, slender, to at least 1.7 cm diam. at nodes; leaves regularly pinnate; sheath 9— 18+ cm; petiole 41—52 cm, broadly channeled adaxially, rounded abaxially; rachis 62-81 cm, oc¬ casionally terminating in a slender filament to at least 1.7 cm; pinnae 25—35(—51) per side, narrowly linear with one prominent raised vein adaxially, one prominent and two less prominent raised veins abaxially, 9—42 X 0.4—1.5(—3.1) cm, glabrous or essentially so; inflorescence infrafoliar, 2—4 nodes below current leaves, paniculate, branched to three orders, 24—32 X 18-38 cm; peduncle 6-17 cm; prophyll and peduncular bract inserted close to¬ gether at base of peduncle; prophyll 11—15.5(-21) X 2.8—5.0 cm, slightly swollen in bud, obscurely striate; peduncular bract slightly smaller than pro¬ phyll, included in it; inflorescence rachis 6-20.5 cm, with 4—7 primary branches; rachillae ca. 110- 165, reddish, 13—19 cm X 0.5—1 mm, sparsely to moderately setulose at anthesis (later glabrescent). drying rugose; floral pits spirally arranged, ca. 1—2 mm apart, the lips prominently exserted from the rachillae, the lower lip bifid, the orifice ca. 1 X 1 mm; staminate flowers ca. 3 mm, sepals and petals subequal, free, erect at anthesis, slightly imbricate below; stamens 6, filaments spreading, anthers sharply reflexed from the filaments; pistillate flow¬ ers ca. 2 mm, sepals and petals subequal, barely exserted, styles 3, reflexed, staminodial tube trun¬ cate; fruit oblong to globose, green, drying brown, 5-6 X 4—5 mm, surface minutely verrucose (peb¬ bled); seed black, oblong, 4—5 mm; germination un¬ known. Figure 1. Geonoma mooreana occurs in wet forests on the Atlantic slope of western Panama, at elevations of ca. 100-1000 m. It has been collected about 30 km from the Costa Rican border in Bocas del Toro Province. The apparent geographic discontinuity between the Bocas del Toro and Veraguas stations is probably spurious, as the area in question is one of the most poorly collected in Panama. Geonoma mooreana is most similar phenetically to G. scoparia Grayum & de Nevers, G. concinna Burret, and G. tenuissima H. E. Moore. Geonoma scoparia, with which specimens representing G. mooreana were compared by Grayum and de Ne¬ vers (1988: 113), shares the highly branched, broomlike inflorescence and pebbled fruit, but dif¬ fers in its consistently solitary habit, generally smaller dimension?, trijugate leaves, and crenate (vs. truncate) staminodial tube. Geonoma concinna has the truncate staminodial tube and pebbled fruits of G. mooreana, but differs in its consistently trijugate leaves, much smaller inflorescence bracts, and fewer, thicker rachillae. The inflorescence of G. mooreana also resembles that of G. tenuissima of Ecuador, which differs in its simple leaf blades, shorter, more delicate inflorescence rachillae, and floral pits with the lower lip entire rather than bifid. Several collections of Geonoma mooreana bear incomplete or partial inflorescences. Bulky palm inflorescences usually must be pruned for pressing, and the severed branches are often used to make Novon 5: 354-356. 1995. Volume 5, Number 4 1995 de Nevers & Grayum Geonoma mooreana 355 Figure 1. Geonoma mooreana de Nevers & Grayum (A [except fruits]. Nee 11252; B, Hammel 3459; A [fruits], C, Kirkbride & Duke 649). —A. Section of stem with leaf base and inflorescence (many rachillae removed for clarity) (X0.50). —B. Medial portion of leaf (X0.75). —C. Section of infructescence rachilla with fruits (X1.75). 356 Novon duplicate specimens. Thus, the PMA and CAS is¬ otypes of G. mooreana include the main basal and suprabasal branches (respectively) that are missing from the holotype infructescence. Isolated branches may easily be mistaken for entire inflorescences or infructescences in the herbarium, yielding substan¬ tially false impressions of such attributes as pedun¬ cle length, overall inflorescence size, and the num¬ ber of rachillae. The specific epithet commemorates the late Har¬ old E. Moore, Jr. (1917—1980), leading palm au¬ thority of his generation, who realized this was a new species and annotated two of the paratypes (Hammel 3459 and Nee 11252) as “Geonoma sp. nov.” Paratypes. PANAMA. Bocas del Toro: between Q. Lugron and Cerro Bonyic, nr. Rfo Teribe, 90-270 m, 13 Apr. 1968 (fmit), Kirkbride & Duke 649 (MO, NY). Code: Atlantic slope along Rfo San Juan above its fork with Rfo Tife, ca. 1200 ft., 12 June 1978 (fruit), Hammel 3459 (MO). Veraguas: Caribbean slope above Rfo Primero Bra- zo 5 mi. NW of Santa Fe, 700-1200 m, 18-19 Mar. 1973 (flower), Croat 23120 (MO 2 sheets); along road between Escuela Agrfcola and Alto Piedra (above Santa Fe) and Rfo Dos Bocas ca. 5-8 km from Escuela, 730—770 m, 26 July 1974 (bud), Croat 25931 (MO); rd. from Santa Fe to (ialovehora. creek about 5 km beyond Escuela Agrfcola, ca. 500 m, 8°40'N, 81°5'W, 1 Mar. 1995 (past fruiting), de Nevers et al. 10601 (CAS); lower montane wet forest 7 km W of Santa Fe on new road past agricultural school, ca. 2900 ft., 12 Apr. 1974 (flower/young fruit). Nee 11252 (MO). Acknowledgments. We are indebted to Kathy Bell for the excellent line drawings of Geonoma mooreana, and to John Dransfield and an anony¬ mous reviewer for valuable comments on the manu¬ script. Literature Cited Grayum, M. H. & G. C. de Nevers. 1988. New and rare understory palms from the Peninsula de Osa, Costa Rica, and adjacent regions. Principes 32: 101—114. Nomenclatural Notes on the Myrsinaceae of China John J. Pipoly III Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, Texas 76102-4060, U.S.A. Chen Chieh Kunming Institute of Botany, Academia Sinica, Kunming, People’s Republic of China Abstract. The following nomenclatural changes are proposed: Maesa brevipaniculata (C. Y. Wu & C. Chen) Pipoly & C. Chen, M. confusa (C. M. Hu) Pipoly & C. Chen, Embelia laeta (L.) Mez subsp. papilligera (Nakai) Pipoly & C. Chen, E. ribes Bur- man f. subsp. pachyphylla (C. Y. Wu & C. Chen) Pipoly & C. Chen, Ardisia obtusa Mez subsp. pa¬ chyphylla (Dunn) Pipoly & C. Chen, Myrsine cica- tricosa (C. Chen) Pipoly & C. Chen, M. verruculosa (C. Chen) Pipoly & C. Chen, M.faberi (Mez) Pipoly & C. Chen, and M. kwangsiensis (E. H. Walker) Pipoly & C. Chen. Twenty-three taxa in Embelia and Ardisia are reduced to synonymy for the first time. Twelve names in Embelia, Ardisia, and Myr¬ sine are lectotypified. In order to make the names available for the forthcoming treatment of Myrsinaceae for the Flora of China, the following nomenclatural changes in the genera Maesa, Embelia, Ardisia, and Myrsine (including Rapanea) are herein proposed. The au¬ thors have studied type or authentic material from China, the Pacific, and eastern Malesia and are broadening species concepts based on those stud¬ ies. Type specimens seen from KUN have the ac¬ ronym HY marked on them. Maesa ForsskAl Maesa is the only genus in the Myrsinaceae with a semi-inferior to inferior ovary and a berrylike fruit. This paleotropical genus contains nearly 200 species, of which 28 are known from China. Maesa brevipaniculata (C. Y. Wu & C. Chen) Pi¬ poly & C. Chen, comb. nov. Basionym: Maesa parvifolia A. DC. var. brevipaniculata C. Y. Wu & C. Chen, FI. Yunnan. 1: 324. 1977. TYPE: China. Yunnan: Malipo, 26 Aug. 1962, K. M. Feng 22805 (holotype, KUN). Maesa brevipaniculata is distinguished from M. parvifolia by having a velutinous brown tomentum. instead of a brown hirtellous-tomentose and rufous glandular-granulose tomentum on the branchlets, petioles, and inflorescence rachises. Therefore, it is more consistent to recognize M. brevipaniculata as a distinct species. Maesa confusa (C. M. Hu) Pipoly & C. Chen, comb. nov. Basionym: Maesa consanguinea Merrill var. confusa C. M. Hu, Bot. J. South China 1: 12. 1992. TYPE: China. Hainan: Bao-ting Xian, Tong-za Mao-an, 23 Feb. 1957, L. Tang 3789 (holotype, IBSC). Maesa confusa is a poorly known taxon, but its densely and minutely glandular-granulose inflores¬ cence rachises, long pedicels, highly angulate branchlets, and densely black punctate-lineate fruits readily separate it from M. consanguinea. Embeua Burman f. Embelia contains approximately 150 species of lianas (at maturity) that are distributed in the Pa- leotropics from southern China to Malesia, west through Indochina to India, and on to Madagascar and continental tropical Africa. Embelia is a mem¬ ber of the tribe Myrsineae and is the sister genus of Cybianthus Martius (Pipoly, 1987). It is the only lianous member of the family and is distinguished by its plagiotropic branches and free petals and stamens. Walker (1940) recognized 18 species in China, of which 12, as well as 2 additional species hitherto unknown in China, are recognized by the authors. Embelia pauciflora Diels, Bot. Jahrb. 29: 517. 1900. TYPE: China. Sichuan: Nanchuan, Bock & von Rosthom 25 (holotype, B presumed de¬ stroyed; fragment, US). Embelia blinii H. L£veill6, Repert. Spec. Nov. Regni Veg. 10: 375. 1912. Syn. nov. TYPE: China. Guizhou: near Mt. College, to Kematong, Feb. 1898 (pistillate fl), J. Chattanjon & E. Bodinier 2081 (holotype, E; isotypes. A, P). Novon 5: 357-361. 1995. 358 Novon Embelia dielsii H. L6veill6, Repert. Spec. Nov. Regni Veg. 10: 374. 1912. Syn. nov. TYPE: China. Guizhou: Pin-Fa, 17 Feb. 1903 (staminate fl),./. Cavalerie (ho- lotype, E; isotypes. A, P). Embelia kaopoensis H. I/veiHc, Repert. Spec. Nov. Regni Veg. 12: 186. 1913. Syn. nov. TYPE: China. Guizh¬ ou: Kao-Po, 15 Dec. 1911 (pistillate fl, fr), E. Bod- inier s.n. (holotype, E). Embelia pauciflora apparently exhibits extreme local variation in its precocious staminate flower¬ ing, which causes abrupt quantitative variation for those plants bearing essentially “juvenile” leaves. Differences among the populations represented by the type specimens of Embelia blinii, E. dielsii, E. kaopoensis, and E. schlechteri are entirely quanti¬ tative and represent at best localized variation. Embelia laeta (L.) Mez in Engler, Pflanzenr. 9(IV. 236): 326. 1902. Embelia laeta is readily recognized from the oth¬ er members of Embelia by the minute emargination at the leaf apex and gray to reddish bark. It consists of two subspecies, of which subspecies papilligera is distributed in China (Guangdong, Jiangxi, Tai¬ wan) and Japan, whereas subspecies laeta is dis¬ tributed in China (Fujian, Guangdong, Guangxi, Hainan, Jiangxi, Taiwan, Yunnan), Cambodia, Laos, Thailand, and Vietnam. It appears that subspecies papilligera grows in somewhat more humid habitats than subspecies laeta. The two subspecies may be easily distinguished by the following key. la. Branchlets straight to somewhat flexuous, gla¬ brous; leaves chartaceous, margins flat; pedicels and calyx glabrous; petals glandular-ciliate along the margin.subsp. laeta lb. Branchlets geniculate, minutely and densely ru¬ fous glandular papillose-tomentose; leaves cori¬ aceous, margins revolute; pedicels and calyx glandular-puberulent; petals glabrous along the margin.subsp. papilligera Embelia laeta (L.) Mez subsp. papilligera (Nakai) Pipoly & C. Chen, stat. nov. Basionym: Sam¬ ara laeta var. papilligera Nakai in Nakai & Honda, Nov. Fl. Japan 9: 24. 1943. TYPE: Ja¬ pan. Mt. Arisan, between Heishana & Niman- daira, to 7000 ft. alt., Apr. 1914 (fl, fr), Hayata s.n. (holotype, TI). Embelia parviflora Wallich ex A. de Candolle, Trans. Linn. Soc. 17: 130. 1834. TYPE: India. Assam: Wallich 2307 (holotype, K-W). Embelia pulchella Mez in Engler, Pflanzenr. 9(IV. 236): 324. 1902. Syn. nov. SYNTYPES: India. Assam: Griffith s.n. (B destroyed, K); Manipur: Watt 7277 (lectotype, here designated, K). Embelia myrtifolia Hemsley & Mez, Notizbl. Bot. Gart. Berlin 3: 107. 1901. SYNTYPES: China. Yunnan: Mentze, S mountains, (pist. fl), A. Henry 12826; Chi¬ na. Yunnan: Mentze, S mountains, (stam. and pist. fls), A. Henry 9384 (lectotype, here designated. A; isolectotypes, MO, US (pist. fl)). Embelia ribes Burman f., Fl. Ind. 62. pi. 23. 1768. As a result of a detailed study of Embelia ribes throughout its geographic range, it became appar¬ ent that two distinct entities exist. One grows in forests on well-drained soils, while the other is re¬ stricted to water-logged habitats. These are recog¬ nized as subspecies that can be distinguished as follows: la. Branchlets lenticellate; leaf blade chartaceous, sordid adaxially; floral bracts subulate; petals erect in anthesis; stamens and staminodes equal in size, subequaling petals; anthers longer than wide; fruit 3—4(—5) mm diam. when dried; plants of forests, on well-drained sands or sandy loams.subsp. ribes lb. Branchlets without lenticels; leaf blade coria¬ ceous to cartilaginous, nitid adaxially; floral bracts lanceolate; petals highly reflexed in an¬ thesis; stamens slightly exserted ca. 1 mm in an¬ thesis, staminodes ca. Vs petal length; anthers wider than long; fruit 2—3 mm diam. when dried; plants of poorly drained clay areas, swamps, and forests.subsp. pachyphylla Embelia ribes subsp. ribes is distributed in China (Fujian, Guangdong, Guangxi, Guizhou, Hainan, Xizang, and Yunnan), Sri Lanka, India, Myanmar, Laos, Thailand, Cambodia, Vietnam, Malaysia, Su¬ matra, Java, and Kalimantan. Embelia ribes Burman f. subsp. pachyphylla (C. Y. Wu & C. Chen) Pipoly & C. Chen, stat. nov. Basionym: Embelia ribes var. pachyphylla Chun ex C. Y. Wu & C. Chen, Fl. Yunnan. 1: 364. 1977. TYPE: China. Guangdong: Le- chang Xian, Zuo Jing-lie 20451 (holotype, KUN). Embelia tenuis Mez in Engler, Pflanzenr. 9(IV. 236): 314. 1902. Syn. nov. TYPE: Malaysia. Sarawak: Hullet 271 (holotype, K). Subspecies pachyphylla is distributed in China (Guangdong, Guangxi, Hainan, Yunnan), Sarawak, Sumatra, and Vietnam. Embelia vestita Roxburgh, FL Ind. ed. Carey 2: 288. 1824. Volume 5, Number 4 1995 Pipoly & Chen Myrsinaceae of China 359 Embelia prunifolia Mez in Engler, Pflanzenr. IV (236): 316. 1902. SYNTYPES: China. Yunnan: Mentze, 1700-2000 m, (stam. fl), A. Henry 9380C (B de¬ stroyed, A); 10260 (B destroyed, A), 11327A, 11327B (B destroyed. A); China. Yunnan: 5500', A. Henry 9380 (lectotype, here designated. A; isolec- totype, B destroyed). Embelia oblongifolia Hemsley, J. Linn. Soc., Bot. 26: 62. 1882. Syn. nov. TYPE: China. Guangdong: Lofau- shan. Ford 90 (holotype, K). Embelia bodinieri H. Levoillf'. Repert. Spec. Nov. Regni Veg. 9: 327. 1911. Syn. nov. TYPE: China. Guizhou: ./. Esquirol 743 (holotype, E; isotype, P). Embelia rudis Handel-Mazzetti, Anz. Akad. Wiss. Wien. Math.-Nat. 59: 108. 1922. Syn. nov. TYPE: China. Jiangxi: Handel-Mazzetti 157 (holotype, W). Embelia nigroviridis C. Chen, Acta Phytotax. Sin. 16(3): 81. 1978. Syn. nov. TYPE: China. Xizang: Cha-yu, Tsinghai-Tibet Expedition 73-678 (holotype, HP). Embelia vestita is here treated as a polymorphic, broad-ranging species with numerous locally vari¬ able populations. It grows in mixed forests, ever¬ green, broad-leaved forests, shrubby areas, and limestone hillsides in China (Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hunan, Sichuan, Tai¬ wan, Xizang, Yunnan, Zhejiang), India, Myanmar, Nepal, and Vietnam. The type of Embelia rudis represents populations whose tertiary veins are extremely thick and whose adaxial leaf surface is more nitid than the type of E. vestita. The type of Embelia oblongifolia repre¬ sents populations with remotely serrate leaves and fewer, coarse secondary veins than those of the type of E. rudis. The variation noted above precluded segregation of even infraspecific groupings. There¬ fore, the species is treated as a polymorphic och- lospecies (sensu White, 1962; Prance, 1972, 1982; Pipoly, 1983). The highly nitid leaf blades and inconspicuously pellucid or brown punctate sepals easily separate E. vestita from E. undulata and E. scandens, the most closely related congeners. A finisi a Swartz Ardisia is a pantropical genus of about 500 spe¬ cies, including many as yet undescribed from Viet¬ nam, Sabah, Sarawak, Sulawesi, New Caledonia, Madagascar, Colombia, Ecuador, and Peru. It is represented in China by 64 species, most of which are members of subgenus Crispardisia. Ardisia obtusa Mez in Engler, Pflanzenr. 9(IV. 236): 104. 1907. TYPE; China. Hainan: Henry 7990 (holotype, B destroyed; photo and frag¬ ment, US). Ardisia obtusa is a member of subgenus Akosmos Mez, which includes about 50 species and has its centers of diversity in Borneo, Celebes, and Mo¬ luccas. It is represented in China by two subspe¬ cies: subspecies obtusa is endemic to China (Guangdong and Hainan) and grows in thickets, broad-leaf forests, hilly areas, and along creek mar¬ gins; subspecies pachyphylla grows in dense ever¬ green broad-leaf forests, steep valley slopes, and in places with poor drainage in China (Guangxi, Hai¬ nan) and Vietnam. The two subspecies can be sep¬ arated as follows: la. Leaves oblanceolate to obovate; petioles broadly marginate, 1.2—1.5 cm long; sepals 1.2-1.5 mm long; petals 3-6 mm long, inconspicuously pellucid punctate; 0-100 m, lowland forests . . . subsp. obtusa lb. Leaves narrowly oblanceolate or linear-lanceolate; petioles maiginate and canaliculate, 0.6-1 cm long; sepals 0.8-1 mm long; petals 2-2.5 mm long, densely and minutely orange punctate, 400-700 m, montane forests.subsp. pachyphylla Ardisia obtusa Mez subsp. pachyphylla (Dunn) Pi¬ poly & C. Chen, comb, et stat. nov. Basionym: Ardisia pachyphylla Dunn, Bull. Misc. Inform. 1912: 368. 1912. TYPE: China. Hainan: Five Finger Mountains, Katsumata s.n, (HK No. 6680) (holotype, HK; photo and fragment. A, K). Ardisia gracUiflora Pitard in Lecomte, FL Gen. Indo-Chine 3: 832.1930. Syn. nov. TYPE: [Vietnam], Annam: Phu- Hu, prov. Nhatiang, 19 Jan. 1923 (fl), E. Poilane 1362 (holotype, P; isotypes, A 2 sheets). Ardisia graciliflora differs from subspecies pachy¬ phylla by a combination of largely quantitative char¬ acters, and it is considered to be a diminutive form of subspecies pachyphylla, perhaps a juvenile flow¬ ering precociously. Our study of specimens most commonly referred to Ardisia yurmanensis Mez has revealed that the earliest name for the taxon is Ardisia thyrsiflora D. Don. Quantitative differences in leaf length and number of flowers per corymb has caused taxonomic confusion. This species, as now interpreted, is most closely re¬ lated to A. amherstiana A. DC., a taxon most fre¬ quently encountered in Vietnam. The revised synon¬ ymy follows. Ardisia thyrsiflora D. Don, Prodr. Fl. Nep. 148. 1825. TYPE: Nepal, Wallich s.n. (BM). Ardisia tenera Mez in Engler, Pflanzenr. 9(TV. 236): 104. 1902. Syn. nov. SYNTYPES: China. Yunnan: Szemao, 1500 m, A. Henry 12123A (B destroyed, A); China. Yunnan: Szemao, 1500 m, A. Heruy 12094 (holotype, B destroyed; lectotype, here designated. A; isolecto- types, MO, US). Ardisia yurmanensis Mez in Engler, Pflanzenr. 9(TV. 236): 107. 1902. Syn. nov. TYPE: China. Yunnan: Szemao, 5000', tree 30', fls. white, A. Henry 13095 (holotype, B destroyed; lectotype, here designated. A; isolectotypes, MO, NY, US). 360 Novon Ardisia penduliflora Mez in Engler, Pflanzenr. IV(9). 236: 150.1902. Syn. nov. SYNTYPES: China. Hubei: 1885— 1888 (11), A. Henry 4314 (B destroyed); China. Hubei: 1885-1888 (fl), A. Henry 6365 (lectotype, here desig¬ nated, US). Ardisia undulata Mez in Engler, Pflanzenr. 9(IV. 236): 146. 1902. Syn. nov. TYPE: China. Yunnan: Mephra, 1700 m, A Henry 10779 (holotype, B destroyed; lectotype, here designated. A; isolectotypes, MO, US). Later hom¬ onym, not C. B. Clarke (1882). Ardisia thyrsiflora grows on woody hills and in broad-leaf forests, valleys, and damp places in China (Guangxi, Hainan, Xizang, Yunnan), India, Myanmar. Nepal, and Vietnam. It is a highly variable species closely related to A. amherstaiana A. DC., A. waitakii C. M. Hu, A. baotingensis C. M. Hu, and A. quinque- gona Blume. These four taxa, along with Ardisia fordii Hemsley, a diminutive plant, form a natural group within subgenus Akosmos. The separation of these taxa in the flora is at best artificial, and further studies are needed to resolve the limits in this complex. Ardisia corymbifera Mez in Engler, Pflanzenr. 9(IV. 236): 149. 1902. SYNTYPES: China. Yunnan: Simao, 1250 m, (fl), A. Henry 11724 (B de¬ stroyed); China. Yunnan: Simao, (fl), A. Henry 12,000A (holotype, B destroyed; lectotype, here designated. A; isolectotype, MO). Ardisia corymbifera Mez var. tuberifera C. Chen, Acta Phy¬ totax. Sin. 16(3): 80. 1978. Syn. nov. TYPE: China. Guangxi: Lung Chow, S. Chun 13096 (holotype, HK). Ardisia maculosa Mez in Engler, Pflanzenr. 9(IV. 236): 146. 1902. Syn. nov. SYNTYPES: China. Yunnan: Szemao, 6000 ft. elev. (fl, fr), A. Henry 12088D (B destroyed); China. Yunnan: Szemao, shrub 8', fls. pink, fr. red. (fl, fr), A. Henry 12088E (holotype, B destroyed; lectotype, here designated, US 458840; isolectotypes, A 00025251, 00025252; MO 126457, 126458). NOTE: Herbarium accession numbers are included here be¬ cause many sheets contain two accession numbers with two specimens, one of which may not have been cited in the protologue. Ardisia radians Hemsley & Mez in Engler. Pflanzenr. 9(IV. 236): 146. 1902. Syn. nov. Blahfya radians (Hemsley & Mez) Masamune, Short PL Form. 162. 1936. SYN¬ TYPES: China Yunnan: Szemao, 4500 ft. elev., A. Hen¬ ry 535, 9226A (B destroyed, K); China. Yunnan: Sze¬ mao. 4500 ft. elev., shrub 8', fls white (fl), A. Henry 11624C (lectotype, here designated, K; isolectotypes. A, B destroyed, MO). Ardisia patens Mez in Engler, Pflanzenr. 9(IV. 236): 149. 1902. Syn. nov. TYPE: China. Yunnan: Szemao, 5000 ft. elev., shrub 3', fr. red (fr), A. Henry 12088B (holo¬ type, B destroyed; lectotype, here designated, MO; iso¬ lectotype, A). Ardisia tonkinensis A. DC., Repert. Spec. Nov. Regni Veg. 8: 354. 1910. Syn. nov. SYNTYPES: Vietnam. Tonkin: Kirn kibe. in fields of Dong Ham & Dong Bau, H. Bon 2142. 2541. 2912 (G, P); Vo-Xa on Mt. Chuir-Hac, H. Bon 2902 (G, P). Lectotypification of this binomial will be made after further study of all duplicates. Ardisia rectangularis Hyata, J. Coll. Sci. Univ. Tokyo 30: 182. 1911. Syn. nov. SYNTYPES: Japan. Shintiku: Gozhi- zann, June 1906, T. Kawakuimi & U. Mori 1435 (TI); Kusshaku, 1905, S. Nagasawa 358 (TT); Taito: Muri- murisha, Dec. 1899, K. Miyake s.n. (TI). This binomial will be lectotypified when all duplicates of the collec¬ tions cited in the protologue are examined. Ardisia stellifera Pitard, Fl. Indo-Chine 3: 863. 1930. Syn. nov. TYPE: [Vietnam]. Annam: Thua-tien, Thuy-cam, Eberhardt s.n. (holotype, P; fragment. A). Ardisia virens Kurz var. annamensis Pitard, Fl. Indo-Chine 3: 868. 1930. Syn. nov. TYPE: (Vietnam], Annam: Quang- tri: Dent du Tigre, 800 m, Poilane s.n. (holotype, P; fragment, A). Ardisia maculosa Mez var. symplocifolia C. Chen, Act. Phy¬ totax. Sin. 16(3): 79. 1978. Syn. nov. TYPE: China. Yunnan: Hokao, S. Hsuan 610078 (holotype, KUN). Ardisia virens is closely related to A. elegantissima H. Leveille. The separation of the two is based on the presence or absence of papillae on the inflorescence rachises and branchlets. Ardisia virens grows in dense evergreen broad-leaf forests, hillsides, dark damp places, humus-rich soils in China (Guangxi, Guizhou, Hainan, Taiwan, Yunnan), India, Myanmar, Thailand, Vietnam, Kalimantan, Brunei, and Sabah. Myrsine L. As here delimited, Myrsine L. (including Rapanea Aublet) is a pantropical genus of about 300 species, of which 11 grow in China. Recent studies of tropical American taxa (Pipoly, 1991, 1992a, b, c) have shown that many taxa have staminate flowers with anthers on filaments, while the pistillate flowers have the fil¬ aments reduced or barely detectable. Pipoly (1992a) provided a complete rationale for including Rapanea as a synonym of Myrsine. Our treatment of Myrsine to include species formerly placed in Rapanea requires several new combinations. Myrsine cicatricosa (C. Chen) Pipoly & C. Chen, comb. nov. Basionym: Rapanea cicatricosa C. Y. Wu & C. Chen ex C. Chen, Fl. Yunnan. 1: 381. 1977. TYPE: China. Vietnam, Shaba, Huan- glianshan, Sino-Vietnamese Expedition 55 (holo¬ type, KUN). Myrsine verruculosa (C. Chen) Pipoly & C. Chen, comb. nov. Basionym: Rapanea verruculosa C. Y. Wu ex C. Chen, Fl. Yunnan. 1: 380. 1977. TYPE: China. Yunnan: Xichou Xian, 26 Sep. 1947, K. M. Feng 12062 (holotype, KUN). Myrsine faberi (Mez) Pipoly & C. Chen, comb. nov. Basionym: Rapanea faberi Mez in Engler, Pflan¬ zenr. 9(IV. 236): 358. 1902. SYNTYPES: China. Yunnan: Mentze, 2000 m, A. Henry 9173A (B destroyed, W); China. Guangdong: Faber 132 (B destroyed; lectotype, here designated, US; iso¬ lectotype, W). Volume 5, Number 4 1995 Pipoly & Chen Myrsinaceae of China 361 Myrsine kwangsiensis (E. H. Walker) Pipoly & C. Chen, comb. nov. Basionym: Rapanea kwang- siensis E. H. Walker, J. Wash. Acad. Sci. 21: 479. 1931. TYPE: China. Guangxi: Lanlow, E Linyen, Ching 6657 (holotype, US; isotype, A). Rapanea kwangsiensis E. H. Walker var. lanceolata C. Y. Wu & C. Chen, FI. Yunnan. 1: 383. 1977. Syn. nov. TYPE: China. Yunnan: Jinping Xian, 8 Jan. 1933, H. T. Tsai 52550 (holotype, KUN). Acknowledgments. Our studies for the Flora of China have been greatly facilitated by the Missouri Botanical Garden. We thank Ihsan Al-Shehbaz, An¬ thony Brach, and Michael Gilbert for assistance. We also thank the curators and staff of the Harvard Uni¬ versity Herbaria, particularly Emily Wood and David Boufford, and those of the Smithsonian Institution, particularly George Russell, Vicki Funk, and Larry Skog, for hospitality and help during our visits to those institutions. Literature Cited Pipoly, J. 1983. Contributions toward a monograph of Cy- bianthus (Myrsinaceae): III. A revision of subgenus Laxi- ftorus. Brittonia 35: 61-80. -. 1987. A systematic revision of the genus Cybian- thus subgenus Grammadenia (Myrsinaceae). Mem. New York Bot. Gard. 43: 1—76. -. 1991. Systematic studies in the genus Myrsine L. (Myrsinaceae) in Guayana. Novon 1: 204—210. -. 1992a. A further note on Myrsine perpauciflora Pi¬ poly (Myrsinaceae). Novon 2: 176. -. 1992b. Estudios en el g&iero Myrsine (Myrsina¬ ceae) de Colombia. Caldasia 17(1): 3—9. -. 1992c. Notes on the genus Myrsine (Myrsinaceae) in Peru. Novon 2: 392-^107. Prance, G. 1972. Chrysobalanaceae. Flora Neotropica 9: 1-410. -. 1982. Forest refuges: Evidence from woody an- giosperms. Pp. 137-156 in G. Prance (editor), Biolog¬ ical Diversification in the Tropics. Columbia Univ. Press, New York. Walker, E. H. 1940. A revision of the Eastern Asiatic Myrsinaceae. Philipp. J. Sci. 73: 1-258. White, F. 1962. Geographic variation and speciation in Africa with particular reference to Diospyros. Pp. 71- 103 in D. Nichols (editor). Taxonomy and Geography. The Systematics Association Publ. No. 4, London. New Andean Zanthoxylum (Rutaceae) with Distinctive Vegetative Characters Carlos Reynel Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. ABSTRACT. Three new Andean species of Zan¬ thoxylum (= Fagara), Z. gentryi Reynel, Z. lenti- culare Reynel, and Z. lepidopteriphilum Reynel, are described and illustrated. Vegetative characters of taxonomic and practical identification value are briefly discussed. Some of these features relate to insect—plant interactions. RESUMEN. Tres nuevas especies de Zanthoxylum (= Fagara ), Z. gentryi Reynel, Z. lenticulare Reynel y Z. lepidopteriphilum Reynel, son descritas e ilustra- das. Se discuten brevemente caracteres vegetativos de utilidad taxonomica y prdctica en la ideritificaeion de especies en el genera. Algunos de estos caracteres tienen relacidn con interacciones insecto-planta. The mainly pantropical and subtropical genus Zanthoxylum L. s.l. comprises about 200 woody species, mostly trees. Engler (1874, 1896, 1931) regarded Zanthoxylum s. str. as separate from Fa¬ gara L. and proposed a sectional and subsectional scheme for the latter. Waterman (1975a, 1976) combined the two genera and transferred the spe¬ cies from Fagara to Zanthoxylum s.l.; evidence has accumulated in recent years supporting that unifi¬ cation, on the grounds of floral morphology (Bri- zicky, 1962; Hartley, 1966; Beurton, 1994), floral anatomy (Yamazaki, 1988), wood anatomy (Reynel & Miller, in prep.), and phytochemistry (summa¬ rized in Waterman, 1975b, 1977, 1983). In this paper I describe three new Andean species of Zanthoxylum that have distinctive vegetative fea¬ tures: interspaced, short zones of clustered leaf and bud-scale scars (Z. gentryi. Fig. 1A, 2B); hollow myrmecophilous apical branchlets (7. lenticulare, Fig. IB); and twice-revolute basal leaflet margins (Z. lepidopteriphilum, Fig. 2A, 2C). These new species belong to the group of New World species with 5- merous, differentiated perianths placed in Fagara subsection Paniculatae-Neogaeae by Engler (1896). Zanthoxylum gentryi Reynel, sp. nov. TYPE: Co¬ lombia. Valle: Municipio Restrepo, Vereda de Riogrande, abajo del puente, en predios de la familia Reyna, en la via Pavas-Restrepo, 1200 m, 10 Feb. 1990 (fr), J. Ramos & C. Reyna 2551 (holotype, MO; isotype, CUVC). Figure 1A, 2B. Novon 5: 362-367. 1995. Haec species Z. petiolari A. St.-Hilaire & Tulasne af- finis, sed ab eo foliis 15—45 cm longis, interjugis 3.5—9 cm longis, foliolis lateralibus ovatis vel oblongis, sessili- bus, (5-)7—17 cm longis, marginibus integris, panicula ampla, 7-20 X 6—16 cm (infructescentia usque ad 25 X 35 cm); petalis 2.6-3.5 mm longis, coccis 4—7 mm longis et seminibus 4—6 mm longis differt. Dioecious treelets or trees to 10 m tall; apical branchlets 5-10 mm thick, with interspaced, short zones of clustered leaf and bud-scale scars, also with sparse, straight to incurved, compressed-con¬ ical spines to 6 mm long and sparse lenticels to 1 mm long, the branchlets covered with occasional white simple hairs to 0.1 mm long, the periderm usually peeling when dry. Leaves paripinnate or im- paripinnate, 4—7(-9)-foliolate, 15-45 cm long, the terminal leaflet in imparipinnate leaves promptly deciduous; petiole 4—13 cm long; rachis terete, 2- 3 mm thick, glabrescent, the interjuga 3.5-9 cm long, sometimes with an abaxial spine to 5 mm long; leaflets chartaceous, rigid, usually drying ol¬ ive green and shiny on both sides, laterals opposite, sessile, terminal leaflet with petiolule 0—3 cm long, ultimate pair usually larger than rest, all leaflets ovate to oblong, (5—)7—17 X 3.5—9.5 cm, the apex acuminate, with acumen to 10 X 6 mm, base acute to obtuse or truncate; margin entire or subentire; leaflets glabrescent; midvein adaxially prominu- lous, secondary veins 10—16 pairs, prominulous on both sides, tertiary veins subreticulate. Inflores¬ cences terminal or subterminal, multiflorous pani¬ cles 7-20 X 6-16 cm (-25 X 35 cm in fruit), branched 3-4 times; main axis 2—3 mm thick (-4 mm in fruit), secondary axes to 3—5 cm long (—20 cm in fruit), mostly forming a 60°-90° angle with main axis, the inflorescence axes covered with white, sparse simple hairs to 0.1 mm long; inflo¬ rescence bracts ovate to deltoid, 0.6-1 (-3) X 0.7— 1.2(—2) mm, glabrescent. Flowers 5-merous; floral bracts 1-3 at base of pedicel, deltoid to ovate, 0.6- 1 X 0.5-1 mm, covered with indument similar to inflorescence axes or glabrescent; pedicel 0-1 X 1.2—2 mm (-4 mm long in fruit); calyx 0.5—1.4 X 1—3 mm, sepals free or connate to Vi their total length, ovate to deltoid, 0.8-1.2 mm broad, acute, glabrescent; petals saccate, ovate, 2.6-3.5 X 1.2— Volume 5, Number 4 1995 Reynel New Andean Zanthoxylum 363 Figure 1. -A. Zanthoxylum gentryi Reynel, apical branchlet with leaf and inflorescence (Ramos 2334). -B. Zan¬ thoxylum lenticulare Reynel, apical branchlet with leaf and infructescence (Soeyarto et al. 4058). 1 .4 mm, the apex rounded to acute, strongly in- flexed, usually with an ellipsoid resin gland to 0.2 mm diam., the petals glabrous. Staminate flowers with 5 stamens, exserted Vy-Vi their total length; filaments slender, 2.8-4 mm long, anthers 1.2—1.8 mm long, thecae oblong, the connective with an apical ellipsoid gland 0.2-0.3 mm long; nectary discoid to conical-truncate, 5-lobate, 0.4—0.5 X 1- 1.3 mm; pistillodes 1-3, distinct to totally connate, elongate-pyriform, 0.7—1.5 X 0.2—0.5 mm. Pistil- 364 Novon late flowers with 0—5 staminodes, membranous, ovate to oblong; gynophore conical, 1.2—1.8 X 0.7— 1.6 mm; carpels 3—5, free except in the common stigma, ovaries obovoid, 0.5—0.8 X 1—1.5 mm, styles attached at adaxial upper half of ovaries, to 0.2 X 0.2 mm, stigma discoid, 3—5-lobed, 1.8-2.1 mm diam., 0.2-0.4 mm thick. Fruit apocarpous, (2—)3—4(—5)-follicular, frequently 1—3 follicles aborting; follicles subovoid, slightly laterally com¬ pressed, 4—7 mm long, glabrescent; seed ellipsoid to subreniform, 4-6 X 2.5—5 mm, the funicular scar rounded, 1.5-2 X 0.8-1.2 mm. Distribution. Endemic to the Department of Valle in Colombia, at 600-1100 m, in wet primary or secondary forests. Within neotropical species of Zanthoxylum with differentiated perianths, the presence of inter¬ spaced, short zones of clustered leaf and bud-scale scars on apical branchlets (Fig. 2B) is character¬ istic of a small group: Z. petiolare A. St. Hilaire & Tulasne (syn. Z. naranjillo Grisebach), a common species from southern Brazil, Bolivia, Argentina, and Paraguay; Z. pentandrum (Aublet) R. A. How¬ ard (syn. Z. hermaphroditum Willdenow) from northern Brazil, Venezuela, and the Guianas; Z. verrucosum (Cuatrecasas) P. G. Waterman from Co¬ lombia; and Z. pucro from Panama. Zanthoxylum, gentryi can be distinguished from the species of the Zanthoxylum petiolare group by the following key: Key to Species in the Zanthoxylum petiolare Group la. Leaves with lateral leaflets sessile. Z. gentryi lb. Leaves with lateral leaflets petiolulate, petiolules 2—10 mm long. 2a. Interjuga 5-12 cm long; leaflets 12-22 cm long. 3a. Leaves imparipinnate, with ovate leaf¬ lets; inflorescence axes glabrous; pistil¬ late flowers with petals 3-3.5 mm long; follicles 5-6 mm long .... Z. verrucosum 3b. Leaves paripinnate, with oblong leaflets; inflorescence axes densely and finely pubescent; pistillate flowers with petals 1.7-2.4 mm long; follicles 6-9 mm long. Z. pucro 2b. Interjuga 1.5-5 cm long; leaflets 3—12(—16) cm long. 4a. Petals 1.3—1.7 mm long; anthers 0.4— 0.6 mm long. Z. pentandrum 4b. Petals 1.9—2.2 mm long; anthers 0.9- 1.1 mm long.Z. petiolare The new species is named after Alwyn H. Gentry (1947—1993), botanist and ecologist, whose work and knowledge of the Andean-Amazonian flora and its diversity inspired a generation of tropical biologists. Paratypes. COLOMBIA. Valle: Municipio Toro, que- brada San Lazaro (fr), Devia 328 (MO, T1JLV); near Yo- toco, N of Cali, 3°45'N, 76°30'W, Gentry et al. 54073 (COL, MO); El Silencio, Hacienda Himalaya, W of Yum- bo, 3°50'N, 76°40'W (fr). Gentry et al. 65516 (COL, MO); Finca El Espinal, 25 km N of Cali (fr), Hughes 129 (FHO, MO); Dagua Valley, 15.5 km below Dagua and 1.5 km below Lobo Guerrero, quebrada de la Chapa, upper end El Boquerfin of rfo Dagua (fl), Hutchinson et al. 3102 (COL, F, G, K, MO, NY, UC, US); Municipio Restrepo, Ramos 2334 ( 5), 2281 (d) (MO, CUVC); Municipio El Cerrito, pueblo de Guabas, cerca de la frontera con Mun¬ icipio Guacari, Silverstone-Sopkin et al. 2294, 3662 (fr) (CUVC, MO). Zanthoxylum lenticulare Reynel, sp. nov. TYPE: Colombia. Antioquia: Municipio de Anorf, Corregimiento de Providencia, entre Dos Bo- cas y Anorf, Buenos Aires, 4 km from Provi¬ dencia, 400-900 m, aprox. 7°N, 75°W, May 1973 (fr), Soejarto et al. 4058 (holotype, COL; isotypes, F, GH, MO). Figure IB. Haec species Z. compacto (Huber ex Albuquerque) P. G. Waterman affinis, sed ab eo foliis 40-70 cm longis 13- 21-foliolatis, foliolis 8-12 cm longis, subcaudatis, margi- nibus minute crenulatis; paniculis amplissimis congestis 15-35 cm longis, 3( -4)-ramificatis, axibus primariis non fistulosis, axibus secundariis 4—13 cm longis; petalis 1.5- 1.9 mm longis; coccis discoideis laevibus 3-4 mm longis, seminibus 2.5—3 mm longis differt. Dioecious, myrmecophilous treelet or tree to 12 m tall; apical branchlets hollowed, 4—8 mm thick, with sparse, straight, compressed-conical spines to 6 mm long and lenticels to 0.5 mm long, the branchlets cov¬ ered with sparse to dense, white, simple hairs to 0.1 mm long or glabrescent. Leaves evenly distributed on branchlets, imparipinnate, (11—)13—21(—25)-foliolate, 40-70 cm long; petiole (5.5—)7-10(—15) cm long; ra- chis terete, narrowly and deeply canaliculate, 2-4 mm thick, interjuga (2.5-)3-4.5 cm long, sometimes with 1(2) abaxial spines to 5 mm long, the rachis covered with indument similar to the branchlets; leaflets char- taceous to submembranaceous, drying pale green to reddish brown above, pale green beneath, laterals op¬ posite to alternate, sessile, terminal leaflet with peti- olule 2—10 mm long, ultimate pair or terminal leaflet usually larger than rest, all leaflets oblong to subovate, (2.5—)8—12(—15) X 2-4.5 cm, the apex acuminate to caudate, the acumen to 15 X 5 mm, base acute; mar¬ gin crenate, the teeth separated 1—1.5 mm at the mid¬ dle of leaflets; leaflets without hairs; midvein adaxially prominulous; few rounded glands 0.2-0.3 mm diam. usually present on abaxial blade near midvein; sec¬ ondary veins 16-22 pairs, prominulous on both sides, tertiary veins usually prominulous and subreticulate on both sides. Inflorescences terminal or subterminal, multiflorous, congested panicles (8-)15-35 X (8-)15- 22 cm, branched 3(-4) times; main axis 2-4 mm thick, secondary axes 4—13 cm long, mostly forming Volume 5, Number 4 1995 Reynel New Andean Zanthoxylum 365 a 60°-90° angle with main axis, the inflorescence axes covered with sparse to dense, white, simple hairs to 0.05(-0.1) mm long or glabrescent; inflorescence bracts deltoid to ovate, 0.2-0.5(-l) X 0.3-0.5 mm, covered with indument similar to inflorescence axes, usually ciliate. Flowers 5-merous; floral bracts (1)2—5 at base of pedicel, ovate to lanceolate, 2-3 X 1 mm, covered with indument similar to inflorescence axes or glabrescent; pedicel terete, 0.2-0.7 X 0.1-0.2 mm (—2.5 X 0.5 mm in fruit), glabrescent; calyx 0.5-0.7 X 0.&-1 mm, sepals connate Vi their total length, ovate, 0.3-0.4 mm broad, rounded to acute, glabres¬ cent; petals saccate, ovate, 1.5—1.9 X 0.8-1.1 mm, the apex rounded, usually apiculate with apiculum to 0.2 mm long, also with an ellipsoid gland to 0.2 mm diam., the petals glabrous. Slaminate flowers with sta¬ mens 5, exserted Vs to Vi their length, the filaments slender, 2—3 mm long, anthers 0.8-1 mm long, thecae oblong; nectary conical-truncate, 5-lobate, 0.3-0.4 X 0.3-0.6 mm; pistillodes 1-3, elongate-pyriform, dis¬ tinct to totally connate, 0.2-0.5 X 0.2 mm. Pistillate flowers with no staminodes; gynophore conical, 0.2- 0.3 X 0.4—0.6 mm; carpels (2)3, distinct except in the common stigma, ovaries oblongoid, 0.4—0.6 X 0.4—0.6 mm; styles terete, attached at adaxial upper half of ovaries, 0.1-0.2 X 0.1-0.2 mm; stigma dis¬ coid. Fruit apocarpous, (2-)3-follicular, frequently 1- 2 follicles aborting; follicles discoid, 3-4 X 3 mm, glabrous; seed subdiscoid, 2.5-3 X 0.6-0.9 mm, the funicular scar linear, 1—2 X 0.3-0.4 mm. Common name: “Sando.” Distribution. Colombia (Antioquia) and Vene¬ zuela (TAchira and Zulia), between 90 and 900 m, in wet primary forests. Three other New World species of Zanthoxylum also have hollow, myrmecophilous apical branch- lets, which are very consistent and of practical val¬ ue for identification. The following key allows the comparison of related species. Key to Myrmecophilous Neotropical Species of Zan- thoxylum la. Apical branchlets with enlarged, interspaced zones of clustered leaf scars; panicles clustered at the apex of branchlets . . Zanthoxylum setulosum lb. Apical branchlets uniformly terete; panicles sol¬ itary, terminal. 2a. leaves velutinous-pubescent; leaflets 5-12 cm broad. Zanthoxylum formiciferum 2b. leaves glabrous or glabrescent; leaflets 1—5 cm broad. 3a. Panicles compact, 3—10 cm broad; fruit with follicles ovoid to obovoid, 6-9 mm long. Zanthoxylum compactum 3a. Panicles expanded, 15-22 cm broad; fruit with follicles discoid, 4—4.5 mm long. Zanthoxylum lenticulare Paratypes. COLOMBIA. Antioquia: Municipio San Luis, carton del rfo Claro (3), Cogollo 831 (COL, MO); Municipio lani/a. Rfo Cauca, right margin on road from El Doce to Barro Blanco, 8°35'N, 75°25'W (fr), Daly et al. 5253 (COL, F, NY); Rio Claro, camino al Refugio (9, fr), Renteria et al. 2855 (COL, MO); Municipio Anori, Rio An- ori valley, near Planta Providencia, 7°30'N, 74°50'W, Shep¬ herd 374 (COL, MO, WIS), 6 Aug. 1977, 913 (COL, MO, WIS); Corregimiento de providencia, entre Dos Bocas y An¬ ori, Buenos Aires, ca. 7°N, 75°W (3), Soejarto 3922 (COL, F(2), MO). VENEZUELA. Carabobo: vicinity of Las Trincheras, near Valencia, H. Pittier 7655 (GH). Tachira: ca. 10.5 km NE of La Fria, 90 m, 8°16'N, 72‘TO'W, 16 Nov. 1979 (3), J. Steyermark et al. 120511 (MO, NY, VEN). Zulia: Distrito of Coldn, aldea Querrequerre, 12—15 km N of Casigua (fr). Bunting et al. 7427 (NY, VEN). Zanthoxylum lepidopteriphilum Reynel, sp. nov. TYPE: Peru. Piura: Prov. Huancabamba, Canc- haque, between “Chorro Blanco” and “War- war,” 2000-2500 m, 18 Jan. 1989 (fr), C. Diaz, T. D. Pennington & C. Reynel 3192 (holotype, MO; isotypes, K, MOL, USM). Figure 2A, C. Haec species Z. mantaro (J. F. Macbride) J. F. Mac- bride affinis, sed ab eo foliis 32 45 cm longis, foliolis 9— 17(—25), 7— 10(—13) cm longis, basibus abaxialibus bis re- volutis cylindrum marginalem formantibus 8-10(—14) mm longis; panicula ampla 9—14 cm longa (infructescentia usque ad 20 cm longa); petalis 2—3 mm longis; coccis 5- 8(-10) mm longis, seminibus 5-6 mm longis differt. Dioecious trees to 20 m tall; apical branchlets terete, 7-10 mm thick, unarmed, covered with sparse to dense, white or yellow, stellate, bifid and simple hairs 0.2-0.6 mm long. Leaves evenly dis¬ tributed on branchlets, imparipinnate, 32—45 cm long, 9-17(-25)-foliolate; petiole (3.5-)4—10 cm long; rachis terete, obscurely canaliculate, 2-4 mm thick, interjuga 3-5 cm long, the rachis covered with indument similar to branchlets or glabrescent; peti- olules 3-5 mm long; leaflets chartaceous, drying ol¬ ive green to reddish brown on both sides, laterals opposite, ultimate or medial pair larger than rest, all leaflets oblong, 7—10(—13) X 2.5—3(—3.5) cm, the apex emarginate, rounded or acuminate, with acu¬ men to 6 X 4 mm, base acute, oblique in laterals, tubular-twice-revolute 8-10(-14) mm or more along margin, each side of the midvein; margin entire to slightly crenate, with teeth separated 2-3 mm at the middle of leaflets; leaflets covered with sparse to dense, white to yellow, stellate, bifid and simple hairs to 0.2-0.5 mm especially along midvein and abaxial base, or glabrescent; midvein adaxially nar¬ rowly and deeply impressed; secondary veins 12-16 pairs, prominulous on both sides, tertiary veins usu¬ ally prominulous and subreticulate on both sides. Inflorescences terminal or subterminal panicles, branched 1-2 times, pauciflorous to submultiflorous, 9-14 X 2.S-5.5 cm (-20 X 20 cm in fruit); main 366 Novon 3 cm -B r zLwfrtt7 /U R R ^ el - a P' ca ^ branchlet with leaf and infructescence (Diaz et al. 3192). (sLfs“^SoaZ!tai^2^i n ^r aP 7 ! )ra,, ; et , Sh r ng short zones of clus '™ d «" .<0 4 L? }q |( |i IV CO 7Cfl ?? ij 7 { >( 3! H ?c ? I If f f •» < )\ « *'» Figure 2. Karyotype of Kengyilia guidenensis Yen, J. L. Yang & Baum. First row illustrates the P haplome; second and third rows illustrate the S and Y haplomes mixed. Literature Cited Baum, B. R., C. Yen & J. L. Yang. 1990. Kengyilia ha- bahenensis (Poaceae: Triticeae)—A new species from the Altai mountains, China. PI. Syst. Evol. 174: 103- 108. Yang, J. L., C. Yen & B. R. Baum. 1992. Three new species of the genus Kengyilia (Poaceae: Triticeae) from West China and new combinations of related species. Canad. J. Bot. 71: 339-345. Yen, C. & J. L. Yang. 1990. Kengyilia gobicola, a new taxon from West China. Canad. J. Bot. 68: 1894—1897. Volume 5, Number 1, pp. 1-118 of NOVON was published on 27 March 1995. Volume 5, Number 2, pp. 119-208 of NOVON was published on 21 June 1995. Volume 5, Number 3, pp. 209-304 of NOVON was published on 05 September 1995. Volume 5, Number 4, pp. 305-398 of NOVON was published on 15 December 1995.