Volume 14 Number 1 2004 NOVON Suaeda neuquenensis (Chenopodiaceae), a New Species from Argentina M. A. Alonso GIB (Institute Universitario de la Biodiversidad), Universidad de Alicante, Apartado 99, E-03080 Alicante, Spain, ma.alonso@ua.es L. Conticello and M. B. Cerazo Departamento de Botanica Aplicada, Universidad Nacional del Comahue, Rio Negro, Argentina Abstract. Suaeda neuquenensis , a new species of Suaeda (Chenopodiaceae), is described from Ane- lo’s Bowl near Anelo, Neuquen Province, Argenti¬ na. It differs from other perennial Argentinean spe¬ cies in being a decumbent shrub with creeping stems that root at the lower nodes bearing short, oblong, often reflexed leaves and a large seed. Key words: Argentina, Chenopodiaceae, Neu¬ quen, Suaeda. Two perennial species of Suaeda Forsskal ex J. F. Gmelin (Chenopodiaceae), S. divaricata Moquin- Tandon and .S', argentinensis Soriano, are currently known from Argentina (Soriano, 1942; Giusti, 1997). Suaeda fruticosa (L.) Forsskal was reported from the Boca district of Buenos Aires by Soriano (1945), who nonetheless suggested it was a recent introduction and not likely to persist; there is now no evidence that it did. Species of the genus are highly variable as to succulence, habit, and size of leaves (Morello, 1958; Hopkins & Blackwell, 1977; Bassett & Crompton, 1977; Pedrol & Castroviejo, 1988). Other features are more constant: herba¬ ceous versus woody habit, type of inflorescence (in¬ cluding the density of the glomerules), relationship of leaf length and internode length, shape of the calyx, stigma shape and length, and seed size (Hop¬ kins & Blackwell, 1977; Pedrol & Castroviejo, 1988). Material and Methods Dried specimens of Suaeda at ARC, CORD, and MERI , were studied, as well as fresh material col¬ lected in the Neuquen Province. Taxonomic and nomenclatural treatments follow Standley (1916), Morello (1958), Bassett and Crompton (1977), Hopkins and Blackwell (1977), Correa (1984), Pedrol and Castroviejo (1988), Teill- er (1996), Guisti (1997), Greuter el al. (2000), and Schenk and Ferren (2000, 2001). Herbarium sheets of Suaeda neuquenensis are conserved at ABH of the University of Alicante (Holmgren & Holmgren, 1993). Results and Discussion Suaeda neuquenensis M. A. Alonso, Conticello & Cerazo, sp. nov. TYPE: Argentina. Neuquen Province: Anelo’s Bowl near Anelo, 38°11'56"S, 69°00'46”W, 350 m.s.m. on sa- line-sodic soil, 24 Mar. 1998, L. Conticello & M. A. Alonso s.n. (holotype, ABH 41516; iso¬ type, ABH 42972). Figure 1. Nanophanerophitus in fructicibus pulvinatis usque ad 50 cm altitudinis et 2.50 m diametri se extendentihus. Caulis atramineus leviler strialus, glaber cum internodiis 1 mm, interdum inaestimabilis in gemmis recentibus, 2- 3 mm longis in ramis adultis. Folia adulta patentia aut reflexa, iuvenilia perpendicularia, alterna oblonga, sue- culenta cum apiee obtuso ac leviter mucronato, parum aut Novon 14: 1-5. 2004. 2 Novon Figure 1. Suaeda neuquenensis M. A. Alonso, Conticello & Cerazo. — A. Habit (holotype, ABH 41516). — B, C. Flowering branches. — I). Young sterile shoot. — E. Detail of an axillary floral glomerule from the holotype. nihil ad basim attenuata, dorsi ventral iter compressa, mar- gines integri et rotundi, cum nitore virentia ant glauea vel albescentia propter salina eorpuscula (|uae in superficie deponuntur, 5—15 mm longa et 3—8 mm lata. Gynodioe- cius. sessiles llores iuncti in glomerulis axillaribus 2-6, hermaphroditi cum sterili gynaecio, 5 staminibus et calyx 5 sepalorum. Flores feminei cum gynaecio fecundo unil- oculari, stylis 3 ad 0.5 mm longis, stigmatibus papillis elongatis atque pilis acutis vestitis. Flores isti minores sunt quam hermaphroditi, sed dia parte eius, et cum apice Volume 14, Number 1 2004 Alonso et al. Suaeda neuquenensis from Argentina 3 abtuso glabris ac inflexis, 2—3 mm longi. Fructus utricu- lalus, ovoideus, 3X4 mm, in calycis segmentis inclusus sed illis non affixus. Semen vertieale, nigrum, lenticulare, testa reticulata. Plants woody, cushion-shaped, dwarf shrubs with creeping stems, forming mats 20—50 cm high and 25 dm across, the decumbent branches rooting at the nodes. Stems light brown, weakly striated, the internodes of new branches 1 mm long or less, 2— 3 mm long in mature branches. Leaves alternate and positioned perpendicular on new branches, be¬ coming reflexed on mature branches, succulent, ob¬ long, 5—15 mm long, 3—8 mm wide, entire, shining green to glaucous or white due to saline deposited on the surfaces, the apices obtuse to rounded or slightly mucronate, the bases attenuated. Flowers gynodioecious, axillary, sessile, 2 to 6 per glomer- ule, the perianth composed of 5 sepaloid perianth segments, 2—3 mm long, these inflated in the mid¬ dle, apically obtuse and indexed, the hermaphro¬ dite flower with 5 stamens and a sessile, sterile gynoeeium, the female flowers with a solitary, 1- loeular gynoeeium with 3 stigmas up to 0.5 mm long, bearing elongate papillae and acute hairs. Fruit an ovoid utricle, 3 X 4 mm, enclosed by the perianth segments, which are not closely attached to it; seeds vertical, black, lens-shaped, reticulate on the testa. Ecology. Suaeda neuquenensis is known only from Anelo’s Bowl near Anelo in the province of Neuquen, where it occurs on saline-sodic soils in monospecific communities or in association with Atriplex undulata (Moquin) Dietrich, Distichlis spi- cala (L.) Greene. Halophytum ameghinoi Spegaz- zini, Sesuvium portulacastrum (I,.) L., and Suaeda divaricata. Taxonomic Discussion In a recent paper on the nomenclature of Suae¬ da, Schrenk and Ferren (2001) stressed features characterizing sections in the genus. According to their arrangement, Suaeda neuquenensis belongs to section Limbogermen lljin (sect. Salsina Moquin, pro parte), which includes American shrubs and subshrubs, with radial flowers, stigmas from a pit at the top of the ovary, and leaves with suaedoid C4 anatomy, consisting of a ring of dark green palisade and Kranz cells surrounding internal aqueous tis¬ sue, among other characters. Relationships among taxa of section Limboger¬ men are shown in Table 1. Morphologically, S. neu- quenensis shows a syndrome of characters that is not found in other South American taxa in the sec¬ tion. Stems are decumbent and root at the basal nodes, a feature that is absent in both S. argenti- nensis and S. divaricata ; seeds are vertical and have a reticulate testa (neither horizontal and smooth as in S. argentinensis, nor tuberculate as in S. divaricata). With regard to other American spe¬ cies, S. neuquenensis resembles ,S. conferla (J. K. Small) 1. M. Johnston by its delicate cushion¬ shaped habit, though it differs by its robust, strong¬ ly woody stems, large fruits up to 4 X 3 mm (the largest in the section), and its narrower and shorter leaves, among other features. Connections to other Central and North American taxa are very weak. It is unlikely that Suaeda neuquenensis is an ex¬ otic, introduced taxon. Its natural habitat is quite far from populated or frequented areas, and the soil in which it grows has a high salt content, making it unsuitable for agriculture. It is difficult therefore to support an alien origin for this plant, since no taxon is currently known to which it could be re¬ ferred. Key to Related Species ok Suaeda in the Argentine Republic The following is part of Giusti's (1997) kev for the genus Suaeda, to which S. neuquenensis has been added. la. Shrubs or small shrubs. 2a. Plants with ramification in acute angles. Leaves appressed to stem. Flowers in axil¬ lary, pedicellate glomerules. Seeds usually horizontal . Suaeda argentinensis 2b. Plants with ramification in divergent angles. Leaves divergent, not appressed to stem. Flowers in axillary, sessile glomerules. Seeds vertical. 3a. Shrubs erect, 1—3 m high. Leaves sem- icylindric, perpendicular to the stem. Young stems downy. Stems brown to reddish . Suaeda divaricata 3b. Sbrubs decumbent, 0.2— 0.5 m high. Leaves flat, reflexed on the stem. Young stems glabrous. Stems light brown . . . . Suaeda neuquenensis lb. Annual herbs . . Suaeda patagoniea, Suaeda altisima, and Suaeda densiflora Acknowledgments. We thank Fidel Roig and Ma¬ nuel B. Crespo for comments on the manuscript, Ar- eadio Alonso for his assistance with the Latin descrip¬ tion, and Pedro Marco for the illustration of this species. Literature Cited Bassett, I. J. & C. W. Crompton. 1977. lhe genus Suaeda (Chenopodiaceae) in Canada. Canad. .1. Bot. 56: 581—591. Correa, M. N. 1984. Flora Patagoniea. Parte I\ a. Tomo \ 111. Coleccion Cientffica del INTA. Buenos Aires. Ferren. W. R. & H. J. Schenk. 2000. Flora of North America. (http://hua.huh.harvard.edu/FNA/sueda 1 1 .16.00.html) Gehu, J. M. & S. Rivas-Martfnez. 1982. Notions fondamen- Table 1 . Comparison of morphological features among the species of Suaeda sect. Limbogt 4 Novon 15 CA) 3 — $1 CM ^ 53 cd .Jc 3 u O r/T -§ g bjj £ Southeast Asia (2 spp.) INovoin 14: 12-16. 2004. Volume 14, Number 1 2004 Arbelaez Meliosma from Bolivia 13 Meliosma subg. Meliosma sect. Meliosma — > Southeast Asia (12 spp.) sect. Lorenzanea (Liebmann) Beusekom — > Neo¬ tropics (ca. 50 spp.) Fifteen species were recognized by Van Beuse¬ kom (1971) in the revision of the Southeast Asian sections of Meliosma. Section Lorenzanea is re¬ stricted to the Neotropics and has never been monographed. All but one of the New World spe¬ cies, M. alba (Schlechtendal) Walpers of Mexico, belong to subgenus Kingsboroughia. Meliosma alba also occurs in south-central China, and it is the only New World species of the genus with com¬ pound leaves. A few revisions of the genus in some countries exist, e.g.. Flora lalis, petiolis 4—5 mm longis et bracteolis pilis 1.5-2 mm longis. 8— It) seplatis obtectis differt. Shrub or small tree to 6 m tall; branches and 5 cm 5 mm Figure 2. Aegiphila hystrieina Aymard & Cuello. — A. Flowering branch with detail of lower leaf surface. — B. Flower subtended by the three bracteoles covered by the longer triehomes, ovary, pistil, and stigma represented by dotted lines. — C. Medial section of flower showing the stamens with pistil removed. Drawn from the holotype, Koyama & Agostini 7254 (VEN). Volume 14, Number 1 2004 Aymard & Cuello New Species of Aegiphila 23 Table 2. Comparison of the distinguishing character states of Aegiphila hystricina and A. arcta. Characters Aegiphila hystricina Aegiphila arcta Branches, branchlets, and peduncles I, eaves Petiole Bracteole vestiture Fruit yellow-gray pubescent elliptic or ovate-elliptic, 8—13 cm long, 3—6 cm wide, glandulose vil¬ lose on both surfaces, glandulose triehomes, ca. 2 mm long, 8— 9-sep- tatc, margin densely ciliate 4— 5 mm triehomes 1.5—2 mm long, 8— 10-sep- tate 5— 6 mm long, sparsely glandular-pu¬ bescent brown pubescent oblanceolate or obovate, 10—18 cm long, 5—8 cm wide, sparsely glandu lose villose on both surfaces, glan¬ dulose triehomes, ca. 0.4 mm long, 2— 4-septate, margin sparsely ciliate 0.5—1 mm triehomes ca. 5 mm long, 3— 4-septate ca. 10 mm long, glabrescent branchlets obtusely tetragonal, densely appressed- villous w ith yellow-gray hairs on the younger parts, glabrescent when mature. Leaves membranous, el¬ liptic or ovate-elliptic, 8—13 X 3—6 cm, apex acu¬ minate, base acute, margin entire, densely ciliate, glandular villose on adaxial and abaxial surfaces, denser on the midrib and secondary veins, glan¬ dular triehomes ca. 2 mm long, 8-9-septate; sec¬ ondary veins 8 to 10 per side, elevated on the ab¬ axial surface; petioles 4—5 mm long, densely appressed-villous with yellow-gray hairs. Inflores¬ cence terminal thyrses, pendent, very congested, peduncle elongate, 8—1 1 cm long, rachis and ped¬ icels densely appressed-villous with yellow-gray hairs. Flowers subtended by three bracteoles at the base of the pedicels, bracteoles linear, 5—10 mm long, densely covered by glandular triehomes, 1 .5— 2 mm long, 8— 10-septate, pedicel 2—3 mm long; calyx 1—1.5 mm long, sparsely glandular pilose ex¬ ternally, glabrous internally, the lobes 4, ca. 0.2 mm long, triangular-ovate, ciliate-glandular, corolla tube 6—10 mm long, yellow-green, glabrous on both sides, the lobes 4, 2—4 mm long, imbricate, shortly glandular-ciliate at the margins; the stamens 4, in¬ serted ca. 5 mm from the base of the tube, exserted, filaments 8—10 mm long, glabrous, anthers oblong, ca. 1 mm long; ovary obconical, ca. I mm long, glabrous, style ca. 4 mm long, inserted, glabrous, bifid, stigmas 2. Fruit globose, 5—6 mm long, ap- pressed glandular-pubescent, mature calyx mem¬ branous and persistent, 3—4 mm long, sparsely glandular-pubescent, subtended by the three per¬ sistent bracteoles. Distribution and habitat. Aegiphila hystricina is known from savanna and forest ecotone areas in Ve¬ nezuela anil farther southeast in the Rio Araguari, Amapa state, in Brazil. In both regions, the species occurs at elevations between 100 and 450 m. Local names. “Hueraya dad eo” (Piaroa), from ./. Con t reras 117 ( V E N) . Because of its four stamens and four corolla lobes, terminal inflorescence, and calyx not deeply dentate, Aegiphila hystricina belongs to section Ae- giphila, group Paniculatae, subgroup Edentatae (Moldenke, 1934; Ldpez-Palacios, 1977). flic inflo¬ rescence very congested, and the linear bracteoles subtending the flowers, covered by glandulose tri- chomes at the base of the pedicels, places this new species next to A. arcta Moldenke, from Yaracuy state in the Coastal Cordillera in Venezuela (Mol¬ denke, 1972). Aegiphila hystricina can be distin¬ guished from A. arcta by the characters indicated in Table 2. Herbarium specimens of Aegiphila hystricina were previously referred to A. fendleri Moldenke, a species from the mountain forests in the Andes of Colombia and Venezuela. However., the new species differs from A. fendleri in its leaves elliptic or ovate-elliptic and the base acute (vs. lanceolate- oblong to oblong and the base rounded or subcor- date); petiole 4—5 mm long (vs. 7—12 mm long); branches, branchlets, petioles, leaves on the lower surface, and calyces covered by a yellow-gray in- dument (vs. all parts of the plant covered by a fer¬ ruginous indument); and inflorescences thyrses (vs. cymes). Paratypes. VENEZUELA. Amazonas: Atures, alto no Carinagua, 05°45'N, 67°20’W, ./. Contreras 117 (VEN). Bolivar: Cedeno, Maniapure, Hato Pisurca, 06°55'N, 66°32'W, A. Fernandez 16938 (PORT). BRAZIL. Amapa: Rio Araguari, betw. Cachoeiras Travessao & Santa Maria, 01°09'N, 51°52'W, J. M. Pires, W. Rodrigues A 6’. C. Ir¬ vine 50394 (NY). Acknowledgments. We are grateful to H. van der Werff (MO) and B. Manara (\ EN) for revising the Latin descriptions; Victoria C. Hollowell and two anonymous reviewers for comments on the 24 Novon manuscript; B. Manara (VEN) for preparing the il¬ lustrations; and the Missouri Botanical Garden and New York Botanical Garden staff for making their laeilities available for our research. Literature Cited Berry, I’.. (). Huber A B. Holst. 1995. Holistic analysis and phytogeography. Bp. 161—192 in I’. E. Berry, B. Holst & K. Yatskievych (volume editors), Flora of the Venezuelan Guayana, Vol. I (Introduction). Missouri Botanical Garden I’ress, St. Louis. Huber. (). 1994. Recent advances in the phytogeography of the Guyana region. South America. Mem. Soe. Bio- geogr. 4: 53—63. - . 1995. Geographical and physical features. Bp. 1 — 62 in B. E. Berry, B. Holst & K. Yatskievych (volume editors), Flora of the Venezuelan Guayana, Vol. 1 (In¬ troduction). Missouri Botanical Garden Bress, St. Louis. Jansen-Jaeobs, M. J. 1988. Ease. 4. 148. Verbenaceae. In: A. R. A. Corts-van Rijn (editor), Flora of the Guianas. Koeltz, Konigstein. Lopez- Balacios, S. 1977. Verbenaceae. Flora de Venezue¬ la, U.L.A., Merida. Michelangeh. F. 2000. Species composition and species- area relationships in vegetation isolates on the summit of a sandstone mountain in southern Venezuela. J. Trop. Ecol. 16: 69-82. Moldenke, H. 1934. A monograph of the genus Aegiphila. Brittonia I: 245—477. - . 1953. Botanical exploration in Venezuela, Ver¬ benaceae. Field iana (Hot.) 28: 512—516. - . 1972. A new species of Aegiphila from Venezue¬ la. Acta Bot. Venez. 6: 93—94. France, C. T. & I). M. Johnson. 1992. Blant collections from the Blateau of Serra do Araed (Amazonas, Brazil) and their phytogeographic affinities. Kew1 Bull. 47: 1 — 24. Senecio nemiae (Asteraceae, Senecioneae), a New Species from Sierra Grande in Patagonia (Argentina) Adriana Hartal i, 1 Roberta I). Tartosa,'2 and Susana E. Ere ire2' ‘Laboratories de Botanica “Lorenzo R. Parodi,” Facultad de Agronomfa, Universidad de Buenos Aires, Av. San Martin 4453, 1417 Buenos Aires, Argentina. cbartoli@mail.agro.uba.ar; tortosa@mail.agro.uba.ar 2Consejo Nacional de Investigaciones Cientificas y Tecnicas (CONICET) ‘Division Plantas Vaseulares, Museo de La Plata, Paseo del Bosque, 1900 La Plata. Argentina. freire@museo.fcnym. unlp.edu. ar Abstract. A new species, Senecio nemiae, from Sierra Grande, Rio Negro. Argentina, is described and illustrated. Senecio nemiae most closely resem¬ bles S. adenotrichius from Chile, but is clearly dif¬ ferent by the shorter leaves without auricles and smaller eapitula, with a calyculus of few bracts and length shorter than the involucre. A new status, Se¬ necio sect. Senecio ser. Corymbosi subser. Adenotri- chi, is also proposed to accommodate this taxon in the previous classification of Senecio for Argentina. Included is a key to those species of Senecio subser. Adenotrichi with glabrous achenes. key words: Argentina, Asteraceae, Compositae, Patagonia, Senecio, Senecioneae. Senecio L. has been subject to diverse taxonomic interpretations varying from a single, broad genus (Barkley, 1985) to numerous segregate genera (e.g., Nordenstam, 1978; Robinson & Bretell, 1973, 1974; Jeffrey, 1992). These new taxa were estab¬ lished based on microfeatures such as stylar ap¬ pendages, stigmatie surfaces, shape of filament col¬ lars, anther apex, anther base, and endothecial tissue (Wetter, 1983; Vincent, 1996). In the present paper, Senecio is treated in its wide concept, containing about 3000 species world¬ wide, with the most diversity in the mountainous areas or deserts of southern South America, tropical South Africa, as well as central and eastern Asia. Senecio s.l. includes monoecious herbs, subshrubs, shrubs, vines, and small trees, with eapitula radi¬ ate, discoid, or rarely disciform; phyllaries of equal length in one row, usually subtended by a calycu¬ lus; ray florets, if present, female with ligulate or short filiform corollas; disc florets hermaphroditic with tubular 5-dentate or 5-lobed corollas; the an¬ thers ecaudate or slightly tailed, filament collar slender or dilated; pappus of many slender and sca- brid bristles, and achenes 5- to 10-ribbed, glabrous or pubescent. On a regional basis, a large number of species of Senecio has been treated by Cabrera in southern South America, i.e., 208 from Chile (Cabrera, 1949), 90 species from Brazil, Paraguay, and Uru¬ guay (Cabrera, 1957), and 1 14 from Bolivia (Ca¬ brera, 1985). In Argentina Senecio is represented by 266 native species, arranged in 3 sections and 16 series, with 1 14 species endemic to this country (Cabrera el ah. 1999). Since the publication of this treatment for Argentinian species, only one new species was added (S. calingastensis Tombesi, 2001). The new species described here belongs to sec¬ tion Senecio, series Corymbosi, which is character¬ ized by the eapitula disposed in terminal corymbs and leaves sessile, attenuate into a pseudopetiole or auriculate at the base. We include it within the subsection Adenotrichi, established by Cabrera (1949) for Chilean species of the genus, not cited until now for Argentina. We propose the following new status to accommodate this taxon in the clas¬ sification sustained by Cabrera et al. (1999). Senecio sect. Senecio ser. Corymbosi subser. Adenotrichi (Lindley) A. Bartoli, Tortosa & S. E. Freire, stat. nov. Basionym: Adenotrichia Lindley, Bot. Reg. 14: 1 19(1. 1828. Senecio subsect. Corymbocephalus subsect. Adenotrichi (Lindley) Cabrera, Lilloa 15: 380. 1949. TYPE: Senecio adenotrichius DC. The subseries Adenotrichi within Senecio appears to be closely related to series Corymbosi subser. Viscosi. Both subseries share glandular pubescence, whereas the rest of the subseries of series Corym¬ bosi include species either tomentose or glabrous. Subseries Adenotrichi differs from subseries Viscosi in having deeply lobed to pinnatisect leaves (vs. entire, crenate, or serrate leaves). In addition, sub¬ series Viscosi ranges from Brazil to northeastern Ar- Novon 14: 25-28. 2004. 26 Novon Figure 1. Senecio nemiae A. Bartoli, Tortosa & S. E. Freire. —A. Habit. — B, C. Leaves. — I). Phyllary. — E. Trichome. — F. Capitulum. — G. Bay lloret. — H. Disk floret. — I. Anther. — J. Style branches. Scale bars: A = 3 cm; B-D = I cm; E = 0.1 mm; F = 5 mm; G, H = 2 mm; I, J = 1 mm. Drawn from holotype, Correa el al. 7090 (BAB). Volume 14, Number 1 2004 Bartoli et al. Senecio nemiae from Patagonia 27 gentina, whereas subseries Adenotrichi occurs in Chile and is now reported from southern Argentina. Senecio nemiae A. Bartoli, Tortosa & S. K. Freire, sp. nov. T\ PE: Argentina. Rio Negro: San An¬ tonio, Sierra Grande, 20 Oct. 1979, Correa el al. 7090 (holotype, BAB). Figure I. Quoad formam foliorum, fabricam capitulorum et in¬ dumentum ad Senecionem adenotrichium accedit, setl ah ea differt capitulis, calyculo et foliis minoribus, bracteis involucri in numero minori, foliis non aurieulatis et pilis capitatis brevioribus. Shrublet 20—25 cm, much branched at tin* base, densely hirsute glandulose on stems, leaves, and phyllaries, with glandular capitate trichomes, most¬ ly 0.5— 0.8 mm long; stems ascending. Leaves al¬ ternate, sessile, deeply pinnatilobed, 2.5—4 X 0.4— 0.6 cm, linear-elliptic, obtuse at the apex and slightly dilated at the base, margin often revolute, with midvein prominent, pubescent on both surfac¬ es; lobes 5 or 6 at each side, irregularly dentate, acute and mucronate at the apex. Capitula 2 or 3 pedunculate, peduncles 1—2 cm long, arranged in terminal corymbs; involucre calyculate, cylindric, 8-9 X ea. 8 mm; phyllaries 28 to 30, linear ovate, 8—9 X ea. 1 mm, dorsally pubescent, apex acu¬ minate with a tuft ol short trichomes; ealycular bracts ca. 9, linear, ea. 6 mm long, dorsally pubes¬ cent. Ray florets ca. 15, female, corollas yellow, ligulate, 3-dentate, ca. 6 mm long, the basal tube pilose, 2—4 mm long, the limb 3-dentate, ca. 4 mm long; disc florets ca. 50, hermaphroditic, corolla yellow, tubular, 5-lobed, ca. 7 mm long, glabrous, tube ca. 4 mm long, not basally broadened, limb narrowly campannlate, only slightly broader than the tube and not markedly distinct, throat ca. 2.2 mm long, lobes broadly triangular, ca. 0.80 mm long, at base 0.5— 0.6 mm wide; anthers minutely tailed, ca. 3 mm long; filament collars balusterform, ca. 0.5 mm long; apical anther appendage ovate- lanceolate, ca. 0.4 mm long; the basal tails ca. 0.2 mm long. Style branches penicillate, ca. 1.5 mm long, apically truncate. Achenes oblong-obovate, glabrous, ca. 1.8 mm (immature length), slightly costate. Pappus bristles many, slender and scabrid, ca. 7 mm long, white. The specific epithet honors Maevia Noemi Cor¬ rea, Argentinian botanist, in recognition of her work as director of Flora Patagdnica. Distribution and habitat. Senecio nemiae is known only from Sierra Grande, in the political province Rio Negro of southern Argentina. Sierra Grande represents a mountain range ol ca. 500 m altitude, at approximately 42s south latitude, and together with the Meseta de Somuncura constitutes a particular district of the vegetation of Patagonia since both regions contain a number of local en¬ demics, e.g., Adesmia serrana M. N. Correa (Fa- baceae), Boopis raffaelli Spegazzini (Calyceraceae), Conyza magnimontana Cabrera and Senecio rne- gaoreinus Zardini (Asteraeeae), and Silene melan- opotamica var. agrostophylla Pedersen (Caryophyl- laceae). Senecio nemiae is closely related to S. adenotri- chius DC. from Chile. Both species have deeply pinnatilobed leaves with dentate-acute lobes and ligulate corollas with trichomes on the tube. How¬ ever. the new taxon has shorter leaves without au¬ ricles and smaller capitula with fewer phyllaries and ealycular bracts. Kk'i to Species w ith Glabrous Achenes ok Senecio sect. Senecio ser. Corymbosi subser. Adenotrichi la. Phyllaries few (ca. 13) . . Senecio rivularis J. Bcmy l b. Phyllaries numerous (ca. 30 to 40). 2a. Leaves auriculate, 4—10 cm long; calyeulus of many bracts as long as the phyllaries. 3a. Leaves deeply lobed, lobes dentate- acute . Senecio adenotrichius DC. 3b. Leaves lobed, lobes entire or dentate, obtuse to subacule . . Senecio murinus Philippi 2b. Leaves without auricles, 2. ,5-4 cm long; ca- lyculus of few bracts shorter than the phyl¬ laries . Senecio nemiae A. Bartoli, Tortosa & S. E. Freire Acknowledgments. We thank John Pruski and an anonymous reviewer for reviewing the manu¬ script and for useful comments. Thanks to the cu¬ rator of BAB for the loan of specimens that made this study possible, and to Victor H. Calvetti for inking the illustration. We also thank Victoria C. Hollowell for her accurate editorial work. This study was supported by the Consejo Nacional de Investigaciones Cientfficas y Tecnicas (CONICET), Argentina. Literature Cited Barkley, T. M. 1985. Generic boundaries in Senecioneae. Taxon 34: 17-21. Cabrera, A. L. 1949. El genero Senecio (Compositae) en Chile. Lilloa 15: 27—501. - . 1957. El genero Senecio (Compositae) en Brasil, Paraguay y Uruguay. Arch. Jard. Bol. Rio de Janeiro 15: 163-269. - . 1985. El genero Senecio (Compositae) en Bolivia. Darwiniana 26(1—4): 79—217. - . S. E. Freire & L. Ariza Espinar. 1999. Tribu VIII. Senecioneae. Tribu VIII bis. Liabeae, in A. T. Hunziker (editor), Flora Eanerogamica Argentina 62: 3—180. Jeffrey, C. 1992. The tribe Senecioneae (Compositae) in the Mascarene Islands with an annotated world check- 28 Novon list of the genera of ihe tribe, Notes on Compositae: VI. Kew Bull. 47: 49-109. Nordenstam, B. 1978. Taxonomic studies in the tribe Se- necioneae (Compositae). Opera Bot. 44: 1-88. Robinson, H. & R. I). Bretell. 1973. Studies in the Se- necioneae (Asteraeeae), IV. The genera Mesadenia, Sy- neilesis, Miricacalia, Koyamacacalia and Sinacalia. Phytologia 27: 265—276. - & - . 1974. Studies in the Senecioneae (As¬ teraeeae), V. The genera Psacaliopsis, Barkley anthus, Telanthophora and Roldana. Phytologia 27: 402—439. Tombesi, T. 2001. Novedades en Senecio (Senecioneae, Asteraeeae). Hickenia 3(24—31): I I 1 — 1 14. Vincent, P. L. I). 1996. Progress on clarifying the generic concept of Senecio based on an extensive worldwide sample of taxa. Pp. 597—611 in I). J. N. Hind & H. J. Beentje (editors), Compositae Systematics. Proceedings of the International Compositae Conference, Kew, Vol. 1. Royal Botanic Gardens, Kew. Wetter, M. A. 1983. Micromorphological characters and generic delimitation of some New World Senecioneae (Asteraeeae). Brittonia 35: 1—27. Bulbophyllum filifolium (Orchidaceae), a New Species from Southeastern Brazil Eduardo L. Borba* and Eric C. Smidt Laboratorio < le Taxonomia Vegetal, Departamento de Ciencias Biologicas, l) n i vers it lade Estadual de Feira de Santana, Rod. BR 116, km 03, 44031-460, Feira de Santana-BA, Brazil. ^Author for correspondence: borba@gmx.net ABSTRACT. Bulbophyllum filifolium, a new orchid species from Brazil belonging to section Xiphizusa , is described and illustrated. The new species is rare and endemic to the municipality of Grao Mo- gol, Minas Gerais state, growing on rocks in campo rupestre vegetation. Bulbophyllum filifolium is eas¬ ily recognized by its thread-like leaves, one-flow¬ ered inflorescences, and ovate petals with long club-like appendages on the margin. hey words: Brazil, Bulbophyllum, Orchidaceae. Bulbophyllum Thouars is the largest orchid ge¬ nus, comprising over 1000 species throughout the tropics (Vermeulen, 1991; Dressier, 1993). The species occur mostly in the Old World, although nearly 70 species are present in the New World. Pabst and Dungs (1975, 1977) cited 54 species for Brazil, and with the addition of some recently de¬ scribed species and new occurrences for this coun¬ try (e.g., Borba et al., 1998; Fraga, 1999; Toscano de Brito, 2000), this number is increased to about 60. Twelve of the Brazilian Bulbophyllum species be¬ long to section Xiphizusa (Reichenbach f.) Cog- niaux, characterized mainly by the long fused se¬ pals. Those species are mainly epiphytic in forest areas in southeastern Brazil, and occasionally lith- ophytic in campo rupestre vegetation, as B. plu- mosum (Barbosa Rodrigues) Cogniaux. The campo rupestre of the Espinhago Range is the Brazilian vegetation with the highest number of endemisms and greatest diversity (Joly, 1970; Giulietti & Pir- ani, 1988). Due to the discontinuity of these moun¬ tain ranges and outcroppings, many species, mainly the 1 it hophy tic ones, are distributed in disjunct populations. Because of that, some sites are espe¬ cially rich in the number of endemic species and/ or present populations differentiated (e.g., geneti¬ cally, morphologically, chemically) from the core area of the species, as in the region of Grao Mogol in the north of Minas Gerais (Giulietti & Pirani, 1988; Borba et ah, 2001a, 2001b, 2002). In the course of a revision of Brazilian Bulbophyllum and the population genetics and reproductive biology studies of the orchids of Grao Mogol, we found a newT species of Bulbophyllum in section Xiphizusa with remarkable leathery thread-like leaves, de¬ scribed as follows: Riilhopliyllum filifolium Borba & Smidt, sp. nov. TYPE: Brazil. Minas Gerais: Grao Mogol. Ser- ra do Barao, 16°33'S, 42°54'W, 7 Jan. 2002 (fl), E. L. Borba 1999 & C. van den Berg (ho- lotype, IIUEFS). Figure 1. Species liaee Bulbophyllo pabstii, B. bidentato , B. plu- rnoso et aliis speeiebus sectionis Xiphizusa similis; ab om- nis speeiebus folio filiforme inflorescenlia uniflora et com- binatione labelli lobo medio carnoso. lobis marginibus lateralibus dense curto-ciliatis ornatis et petalis ovatis ap- pendicibus claviformibus ornatis longioribusque ad mar- ginem differt. Eithophytie herbs; roots 20--40 mm long, thread-like, fasciculate; rhizome 2—8 X 1 mm, creeping; pseudobulbs 4—8 X 6—7 nun, aggregate, ovoid, slightly 3- to 4-angular, monophyllous. Leaves 25—30 mm long, 1 nun diam., thread-like, leathery, folded, erect, apex acute, base canalic¬ ulate and attenuate. Inflorescence basal, one-flow¬ ered; scape 40—50 mm long, curved, with 4 sheath-like bracts, 3—4 nun long, adpressed, not imbricate; floral bracts 4X3 mm, ovate-lanceo¬ late, glah rous, apex acute; flower pendent; ovary 3 mm long, suleate; sepals membranaceous, sub¬ equal, erect-patent, boat-shaped, glabrous; dorsal sepal 14 X 3 mm, linear-lanceolate, 3-nerved, apex acute; lateral sepals fused up to the apex, synsepal 14X5 mm, lanceolate, 7-nerved, apex acute, slightly gibbous at base; petals 4X3 mm, oblique-ovate, erect, 1-nerved, margins eiliate at the base, with club-like appendages up to 1 mm long increasing in size from base to apex; 1 i p 9 X 1.7 mm, fleshy, erect, parallel to column, movable; lateral lobes 4 mm long, erect, auriculate, densely Novon 14: 29-32. 2004. 30 Novon XIaah E O id o E o Figure 1. Bulbophyllum Jilifolium Borba & Smidt. — A. Dorsal sepal. — B. Fused lateral sepals. — C. Petal. — D. Lip, lower view. — E. Lip, upper view. — F. Lip (below) and column (above), in side view. — G. Habit. Scale bars: 0.5 cm, A-F; 1 cm, G. Drawn from the holotype, Borba 1999 & ran den Berg, by Alarm Duraes. Volume 14, Number 1 2004 Borba & Smidt Bulbophyllum filifolium from Brazil 31 short-ciliate; mid-lobe 5 mm long, linear-elliptic, adaxially with one distinct longitudinally sulcate ridge, adaxial surface glabrous, apex rounded; col¬ umn 3 X 1.1 mm, with two sinuate stellidia at apex and two falcate teeth on the ventral face; anther versatile, 3-globose. Etymology. This species is named for the ex¬ tremely narrow, linear (thread-like) leaves, the narrowest among the Brazilian Bulbophyllum spe¬ cies. Distribution. Bulbophyllum filifolium grows on rock outcrops of quartzite in the eampo rupestre vegetation of Serra do Barao, in the municipality of Grao Mogol, in the state of Minas Gerais, Brazil, a disjunct area of the main Espinhayo Range. The eampo rupestre vegetation is characterized by open, herbaceous vegetation on sandy, stony soils mixed with herbs and shrubs growing on outcrop¬ pings of quartzite, sandstone, gneiss or “canga” (iron) rocks (Giulietti & Pirani, 1988; Borba et ah, 2001a). This species appears to be endemic to the Grao Mogol region and is rare, as only three in¬ dividuals have been found despite several field trips made to the type locality. A fragment of the same individual as the holotype is currently under cultivation at the Universidade Estadual de Feira Santana (another fragment of the same individual and the other two individuals remain undisturbed at the original locality). The Serra do Barao was recently defined as a State Reserve (Parque Es¬ tadual de Grao Mogol), and B. filifolium possibly occurs at other sites there. Further studies are necessary to assess the conservation status of the species. Bulbophyllum filifolium belongs to section Xiphizusa as suggested by its floral and vegetative morphology, such as the erect rachis not genicu¬ late and not thickened, lateral sepals long and connate to the apex forming a synsepal, a column with two long stellidia at the apex and two small teeth on the ventral face, and pseudobulbs aggre¬ gate and sulcate. The movable lip is a common feature in the genus and probably plays an im¬ portant role in the pollination mechanism of this species (wind-assisted fix pollination; Borba & Se- mir, 1998), as observed in other species of section Xiphizusa presenting similar flower structure (e.g., B. plumosum (Barbosa Rodrigues) Cogniaux and B. bidentatum (Barbosa Rodrigues) Cogniaux; Ver- ola, 2002). This species is easily recognizable by its thread-like leaves and one-flowered inflores¬ cences, unique among the members of B. sect. Xiphizusa. The other two Brazilian Bulbophyllum species with narrow leaves are B. insectiferum Barbosa Rodrigues and B. adiamantinum Brade, belonging to sections Bulbophyllaria and Micran- tha, respectively. However, conversely to B. fili¬ folium , these two species have succulent cylindri¬ cal leaves and multi-flowered inflorescences with very distinct flowers; they also occur in Minas Gerais, but are not sympatric with B. filifolium. The long club-like appendages on the petal mar¬ gins are uncommon in other species of the section, being observed only in B. pabstii Garay. However, B. filifolium has ovate petals and the mid-lobe of the lip is fleshy, whereas B. pabstii has ovate-lan¬ ceolate petals and the mid-lobe of the lip is mem¬ branaceous; it also differs by the oblong-lanceo¬ late leaf (ea. 10 mm w ide) and the multi -flowered inflorescence. The lip mid-lobe of B. filifolium is similar to that of B. bidentatum, but the latter dif¬ fers by the petals without the club-like append¬ ages, the lateral lobes of the lip not ciliate, the oblong-lanceolate leaves (ca. 8 mm wide), and the multi -flowered inflorescence. Because we found only plants with old (lowers we cannot describe the flower color accurately, but they appear to be entirely purple or dark red and similar to those of B. bidentatum. Acknowledgments. Thanks are due to Cassio van den Berg for the Latin diagnosis and other im¬ provements to the manuscript. Literature Cited Borba, E. I,. & J. Semir. 1998. Wind-assisted fly polli¬ nation in three Bulbophyllum (Orchidaceae) species oc¬ curring in the Brazilian campus rupestres. Lindleyana 13: 203-218. - . - & F. Barros. 1998. Bulbophyllum ano¬ int urn Borba, Semir & F. Barros (Orchidaceae), a new species from the Brazilian campus rupestres. Novon 8: 225-229. - . J. M. Felix, V. N. Solferini & J. Semir. 2001a. Fly-pollinated Pleurothallis (Orchidaceae) species have high genetic variability: Evidence from isozyme mark¬ ers. Amer. J. But. 88: 419-428. - . ,1. R. Trigo & .1. Semir. 2(H)] b. Variation of dia- stereoisomeric pyrrolizidine alkaloids in Pleurothallis (Orchidaceae). Biochem. Syst. Ecol. 29: 45—52. - , G. .1. Shepherd, C. van den Berg & J. Semir. 2002. Floral and vegetative morphometries of five Pleu¬ rothallis (Orchidaceae) species: Correlation with tax¬ onomy, phylogeny, genetic variability and pollination systems. Ann. Bot. 90: 219—230. Dressier, B. I.. 1993. Phylogeny and Classification ol the Orchid Family. Cambridge Univ. Press, Cambridge. Fraga, C. N. 1999. Bulbophyllum gomesii Fraga (Orchi¬ daceae), uma nova espdcie da floresta atlantica do Es- pfrito Santo. Brasil. Bradea 8: 135—138. Giulietti. A. M. & J. R. Pirani. 1988. Patterns of geo¬ graphic distribution of some plant species from the Fspinhayo Range, Minas Gerais and Bahia, Brazil. Pp. 39—69 in W. R. Heyer & P. F. Vanzolini (editors). Pro- 32 Novon ceedings of a Workshop on Neotropical Distribution Patterns. Academia Brasileira de Cieneias. Rio de Ja¬ neiro. Joly, A. B. 1970. Conhecja a Vegeta^ao Brasileira. EDUSP, Sao Paulo. Pabst, G. K. .). & K Dungs. 1975. Orehidaceae Brasilien- sis, Vol. I. Briicke-Verlag, Hildesheim. - & - . 1977. Orehidaceae Brasiliensis, Vol. 2. Briicke-Verlag, Hildesheim. Toscano de Brito, A. L. V. 2000. Two new species of Or- chidaeeae from Brazil. Lindleyana 15: 184—188. Vermeulen, .1. .). 1991. Orchids of Borneo, Vol. 2 — Btil- bophyllum. Royal Botanic Gardens, kew. Verola, C. F. 2002. Biologia Floral e Sistemas de Reprod- m,ao em Fspeeies de Bulbophyllum (Orehidaceae) Oeorrentes em Mata de (Valeria, Campo Rupestre e Flo- resla Estaeional. M.Sc. Thesis, Universidade E^stadual de Campinas. Campinas. Paronychia revoluta , a New Species of Caryophyl laceae from Brazil Claudia Ciena Carneiro Universidade Estadual de Feira de Santana, Departamento de Cieneias Biologicas, Laboratorio de Mieromorfologia Vegetal (LAJV1 IV), BR I 16, km 3, Campus U niversitario, CEP 44031-460, Feira de Santana. Bahia, Brazil, cameiro@uefs.br Antonio Furl an Universidade Estadual Paulista, Instituto de Biociencias, Departamento de Botanica, Av. 24A, 1515, Caixa Postal 199, Bela Vista, CEP 13506-900, Rio Claro, Sao Paulo, Brazil. afurlan@rc.unesp.br ABSTRACT. A new species of Paronychia, P. re¬ voluta, from the state of Rio Grande do Sul, Brazil, is described and illustrated. This species is closely related to P. camphor osmoides, distinguished from it by linear-subulate leaves with a revolute margin and prominent midribs. It is also allied to P. Jas- ciculata, from which it differs in the lanceolate, acuminate, fimbriate, and whitish stipules, and lin¬ ear-subulate leaves. Key words: Brazil, Caryophyl laceae. Paronych¬ ia. Paronychia Miller (Caryophyllaceae) comprises about 1 10 species distributed among three subgen¬ era (Chaudhri, 1968; Bittrich, 1993). This genus is characterized by the presence of stipules and bracts, very small and clustered flowers often con¬ cealed by the bracts, filiform petals, episepalous stamens, uniovulate ovary, utricle rupturing irreg¬ ularly at base, globose seeds, and a curved embryo. Although occurring mainly in the Mediterranean region, the species of Paronychia are widely dis¬ tributed throughout the world, inhabiting warm- temperate, montane, and occasionally alpine re¬ gions. They may be found in dry stony or rocky places, calcareous soils, mountain steppes or along sea coasts and river banks. In this paper a new Paronychia species, discov¬ ered during a study of Caryophyllaceae from Brazil, is described and illustrated. Paronychia revoluta C. E. Carneiro s and S. sipapoanus Maguire. There¬ fore, I have elevated S. duidae subsp. neblinae to the species level. Additional specimens examined. VENEZUELA. Ama¬ zonas: Dept. Rio Negro, Cerro Aracamuni, summit, Proa Camp, 31 Oct. 1987, A'. Liesner A' G. Carnevali 22081 (MO, U); Sierra de la Nehlina, Rfo Yatua, escarpment above Camp 4, 30 Dee. 1953, B. Maguire et at. .37007 (NY); Sierra de la Nehlina, Rfo Yatua, escarpment edge betw. Cumbre Camp & N Escarpment, 18 Jan. 1954. B. Maguire et al. .37240 (NY); Sierra de la Nehlina. Rfo Ya¬ tua, W of Cumbre Camp, 1-2 Dec. 1957. B. Maguire et id. 4228.3 (NY); Dept. Rfo Negro, Sierra de la Nehlina. above Rfo Marawinuma, E of Puerto Chimo camp, 26 Apr. 1984, W. Thomas 32.37 (CAS). Styrax nunezii P. W. Eritscli, sp. nov. TYPE: Peru. Cuzco: provineia La Convencidn, distrito Eeharati, E Rfo Apurimac, NE Pueblo Libre, Anchihuay and Bellavista Mountains, S Cor¬ dillera Vilcabaniba, 2445 m, 12°51'S, 73°30'W, 3 Aug. 1998 ((I), T. V. Nunez, W. Nan ray, R. de la Colina & S. Udvardy 23320 (holotype, CAS; isotypes, AAU, RR. COL, F, GH, K. LOJA, LPB, MICH. MO, NY, TEX. US, W). Figure 3. Arbor sempervirens. Lamina 7. 0-9. 3 X 3. 5-4.0 cm crassi-chartacea ovata, apiee aeuminato, subtus pilis den- sis minutis albi-stellato-tomentosis instrueta, magis dis- persis sed densis lanatis atro-ferrugineo-stellatis braehiis usque 0.2 mm longis intermixtis, venis quaternariis leviler vel baud elevatis, paginis venarum non manifestis subter indumentum, ubi veteribus pilis ferrugineis absentibus et margine revoluto. Flos 10 mm longus; calyx 3 X 3—4 mm. dense pallidus fusco-stellato-tomentosus, ad medium bra- eliiis pilorum usque 0.2 mm longis; stamina 10; filament a auriculata pilis albi-stellatis undulatis vel aliquantum rig- idis braehiis usque 0.8 mm longis instrueta; anthera 3.5 mm longa loeulis lineari-triangularibus ad apicem con- traetis; connectivum locula leviter superante; stylus filifor- mis; ovula multa. Evergreen tree 6 m tall; young twigs densely brown stellate-tomentose. Petiole 10—1 I mm long; lamina 7.0— 9.3 X 3. 5-4.0 cm, 1.8— 2.5 times as long as wide, thiek-chartaeeous. ovate, secondary veins 7 to 10 on each side of midvein; apex acu¬ minate; base broadly cuneate to subrounded; adax- ially glabrous except on midrib; abaxially appear¬ ing light brown to the naked eye, with a dense base tomentum of minute white stellate trichomes, and more scattered but dense dark ferrugineous woolly stellate trichomes with arms to 0.2 mm long, sur¬ face ol the veins not visible beneath the pubes¬ cence. ferrugineous trichomes often absent on older leaves, tertiary veins slightly raised, quaternary veins faintly or not at all raised; margin entire, rev- olute on old leaves. Inflorescences axillary or ter¬ minal, racemose (or I - or 2-llowered or rarely pa¬ niculate), I to 3 per node, 2-3 cm long, 3- to 14-flowered, densely light brown stellate-tomen¬ tose; lower pedicels 4—6 mm long. Flowers her¬ maphroditic. 10 mm long; calyx 3 X 3-4 mm, eu- puliform, densely light brown stellate-tomentose, trichomes at mid-calyx with arms to 0.2 mm long, margin truncate between the minute teeth, inner margin eglandular; corolla 9 mm long; tube 1 mm long, extending 1—2 mm beyond the calyx margin; lobes 5, 9 X 1.5 mm, spreading, thickened; sta¬ mens 10; free portion ol stamen tube none; distinct portion of filaments 1 mm long, ventrally with au¬ ricles bearing white, undulating to somewhat still stellate trichomes with arms to 0.8 mm long, tri¬ chomes nearest the proximal end of the filament with arms predominantly pointing downward, those nearest the distal end with arms predominantly pointing upward, also stellate-pubescent in sinuses of filaments and dorsally, trichomes with arms to 1.3 mm long; anthers 3.5 mm long, the glabrous connectives slightly exceeding the linear-triangular, apically tapered, stellate-pubescent thecae; ovary densely gray-green stellate-tomentose; stvle fili¬ form, glabrous; ovules many. Fruit unknown. Habitat, distribution , and phenology: Known only from the type collection (I have only seen flow¬ ers, but the specimen label states “young flowres | sic ) and fruits”) in the cloud forest zone of the Cordillera Vilcabaniba in southern Peru; flowering August. This species resembles Styrax rigidifolius Idrobo & R. E. Schultes and several species of Styrax from the G uayana Highland, such as .S’, guanayanus and S. sipapoanus, in its ahaxial leaf surfaces. These are abundantly covered with woolly ferrugineous stellate trichomes in addition to the white stellate base tomentum, rendering the surface light to dark brown to the naked eye. Styrax nunezii is easily distinguished from these species, however, by the presence (vs. absence) of auricles on the ventral portion of the stamen filaments. These auricles, as well as the presence of linear-triangular (vs. linear) anthers, allies S. nunezii with a group of Andean species with the same combination of features (e.g.. S. pavonii A. DC., S. pentlandianus .). Remy, S. peruvianas Zahlbruekner). No other species of this group is known to have the type of abaxial leaf surface possessed by S. nunezii. In addition, the small (10 mm long) flowers and short (2-3 cm) in¬ florescences are unusual in the group. The species is named for my colleague Percy Nunez Vargas, curator of Herbario Vargas (CUZ) at the Universidad Nacional San Antonio Abad del Cusco, Cuzco, Peru, who collected the type and Volume 14, Number 1 2004 Fritsch Sty rax from South America 51 Figure 3. Styrax nunezii P. W. Fritsch. — A. Flowering branch. — B. Leaf, abaxial view. — C. Leaf surface, abaxial view. — D. Stellate trichome from the base tomentum of the abaxial side of the leaf. — E. Larger stellate triehome from the abaxial side of the leaf. — F. Flower. — G. Flower, median long-section. — H. Part of corolla + androecium, opened. — I. Stamen, lateral view. Based on the holotype, Nunez el al. 23320 (CAS). 52 Novon generously made specimens available to me for ex¬ amination. Additional isotypes are to be expected at CUZ and USM. Styrax rotundatus (Perkins) P. W. Fritsch, slat, nov. Basionym: Styrax martii var. rotundatus Perkins, Pflanzenr. 4. 241: 40. 1907. TYPE: Brazil. Bahia: Jaeobina, ./. S. Blanchet 3614 (leetotype, selected here, BB: isotypes, C, F[2], W; photo of C at F, MICH, MO). Evergreen shrub to 3(4) m tall: young twigs densely tawny, gray-brown, dark ferrugineous, or brown stellate-tomentose. Petiole (2-)4— 8 mm long; lamina 3.3— 5.8(-6.6) X 2. 1-4.3 cm, 1.3-2 times as long as wide, thick-chartaceous to subcoriaceous, elliptic, ovate-elliptic, ovate, or suborbicular, sec¬ ondary veins 4 to 8 on each side of midvein; apex acute to subrounded; base broadly euneate, round¬ ed, truncate, or cordate; adaxially densely brown stellate-tomentose when young, glabreseent except on major veins; abaxially dark ferrugineous or brown to the naked eye, with a dense base tomen- tum of green-white minute stellate trichomes, and abundant but more scattered woolly ferrugineous trichomes with arms to 0.2— 0.3 mm long that are sparse to absent in old leaves, tertiary and quater¬ nary veins distinctly differentiated from one anoth¬ er, distinctly raised, their surfaces visible through the pubescence to completely obscured; margin en¬ tire, sometimes revolute. Inflorescences axillary or terminal, racemose or paniculate (axillary inflores¬ cences occasionally 1- or 2-flowered), 1 or 2 per node, 2—6 cm long, 3- to 8-flowered, subtending leaves occasionally strongly reduced, densely gray, dark ferrugineous, or brown stellate-tomentose; lower pedicels 4—7 mm long. Flowers hermaphro¬ ditic, 10—13 mm long; calyx 3.5—5 X 2.5— 3.5 mm. cupuliform to narrow-cupuliform, densely gray- green stellate-pubescent mixed with larger evenly scattered woolly golden or brown stellate trichomes with arms at mid-calyx to 0.5 mm long, margin truncate between the minute teeth, inner margin eglandular; corolla 8-1 I mm long, white; tube 2-3 mm long, extending up to the calyx margin; lobes 5, 5—10 X 0.5— 1.5 mm, recurved to reflexed, thin; stamens 10; free portion of stamen tube none; dis¬ tinct portion of filaments 2 mm long, ventrally with¬ out auricles, bearing a dense mass of white stellate trichomes with thin, ± straight arms to 1.5 mm long, trichomes nearest the distal end of the fila¬ ment with arms predominantly pointing upward, those nearest the proximal end with arms predom¬ inantly pointing downward, dorsally glabrous, tri¬ chomes along the edges of the filament and at the sinus between filaments with arms to 2 mm long; anthers 3—6 mm long, the glabrous connectives slightly exceeding the linear, apieally tapered, stel¬ late-pubescent thecae; ovary densely golden yellow stellate-tomentose; style filiform, glabrous; ovules many. Drupe 5—7 X 4—6 mm, gray-green to dull orange, subglobose, apex slightly to strongly de¬ pressed, annular-suleate around the remains of the style base; wall irregularly and coarsely rugose on herbarium specimens; fruiting calyx 3—5 X 4—6 mm, 0.4— 0.8 times the length ol the drupe proper. Habitat, distribution , and phenology. Endemic to Brazil, occurring in the states ol Bahia and Minas Gerais in campo rupestre, among rocks, rarely on sandy soil at 900— 1850(— ?2000) m; flowering No- vember-May, fruiting March-June. Styrax rotundatus was originally described as a variety ol S. martii by Perkins (1907; as S. martii var. rotundatus), who distinguished it from the typ¬ ical variety by its orbicular leaves with rounded or broad-acuminate apices. In my view, the two taxa differ consistently not only through the shape of the leaves (although more subtly than Perkins has sug¬ gested, as indicated in the descriptions herein), but also through numerous other characters that, taken together, have prompted me to recognize .S', martii var. rotundatus at the specific level. The following characters serve to delimit S. rotundatus from .S'. martii: petiole (7-) 10— 19 mm long (vs. (2— )4— 8 mm long), lamina 3.3-5.8(-6.6) cm long (vs. 5.9-11.7 cm long), tertiary and quaternary veins distinctly differentiated from each other and conspicuously raised (vs. not well differentiated from each other and moderately raised), lower pedicels 4—7 mm long (vs. 1—5 mm long or nearly absent), calyx 3.5— 5 X 2.5— 3.5 mm (vs. 5—8 X 4—5 mm), not striate (vs. striate) proximally. flowers 10-13 mm long (vs. 13—18 mm long), corolla tube 2—3 mm long (vs. 5— 8 mm long), corolla lobes 0.5— 1.5 mm wide (vs. 1 .5— 2.5 mm wide) and recurved to reflexed (vs. spreading), and drupes 5—7 X 4-6 mm (vs. 9—16 X 6—7 mm), gray-green to dull orange (vs. olive). In addition, S. rotundatus is always a shrub, where¬ as S. martii can be either a shrub or a tree; S. rotundatus occurs exclusively in campo rupestre vegetation, whereas S. martii can occur in campo rupestre but also occurs in forests, forest edges, and campo de altitude. The ranges of the two species are nearly non-overlapping, with .S’, rotundatus oc¬ curring from extreme northern Bahia to south-cen¬ tral Minas Gerais, and S. martii occurring from south-central Minas Gerais to northern Rio Grande do Sul. The two species are apparently sympatric Volume 14, Number 1 2004 Fritsch Styrax from South America 53 only in the Serra do Cipd in south-central Minas Gerais, Brazil. Perkins (1907) listed two collections in the pro- tologue of Styrax martii var. rotundatus: Blanchet 234 and Blanchet 3614. 1 have leetotypified this taxon on the BR specimen of Blanchet 3614 be¬ cause it was annotated by Perkins, and because duplicates of Blanchet 3614 are apparently more widely distributed than those of Blanchet 234 (1 have detected only two sheets of Blanchet 234. from P and W, whereas Blanchet 3614 is housed at BR, C, F, and W). Additional specimens examined. BRAZIL. State not specified: Glocker 234 (BM (the specimen has the same appearance and is at the same stage of development as collections of Blanchet 234 ; the numeral “ 234 ” is written on a label separate from another containing the words “Glocker. Brazil. Ex Herb. Shuttleworth.,” suggesting that this is actually a duplicate of Blanchet 234 that has been incorrectly attributed to Glocker)); C. F. P. von Martins s.n. (M no. 47). Bahia: Jacobina, ra. 24 km from Jacobi na to Morro do Chapeu, 28 Oct. 1995, A. M. A. Amorim el al. IBOI (CAS[3]); Palmeiras, Morro do Pai Inayio, 25 Jan. 1998, A. M. A. Amorim et al. 2137 (CAS); Utinga. Morro Felix Joao (Morro da Torre da Embratel), 14 km from Utin¬ ga. 16 Mar. 1985, A. P. de Araujo & IE N. da Fonseca 404 (UB); Morro do Chapeu. 27 Aug. 1980, //. P. Bautista 414 (K). 5 Apr. 1984. H. P. Bautista & O. A. Salgado 941 (GLIA. MBM, RB), ./. S. Blanchet 234 (P. W); Bomfim, Mount Taboa, 8 May 1918. II. M. Curran 174 (GH. NY, US); 34 km E of Morro do Chapeu along Highway BA- 052, Chapada da Diamantina, 3 Apr. 1976, G. Davidse et al. 1 1303 (MO); Morro do Chapeu, 2 Mar. 1997, F. Franga et al. 5924 (k, W); 11-13 km from Mucuge along the new A ndaraf— Mucuge road. 8 Sep. 1981, A. Furlan et al. 2120 (k); Rio tie Contas, Pico das Almas, 14 Dec. 1984,4. M. Giulietti et ex of all five anthers. The two novelties bear apical triehomes only on the ventral pair of anthers. In point ol fact, there is no a priori reason to restrict the search for relatives of C. duvalliorum and C. mar¬ itae to species with dissected adult foliage. The avail¬ able evidence suggests that deeply dissected adult fo¬ liage has evolved several times within Cyanea (Carlquist, 1974; Lammers, 1990a, 1990b; Givnish et ah, 1994, 1995). Cyanea aspleniifolia and C. ship¬ manii are related to species with toothed leaves, in¬ cluding C. dunhariae Rock and C. profuga C. N. Forbes (Lammers, 1990b), while the closest relative ol the C. grimesiana complex appears to be C. loliata H. Mann, in which the adult leaves are merely lolled (Lammers, 1990a). For these reasons, species with non-dissected foliage must also be considered as po¬ tential relatives of the novelties. Detailed consideration of the entire genus re¬ vealed that there are two informal species groups in addition to the C. grimesiana complex, which are characterized by many of the same features found in C. duvalliorum and C. maritae: frequent produc¬ tion of prickles; relatively well developed bracts; relatively large white flowers; corolla tubes stout, arcuate, laterally compressed, and tallest above the middle; and orange or yellow berries crowned by the persistent calyx lobes. The first consists of the ten species formerly seg¬ regated as the genus Rollandia (Lammers et ah, 1995; Lammers. 1998a): C. calycina (Chamisso) Lammers; C. crispa (Gaudichaud) Lammers, Giv¬ nish & Sytsma; C. humboldtiana (Gaudichaud) Lammers. Givnish & Sytsma; C. koolauensis Lam¬ mers, Givnish Sytsma; C. lanceolata (Gaudi¬ chaud) Lammers. Givnish N Sytsma; C. longiflora (Wawra) Lammers, Givnish & Sytsma; C. parvifolia (C. N. Forbes) Lammers. Givnish & Sytsma; C. pur- purellifolia (Rock) Lammers, Givnish cK Sytsma; C. sessilifolia (0. Degener) Lammers; and C. st.-johnii (Hosaka) Lammers, Givnish & Sytsma. These plants are unique within the genus in the dorsal adnation of the filament lube to the corolla tube and further differ from the C. grimesiana complex in their lack of dissection of the adult foliage; mar¬ gins are entire or more often merely toothed. How¬ ever, this complex does show some developmental dimorphism. Four species (C. calycina, C. lanceo¬ lata, C. longiflora, and C. sessilifolia ) produce dis¬ tinctive juveniles, though in these the margins of the juvenile leaves are merely lohed rather than parted or divided. Biogeographieally, the species of the Rollandia complex are all endemic to O'ahu, except for C. parvifolia, which is endemic to kaua'i; no representatives occur on or near East Maui. The second informal group consists of five spe¬ cies found on the two youngest islands of the ar¬ chipelago: C. glabra (E. Wimmer) St. John, C. lon- gissima (Rock) H. St. John, C. mceldowneyi Rock (all from East Maui), C. platyphylla (A. Gray) 1 1 i I- lebrand (windward Hawai'i), and C. scabra Hille- brand (West Maui). Like the Rollandia complex, the C. scabra complex differs from the C. grimesi- ana complex in its lack ol dissection of adult fo¬ liage and the presence of developmental dimor- Volume 14, Number 1 2004 Lammers Endemic Hawaiian Genus Cyanea 89 phism. As before, llie species that do produce distinctive juveniles ( C . glabra, C. mceldowneyi, and C. platyphylla ) bear leaves that are merely lobed rather than parted or divided. The relationships of C. duvalliorum and C. maritae lie somewhere among these three informal groups. 01 them, the Rollandia complex seems the most distantly related to C. duvalliorum and C. maritae. None of its species has leaves, adult or juvenile, that even ap¬ proach the two novelties in degree of dissection, while the novelties lack the unique adnation of the staminal column that characterizes the group; furthermore, the species of the Rollandia complex occur three or four islands away in a very' different biogeographic setting. The C. grimesiana and C. scabra complexes lack this adnation of the staminal column and are represented on East Maui. It is more difficult to decide between the C. gri¬ mesiana and C. scabra complexes, because their distinctness is blurred to some extent by C. duval¬ liorum and C. maritae. Though the C. grimesiana complex would seem more similar to the novelties due to its dissected adult loliage, it does not show any tendency toward developmental dimorphism. This seems to me to reflect a fundamental differ¬ ence in life history. Furthermore, the novelties dif¬ fer from C. grimesiana and most of its allies in their lack of apical pubescence on the three dorsal an¬ thers. Eli is feature is unique to these species within Cyanea (and very rare in the subfamily) and seems a valuable indicator of affinity. Biogeographically, this complex is represented on East Maui by just one of its five species. Though the C. scabra complex shows no dissec¬ tion of the adult lamina, three of its five species do produce distinctive juveniles. In all its species, the anthers bear triehomes only on the ventral pair, as in the novelties. Furthermore, East Maui is the cen¬ ter of diversity for this complex, with three of its five species endemic there. Overall, C. duvalliorum and C. maritae do seem more closely related to the members of the C. scabra complex. Within this complex, one would assume that the two new species are each others closest relative, as they are the only members of the group with parted or divided adult leaves. 01 the species described pre¬ viously, C. scabra and C. mceldowneyi seem most closely related to the two novelties, based on their common possession of persistent narrowly oblong bracts and calyx lobes, rounded at apex (see key be¬ low). It is noteworthy that C. duvalliorum possesses a feature unique in the genus: the concrescence of the apical triehomes on the ventral anthers into a triangular scale. An identical scale, formed in the same manner, characterizes a Latin American taxon of Lobelioideae, Centropogon C. Presl subg. Cen¬ tro/ logon (Lammers, 1998b). Like the branched tri- chomes found in Cyanea calycina and Centropogon sect. Siphocampyloides Bentham (Lammers, 1998b, 2002; Batterman & Lammers, in press), this is ap¬ parently a case of parallel evolution. Although the plants discovered at WaLoltiwi Stream could not be equated with any named spe¬ cies, they do appear to match a number of speci¬ mens collected on East Maui by earlier botanists. These specimens were sterile or otherwise inade¬ quate for confident identification, and languished in herbaria under various names for many years. Of these earlier specimens, Rock 8797, 8797a, and 8798 served as paratypes of C. scabra var. var- iabilis Rock. Although these gatherings are referable to C. duvalliorum (8797) and C. maritae (8797a, 8798). the holotype of C. scabra var. variabilis is a specimen of C. longissima and that name is properly treated as a synonym of this species. Previously (Lammers, 1990a), all these paratypes were regarded as juveniles of C. glabra. However, I subsequently realized that the leaf margin of true juvenile C. gla¬ bra (e.g., Forbes I657.M, BISH, K. L. RSA. S) is only shallowly lobed, not deeply dissected. The remainder of the early sterile collections ol C. duvalliorum had been identified tentatively as C. horrida. Though these two species are very sim¬ ilar in morphology of both juvenile and adult leaves, sterile specimens may be distinguished on the basis of pubescence. In C. horrida, the leaves are sparsely pubescent adaxially and more densely so abaxially; in C. duvalliorum, pubescence is con¬ fined to the abaxial venation or totally absent. These earlier collections considerably extend the distributional range of both C. duvalliorum and C. maritae on the windward slopes of Haleakala. Some were collected 7—12 km east of Wai’ohiwi Stream, from Honomanu Stream to Wailua Iki Stream; oth¬ ers were obtained approximately 27> km east, at Kt- pahulu Valley. Efforts should be made to ascertain whether any of these populations is still extant. Cyanea duvalliorum Lammers & H. Oppenheimer. sp. nov. Type: U.S.A. Hawaiian Islands: East Maui, Makawao District, ko'olau Forest Re¬ serve, Wai'ohivvi. 20°50'04”N, 156°15'55"W, mesic forest, steep shady slope, 2800 ft.. 15 Oct. 2000. //. Oppenheimer & F. Duvall H100026 (holotype, BISH [sheet plus liquid); isotypes, B. k, MO, NY, OSH. ITBG, US). Fig¬ ure 2. Species of) marginem foliorum maturorum pinnatim (is- 90 Novon Figure 2. Cyanea duvalliorum Lammers & H. Oppenheimer. — A. Adult leaf. — B. Juvenile leaf. — C. Flower. — I). Fruil. (A, C drawn from the holotype, Oppenheimer & Duvall H 100026: B. I) drawn from a paratype, Oppenheimer el al. HI 0003.) sum partition vel divisum el corollae tubum compressum arcuatum supra medium altissimum affinitatem Cyaneae grimesianae simulans, sed ob plantas juveniles proprias et tubum antherarum cum trichomatibus apiealibus solum in pari ventrali gregem C. seahrae vero pertinet; inter C. scabram et species affines habitu arboreo 4—6 m alto, rtiar- gine foliorum maturorum pinnatim partito vel fisso basin versus pinnato, bracteolis longioribus 4—5 mm longis, bracteis longioribus 10—13 mm longis, ealyeis lobis lon¬ gioribus 13—19 mm longis, corolla hypanthio triplo vel quadruplo longiore, staminali columna pubescenti, tubo filamentorum antheris dorsalibus triplo longiore, antheris dorsalibus longioribus 1 1-12 mm longis, antheris ven- tralibus ad apicem squama triangularis formata trichom- Volume 14, Number 1 2004 Lammers Endemic Hawaiian Genus Cyanea 91 atihus concrescentibus, et baccis longioribus 17-21 mm longis distinguenda. Trees, 4—6 m tall, producing distinctive juveniles but these not flowering precociously; stems of ju¬ veniles 0.7— 1.2 cm diam., unbranched, armed with numerous prickles consisting of a sharp trichome atop a bulbous multicellular base; stems of adults 1.6—2 cm diam. below the leaves and inflorescenc¬ es, 4—21 cm diam. at base, branching about 1 m above ground, the 4 to II branches diverging from trunk at a 45°— 60° angle, ascending, sometimes branching sparingly once again, armed with stout conic prickles below, becoming smooth toward apex, sometimes producing short shoots with juve¬ nile foliage; latex white. Juveniles with lamina el¬ liptic or oblanceolate, 13—27 cm long, 7—17 cm wide across the segments, 0.2— 0.4 cm wide be¬ tween the segments; upper surface dark green and glabrous; lower surface light green, glabrous, armed with prickles consisting of a sharp trichome atop a bulbous multicellular base; margin pinnately divid¬ ed. becoming pinnately compound at base, the seg¬ ments oblong or elliptic, 1—9 X 0.7— 2.8 cm, 8 to 10 per side, separated by sinuses, their margin pin¬ nately lobed, cleft, parted, or divided (these sec¬ ondary segments oblong or narrowly triangular, 2— 18 X 2—10 mm, the apex obtuse); apex obtuse; petiole 3—4 cm long, 0.2— 0.3 mm diam., X/-—Vi as long as the lamina, armed with prickles consisting of a sharp trichome atop a bulbous multicellular base. Adults with lamina elliptic or oblanceolate, 31—53 cm long, 10—20 cm wide across the seg¬ ments, 0.5— 4.7 cm wide between the segments; up¬ per surface dark green and glabrous; lower surface light green and pubescent on veins with short stiff curved hairs; margin pinnately parted or divided, becoming pinnately compound at base, the seg¬ ments oblong or elliptic, 1.5—12.5 X 0.5—4 cm, 10 to 14 per side, separated by sinuses, their margin pinnately cleft or parted (these secondary segments deflate or triangular, 2—22 X 2—18 mm, the apex acute or acuminate); apex acute or acuminate; pet¬ iole 4.5— 6.5 cm long, 3.5—9 mm diam., Yu—Vs as long as the lamina, minutely pubescent. Racemes 6- to 10-flowered, arising below the apical rosette of leaves, the flowers all blooming ± simultaneous¬ ly; peduncle and raehis horizontal, 35—54 mm long, 4—9 mm diam., glabrous; bracts narrowly oblong, 10-13 X 1.5-1. 6 mm, subtending the pedicels, sometimes a few empty ones on the peduncle, de¬ ciduous, rounded at apex; pedicels ascending, 9— 14 mm long, 0.8—1 mm diam., medially or suba- pically bibracteolate, glabrous; bracteoles narrowly oblong, 4—5 X 0.8— 1.2 mm. Hypanthium obconic. 10—12 mm long, 5-9 mm diam., Vg-Vs as long as the corolla, glabrous, attenuate at base; calyx lobes narrowly oblong, 13—19 X 2.6—5 mm, abutting at base but distinct, erect, glabrous, the margin entire, rounded and apiculate at apex; corolla white, 38— 47 mm long, glabrous; corolla tube arcuate, later¬ ally compressed, 23—28 mm long, 5—6 mm tall at base, narrowing gradually to 3—4 mm tall at mouth, cleft dorsal ly for ca. half its length; dorsal corolla lobes spreading, 16—19 X 1 mm, ca. % as long as tube; ventral corolla lobes deflexed. 13—15 X 1 mm; filament tube 32—40 mm long, 1.3— 1.5 mm diam., pubescent with long white hairs, free from the corolla; anther tube gently decurved, 2.3— 2.7 mm diam., densely pubescent at base and on the connectives; dorsal anthers 11 — 12 mm long, ca. Vi as long as the filament tube; ventral anthers 7.5— 8.5 mm long, their apex crowned by a triangular scale 3—4.5 mm long, formed from loosely eoneres- eent white trichomes. Berries bright orange, ellip¬ soid, 17-21 mm long, 10—15 mm diam., crowned by the persistent calyx lobes; seeds brown, broadly ellipsoid, 0.7 mm long, 0.5 mm diam., the testa Type C of Murata (1992. 1995), smooth, shiny. Distribution, habitat, and phenology. Endemic to windward East Maui in the Hawaiian Islands. Recent fertile collections are from rnesic forests on the northwestern slopes of Haleakala in the Wai‘ohiwi watershed at an elevation of 855 m; more than 50 mature trees and numerous juveniles were observed here during 2000—2002. Sterile collec¬ tions were obtained from the same general region and points to the east at slightly higher elevations (1070-1522 m) in 1911, 1920, 1927, and 1980. Sterile specimens were also collected farther east, at Klpahulu Valley, in 1919. This species flowers during October, and the fruits ripen during Decem¬ ber and January. Etymology. Cyanea duvalliorum is named for two of its discoverers, ornithologists Renate Gass- mann-Duvall (b. 1946) and her husband Fern Du¬ vall (b. 1953), who are extremely active in the Ha¬ waiian conservation community. They have brought the new species (and others) into cultivation and taken the lead in securing its preservation. Paratypes. U.S.A. Hawaiian Islands. Flast Maui. !Y1ak- awao: above Kula pipe line, Munro 724A (BISH): Hale¬ akala pipe line, Munro 724H (BISH. NY, FS), 6. C. Munro 725 (BISH); Ha‘iku Uka, Wai'ohiwi Stream, Oppenheimer et al. HI 1)005 (BISH [2 sheets plus liquid . MO. NY, OSH [sheet plus liquid], FTBG, US); Ha'iku l ka, tributary of Wai‘ohiwi Stream, Oppenheimer et al. H10012 (BISH. OSH, PTBG); Vt of "Opana Gulch across stream, War- shauer & McEldowney 2754 (BISH). liana: ridge left kl- pahulu, Forbes I72H.M (BISH), Forbes I729.M (BISH); 92 Novon trail K ol Ke‘anae & up ridge between W & E Wailua lki. Forbes 2659. M (I5I5H); Ke'anae Valley, Rock 8797 (BISH). Cyuncu maritae Lammers & H. Oppenheimer, sp. nov. TYPE: U .S.A. Hawaiian Islands: East Maui. Makawao District, Ko‘olau F. R., Wai‘ohiwi drainage basin, 2800 ft., growing above the confluence of two intermittent trib¬ utaries, near the main stream, 20°50'06"N, 156°15'52"W, degraded mesic forest amidst forestry plantings of Eucalyptus robusta & E. globulus, 26 Nov. 2000, H. Oppenheimer & F. Duvall HI 10029 (holotype, BISH; isotypes, OSH, PTBG). Figure 3. Species ob marginem foliorum maturorum pi i mat i m lis- sum partitum vel divisum, calycis lobum et fforum brac- team oblongum, et corollae tubum arcuatum compressum af finis Cyaneae duvalliorum , sed ab hac specie habit u fru- ticoso, ramis basalibus paucioribus 2-4 valde assurgen- tibus, foliis juvenilibus et maturis cum segmentis trian¬ gularis serrulatis 5—8 utrobique, petiolis foliorum maturorum proportione longioribus admodum Vs—Yz lami- nam aequantibus, hypanthio proportione breviore modum Yf— Zb corollam aequanti, calycis lobis f)revioribus 3. 5-9.5 mm longis, corolla longiore 59—61 mm longa cum lobis dorsalibus Vi—% tubum aequantibus, filamentorum tubo antheris dorsalibus quadruplo vel quintuplo longiore, et baccis parvioribus 12—13 mm longis bene distinguenda. Shrubs, 2—2.4 m tall, producing distinctive ju¬ veniles but these not flowering precociously; stems of juveniles unbranched, armed with numerous [trickles consisting of a sharp triehome atop a bul¬ bous multicellular base; stems of adults 0.7—1 .5 cm diam. toward apex, 6.3-7. 4 cm diatn. at base, pro¬ ducing 2 to 4 strongly ascending branches from near the base (sometimes unbranched), armed with stout conic prickles below, becoming smooth toward apex; latex white. Juveniles with lamina broadly el¬ liptic, 6.8—15 cm long, 4.6— 8.7 cm wide across the segments, 2.3— 3.8 cm wide between the segments, both surfaces armed with prickles consisting of a sharp triehome atop a bulbous multicellular base; upper surface dark green; lower surface light green; margin pinnately parted; segments triangular, 0.5— 2 X 0.3— 1.8 cm, 5 to 8 per side, separated by sinuses, their margin minutely serrulate, acuminate at apex; apex acuminate; base truncate; petiole 2— 4.7 cm long, 1.2—2 mm diam., ca. Vs as long as the lamina, armed with prickles consisting of a sharp triehome atop a bulbous multicellular base. Adults with lamina 17—37 cm long, 8.5—10 cm wide across the segments, 1.3—10 cm wide between the seg¬ ments, elliptic or oblong, both surfaces armed with [trickles consisting of a sharp triehome atop a bul¬ bous multicellular base; upper surface dark green; lower surface light green, glabrous or pubescent on veins with short stiff curved hairs; margin pinnately cleft, parted, or divided; segments narrowly trian¬ gular, triangular, or deflate, or shallowly triangular, often falcate, 0.6-10 X 0.8-4. 5 cm wide, 6 to 8 [ter side, separated by sinuses, their margin mi¬ nutely serrulate, acuminate or acute and apiculate at apex; apex acuminate; base cuneate or obtuse; petiole 10—13.5 cm long, 3—4.3 mm diam., V3—Y2 as long as tilt' lamina, sparsely armed with prickles consisting of a sharp triehome atop a bulbous mul¬ ticellular base. Racemes 9- to 45-flowered over their lifetime, but only 4 to 8 flowers present at any given time, arising within the apical rosette of leaves; peduncle and rachis horizontal, 45—90 mm long, 2.5—4 mm diam., sparsely armed with prickles consisting of a sharp triehome atop a bulbous mul¬ ticellular base; bracts oblong. 1.5— 2.2 X 0.8-1 mm, subtending the pedicels, deciduous, rounded or ob¬ tuse at apex; pedicels spreading, 12—25 mm long, 0.9—1 mm diam., ebracteolate, glabrous. Hypanthi- um campanulate or obconic, 9—9.5 mm long, 7-9 mm diam., */7— V6 as long as the corolla, glabrous, cuneate at base; calyx lobes oblong, 3.5— 9.5 X 1.5— 3 mm, abutting at base but distinct, erect, glabrous, the margin entire, acute or rounded and apiculate at apex; corolla white tinged with pink, 59-61 mm long, sparsely pubescent, especially on the lobes; corolla tube arcuate, laterally compressed, 42—46 mm long, 5—6.3 mm tall at base, increasing to 7— 9.2 mm tall above the middle before decreasing to 3.8— 5.5 mm tall at mouth, cleft dorsal ly for ca. Yi its length; dorsal corolla lobes spreading. 14-18 X 1.5—2 mm, Yi—Y5 as long as tube; ventral corolla lobes deflexed, I 1-14 X 0.9-1. 7 mm, connate ba- sally for 5—6 mm; filament tube 50—51 mm long, 1.8— 2.1 mm diam., glabrous, Iree from the corolla; anther tube gently decurved, 2.8— 3.4 mm diam. at base, tapering to 1.5—2 mm diam. at mouth, gla¬ brous; dorsal anthers I 1—12.5 mm long, Vc—Ya as long as the filament tube; ventral anthers 8-9 mm long, their apex crowned by a tuft of white tri - ebonies 2.5—3 mm long. Berries yellowish orange, broadly ellipsoid or obovoid, 12—13 mm long, 8.5— 10.5 mm diam., crowned by the persistent calyx lobes; seeds brown, broadly ellipsoid, 0.7 mm long, 0.5 mm diam., the testa Type C 4 >1 Murata (1992, 1995), smooth, shiny. Distribution, habitat, and phenology. Endemic to windward East Maui in the Hawaiian Islands. Recent fertile collections are from mesic forests on the northwestern slopes of Haleakala in the Wai‘ohiwi watershed at an elevation of 855 m; less than 20 adult and juvenile plants were observed here during 2000—2002. Sterile specimens were collected from the same general region and points Volume 14, Number 1 2004 Lammers Endemic Hawaiian Genus Cyanea 93 Figure 3. Cyanea marline Lammers & II. Oppenheimer. — A. Adult leaf. — B. Juvenile leaf. C. flower. (A. (, drawn from the holotype, Oppenheimer & Duvall HI 10029; B drawn from a paratype, Oppenheimer A Duvall HI 0005.) to the east in 191 1, and farther east at Klpahulu Valley in 1919. This species flowers during Novem¬ ber and December, and the berries ripen by May. Etymology. This species is named in honor of one of its discoverers, Marita Oppenheimer nee Lewis (b. 1957), long-time Maui resident and avid hiker. Paratypes. U.S.A. Hawaiian Islands. Last Maui. Mak- awao: Ha'iku Uka, Wai'ohiw i Stream, Oppenheimer & Du¬ vall HI0005 (BISH. OSH. PTBG); Ha'iku Uka, Wai'ohiwi 94 Novon Stream, Oppenheimer & Duvall H10005B (BISH, K, NY, OSH, ITBG). liana: ridge left side Klpahulu Valley, Forbes I636.M (BISH), Forbes 1 680a. M (BISH); Hono- manu Valley, Rock 8707a (BISH, (ill); ke'anae Valley, Rock 8708 (BISH, GH). I he species of the Cyanea scabra complex as defined above may be distinguished using the fol¬ lowing key. In the most recent key to the genus (Gammers, 1990a), this may be substituted for cou¬ plets 48—51 on p. 443. Key to the Species ok the Hawaiian Cyanea scabra Complex la. Bracts oblong, their apex obtuse; calyx lobes ob¬ long, their apex rounded, truncate, or emargin- ate, often apiculate. 2a. Margin of adult lamina cleft, parted, or di¬ vided. da. Trees, 4—6 m tall; stems branching at least 1 m above the base, producing 4 to 1 I spreading branches; juvenile leaves with 8 to 10 oblong or elliptic segments per side, the margin of these segments lobed, cleft, parted, or divid¬ ed; adult leaves with 10 to 14 oblong or elliptic segments per side, the margin of these segments cleft or parted; petiole of adult leaves 4.5— 6.5 cm long, y,o-K as long as the lamina; bracts 10—13 X 1.5— 1.6 mm; hypanthium 'A—Va as long as corolla; calyx lobes 13—19 mm long; corolla 38-47 mm long, the dorsal lobes ca. Va as long as the tube; filament tube 32—40 mm long, ca. 3 times longer than the dorsal anthers, pubescent; anther tube 2.3— 2.7 mm diam., densely pubes¬ cent at base and on connectives; ventral anthers tipped by a triangular scale of concrescent trichomes 3—4.5 mm long; berries 17—21 mm long, 10—15 mm diam. (bast Maui) . C. duvalliorum 3b. Shrubs, 2—2.4 m tall; stems branching near base, producing 2 to 4 strongly as¬ cending branches (rarely unbranched); juvenile leaves with 5 to 8 triangular segments per side, the margin of these segments serrulate; adult leaves with 6 to 8 narrowly triangular, triangular, del- tate, or shallowly triangular segments per side, the margin of these segments serrulate; petiole of adult leaves 10— 13.5 cm long, Va-Va as long as the lam¬ ina; bracts 1.5-2. 2 X 0.8-1 mm; hy¬ panthium 14— ‘/() as long as corolla; calyx lobes 3. 5-9.5 mm long; corolla 59-61 mm long, the dorsal lobes Va— % as long as the tube; filament tube 50—51 mm long, 4—5 times longer than the dorsal anthers, glabrous; anther tube 2.8-3.1 mm diam., glabrous; ventral anthers tipped by a tuft of free trichomes 2.5-3 mm long; berries 12—13 mm long, 8.5— 10.5 mm diam. (East Maui) . . C. maritae 2b. Margin of adult lamina dentate or denticu¬ late. 4a. Lamina elliptic' or obovate, 2. 5-4. 5 times as long as the petiole, ihe base obtuse or rounded; corolla 50—60 mm long, the tube 37—45 mm long, 3—3.8 mm tall at mouth and 4.7— 7.3 mm tall near middle; filament tube 43—55 mm long; dorsal anthers 8-8.5 mm long, '/7- V- as long as filament tube; ventral an¬ thers 5.5— 6.5 mm long, their apical hairs 3—3.8 mm long (West Maui) . . . . C. scabra 4b. Lamina narrowly oblong or oblanceo- late, 5—9 times as long as the petiole, the base cuneate; corolla 62-65 mm long, the tube 48—51 mm long, 5.5— 6.5 mm tall at mouth and 8.2-1 1 mm tall near middle; filament tube 56-60 mm long; dorsal anthers 5 mm long, '/, 2 — , , as long as filament tube; ventral anthers 4 mm long, their apical hairs 1.5—2 mm long (Kast Maui) . C. mceldowneyi lb. Bracts linear, linear triangular, or narrowly tri¬ angular, their apex acuminate; calyx lobes tri¬ angular or narrowly triangular, their apex acu¬ minate. 5a. Leal base long attenuate, narrowing imper¬ ceptibly into a winged petiole; corolla tube 45-49 mm long; dorsal anthers 5.3-6 mm long, about VH) as long as the filament tube; ventral anthers 4.2— 4.9 mm long, their api¬ cal hairs 1-1.5 mm long (East Maui) .... . C. longissima 5b. Leaf base cuneate or obtuse, rarely rounded, the petiole not winged; corolla tube 26—45 mm long; dorsal anthers 7-10 mm long, Vs- Va as long as the filament tube; ventral an¬ thers 5-8 mm long, their apical hairs 1.7- 3.5 mm long. 6a. Corolla 55—64 mm long, the tube 43— 45 mm long, 5.5—8 mm tall at base, 9— 1 1 mm tall near middle, the lobes Va— % as long as the tube; filament tube 48- 57 mm long; hairs at apex of ventral anthers 1.7—2 mm long (East Maui) . . . 6’. glabra 6b. Corolla 43-56 mm long, the tube 26- 39 mm long, 3—5 mm tall at base, 4—9 mm tall near middle, the lobes 2/--7s as long as the tube; filament tube 36—50 mm long; hairs at apex of ventral an¬ thers 2—3,5 mm long (windward Ha¬ waii) . platyphylla Cyanea sect. Deusseoideae As circumscribed by Rock (1919), Cyanea sect. Delisseoideae (Hillebrand) Rock comprises those members of the genus that are shrubs or treelets 0.3—5 m tall (rarely small branched trees to 7 m tall) with stems 0.5— 2.5 cm in diameter just below the apex, totally lacking prickles, with entire or toothed leaves, minute deciduous triangular bracts, bibracteolate pedicels, minute calyx lobes, hypan¬ thium only I/,,— V- as long as the relatively small, slender, suberect or gently curved white corolla. Volume 14, Number 1 2004 Lammers Endemic Hawaiian Genus Cyanea 95 purple berries, and small minutely papillose seeds (Type F testa; Buss et al., 2001). Of the species recognized in the most recent treatment of the ge¬ nus (Lammers, 1990a) and its supplements (Lam¬ mers & Lorence, 1993; Lammers, 1998a), the fol¬ lowing are referable to section Delisseoideae: C. angustifolia (Chamisso) Hillebrand (lectotype of the section; St. John, 1969), C. asarifolia H. St. John, C. coriacea (A. Gray) Hillebrand, C. dolichopoda Lammers & Lorence, C. elliptica (Rock) hammers, C. fauriei H. Leveille, C. habenata (H. St. John) Lammers, C. hardyi Rock, C. kahiliensis (H. St. John) Lammers, C. linearifolia Rock, C. mannii Brigham, C. membranacea Rock, C. obtusa (A. Gray) Hillebrand, and C. spathulata (Hillebrand) A. Heller. In my earlier treatment, 1 followed Rock (1917. 1919) in recognizing C. coriacea and C. fauriei as distinct anti largely allopatric species, distin¬ guished primarily by inflorescence length: plants of northern Kaua‘i with short spreading or ascending condensed racemes were called C. coriacea, while plants of southern Kaua'i with long pendent elon¬ gate racemes were called C. fauriei. However, field studies on Kaua‘i in 1991 and detailed examination of a larger body of herbarium specimens convinced me that many of the long-racemed plants I had identified as C. fauriei were otherwise indistin¬ guishable from C. coriacea. Variation in raceme length among these plants is continuous and essen¬ tially clinal. In northern Kaua‘i (e.g., on the Napali Coast) are found populations with relatively short often ascending racemes (cf. Rock. 1919, pi. 109). These lengthen as one proceeds south, reaching their greatest development with the highly elongate and pendent racemes found in Olokele Canyon (cf. Rock, 1919, pi. 1 10). It is worth noting that Rock himself had earlier (Rock, 1913. 1914) treated the two names as synonymous, while Wimmer (1943) treated C. fauriei as a variety of C. coriacea, a clas¬ sification embraced by Degener and Degener (1956) and St. John (1973. 1980). However, not all specimens that I had identified as C. fauriei resemble C. coriacea. Many of the plants with relatively long pendent racemes differ from C. coriacea in a number of characteristics, in¬ cluding slimmer twigs and smaller leaves. In these features, they more closely resemble C. hardyi ; in fact, some of these specimens were previously iden¬ tified as that species. However, these smaller- leaved plants differ from C. hardyi in several fea¬ tures, including their broader leaf base, shorter rachis and pedicels, shorter hypanthium and calyx lobes, and shorter filament tube. Furthermore, one specimen previously identified as C. fauriei differs from all these plants in its extremely small flowers, e.g., hypanthium only 2.3— 2.7 mm long, corolla just 15—16.5 mm long, filament tube 12—14 mm long, and anthers (dorsal pair) 4.5—5 mm long. In fact, these are the smallest known flowers in the entire genus, by a significant margin. It thus appears that three distinct species were confounded under the name C. fauriei: the long-racemed southern popu¬ lations of C. coriacea, plants with slimmer tw igs and smaller leaves resembling C. hardyi, and a plant with the smallest flowers in the genus. In order to determine which of these takes the name C. fauriei, it was first necessary to clarify the typification of that name. In the protologue, Leveille cited two gatherings: “Kauai: Koloa; Waimea, 1000 m, dec. 1909, mart. 1910, (Faurie, 565, 591).” These are thus syntypes from which a lectotype must be selected. Rock (1913, 1914, 1919) appar¬ ently only saw Faurie 565, but never referred to it as the type. Wimmer (1943) saw both gatherings and specifically excluded Faurie 591 from C. cor¬ iacea var. fauriei by referring it to C. coriacea var. hardyi (Rock) E. Wimmer, a homotypic synonym of C. hardyi. Like Rock, he did not specifically state that Faurie 565 was the type of C. fauriei. As a result, neither action can be interpreted as an ef¬ fective lectotypification under Article 7.11 of the ICBN (Greuter et al., 2000). The specimens upon which Leveille based his new taxa were deposited in his personal herbarium, which was sold to E in 1919 (Stafleu & Cowan. 1979). Staff members of that institution have com¬ piled a catalogue of all names ever published by Leveille, together with information on typification (cf. Lauener, 1983). In this catalogue, even' one of the Hawaiian Lobelioideae is listed as “typus — n[on]. v [idi].,” even though types for nearly all Chi¬ nese Campanulaceae were seen (Chamberlain, 1977). Indeed, when I requested a loan of Levei lie's Hawaiian lobelioid types from E in 1985. I received only a set of photographs showing specimens at BM. and a note stating that no actual specimens were to be found at E. Lauener (1980) referred to the specimens at BM as isotypes (or syntypes, if two or more numbers had been cited in the proto¬ logue), but they appear to be the original specimens from Leveille’s personal herbarium. I have been un¬ able to discover why these specimens are at BM and not E with the remainder of Leveille's personal herbarium. In any event, both Faurie 565 and Faurie 591 were found at BM. each represented by a photo¬ graph at E. As noted bv Wimmer (1943), the latter gathering does not conform to the original diagnosis of C. fauriei and is referable to C. hardyi. For this 96 Novon reason, C. fauriei is here leetotypified on the basis of Faurie .565, which does conform to the diagnosis. Because lliis specimen represents the long-raeemed southern populations of C. coriacea, this binomial is here treated as a synonym of C. coriacea (follow¬ ing Rock, 1913, 1914), along with several varieties of that species that do not merit recognition (cf. hammers. 1990a). All are listed here for clarity. Cyanea coriacea (A. Gray) Hillebrand, FI. Ha¬ waiian I si. 254. 1888. Delissea coriacea A. Gray, Proc. Amer. Acad. Arts 5: 147. 1861. TYPE: U.S.A. Hawaiian Islands: Kaua‘i, 1851-1855, ./. Remy 302 (holotype, GH; iso¬ types, A, L, P[5], /). Cyanea fauriei H. Reveille, Repert. Spec. Nov. Regni Veg. 10: 156. 191 1. Cyanea coriacea var. fauriei (H. Rev¬ eille) K. Wimmer, Pflanzenr. IV.276b: 72. 1943. De¬ lissea fauriei (H. Reveille) II. St. John, Phytologia 63: 342. 1987; non H. Reveille, Repert. Spec. Nov. Regni Veg. 12: 505. 1913. TYPE: U.S.A. Hawaiian Islands: Kaua‘i, Koloa, Dec. 1909, V. Faurie 565 (lectotype, selected here, BM, photograph, E; isolec- totypes. A, BISH, G, P, W). Cyanea coriacea var. degeneriana E. Wimmer, Pflanzenr. 1V.276I): 761. 1953. Delissea coriacea var. degener¬ iana (E. Wimmer) H. St. John, Phytologia 63: 82. 1987. TYPE: U.S.A. Hawaiian Islands: Kaua'i. ridge behind Papa'a, It) Jan. 1952, O. Degener & A. Greenwell 21736a (holotype, W; isotypes, B[2|). Delissea coriacea var. deltoidea II. St. John, Phytologia 63: 340. 1987. Syn. nov. Delissea perlmanii II. St. John, Phytologia 63: 346. 1987 (as 'perlmannii'). TYPE: U.S.A. Hawaiian Islands: Kaua’i. Eimahuli Valley, common, 8 Sep. 1977, S. Perlman I (holotype, BISH; isotypes, BISII[2]). Delissea coriacea var. haupuensis H. St. John, Phytologia 63: 340. 1987. Syn. nov. TYPE: U.S.A. Hawaiian Islands: Kaua'i, Hoary Head Range, near 1286 ft. peak N of Ra'aukahi, E side of range, dryland forest, I 100 It.. 3 Nov. 1976, C. Christensen 91 (holotype, BISH; isotype. BISH). Delissea coriacea var. lumahaiensis II. St. John, Phytologia 63: 340. 1987. TYPE: U.S.A. Hawaiian Islands: Kaua'i, road between Hanalei & Lumaha'i, Sep. 1913, C. I\. Forbes & II. ./. Forbes 464. K (holotype, BISH; isotypes, BISH. GB, K. NY, P, UC, US). The two sets of populations that were formerly identified as C. fauriei hut are not referable to C. coriacea are described here as new species. Though no names at species rank are available, a name at the rank of forma is referable to one species as here circumscribed. However, I explicitly decline to take up this epithet. Though its type falls within the cir¬ cumscription of the species, its original description does not address the characters emphasized here. For the sake of clarity and precision, I prefer to exercise my prerogative and treat the species as new, with its own diagnosis. Cyanea pseudofauriei Rammers, sp. nov. T\ BE: U.S.A. Hawaiian Islands: Kaua'i. Koloa Dis¬ trict. Rlhu'e-Koloa Forest Reserve, along ridge ESE of Pu'u Kolo, lower ridge in diverse forest with rather stunted vegetation on the windward slopes of the ridge, common on windward slope, elev. 1200-1700 ft.. 21 Sep. 1988, T. Flynn. G. Kawakami & R. Nagata 3144 (ho¬ lotype. F; isotype, PTBG). Figure 4A, B. Cyanea coriacea f. gratiosa E. Wimmer, Pflanzenr. IV.276R: 760. 1953. Delissea coriacea var. gratiosa (E. Wimmer) II. Si. John, Phytologia 63: 82. 1987. TYPE: U.S.A. II awaiian Islands: Kaua'i, ridge be¬ hind Papa'a, Moloa'a Forest Reserve, 16 Jan. 1952. 0. Degener & A. Greenwell 21737 (lectotype, select¬ ed here, B; isolectotype, W). I'he protologue of Cyanea coriacea I. gratiosa cit¬ ed "Kauai: Ridges behind Papaa in lower forest, lf>. 1. 1952 (lib. Deg. n. 21736! common form, 21737! rare form intermediate in coloration betw. n. 21736 and n. 21738).” I have seen sheets of Degener c, I). Stone, P. Manos & A. L Thomas: DKH 576 (HN, LE, MO); Moc Chau Distr., Cho Long, Vu Xuan Phuong 79 (1IN). Thanh Iloa Prov.: Ba Thuoc Distr., Co Lung Mun., N. T. Hiep, L. Averyanov & P. K. Ijoc: HAL 948 (HN, LE, MO). Acknowledgments. The study was supported by the Institute of Ecology and Biological Resources, National Center for Science and Technology, Hanoi, Vietnam. I am grateful to Le Xuan Canh and Nguy¬ en l ien Hiep for their encouragement and advice. I thank Roy Gereau for the Latin diagnosis, Dan Harder, Ihsan Al-Shehbaz, and Charlotte M. Taylor for helping with the English, and Ruth Kiew for providing distribution information. The United States National Science Foundation is thanked for a grant (DEB-9870231 to 1). k. Harder) that sup¬ ported the fieldwork. I .iterature Cited Doorenbos, J., M. S. M. Sosef & J. J. F. I',, de Wilde. 1998. The Sections of Begonia including Descriptions, Keys and Species Lists (Studies in Begoniaceae VI). Wag- eningen Agricultural University, Netherlands. Gagnepain, F. 1919. Nouveaux Begonia d’Asie; quelques synonymes. Bull. Mus. llist. Nat. Baris 25: 194—201, 276-283. - . 1921. Begun iacties. In: F. Gagnepain (editor), Flore Generate de L'lndoehine 2(8): 1095—1120. Mas¬ son et Cie, Paris. Ku. K. 1999. Begoniaceae. FI. Reipublicae Popul. Sin. 52(1): 126-268. Science Press, Beijing. Pham, H. H. 1999. Begoniaceae. In: H. H. Pham (editor). Cay co Viet Nam. I: 577-588. Tre Publishing House, Vietnam. Slnii Yu Min, Ching I Peng & Cheng Yih Wu. 2002. Syn¬ opsis of the Chinese species of Begonia (Begoniaceae), with a reappraisal of sectional delimitation. Bot. Bull. Acad. Sin. 43: 313-327. Swenson. S. M. & M. C. Tebbitt. 2001. Evolution, bioge¬ ography and systematics of Begoniaceae. Botany 2001: Plants and People. Albuquerque: 93. | Abstract. | - . W. L. Clement, L. L. Forrest & M. C. Tebbitt. 2001. Hillebrandia sandwichensis : Evolutionary rela¬ tionships and biogeography. Botany 2001: Plants and People. Albuquerque: 95. [Abstract. | Wan Y. & B. N Chang. 1987. A new species of the genus Begonia from Guangxi. Acta Phytotax. Sin. 25(4): 322— 323. A New Species of Ceratolacis (Podostemaceae) from Minas Gerais, Brazil C. Thomas Philhrick Department of Biological and Environmental Sciences, Western Connecticut State University, Danbury, Connecticut 06810, U.S.A. philbrickt@wcsu.edu Alejandro Novelo R. Departamento de Botanica, Institute de Biologfa, Universidad Nacional Autonoma de Mexico, Mexico. D.F., 04510, Mexico, lanovelo@servidor.unam.mx Bruno E. / r gang Departamento de Botanica, Universidad Federal do Rio Grande do Sul. Avenida Bento Goncalves, 9500-Bairro Agronomia, 91570-950, Porto Alegre, Rio Grande do Sul. Brasil Abstract. A new species ol Ceratolacis ( C . pe- dunculatum, Podostemaceae) is illustrated and de¬ scribed. I lie new species is known from four lo¬ cations in central Minas Gerais, Brazil. Ceratolacis pedunculatum is distinguished from the only other species of the genus based on the presence of a prominent peduncle and the oblique orientation of the ovary on the pedicel. Rksumo. Uma nova especie de Ceratolacis ( C. pe¬ dunculatum, Podostemaceae) e descrita e ilustrada. Ksta especie nova e conhecida de quatro local ida- des no centre plo di¬ visa, rare subdichotoma vel pinnata; stipulae asymmetri- cae, ex folii base amplexicauli cymbiformi constatae. In- florescentia uniflora, pedunculata. Flores solitarii in axillis fnliarihus. Spat In 41a claviformis. Pedunculus et pedicellus post spathellae rupturam elongautes. Tepala 3, uno in an- dropodii quoque latere, tertio tepalo ad andropodii apicem portato. Stamina 2. Andropodium et filamenta peranthesin elongantia. Pollinis grana in dyadibus. Ovarium ex 2 car- pellis aequalibus compositum, 2-loculare, in pedicello oblique disposilum. Stigmata 2, libera, stigmatum base in fructu indurata, persistenti. Fructus capsularis, 2-locular- is, apice emarginato, duabus valvis aequalibus, persisten- tibus, 3-costatis, marginibus suturalibus incrassatis. Pe- dunculus a pedicello in fructu non distinguibilis. Aquatic herbs, perennial. Roots green, elongate and cylindrical when young, becoming broad and crust-like, irregular in outline when older, branched, prostrate, flattened to elliptical in cross section, median = 0.9 (0.3-5. 2) mm |N = 50] wide, tightly attached to rocks via adhesive hairs, growing intertwined. Stems monomorphic, arising opposite or subopposite along the Hanks of roots, 4.9 (0.9- Volume 14, Number 1 2004 Philbrick et al. 109 Ceratolacis pedunculatum from Minas Gerais 12) mm | IN = 50] apart, obscured by leaf bases, erect, less than 0.1 mm high, unbranched. Leaves petiolate, distichous, stipulate, double-sheathed, upright, compound, 4 (1-6) [N = 50] times divided, rarely subdiehotomous or pinnate, 17 (2.9—35.2) mm [N = 100] long, ultimate leaf segments linear to spatulate, 3.1 (0.2-9.5) mm |N = 100] long X 0.3 (0.1-1) mm [N = 100| wide, midvein faint or lacking, apices acute or obtuse; petiole 3.9 (0.4— 14) mm | IN = 100] long, elliptical in cross section; stipules asymmetric, composed of an extension of the amplexicaul leal base, 0.5 (0.1— 5.6) mm [N = 50] long, larger on leaves subtending flowers, apex entire. Inflorescence with flowers pedunculate and solitary. Flowers 2 (1—6) [N = 50] per stem, her¬ maphroditic, zygomorphic, borne singly in a leaf axil, covered by a spathella, pedicellate; peduncle (below attachment point of spathella) 0.8 (0.1— 3.1) mm [N = 100] long prior to spathella rupture, elon¬ gating to 2.6 (0.3— 8.5) mm |N = 100] long after spathella rupture; spathella clavate, 2.4 (1-4.5) nun [N = 100] long X l.l (0.6— 1.7) mm |N = 100] wide, rupturing apically, falling away completely in fruit; pedicels 0.8 (0.1— 3.1) mm [IN = 100] long prior to spathella rupture, elongating to 5.1 (1.8 — 8.5) mm [N — 50] long after spathella rupture; te- pals 3, seale-like, linear, acute, two (one on either side) at the base of the' stalk from which the sta¬ mens arise (andropodium), 0.6 (0.2— 1.8) mm [N = 50] long; a third tepal occurring at the apex of the andropodium, andropodial tepal 0.4 (0.3— 0.5) mm |N = 50] long; stamens 2. deciduous; andropodium I (0.6—2) mm [N — 50] long prior to spathella rup¬ ture, during anthesis elongating to 4.3 (2-7) mm [N = 50] long; filaments 0.4 (0.2-1. 8) mm [N = 50] long prior to spathella rupture, during anthesis elongating to 0.9 (0.4-1. 8) mm [N — 50] long; an¬ thers quadrangular, with parallel sides, 1.1 (0.7— 1.4) mm [N = 50] long, I (0.6-1. 2) mm [N = 50] wide, dehiscing introrsely anti longitudinally, inner and outer thecae of equal length; pollen in dyads, tricolpate, 35 (31-40) /am |N = 100] long. 24 (19— 28) /tun [N = 100] wide; ovary with 2 equal carpels, 2-loeular. oval in shape, oriented obliquely on the pedicel, 1.5 (0.7— 2.6) mm [N = 50] long, 1 (0.6— 1.7) mm |N = 50] wide, with 6 longitudinal dark lines on the ovary wall; ovules 20 (12—32) [N = 25]; stigmas 2, free, each stigma triangular in out¬ line, widest at base, upright prior to spathella rup¬ ture, 0.5 (0.3—1) mm [N = 50] long, during anthesis upright or divergent, elongating to 0.9 (0.4— 1 . 1) nun [N = 50] long, base of stigmas becoming hardened and persisting in fruit. Fruit a 2-locular capsule, 1.8 (1.4— 2.1) mm [N = 50] long, 1.3 (0.9— 1.5) mm |N = 50] wide, capsule apex emarginate due to persistent stigma bases, with two equal valves, per¬ sistent, valves 3-ribbed, suture margins thickened and rib-like; peduncle not distinguishable from pedicel in capsule (due to shedding of spathella and parenchymatous tissues of peduncle and ped¬ icel); length of peduncle and pedicel combined 6.8 (1.5—17) mm [N = 100] long. Mature seeds un¬ known. Ceratolacis pedunculatum is known from four lo¬ cations in the central part of the state of Minas Gerais, Brazil (Fig. 2), ranging from about 500 to 650 m elevation. The related C. erythrolichen is known only from the type collection (rio Tocantins), which is approximately 1500 km north of the lo¬ cations where C. pedunculatum was collected. Al¬ though the type locality of C. erythrolichen was in the state of Goias at the time the species was de¬ scribed in 1849, the location is presently in the state of Tocantins. Additional observations. Two reproductive char¬ acteristics distinguish Ceratolacis pedunculatum from C. erythrolichen. Ceratolacis pedunculatum possesses a peduncle (Fig. lb, le, lg, lb), and the axis of the ovary is offset (oblique) from that of the pedicel by ea. 30—45° (Fig. If— h). In contrast. C. erythrolichen lacks a peduncle and the ovary axis is parallel to that of the pedicel. One characteristic of the root also seems to differ between C. erythrolichen and C. pedunculatum. Tu- lasne (1852: 126) and Engler (1930: 52) referred to the roots of C. erythrolichen as being red while living and turning blackish or whitish upon drying. This character is not evident in C. pedunculatum. in which the roots are dark green in living and dried specimens. Justification for recognizing Ceratolacis pedun¬ culatum as distinct from C. erythrolichen warrants some discussion of the type material of the latter species. Tulasne (1849) based his description of C. erythrolichen on a collection of Weddell. As Wed¬ dell worked at Paris and his collections are housed there, it has been reported (Tulasne, 1852: 127; Van Royen, 1954: 224) that the type material of C. erythrolichen is deposited in P. Recently, however, the type specimen has not been located there. There is one specimen located in G that seems to be the holotype. Isotypes are located in K and M. The holotype and isotypes are comprised of only vegetative material (roots, stems, leaves). As a con¬ sequence, it has not been possible to study the re¬ productive structures on the type material for C. erythrolichen. Van Royen (1954) also noted that the type material was incomplete. Ceratolacis erythrolichen is only known from the 110 Novon Figure 1. CeratolacLs pedunculatum C. T. Philbrick, Novelo & Irgang. Drawings based on the holotype. — a. General habit of plant showing cylindrical root from which leaves arise; steins are obscured by base of leaves. — b. General habit of plant showing cylindrical root, short stem (obscured by sheathing leaf liases), and two flowers — one at anthesis, the other enclosed within an intact spathella. Also showing dichotomously and subdiehotomously divided leaves. — c. Details of a double-sheathed leaf with two basal stipules (left) and two simple leaves on a short stem (right). — d. Detail of three leaves showing asymmetric stipule on left side of leaf base and dichotomous to subdiehotomous divisions. A detail of the apical segment of a leaf is included. — e. Flower enclosed within a spathella, which is partially ruptured at the apex. Apex of the peduncle is indicated by the arrow. — f. Two views of a flower at anthesis showing the Volume 14, Number 1 2004 Philbrick et al. Ill Ceratolacis pedunculatum from Minas Gerais type collection. Thus, the present authors have had to rely on t lie descriptions and illustrations in Tu- lasne (1849, 1852. 1855) for their understanding of the reproductive morphology of the species. In the protologue for Ceratolacis erythrolichen (as Di- craea erythrolichen) Tulasne (1849: 102) did not make reference to a peduncle (a stalk below the attachment point of the spathella), nor is one in¬ dicated in the illustration for the species (his PL. X. 1). Rather, Tulasne (1849) referred only to a pedicel. (In Podostemaceae the pedicel is the stalk of the ovary that arises above the attachment point of the spathella.) Subsequently, Tulasne provided an expanded description of the species in his monograph of Podostemaceae (Tulasne, 1852: 12b— 128) and again in Flora brasiliensis (Tulasne, 1855: 253—254), but in neither case is any reference to a peduncle made. Although illustrations of the spe¬ cies are included in these two later works, they are the same illustrations that accompanied the 1849 description of the species. Given the prominent na¬ ture of the peduncle, the authors contended that Tulasne would have noted a peduncle if one had been present in the type material. Thus, the authors are confident that C. erythrolichen lacks a pedun¬ cle. Ceratolacis pedunculatum possesses a distinct peduncle. The peduncle occurs below the attach¬ ment of the spathella (arrows in Fig. le, lg). while the pedicel occurs above. Lastly, Tulasne (1849, 1852, 1855) did not make reference to the ovary of C. erythrolichen being oblique on the pedicel either in his species description or in the accompanying illustrations. Based on the presence of a peduncle, the oblique orientation of the ovary, and to a lesser degree on the green roots, the authors conclude that C. pedunculatum is distinct from C. erythrolichen. Species in three genera of Podostemaceae in the New World possess two stamens, an andropodium, and an asymmetrical sheathing leaf base (Table 1): two species of Ceratolacis , all species of Crenias, and one species of Podostemum (P. muelleri Warm¬ ing). It is thus appropriate to focus on these eight species in a discussion of the characteristics of Cer¬ atolacis pedunculatum. The nature of the leaf base is partially correlated with stipule form among species of Ceratolacis, Crenias, and Podostemum. A single asymmetrical Distrito Federal; RS, Rio de Janeiro; ES, Espfrito Santo. stipule (a stipule on one side of the leaf base) and subdichotomous or pinnate leaves occur in species of Ceratolacis, Crenias, and one species of Podos¬ temum (P. muelleri). In contrast, a symmetrical boat-shaped stipule (entire or divided into apical teeth) occurs in all species of Podostemum (except P. muelleri and P. irgangii C. T. Philbrick & Nov- elo); these species all have dichotomously divided leaves. Podostemum muelleri deviates from this cor¬ relation with its dichotomously divided leaves and asymmetrical stipule, whereas leaves of P. irgangii are uniformly verticillate. Ceratolacis pedunculatum is the only New World species that has an asymmetrical stipule, an andro¬ podium, two stamens, a pedicel, and a peduncle. The peduncle (0. 1-8.5 mm long) occurs below the location where the sac-like spathella attaches to the pedicel (arrows in Fig. le, lg). In contrast, the spa- thellas of species ol Podostemum and Crenias, in¬ deed most other species of Podostemoideae in the New World, attach at the base ol the pedicel — at the point of attachment to the stem. Thus a pedun¬ cle is lacking. A peduncle is known from only three species of Podostemaceae in the New World: Cer¬ atolacis pedunculatum, Macarenia clavigera P. Roy- en, and Vanroyenella plumosa Novelo & C. T. Phil¬ brick. Characteristics of the capsule distinguish species andropodium (stalk from which stamens arise), two anthers, two basal tepals, and one tepal arising from the apex of the andropodium. A detail of one anther is also shown. — g. A flower at anthesis showing its orientation relative to the root. Arrow indicates the apex of the peduncle. — h. A post-anthesal flower showing the remnants of the spathella arising from the apex of the peduncle. — i. Mature capsule prior to dehiscence showing the persistent and divergent stigma bases and the isolobous capsule. — -j. Mature capsule after dehiscence (seeds have been dispersed) showing the two persistent capsule valves. 112 Novon of Ceratolacis from Crenias and Podostemum. The capsule in the latter two genera is cylindrical in cross section, with a rounded or hlunt apex. In con¬ trast, the capsule of Ceratolacis is somewhat flat¬ tened with an emarginate apex, a character that is largely a consequence of the persistent stigma lia¬ ses. The stigma liases are deciduous in species of Crenias and Podostemum. In addition, species in these two genera possess capsules in which one valve is deciduous, the other persists. In contrast, I nit li valves of die capsule of species of Ceratolacis persist. Thus, with capsules alone it is possible to distinguish plants of die genus Ceratolacis from those of other genera of Podostemaceae. Species of Ceratolacis possess an isolobous ova¬ ry, while die ovaries of species of Crenias and Po¬ dostemum are anisolobous. The presence of an an- isolobous ovary is usually associated with the ovary set at an angle (oblique) on the pedicel. However, Ceratolacis peduncidatum is unusual in that the is¬ olobous ovary is clearly oblique on the pedicel (Fig. If-h). There is no indication that C. erythrolichen shares this character. Variation in stem and root form also helps dis¬ tinguish species of Ceratolacis. Elongate stems characterize most species of Podostemum and Cren¬ ias, although stems of some species can be ob¬ scured by the sheathing leaf bases. Ceratolacis pe- dunculatum lacks an obvious upright stem; the diminutive stem is obscured among the leaf bases. Roots of species of Ceratolacis, Crenias, and Po¬ dostemum are elongate and cylindrical when young. Roots of mature plants retain their cylindrical form in species of Podostemum and Crenias. In contrast, mature roots of both species of Ceratolacis often become broadened, irregular in outline, flattened, and can resemble a crust. These flattened roots are difficult to remove from the rocks. Thus, to docu¬ ment this character it is necessary to collect spec¬ imens and the rocks to which they are attached. Species of Crenias and Podostemum possess dvad pollen. Dyad pollen also characterizes Cera¬ tolacis pedunculatum. ’file authors predict that C. erythrolichen has dyad pollen as well, although the absence of adequate type material does not allow this prediction to be confirmed. Paratypes. BRAZIL. IVIinas Gerais: on road from l)i- amantina (Gouveia) to Curvelo, confluence of rio Parauna and rio CipA, 0 July 2002, C. T. Philbrick, A. Novelo A'., It. P. Irgang & C. V. de Senna Gastal 5649 (BHCB. IGN, MKXU, \1(). WCSU); ca. 10 km W of Sao Sebasliao, Route BR-420, on road to Pompeu, 0 July 2001, C. T. Pliilbriek. A. Novell) R., R. E. Irgang & C. V. de Senna Gasta! 5651 (BHCB, 1CN, MKXU, MO. WCSU); ca. 16 km SW of Pompeu, along BR-262 on road to Martinhos Campos, rio Para. 16 July 2001, C. T. Philbrick. A. Novell > Volume 14, Number 1 2004 Philbrick et al. 113 Ceratolacis pedunculatum from Minas Gerais R., B. E. Irgang & C. V. de Senna Cast a l 5678 (BHCB, ICN, MKXU, MO, NY, WCSU); 2 Aug. 2002, C. T. Phil- brick & A. Nor do R. 5781 (BHCB. ICN, MKXU. MO. NY. WCSU); 30 km N of Corinto on BR-496, ca. 200 m up¬ stream of Bridge, rio Bicudo, 28 July 2002, C. T. Philbrick & A. Novelo R. 5761 (BHCB, ICN, MKXU, MO, NY. WCSU). Acknowledgments. We thank Fernando Chiang for the Latin description and Albino Luna for draw¬ ing Figure I. Rolf Rutishauser and an anonymous reviewer are thanked for their helpful comments on an earlier version of the manuscript. Claudio V. de Senna Gastal is thanked for his assistance and en¬ couragement while conducting field studies in southern Brazil. This work was supported by Na¬ tional Science Foundation Grant DEB-9629767 to C.T.R and Donald H. Les, and research grants from the Connecticut State University. The follow ing her¬ baria are thanked for providing access to, or loans of, specimens: B, BHCB, C, ICN, K, L, M, MFXU, NY, P, S, US, WCSU. We offer special thanks to the curatorial staff ol C, K. L, and P for their as¬ sistance. Literature Cited Engler, A. 1930. Podostemonaceae. In: A. Kngler & K. Prantl (editors), Nat. Pffanzenfam. ed. 2 18a. 1-68. 483-484. Tulasne, I.. R. 1849. Podostemacearum synopsis mono- graphica. Ann. Sci. Nat. But. Ser. 3, It: 87—1 14. - . 1852. Monographia Podostemacearum. Arch. Mas. Hist. Nat. 6: 1-208. - . 1855. Podostemaceae. In: Martius, FI. bras. 4(1): 229-276. Royen, P. Van. 1954. The Podostemaceae of the New World. Iff. Acta Bot. Need. 3: 215-263. Weddell. 11. A. 1873. Podostemaceae. In: DC., Prodr. 17: 39-89, 300-301. Validation of Ardisia Section Auriculardisia (Myrsinaceae) Jon M. Ricketson Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. jon.ricketson@mobot.org John J. Pipoly III Fairchild Tropical Garden, 1 1935 Old Cutler Hoad, Coral Gables (Miami), Florida 33156-4242, U.S.A. jpipoly@faircbildgarden.org ABSTRACT. In the recent revision of Ardisia subg. Auriculardisia, the new status, combination, and basionym for Ardisia sect. Auriculardisia were er¬ roneously omitted. The section is validated here. Key words: Ardisia seel. Auriculardisia, Myrsi- naceae. Because the differences used by l.undell (1981) to separate his segregate genus Auriculardisia from Ardisia were found to be unreliable, we included Auriculardisia within the genus Ardisia and revised its taxonomy. However, in our recent revision of Ardisia subg. Auriculardisia (Ricketson & Pipoly, 2003: 203), the new status, combination, and its basionym for Ardisia sect. Auriculardisia were er¬ roneously omitted. That error is here corrected: Ardisia sect. Auriculardisia (Lundell) Ricketson & Pipoly, comb, et stat. nov. Basionym: Auri¬ culardisia Lundell, Phytologia 49: 341. 1981. TYPE: Ardisia glomerata Lundell. I .iterature Cited l.undell, C. I,. 1981. Neotropical Myrsinaceae — IV. I’liy- tologia 49: 341—354. Ricketson, J. M. & J. J. Pipoly III. 2003. Revision of Ardisia subgenus Auriculardisia (Myrsinaceae). Ann. Missouri Bot. Card. 90: 179—317. Novon 14: 114. 2004. Reinstatement of Bonellia (Theophrastaceae) Her til Stahl Department of Science and Technology, Gotland University, SE 62167 Visby, Sweden. bertil.stahl@hgo.se Mari Kallersjo Molecular Systematics Laboratory, Swedish Museum of Natural History, P.0. Box 50007, SE 10405 Stockholm, Sweden, mari.kallersjo@nrm.se Abstract. Following the results of a recent phy¬ logenetic study showing that the genus Jacquinia is paraphyletic, the following new combinations in the genus Bonellia are made (all by Stahl & Kallersjo): B. albiflora (Lundell), B. brevifolia (Urban), B. flammea (Millspaugh ex Mez), B. frutescens (Miller), B. lippoldii (Lepper), B. longifolia (Standley), B. loeflingii (Carrasquel), B. macrocarpa (Cavanilles), B. macrocarpa (Cavanilles) Stahl & Kallersjo suhsp. pungens (A. Gray), B. macrocarpa (Cavan¬ illes) Stahl & Kallersjo suhsp. panamensis (Lun¬ dell), B. montana (Stahl), B. mucronata (Roemer & Schultes), B. nervosa (Presl), B. nitida (Stahl). B. paludicula (Standley), B. pringlei (Bartlett), B. pau- ciflora (Stahl & F. Axelrod), B. shaferi (Urban), B. seleriana (Urban & Loesner ex Mez), B. sprucei (Mez). B. stenophylla (Urban), and B. umbellata (A. DC.). The genus Bonellia is distinguished from Jac¬ quinia s. str. by its mostly orange flowers and al¬ ternate leaves, flattened seeds that are incompletely covered by placental tissue, and two-layered testa. A key to the genera of the Theophrastaceae is pre¬ sented. Key words: Bonellia , Theophrastaceae. The Theophrastaceae form a small Neotropical family of about 100 species of shrubs and small trees. Recent phylogenetic research (Kallersjo et al., 2000) has shown that Samolus L., a genus tra¬ ditionally included in the Prirnulaceae, is sister group to the Theophrastaceae and therefore should he included within the latter family or recognized as a separate family. The Theophrastaceae s. str. (excluding Samolus) were monographed by Mez (1903), who recognized four genera based mainly on floral morphology, viz. Clavija, Jacquinia, Deherainia, and Theophrasta. Votsch (1904), in a leaf anatomical work, trans¬ ferred one of the species of Deherainia to a new genus, Neomezia, and during more recent taxonom¬ ic work (Stahl, 1993), the genus Votschia was es¬ tablished to accommodate a species previously in¬ cluded in Jacquinia. Following a new analysis based on a combination of morphological and epDNA data (Kallersjo & Stahl, 2003), it has become clear that the genus Jacquinia is not monophyletic and has to be divid¬ ed into two genera. According to that analysis, Jac¬ quinia s. str. should include species with pseudov- erticillate leaves lacking or with a poorly developed niucro, lepidote young shoots, whitish flowers, glo¬ bose seeds completely immersed in placental tis¬ sue, and sunken abaxial foliar sclerenchyma. This circumscription makes Jacquinia a chiefly Carib¬ bean genus of 13 species. Only one of these spe¬ cies. J. armillaris Jacquin, occurs outside of the Caribbean islands, in northern coastal South Amer¬ ica and eastern coastal Brazil (Stahl. 1992. 1995). Jacquinia arborea Vahl, with a wide Caribbean dis¬ tribution, has also been reported a few times from Mesoameriea, but then only from islands off the eastern coast of Mexico and Honduras. The reinstated genus Bonellia is defined by al¬ ternate leaves with a well-developed mucro, gla¬ brous or puberulous young shoots, mostly orange flowers, flattened seeds only partly immersed into placental tissue, and abaxial foliar sclerenchyma located adjacent to the lower epidermis. Two spe¬ cies, B. longifolia and B. paludicola, deviate by having whitish flowers, and it is therefore interest¬ ing to note that both appear at the base of the Bo¬ nellia clade in the phylogeny proposed by Kallersjo and Stahl (2003). Of these, B. paludicola is the most problematical species since it, like most mem¬ bers of Jacquinia s. str. (except J. Iceyensis Mez), has pseudoverticillate leaves. It should also be not¬ ed that one additional species, B. albiflora, has white flowers, but that species is morphologically very similar to B. macrocarpa, indicating a close relationship between these two taxa. The genus Bonellia consists of 22 species dis¬ tributed in Mesoameriea, northern and western Novon 14: 115-118. 2004. 116 Novon South America, and the Greater Antilles. Within the genus two morphologically and geographically distinct groups can be discerned. One of these com¬ prises the South American species B. mucronata (northern Peru), B. sprucei (Ecuador), B. frutescens (northern Colombia— Venezuela), B. loefiingii (north¬ ern Venezuela), and the Mesoamerican B. nervosa. All species in this group have rather large, thick- walled and opaque fruits, comparatively flexible leaves (deciduous in B. nervosa ), and whitened young shoots. The second group consists of species from Cuba, Hispaniola, and Puerto Rico (B. brevi- folia, B. lippoldii, B. pauciflora, B. stenophylla, B. umbellata) with small or very small spine-tipped leaves, small, thin-walled fruits, and tiny flowers. I'he remaining, Mesoamerican species are more dif¬ ficult to define as a natural group, although some species appear to be closely related (Stahl. 1989; Kallersjo & St&hl, 2003). fhe name Bonellia was published by Colla (1824) in a catalog of plants from his private gar¬ den. I’he name was evidently proposed by C. G. F. Bertero, whose material of a plant from Santa Mar¬ tha in northern Colombia was distributed under the name of Bonellia cavanillesii to herbaria and bo¬ tanical gardens in Europe. However, since Colla (1824) cited Jacquinia macrocarpa Cavanilles, a different Mesoamerican species, as a synonym, the name B. cavanillesii has to be treated as an obligate synonym of the latter species. The name Bonellia honors the Italian zoologist Franco Andrea Bonelli (1784—1830), evidently a close friend of Bertero and Colla, but whose con¬ tribution to botany is obscure. Key to the Genera of Theophrastaceae s. str. la. Unbranched or sparsely branched shrubs and trees; fully developed leaves 20-120 cm long, margins often serrate or serrulate. 2a. Corolla rotate, tube smooth on inside; stam- inodes ovoid-gibbous; (lowers often unisex¬ ual, male or functionally male flowers with stamen filaments fused into a permanent tube; anther not or inconspicuously pro¬ duced at apex; leaf margins entire, serrulate, or serrate . Clavija Ruiz & Pav6n 2b. Corolla campanulate or bowl-shaped, irreg¬ ularly puckered on inside of tube; stami- nodes not obovoid; flowers bisexual, stamen filaments fused at base only; anther connec¬ tive distinctly prolonged at apex; leaf mar¬ gins usually spinose-dentate. 3a. Plants erect, the stem spiny; corolla deeply campanulate, pale buff turning black upon drying; staminodes trans¬ versely oblong, inserted within the co¬ rolla tube . Theophrasta L. 3b. Plants semi-decumbent, the stem not spiny; corolla bowl-shaped, orange, not blackening upon drying; staminodes tri¬ angular, minute, inserted at the mouth of the corolla tube . Neomezia Votsch lb. Branched shrubs or small trees; leaves usually less than 20 cm long, margins entire. 4a. Corolla green, > 2 cm diam. at anthesis; staminodes obtrullate to linear; fruits ovoid . Deherainia Decaisne 4b. Corolla orange, whitish, or pale yellow, < 2 cm diam. at anthesis; staminodes flattened, petaloid. 5a. Corolla bowl-shaped, pale yellow; style shorter than the ovary; foliar scleren- chyma of the lower side arranged in a continuous or subcontinuous layer . . . . Votschia StShl 5b. Corolla campanulate or ureeolate, or¬ ange or whitish; style of the same length as the ovary; foliar sclerenchyma ar¬ ranged in distinct bundles. 6a. Young shoots ± lepidote from ir¬ regularly branched, thick-walled hairs; leaves mostly pseudoverticil- late (alternate in ./. keyensis), with¬ out or with a poorly developed api¬ cal mucro; corolla white or whitish; seeds ± globose, covered by pla¬ cental tissue; abaxial foliar scleren¬ chyma often somewhat immersed into the mesenchyma . . . Jacquinia L. 6b. Young shoots puberulous from short uniseriate hairs or glabrous; leaves alternate (pseudoverticillate in B. paludicola ), usually distinctly mu- cronate; corolla orange, sometimes white or whitish; seeds flattened, incompletely covered by placental tissue; abaxial foliar sclerenchyma adjacent to lower epidermis . . . Bonellia Colla Taxonomy and New Comrinations Roiiellia Colla, Hortus Ripul. 21. 1824. TYPE: Bonellia cavanillesii Bertero ex Colla (= Jac- quinia macrocarpa Cavanilles). Bonellia albiflora (Eimdell) Stall I & Kallersjo, comb. nov. Basionym: Jacquinia albiflora Fun¬ ded, Wrightia 2: 60. I960. TYPE: Guatemala. Peten: Tikal, 29 July 1959, Contreras 45 (ho- lotype, FT; isotypes, S. TEX, US). Bonellia brevifolia (Urban) Stahl & Kallersjo, comb. nov. Basionym: Jacquinia brevifolia Ur¬ ban. Symb. Antill. 1: 379. 1899. TYPE: Cuba. Sine loco, Wright 2913 (leetotype, designated in Stcihl (1995), GH; isotypes, G, GOET, M). Bonellia (lannnea (Millspaugh ex Me/.) Sullil & Kallersjo, comb. nov. Basionym: Jacquinia flanunea Millspaugh ex Mez, Pflanzenr. IV. 236a: 40. 1903. TYPE: Mexico. Yucatan: Silam, Gaumer 531 (leetotype, designated in St&hl (1989), G; isotypes. A, BM, BR, C, CAS, I)S, E, F, FHO, GH, K. LE, MO. NY, UPS, US. W). Volume 14, Number 1 2004 Stahl & Kallersjo Reinstatement of Bonellia 117 Bonellia frutescens (Miller) Stahl & Kallersjo, coml). nov. Basionym: Ruscus frutescens Miller, Card. Diet. ed. 8: unpaginated (species no. 8 under Ruscus). 1768. TYPE: Colombia. Boli¬ var: vie. of Cartagena, no date, Heriberto 279 (neotype, designated in Stahl (1995), US). Bonellia lippoldii (Lepper) Staid & Kallersjo. comb. nov. Basionym: Jacquinia lippoldii Lep¬ per, Wiss. Z. Friedrich-Schiller-Univ. Jena. Math.-Naturwiss. Keihe 32: 876. 1983. TYPE: Cuba. Guantanamo: San Antonio del Sur, Abra de Mariana. E of Abra. 100—100 m alt.. 9 leb. 1979, Bisse et al. s.n. (holo- and isotype, II AJB no. 39068). Bonellia loeflingii (Carrasquel) Stahl & Kallersjo, comb. nov. Basionym: Jacquinia loeflingii Car¬ rasquel, Acta Bot. Venez. 4: 348. 1970. TYPE: Venezuela. Anzoategui: about 10 km from Pfr- itu on road to Clarines, Jan. 1968, Aristeguieta et al. 6524 (holotype, VEN). Bonellia longifolia (Standley) Staid & Kallersjo, comb. nov. Basionym: Jacquinia longifolia Standley, J. Wash. Acad. Sei. 14: 241. 1924. TYPE: El Salvador. San Vicente: vie. of San Vicente, alt. 350-500 in, 2-11 Mar. 1922, Standley 21159 (holotype, US; isotypes, F, G, GH. K, NY). Bonellia macrocarpa (Cavanilles) Stahl Kal¬ lersjo, comb. nov. Basionym: Jacquinia macro¬ carpa Cavanilles, Icon. 5: 55, tab. 483. 1799. TYPE: Mexico. Acapulco, Eel). 1791, Ned s.n. (lectotype, designated in D’Arcy (1980), B- WILLD; isotypes, 11BG, MA). Bonellia macrocarpa (Cavanilles) Stahl & Kal¬ lersjo subsp. pungens (A. Cray) Stahl & Kal¬ lersjo. comb. nov. Basionym: Jacquinia pun- gens A. Gray, Proc. Amer. Acad. Arts 5: 325. 1854. TYPE: Mexico. Sonora: hills between Rayon and Ure; 5, 10 Oct. 1851, Thurber 9 03 (lectotype, designated in Stahl (1989), GH; isotypes, GH, NY). Bonellia macrocarpa (Cavanilles) Stahl & Kal¬ lersjo subsp. panamensis ( I. undell) Stahl & Kallersjo, comb. nov. Basionym: Jacquinia panamensis Lundell, in Woodson & Schery, Ann. Missouri Bot. Gard. 27: 329. 1940. TYPE: Panama. Panama: vie. of Bejuco, 18 Oct. 1938, Allen 985 (holotype, MICH; iso¬ types, GH, MO. US). Bonellia montana (Stahl) Stahl & Kallersjo, comb. nov. Basionym: Jacquinia montana Stahl, Nordic J. Bot. 9: 27. 1989. TYPE: Nic¬ aragua. Jinotega: Cerro de la Cruz and vie., W of Jinotega, alt. 1200—1400 rn, 7 July 1947, Standley 10968 (holotype, F). Bonellia inneronata (Roemer & Schultes) Stahl Si Kallersjo, comb. nov. Basionym: Jacquinia mucronata Roemer & Schultes, Syst. Veg. 4: 802. 1819. TYPE: Peru. Colasay, Humboldt 8581 (holotype, B-WIEUD, seen on microfiche; isotypes, HAU. P). Bonellia nervosa (Presl) Stahl & Kallersjo, comb, nov. Basionym: Jacquinia nervosa Presl, Reliq. Haenk. 2: 67. 1835. TYPE: Mexico. Sine loco, Haenke s.n. (holotype, PR). Bonellia nitida (Stahl) Stahl & Kallersjo, comb, nov. Basionym: Jacquinia nitida Stahl, Nordic J. Bot. 9: 24. 1989. TYPE: Honduras. Comay- agua: vie. of Comayagua, alt. 600 m, 12—13 Mar. 1947, Standley & Chacon 6025 (holo¬ type, F). Bonellia paludirnla (Standley) Stahl Kallersjo, comb. nov. Basionym: Jacquinia paludicola Standley, Field Mus. Nat. Hist., Bot. Ser. I l: 138. 1932. TYPE: Belize. Punta Gorda, “For¬ est Home,” alt. 60 m, 1 I Sep. 1932, Schipp 1028 (holotype, F; isotypes. A, BM, G, GH, K, MICH, MO, NY, S). Bonellia pauciflora (Stahl & F. Axelrod) Stahl & Kallersjo. comb. nov. Basionym: Jacquinia pauciflora Stahl & F. Axelrod, Brittonia 50: I. 1998. TYPE: Puerto Rico. Muriicipio Quebra- < 1 i 1 las. Barrio Charcas, 0.7 km SSE of Charcas, 200-350 m alt., 23 June 1994, Proctor et al. 49406 (holotype, SJ; isotypes, GB, NY). Bonellia pringlei (Bartlett) Stahl & Kallersjo, comb. nov. Basionym: Jacquinia pringlei Bart¬ lett, Proc. Amer. Acad. Arts 44: 630. 1909. TYPE: Mexico. Guerrero: Iguala Canyon, 8 Oct. 1908, Pringle 10887 (holotype, GH; iso¬ types, BH, BM, C, CM, E, F, G, GOET, HBG, K, L. LE, M. MICH, MO, MSC, NY, P, PH, PR, US, VT, W). Bonellia seleriana (Urban & Loesner ex Mez) Stahl & Kallersjo, comb. nov. Basionym: Jac¬ quinia seleriana Urban & Loesner ex Mez, Pflanzenr. IV. 236a: 43. 1903. TYPE: Mexico. Oaxaca: above San Carlos Yauhtepec, 5 Jan. 1896, Seler & Seler 1759 (lectotype, designat¬ ed in Stahl (1989), GH; isotvpes. A, NY). Bonellia shaferi (Urban) Stahl & Kallersjo, comb, nov. Basionym: Jacquinia shaferi Urban, Symb. Antill. 7: 320. 1912. TYPE: Cuba. Hol¬ guin: vie. of Holguin, 7 Apr. 1909, Shafer 1231 (lectotype, designated in Stahl (1995), NY; isotypes. A, F, (ill, HAC, K, MO, US). Bonellia sprucei (Mez) Stahl & Kallersjo, comb, nov. Basionym: Jacquinia sprucei Mez, Pflan¬ zenr. IV. 236a: 43. 1903. TYPE: Ecuador. Guavas: Chanduy, Spruce 6478 (lectotype, des¬ ignated in StShl (1990), K; isotypes, BM, BP, CGE, G, LE, OXF, P, W). 118 Novon lioucllia slenoplivlla (Urban) Stahl & Kallersjo. comb. nov. Basionym: Jacquinia stenophylla Urban, Symb. Antill. I: 378. 1899. TYPE: Cuba. “Cuba Occ.,” 1803, Wright 2912 />./>. (lectotype, designated in Staid (1995), C; iso¬ types, BM, GH. GOET, HAC, LE, M. NY, S, US, W). |See Sttlhl (1995) for comments.] lioucllia iimbellata (A. DC.) Staid & Kallersjo, comb. nov. Basionym: Jacquinia umbellata A. DC., Prodr. 8: 150. 1844. TYPE: Puerto Rico. Sine loco, Bertero s.n. (holotype, G-DC, seen on microfiche; isotypes, II. MEE, MO. P). Literature Cited Colla, L. A. 1824. Hortus ripulensis. Torino. Kallersjo, M. A1 8. St&hl. 2003. Pliylogeny oi the Theo- phrastaeeae (Ericales s.l.). hit. J. PI. Sci. 164: 579-591. - . G. Bergqvist & A. A. Anderberg. 2000. Generic realignments in primuloid families of the Ericales s.l.: A phylogenetic analysis based on ON A sequence from three chloroplast genes and morphology. Amer. J. Bot. 87: 132.5-1341. Mez. C. 1903. Theophrastaceae. Pp. 1—21 in A. Engler (editor). Das Pflanzenreich 15 (IV, 236a). St&hl, B. 1989. A synopsis of Central American Theo- phrastaceae. Nordic J. Bot. 9: 15—30. - . 1992. On the identity of Jacquinia armillaris ( I heophrastaeeae) and related species. Brittonia 44: 54-60. - . 1993. Votschia , a new genus of the Theophras¬ taceae from southeastern Panama. Brittonia 45: 204— 207. - . 1995. A synopsis of Jacquinia (Theophrastaceae) in the Antilles and South America. Nordic J. Bot. 15: 493-51 I. Votsch, W. 1904. Neue systematische-anatomische Unter- suchungen von Blatt und Aehse der Theophrastaceen. But. Jahrb. Syst. 33: 502—546. New Taxa and Nomenclatural Notes on the Flora of the Marojejy Massif, Madagascar. VT. Rubiaceae: A New Species of Lemyrea John l\. Stone Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. john.stone@mobot.org Aaron P. Davis The Herbarium, Royal Botanic Gardens, kew, Richmond, Surrey, TW9 3AE, U.k. a.davis@rbgkew.org.uk Abstract. Lemyrea marojejyensis J. R. Stone & A. P. Davis, a new species from t tie Marojejy Na¬ tional Park in Madagascar, is described and illus¬ trated. The thick branchlets, large leaves, shortly pedicellate flowers, and large number of ovules (ca. 32 per locule) distinguish it from other species ol Lemyrea. Systematic observations on the genus Le- myrea are provk led. Key words: Lemyrea , Madagascar, Marojejy, Octotropideae, Rubiaceae. Madagascar is a country rich in endemic species and remains poorly known botanically (Morat & Lowry, 1997). In particular, the isolated massifs of northern Madagascar are extremely diverse and have yielded many new species. The broken chain of upland areas that traverses northern Madagascar contains some of the largest areas of remaining pri¬ mary forest and undisturbed natural areas. While several of the larger mountains are included within protected areas, much of the area in between is unprotected and its vegetation remains threatened. This northern massif region remains poorly known botanically, but the available collections indicate that the region is home to a considerable number of endemic species. The massif at Marojejy has been one of the better collected of these isolated uplands (Humbert, 1955), yet new discoveries are still being published on a regular basis (e.g.. Miller. 1998; Miller & Randrianasolo, 1998; Miller, 2000). 'file central portion of the Marojejy Massif and some of the surrounding lowlands comprise the 60,050-hectare protected area. Formerly known as the Reserve I ntegrale No. 12. established 31 De¬ cember 1927, the status of this protected area was upgraded to a National Park (Marojejy Reserve Na- lurelle Integrale) on 19 May 1998. The middle elevations (ca. 600—1200 m) of the Marojejy Massif seem particularly rich ami diverse in woody plant families. Araliaceae, Arecaceae, Clusiaceae, Cunoniaceae, Myrsinaceae, and Rubi¬ aceae are particularly well represented; the her¬ baceous family Balsaminaceae is represented by more than 30 species (Morat & Lowry, 1997). I hese slopes have been the site of many newly described taxa: e.g., Ardisia marojejyensis J. S. Miller & Pi- poly (Myrsinaceae) (Miller & Pipoly, 1993); Sabicea marojejyensis Razafimandimbison & J. S. Miller (Rubiaceae) (Razafimandimbison & Miller. 1999); Rhodolaena macrocarpa G. K. Schatz, Lowry & A. E. Wolf (Sarcolaenaceae) (Schatz et al., 2000); Weinmannia marojejyensis J. S. Miller & J. Bradford (Cunoniaceae) (Bradford & Miller, 2001). Recent collections from these middle-elevation [tarts of the reserve now yield another new species of Rubi¬ aceae belonging to the genus Dnnyrea (A. Cheva¬ lier) A. Chevalier & Beille. Lemyrea marojejyensis J. R. Stone & A. P. Da¬ vis, sp. nov. TYPE: Madagascar. Prov. Antsi- ranana: Marojejy Reserve Naturellc Integrale, path from Mandena to summit of Marojejy, 700-1000 m, I4°26'S, 49°45'E, 2 Oct. 1994 (fl), Lewis et al. 1249 (holotype, MO; isotypes, BR, G, WAG, K. P, TAN). Figure I. Arbor usque 9 m alta, rarnis crassis et nodos tumidis. Folia coriacea, lamina late elliptica usque late obovata, 7.5-10 cm longa, 4-5.5 cm lata, hdlorescentiae axillares, binae, 1.9-2. 2 cm longae. Pedicelli 1.5— 2.4 mm longi. Corolla late infundibuliformis, earnosa, 1.3— 1.5 cm longa, 5—8 mm lata. Ovarium 2-loculare, ovulis in quoque loculo circa 32. Tree, 5—9 m tall. Branchlets terete to subquad- rangular, swollen at nodes, 3.5—7 mm diam., smooth to somewhat scaly, brown to light gray or whitish. Leaves: petioles distinctly articulated at the junction with the branchlet, 0.3—1 cm long, gla¬ brous, base adaxially canaliculate; lamina broadly elliptic to broadly obovate, 7.5—10 X (3.4— )4— 5.5(— Novon 14: 1 19-123. 2004. 120 Novon Figure 1. lemyrea marojejyensis J. R. Stone & A. I*. Davis. — A. Haliit. — B. Flower, cut to reveal internal corolla surface, anthers, and style. Drawn from the holotype: Irwis et 4 cm wide; floral pedicel ^ 2.4 mm long; ovules > 30 per loeule . A. marojejyensis 2b. Leaves usually < 3.5 cm wide; floral ped¬ icel > 5 mm long; ovules < 10 per loeule. 3a. Corolla throat hairs present on almost the entire surface of the corolla tube; style glabrous, divided to the hase but adnate for most of its length . L ciliolata 3b. Corolla throat hairs present only at the base of the corolla tube; style pu¬ bescent, divided into two lobes but not to the base . A. krugii Acknowledgments. Fieldwork at Marojejy was funded by the National Geographic Society (3647- 87), the World Wildlife Fund-U.S., and the World Wide Kund lor Nature (project #6502). We thank the directors and staff of the following herbaria for placing herbarium material of Irrnyrea at our dis¬ posal: k. MO. P. and TKK. J. Stone thanks Jim Mill¬ er and Armand Randrianasolo for valuable com¬ ments on the manuscript. Bee Gunn provided the illustration, and Roy Gereau provided advice on the Latin description, f ieldwork was conducted under a collaborative agreement between the Missouri Bo¬ tanical Garden and the Parc Botanique et Zoolo- gique de Tsimbazaza. We are grateful for courtesies extended by the Government of Madagascar through the Direction Generale de la Gestion des Resources Forestieres and the Association Nation- ale pour le Gestion des Aires Protegees (ANGAP). A. Davis acknowledges the Leverhulme Trust, which funded part ol the research for this paper under the Rubiaceae of Madagascar Project. Literature Cited Beille, L. 1939. Structure anatomique des Lemyrea. Rev. lut. Bot. Appl. Agric. Trop. 19: 250-255. Bradford. .1. A .1. S. Miller. 2001. New taxa and nomen- clatural notes on the flora of the Marojejy Massif. Mad¬ agascar. V. Cunoniaceae: Weinmannia. Adansonia. ser. 3, 23: 219-236. Capuron, R. 1973 [Unpublished manuscript). Revision des Rubiaof'es de Madagascar et des Comoros. Copies at BR. k, MO. P. and TEF. Chevalier, A. 1938. Fssai d’un groupement systematique des Cafeiers sauvages de Madagascar et des lies Mas- careignes. Rev. Int. Bot. Appl. Trop. 18: 825—843. - . 1939. A propos des Lemyrea de Madagascar. Rev. Int. Bot. Appl. Agric. Trop. 19: 250. Humbert. 11. 1955. Une merveille de la Nature & Mada¬ gascar. Premiere exploration botanique du massif du Marojejy et de ses satellites. Mem. Inst. Sei. Madagas¬ car 6(B): 1—27 1 . Miller, J. S. 1998. New taxa and nomenclatural notes on the flora of the Marojejy Massif, Madagascar — I. Cap- paraceae: A new species of Crataeva. Novon 8: 1 67 — 1 69. - . 2000. Newr taxa and nomenclatural notes on the flora oi the Marojejy Massif, Madagascar IV. Myrtaceae: New species of Eugenia. Adansonia, s£r. 3, 22(1): 1 1 1- 1 16. - & J. J. Pipoly. 1993. A new species of Ardisia (Myrsinaceae) from Madagascar. Novon 3: 63-65. - & A. Randrianasolo. 1998. New taxa and nomen¬ clatural notes on the flora of the Marojejy Massif — II. Volume 14, Number 1 2004 Stone & Davis Lemyrea marojejyensis from Madagascar 123 Anacardiaceae: A new species of Campnosperma. No- von 8: 170-1 72. Morat, P. & P. P. Lowry II. 1997. Floristic richness in llie Africa— Madagascar region: A brief history and prospec¬ tive. Adansonia, ser. 3, 19: 101—115. Razafimandimbison, S. & J. S. Miller. 1999. New taxa and nomenclatural notes on the flora of the Marojejy Massif, Madagascar — Ill. Rubiaceae — A new' species of Sabi- cea. Adansonia, ser. 3, 21: 41—45. Robbrecht, E. 1980. The Hvpobathreae (Rubiaceae— Ixo- roideae). 1. Delimitation and division of a new tribe. Bull. Nat. Pantentuin Belg. 50: 69-77. - . 1988. Tropical Woody Rubiaceae. Opera Hot. Belg. 1: 1-127. - . 1994. Supplement to the 1988 outline of the clas¬ sification of the Rubiaceae. In: E. Robbrecht (editor). Advances in Rubiaceae Macrosysteniatics. Opera But. Belg. 6: 173-196. - , I). M. Bridson & D. B. Deb. 1994. The South Indian genus Octotropis (Rubiaceae). In: E. Robbrecht (editor), Advances in Rubiaceae Macrosysteniatics. Op¬ era Bot. Belg. 6: 81-91. Schatz, 0. E. 2001 . Generic Tree Flora of Madagascar. Royal Botanic Gardens, Kew, and Missouri Botanical Garden, St. Louis. - . P. P Lowry II & A. E. Wolf. 2000. Endemic families of Madagascar. VI. A synoptic revision of Rho¬ dolite tin (Sarcolaenaceae). Adansonia, shr. 3, 22: 239— 252. Resolving a Nomenclatural and Taxonomic Problem in Mexican Oenothera sect. Hartmannia (Tribe Onagreae, Onagraceae) Warren L. Wagner United Stales National Herbarium, Department of Systematic Biology, Botany, MRC-166, P.0. Box 37012, Smithsonian Institution, Washington, DC 20013-7012, U.S.A. wagner.warren@nmnh.si.edu Abstract. Study of type material from the trans- volcanie region of Mexico of two species of Oen¬ othera sect. Hartmannia (tribe Onagreae, Onagra¬ ceae), currently known as Oenothera deserticola (Loesener) Munz and O. purpusii Munz, indicates that they represent the same species. Thus, the lat¬ er name O. purpusii becomes a new synonym of O. deserticola. Also, a neotype is designated herein for 0. deserticola. The oidy available name for the sec¬ ond species, previously incorrectly r ailed O. deser¬ ticola , is Hartmannia montana Hose, but since O. montana is already occupied, a new name, O. ori- zabae, is here proposed. A previously unreported chromosome count of n = 7 is given for O. deser¬ ticola. While examining collections to resolve this problem, a series of specimens from the Sierra Ma- dre Occidental was determined to represent a third species here described as 0. luciae-julianiae. These three species can be distinguished by a suite of characters involving habit, petal color, and cap¬ sules. Oenothera deserticola is characterized by de¬ cumbent stems, morning-opening flowers, rose pur¬ ple petals, and angled, but unwinged capsules with an attenuate apex, while O. luciae-julianiae has erect or ascending stems, evening-opening (lowers, white petals, and narrowly winged capsules with an abruptly acuminate apex. Oenothera orizabae has decumbent stems, evening-opening flowers, white petals, and large-winged capsules with a rounded apex. Resumkn. El estudio de los tipos de Oenothera deserticola y O. purpusii de la seeeion Oenothera seccidn Hartmannia procedentes de la region tras- voleaniea de Mexico, indica que ambos representan la misma especie. Como resultado, 0. purpusii es considerado sindnimo de O. deserticola. IJna segun- da especie, originalmente descrita como Hartman¬ nia montana , pero cuyo epfteto especffico esta ocu- pado on Oenothera , es substituido por el nombre O. orizabae , el eual es aqm propuesto. El estudio de las colecciones procedentes de la Sierra Madre Occidental determine que existe una tereera es¬ pecie, la eual es aqui descrita como 0. luciae-ju¬ lianiae. Estas tres especies pueden ser distinguidas por una serie de caracteres relacionados al hahito, color de petalo y de las capsulas. Oenothera deser¬ ticola es caracterizada por tallos decumbentes, flo- res de apertura matutina, petalos purpureo-rosados y capsulas angulosas, dpteras y atenuadas hacia el apice, mientras que la especie de la Sierra Madre, aquf descrita como O. luciae-julianiae, tiene tallos erectos o ascendentes, flores (pie abren al auoehe- < er, petalos blaneos y capsulas estrechamente ala- das con un apice abruptamente acuminado. Oenot¬ hera orizabae tiene tallos decumbentes, flores que abren al anochecer, petalos blaneos y capsulas ala- das, grandes, con el apice redondeado. Key words: evening-primrose, Mexico, Oeno¬ thera sect. Hartmannia , Onagraceae. Nearly one third of the 120 species of the genus Oenothera L. occur in Mexico. Recent focus on the phylogeny of the family Onagraceae, especially of the most diverse tribe Onagreae (Levin et al., 2003), along with analyses of biogeographic pat¬ terns of tribes Epilobieae, Gongylocarpeae, and Onagreae (Kalinas et al., in press) have resulted in the need to resolve a nomenclatural problem and describe a new species. Two relatively uncommon species from tlu1 trans- volcanie region of Mexico, Oenothera deserticola (Loesener) Munz and O. purpusii Munz, have had a confused application of names since the latter was described in a revision of Oenothera sect. Hart¬ mannia (Munz, 1932). A brief history leading up to the confusion clarifies the problem. The first spe¬ cies was described as Hartmannia montana J. N. Rose (1905). The second was published as Xylo- pleurum deserticolum Loesener (1913). both of these species were included in Munz's revision of Oenothera sect. Hartmannia (Munz, 1932), but for some reason he was confused on the correct appli¬ cation of Xylopleurum deserticolum. Munz's descrip¬ tion ol O. deserticola clearly fits the taxon that .1. N. Rose described as Hartmannia montana (1905). Novon 14: 124-133. 2004. Volume 14, Number 1 2004 Wagner Oenothera Sect. Hartmannia 125 It was characterized by decumbent stems, white petals, and large-winged capsules with a rounded apex. Munz cited a number of collections of it. in¬ cluding the type of Hartmannia montana, and placed Rose’s name in synonymy of 0. deserticola. Munz also cited all four syntype collections given in the original publication ol Xylopleurum deserti- colum (Loesener, 1913). It is not clear if Munz had actually seen any of the syntype collections or if he merely cited them from Loesener’s protologue, while at the same time selecting one of them as the type. For reasons unknown to me, Munz was con¬ fused because at least one ol the syntypes, Seler 5614 (photo US), and likely all four judging from Loesener’s description, is clearly the same taxon as the one he described in the same paper (1932) as the new species O. purpusii Munz. This species was characterized by decumbent stems, rose-purple petals, and smaller, angled, but unwinged capsules with an attenuate apex. A photo of one of the four syntypes was taken before all four were destroyed in the Berlin her¬ barium during World War II. This photo is repro¬ duced here in Figure 1. I had hoped to confirm the characteristics of original material of O. deserticola by either locating duplicates of the leetotype, other syntype collections, or photographs of any of the other three syntypes, but none were found by the collections managers who kindly searched for them at the herbaria known to have received Seler col¬ lections (A, B, CAS, F, GH, K, MEXU, NY. US). Despite not having any of Loesener’s original ma¬ terial, it is clear from the leaves and dehisced cap¬ sule shown in the photograph that O. deserticola represents the same taxon that has gone under the name 0. purpusii since 1932. The name Oenothera deserticola has priority over 0. purpusii , and there¬ fore is the correct name. This conclusion is further- supported by Loesener’s original description (1913), as characterized earlier. The other species in question was first described in 1905 as Hartmannia montana Rose. Since the specific epithet montana is already occupied in Oe¬ nothera, a new7 name, 0. orizabae, is proposed here. During the course of study of these two species using material from a number of primarily Ameri¬ can and Mexican herbaria, it became clear that a significant number of Mexican collections from the Sierra Madre Occidental, from Chihuahua south through Durango to Jalisco and Guanajuato that were identified as O. deserticola by I’. II. Raven, myself, or others over the past three decades rep¬ resents an undescribed species characterized by erect or ascending stems, white petals, and narrow¬ ly winged capsules with an abruptly acuminate apex. It is here described as 0. luciae-julianiae. It has a distribution allopatric from both O. orizabae and 0. deserticola. The following key distinguishes them from one another. Key to Species of Mexican Oenothera sect. Hartman N1A la. Capsules winged, acuminate to rounded at apex; petals white at anlhesis, fading purple. 2a. Stems decumbent, ascending toward the apex; cauline leaf blades 1—3 cm wide, nar¬ rowly ovate, occasionally elliptic, subentire to subdenticulate; capsule body 15—26 mm long, obovoid, apex rounded, the margins winged, the wings 2.3-4' mm wide, the sterile stipe stout, 15—50 mm long; capsule body ap¬ parently dehiscing only at apex ... O. orizabae 2b. Stems erect to ascending; cauline leaf blades 0.5— 1.1 (—3) cm wide, elliptic to lan¬ ceolate or narrowly elliptic-ovate, denticu¬ late to serrulate; capsule body 8— 18(— 25) mm long, clavate or narrowly obovoid, apex obtuse to bluntly acuminate, the wings 1.3— 2 mm wide; the sterile stipe slender, 7—35 mm long; capsule body dehiscing from about half its length to nearly throughout . . . . 0. luciae-julianiae lb. Capsules angled, but not winged, attenuate at apex; petals rose-purple at anlhesis, fading dark¬ er . O. deserticola Oenothera deserticola (Loesener) Munz, Amer. J. But. 19: 758. 1932. Xylopleurum desertico- lum Loesener, Repert. Sp. Nov. 12: 238. 1913. TYPE: M exico. Distrito Federal: Contreras, Lyonnet 532 (neotype, designated here, MFXU-238843; isoneotypes, Gil, K not seen, MEXU [3J, MO, NY, US). Figure 2. Oenothera purpusii Munz, Amer. J. Hot. 19: 759. 1932. Syn. nov. TYPE: Mexico. Ixtaceihuatl, open woods, 1903, C. A. Purpus 1839 (holotype, POM 32827 not seen; isotypes, F, G not seen, GH, MO, NY, UC, US). Perennial herb with usually several or sometimes more, decumbent or sometimes ascending when short, branched or occasionally simple stems 5— 20(— 36) cm long, usually reddish purple or occa¬ sionally pale purple, strigillose, rarely with a few longer erect hairs, often producing new rosettes via rhizomes up to 6-7 cm long, from a thick somewhat woody caudex; root fleshy and thickened in age. Rosette leaf blades 4-6 X 1-1.7 cm, oblanceolate to elliptic, sparsely to moderately strigillose, ap¬ parently quickly deciduous; petiole 0.4- 1.6 cm; cauline leaf blades 1-6.5 X 0.5-1. 8 cm, elliptic to elliptic-oblanceolate or narrowly so, subserrate to weakly sinuate-toothed, apex obtuse lo acute, sparsely to moderately strigillose; petiole 0.1— 1.1 cm long. Flowers opening near sunrise (litis et al. 126 Novon Museum botanicuni Berolinense. Cucc. et Ed. Selcr: 1’lnnt mexicnnae. 1 ^,1 HO/rf T, /?*■(» ..„ 'ir’r-, /*. v>»» , ... t/1 figure I. Reproduced photograph of Seler 5611 , a Berlin ( It) syntype of Xylopleurum deserticolum l.oesener no longer extant, taken as part of the Held Museum’s (f) Types of the Berlin Herbarium project. Copies of the photograph are in MICH, MO. NY, ROM, and US. The photograph was scanned into Photoshop 6.5, and then a dehisced capsule showing the long apex of the valves allowing for unambiguous identification was cut. enlarged, and pasted into the upper left-hand corner (obscuring two small branches evident in the original). A white box on the full photograph indicates the part cut and enlarged, file leaves and part of a stem were removed from the inset for clarity. 202 says evening); floral tube, sepals, and ovary densely strigillose, and ovary usually also densely hirtellous with hairs up to 2 nun long; floral tube 4.5—1 1 mm long; sepals 7.5—16 mm long; free tips in bud absent or rarely present and ca. 0. 1-0.2 mm long; petals {1.5-17 mm long, rose-purple fading darker after pollination; filaments 5.5-8 mm long, rose-purple; anthers 2.5— 4.5 mm long, cream; pol- Volume 14, Number 1 2004 Wagner Oenothera Sect. Hartmannia 127 Figure 2. Oenothera deserticola (Loesener) Munz. — A. Habit, showing decumbent stems from a branched rhizomatous base (Arroyo 182, MO. and Balogh 1005, US). — B. Bud, floral tube, and ovary, showing the lack of free sepal tips and pubescence pattern (Arroyo 182, MO). — C. Flower, showing petal shape and overlap of petals (Hinton 9104, US). — 1). Young capsule showing shape, attenuate apex, and sterile stipe (Lyonnet 532, MEXU). — E. Dehisced capsule showing length of dehiscence and seeds clustered on placentas (Straw & Gregory 1116, OH). — E Dehisced capsule showing shape of open valves (Lyonnet 532, US). len cream, > 90% fertile; style 12-21 mm long; stigma surrounded by the shedding anthers at an- thesis or slightly elevated above it; stigma lobes 2.5—4 mm long. Capsule body 10—22 mm long. rose-purple to green with purple tinge, the body narrowly ellipsoid to clavate, apex attenuate, angled or rarely with a narrow wing up to 0.5 mm wide; sterile stipe slender, 7-17 mm long; capsule body 128 Novon dehiscing nearly throughout; seeds clustered in each loeule, 1—1.2 nun long, 0.4— 0.6 mm thick, brown. Gametic chromosome number, n — 7 (Ha¬ ven, unpublished). Distribution. Oenothera deserticola occurs in open forests of Pinns, Pinus-Quercus, or occasion¬ ally Abies, Alnus, and Salix, on sandy open sites or steep banks, in the mountains of the Trans-Mexican Volcanic region from the northeastern Michoaean and central Hidalgo south to Mexico and Orizaba, Veracruz, from (2100)2500 to 3400 m. The leetotype selected by Munz (1932), but now destroyed, was collected in Mexico, D.K., Kl De- sierto de los Leones, May 1909, IT Hopfer s.n. (C. tV E. Seler 5280) (B destroyed). The leetotype and three additional syntypes (without further locality Uhde 1222; El Desierto de los Leones, IT Hopfer s.n. (labeled as Seler 56 14) [photos: MICH, MO. NY, POM, US]; and San Halael, Tlalmanealo, Seler 5810) were destroyed during World War II (Chris¬ toph Oberprieler, pers. comm. 2001). Oenothera deserticola is a member of section Hartmannia, which can be subdivided into three groups of species based on petal color and time of dowering. With its rose-purple petals and morning flowering, O. deserticola appears to be in a group with other morning-flowering species with purple petals, including O. platanorum I*. H. Haven & 1). H. Parnell, 0. rosea l.’l lerilier de Brutelle ex Aiton, and O. texensis P. H. Haven & I). H. Parnell. These species share angled, but not winged capsules and no free sepal tips in bud or tips free and minute (0. 1-0.2 mm long). The geographic and elevational range of Oenot¬ hera deserticola broadly overlaps the range of 0. orizabae. One collection {Barkley et al. 2488) is a mixed collection of O. deserticola and O. orizabae suggesting that they occur sympatrically at some localities; however, there is no documentation of hybridization. Hybridization must be strongly lim¬ ited by the different timing of flowering, with Mow¬ ers of O. deserticola open during the day and those of O. orizabae open al night. Specimens examined (those marked by * are chromosome determination vouchers). MEXICO. Distrito Federal: Desierto de los Leones, * Raven 15424 (also flavonoid voucher) (TKX); hwy. 190, betw. Puebla & Mexico City, Ahshapanek 701 (TKX); Contreras Canyon, SW of Mexico City, Munz 15000 (GH, MO); Chapultepec III. Valle del Sileneio, Wonderly 217 (MKXU, NMC); Cerro de Esqui- Huil, dclcgacidn de Mipa Alta. Ventura 2804 (MKXU, MO. NY); Kfo Krfo, Bussell & Souviron 82 (US); Volcan Pelado, ladera K, Mondragdn s.n. (MKXU); near Kstacidn La Cima, Serranfa del Ajuseo, Rzedowski 10840 (MICH. MO, TKX); Tlalpan, Volcan Ajuseo, Reyes s.n. (MKXU); lava fields ea. 2 km SSW of La Cima R.B. Station on either side of old Hwy. 95 on top of Serjana de Ajuseo, litis et al. 202 (MICH, MSA); Cerro Tepetlahualo, delegacibn de Milpa Alta, Ventura 1412 (MKXU); Santa Rosa, delega- cidn de A. Obregon, Ventura 2720 (MKXU, MO); I mi. SK of Puerto del Aire, Barkley et al. 2488 (MICH, TKX) | hath sheets mixed collection with 0. orizabae ]. Hidalgo: Cerro de los Pitos, Matilda 21562 (MKXU |2|); Kl Chico, Cerro de Las Ventanas, Gentry et al. 82180 (MO); Parque Nacional Kl Chico, Ortiz 10 (MKXU); Met/.quitillan, Car- pinteros. 20 km K Metzquititlan, Hernandez 6806 (MO); Sierra de Pachuca, Rose & Hay 56/ 1 (US). Mexico: on hwy. Mexico to Puebla, Munz 15064 (POM); Monte do Rio Krfo, km 49, road from Mexico City to Pueblo, Mexia 2606a (IJC); near Salazar, Rose & Painter 7027 (US); Ne¬ vada de Toluca, Rose A Painter 7878 (US); El Crucereo, Temascaltepec, Hinton 521 (POM); Cerro de Ledn, Valle de Mexico, Matuda 10128 (MKXU |2|): Paraje Provincial. Balls 4178 (UC, US); San Rafael, ladera Vi del Iztaeei- luiatl, Sanchez 8 (MKXU); Ixtapaluca, Kstacidn Experi¬ mental de Ensenanza y Investigaeidn de Zoqniapan, Koch & Magana 75242 (MKXU, MO, NY); Llano Grande Gap, near Rfo Krfo, Sharp 4406 (MKXU. RSA); Nat. Park La¬ gunas de Zempoala, Cole 40 (MKXU), McAdams 57 (MICH), Traylor 55 (MKXU); Comunidad, Temascaltepec, Hinton 860 (GH, NY, POM); Cumbre, Temascaltepec, Hinton et al. 7478 ((41. LL, MICH [2|, MO, NY, POM, TKX, UC, US); Cumbre Trojes, Temascaltepec, Hinton et al. 8272 (MKXU, MICH, NY, UC, US); Ml. Popoeatapetl, Nelson 10 (US); 10 km al NE del Entronque a Sultepee, Flores A- Terpdn 820 (MKXU); Vallee de Bravo, 0.8 mi. S ol Cajones, 9500 ft., Balogh 1005 (US); 55 km SK of Mexico City, 10,500 ft.. Weaver 712 (POM. US); near To¬ luca, Schery 64 (MO); Ocuilan. Matuda 82454 (MKXU). IYlichouran: near base of Cerro de Tecolote near Zaoapu, Sharp 45581 (RSA); Cerro del Burro, Santa Clara del Co- bre, Perez 58 (MKXU, MO), Escobedo 1400 (MO); Cerro Prieto, Nuevo Parangarieutiro, Medina 2684 (MKXU); En Kl Salto, a 7 km al E de Ocampo, Melchor Ocampo, Mar¬ tinez & Torres 410 (MKXU): Ocampo, I I km al SK de Ocampo, en el Cerro Kl Chivati, Solo Nunez & Solorzano 12600 (MKXU); Ouerendaro. cerea de San Jose de la Cumbre, Santos 1801 (MKXU); ladera SW del Cerro San Andres, Zinapeeuaro, Zamudio 5520 (MKXU, MO); lado N do La Presa La Gachupina, Jasso 1020 (MKXU); ea. 18 mi. S of Patzcuaro, King & Soderstrom 5180 (TKX, US); near Parieutin volcano, Eggler 84 (US). Morelos: km 48. earretera federal, Vazquez 1871 (MKXU [2]); Lagunas de Zempoala, * St raw & Gregory 1068 ((ill. MKXU, MICH. RSA, UC); Huitzilac. ladera NE del l.ago llueyapan, Par¬ que Nacional Lagunas de Zempoala, Avonce s.n. (MEXU). Puebla: 4 km E of Rfo Krfo. Roe et al. 848 (MICH, US); side ol Popocatepetl, Miranda & Barkley I7M200 (K. I EX); Kl Salto, Rfo Krfo, Boege 8078 (MKXU). Veracruz: Lomogrande. Mt. Orizaba, Balls 4877 (UC, US). Without locality: Gregg 607 (MO). Oenothera orizabae W. L. Wagner, nom. nov. Hartmannia montana Rose, Contr. U.S. Natl. Herb. 8: 329. 1905, non Oenothera montana Nuttall. in Torrey & Cray, FI. N. Amer. I: 500. 1840. TYPE: Mexico. El Cima, on the railroad between Mexico, D.F., and Cuernavaca, Mex¬ ico, 19 Sep. 1903, ,/. N. Rose & ./. II. Painter 7170 (holotype, US-00450745; isotypes. GH, MEXU, NY, US). Figure 3. Volume 14, Number 1 2004 Wagner Oenothera Sect. Hartmannia 129 Figure 3. Oenothera orizahae W. U. Wagner. — A. Habit, showing decumbent steins from a multi-stemmed base ( Straw & Gregory 1120, GH). — B. Bud, floral tube, and ovary, showing the free sepal tips and pubescence pattern ( Straw & Gregory 1120, GH). — C. Flower, showing petal shape and expected non-overlap of petals (Straw & Gregory 1120, MICH). — D. Young capsule, showing shape, wings, rounded apex, and sterile stipe (Rose <& Rainier 7170, holotype, US). — E. Dehisced capsule, showing known length of dehiscence (Rose & Painter 7170, isolype, US). 130 Novon Perennial herb with several decumbent to as¬ cending, simple or usually branched stems, 6—46 cm long, the plant hirtellous throughout. Rosette leaf blades 3—5 X 0.9— 1.5 cm, elliptic to oblan- ceolate, hirtellous; petioles 0.3—1 cm long. Cauline leal blades 3—9 X 1—3 cm, narrowly ovate, occa¬ sionally elliptic, subentire to subdentieulate; peti¬ ole 0.1— 2.2 cm long. Flowers opening near sunset; floral tube, sepals, and ovary densely hirtellous with hairs up to 1.5 mm long; floral tube 9—15 mm long; sepals 15—20 mm long; free tips in bud 1.7- 2 mm long; petals 18-22 mm long, white, fading purple; filaments 15—16 mm long, pale maroon; an¬ thers 3—7 mm long, cream; pollen cream-colored, > 90% fertile; style 20—28 mm long; stigmas held above the anthers at anthesis, the lobes 4—5 mm long. Capsule body 15—26 mm long, obovoid, apex rounded, the margins winged, the wings 2.3—4 mm wide; the sterile stipe stout, 15—50 mm long; cap¬ sule body apparently dehiscing only at apex; seeds 1 — 1.4 mm long, 0.5— 0.7 mm thick, brown. Gametic chromosome number, n = 7 (forming 7 pairs at meiotie metaphase I; Gregory & Klein, 1960; Ha¬ ven, unpublished). Distribution. Oenothera orizabae is scattered in rocky grasslands in Pinus-Quercus forest at middle elevations in the Trans-Mexican Volcanos from Mo¬ relos and Mexico to Kl Chico to the north in Hidalgo, and Pico Orizaba in Veracruz, at 2980-3000 m. Oenothera orizabae appears to be allied to a group of evening-flowering species with white pet¬ als currently assigned to section Hartmannia , in¬ cluding O. kunthiana (Spach) Munz, and 0. tetrap- tera Cavanilles, but based on recent molecular studies this white-petaled group may need to be distinguished as a separate section (Levin et al., 2004). Another while-flowered species, Oenothera dissecta A. Gray ex S. Watson, may also belong in this group, but was most recently assigned to sec¬ tion Gauropsis (Wagner, 1984). A single collection was examined for pollen fertility using Alexander stain (Alexander, 1969). Specimens examined (those marked by * are chromosome determination vouchers and voucher for pollen fertility in¬ dicated by +). MEXICO. Distrito Federal: I mi. SE ol Puerto del Aire, Barkley et al. 2438 (MICH, TEX) [both sheets mixed with 0. desert icola\\ Estacibn I. a Cima. Prin¬ gle 10293 (F, (41, MEXU [2], MICH [2], MO. NY, UC. US), *Raven s.n. (Stanford greenhouse #67-354', ffavonoid voucher) (TEX); Colonia del Valle, Lyonnet 198 (US); ca. 2 km SSW of Ua Cima It. 8. Station, old llwr. 95. litis et al. 200 (MICH. RSA); Volean Pelado. ladera W, Sandoval 230 (MEXU). Ilidal^n: El Chico, Lyonnet 2205 (MEXU). Mexico: S of Mexico City, hnwenworth & Leavenworth 954 (F); 55 mi. SE of Mexico City, Weaver 713 (POM); near Gap, near Kfo Frfo, Sharp 4495 (MEXU, KSA); pub¬ lic park area W of divide betw. Mexico & Puebla al the Estaeionamento, Pennell et al. 125 (MO); 1 km al N de Llano Grande sobre la ladera S del Telapon, Espinosa 542 (MO); Llano Grande, Telapon. + Lyonnet 689 (MEXU |2|, US); near Rio Frfo, Puebla hwy., * Straw & Gregory / 120 ((41, MEXU, MICH [2], RSA); Rio Frfo, Kenoyer 2485 ((ill); just W of Rfo Frfo, 40 mi. E of Mexico City, Man¬ ning A- Manning 53656a (GH). Morelos: Cuernavaca, Kenoyer A422 (E); 6 mi. W of Yautepec, Walther 133 (MICH). Veracruz,: Pico de Orizaba, Rose & Hay 5672 (US). Oenothera luciue-juliuniue W. L. Wagner, sp. nov. TYPE: Mexico. Durango: San Dimas, Mesa del Roble, San Miguel de Cruces, 24.99°N, 1 05.99° W, 2740 m, 26 May 1990, A. Garcia & S. Acevedo 345 (holotype, US- 3441077; isotypes, ANSM, CHAPA, CIIDIR, ENCB, IEB, MEXU, 1) AMI/ (none seen)). Fig¬ ure 4. Ab Oenothera orizabae caulibus erectis ad adscenden- tes, laminis foliorum caulinorum 0.5— 1.1 (—3) cm latis elliptieis ad lanceolatas et denticulatis ad serrulatas, cor- pore capsulae 8 — 1 H( — 25) mm longo clavato vel anguste obovoideo ad apicem obtuso vel acuminato alis ejus 1.3— 2 mm latis stipite sterili tenui 7—35 mm longo. Perennial herb with few to several, ascending to erect, simple or occasionally branched stems, the longer ones becoming decumbent and often devel¬ oping lateral branches, 20-50(-90) cm long, sparsely (very sparsely) to moderately hirtellous, the hairs 1.5— 2.5 mm long, also strigillose through¬ out or only in the upper portions of the stem, the hairs mostly 0.3— 0.4 mm long; root a thickened, often branched taproot. Rosette leaf blades 4—10 X 0.5— 2.5 cm, oblanceolate, denticulate to serrulate, or weakly sinuate-toothed, sparsely to moderately strigillose, apex obtuse to rounded; petiole 0—1.5 cm long; cauline leaf blades (1.8— )3-7 X 0. 5-1.1 (— 3) cm, elliptic to lanceolate or narrowly elliptic- ovate, denticulate to serrulate, apex attenuate, acute, sparsely to moderately strigillose; petiole 0— 0.5 cm long. Flowers opening near sunset; floral tube, sepals, and ovary moderately to occasionally densely strigillose and rarely with scattered hirte¬ llous pubescence with hairs up to 1.5 mm long; floral tube 12-21 mm long; sepals 14—24 mm long; free tips in bud 1-2 mm long; petals 16—26 mm long, white fading deep purplish red; filaments 7— 10 mm long, rose-purple or yellow; anthers 4.8— 6.5 mm long, cream; pollen cream, ca. 50—60% fertile (range 2U) — 94%); style 20—30 mm long; stigma el¬ evated above to surrounded by the shedding an¬ thers at anthesis, the lobes 3.6—6 mm long. Capsule body 8— 18(— 25) mm long, gray-green, clavate or narrowly obovoid, apex obtuse to bluntly acumi¬ nate, winged in the upper part or throughout the Volume 14, Number 1 2004 Wagner Oenothera Sect. Hartmannia 131 Figure 4. Oenothera luciae-julianiae W. L. Wagner. — A. Habit, showing ascending to erect stems from a several¬ stemmed base ( Garcui & Acevedo 345, US). — B. Bud, floral tube, and ovary, showing the free sepal tips and pubescence pattern (Reveal & Harley 4095 , NY). — C. Flower, showing petal shape and apparent slight overlap (Garcia & Acevedo 345, US). — I). Young capsule, showing shape, wings, obtuse to bluntly acuminate apex, and sterile stipe (Reveal <0: Harley 4095, TEX). — E. Dehisced capsule, showing known length of dehiscence, and valves (although they can be spreading) (Garcia & Acevedo 345, US). 132 Novon body, the wings 1.3—2 mm wide; sterile stipe slen¬ der, 7—35 mm long; capsule body dehiscing in up¬ per half to nearly throughout; seeds clustered in each locule, 1.4— 1.5 mm long, brown. Distribution. Oenothera luciae-julianiae is scat¬ tered to common in open Pinas or Pinus-Quercus forests or mixed forests of Abies, Pseudotsuga, Pi¬ nas, Quercus, and sometimes with Arbutus, Juni- penis, or Cupressus, on rocky or open sites, steep slopes, or moist flats or along streams in the Mex¬ ican mountains of the Sierra Madre Occidental from Chihuahua south through Durango, Guanajuato, Nayarit, Jalisco, Michoacdn, and Q ue re taro, from 21(H) to 2800(— 3250) m. Flowering nearly through¬ out the year, and documented from February through November. Etymology. It is a pleasure to name this attrac¬ tive Sierra Madre Occidental species in honor of my wife, Lucy C. Julian, on the occasion of our 10th wedding anniversary. Oenothera luciae-julianiae has gone undetected for decades of collecting in t he Sierra Madre Oc¬ cidental. The collections cited herein were deter¬ mined most commonly as O. deserticola by P. II. Haven, myself, or others, but also occasionally as O. purpusii, O. tetraptera, or O. kunthiana. It was treated as part of 0. deserticola (now correctly known as O. orizabae) by Haven and 1). Parnell (pers. comm.) in their unpublished revision of sec¬ tion Hartmannia. Munz, who revised the section in 1932, never annotated any of the collections, as very few were made before 1960. .1. N. Rose, how¬ ever, annotated a single collection from Chihuahua (Nelson 4848) as a new species, but he never pub¬ lished it. Oenothera luciae-julianiae appears to be allied to a group of evening-flowering species with white petals currently assigned to section Hartman- nia, including Oenothera orizabae, O. kunthiana (Spach) Munz, and O. tetraptera Cavanilles. Like Oenothera kunthiana of this white-flowered species group of section Hartmannia, O. luciae-ju¬ lianiae appears to be a permanent translocation heterozygote (I XFI 1 ) species. In PTH species a ring of 14 chromosomes is usually formed in meiotie metaphase. I have not yet been able to obtain cy- lological material to examine meiosis in this spe¬ cies, and thus current data are only suggestive of this condition in O. luciae-julianiae. PTH species in the Onagraceae are nearly always autogamous (Haven, 1979). The stigma is usually surrounded by the shedding anthers at anthesis in O. luciae- julianiae, suggesting that it is predominately autog¬ amous. Another feature of Hi ll species is the bal¬ anced lethals in pollen and ovules, which can be observed in lowered pollen fertility and fewer ma¬ ture seeds (see Cleland, 1972). I examined pollen fertility of as many collections as possible of O. luciae-julianiae. In the 15 collections I examined with Alexander stain (Alexander, 1969; vouchers noted with brackets in the exsiccatae) pollen fer¬ tility ranged from 26% to 94%, a very' wide range, but suggestive nevertheless that O. luciae-julianiae is probably a PTH species. Typically, PTH species have about 50% pollen fertility on average, but all exhibit a range of fertility. For example, Oenothera nutans G. F. Atkinson & Bartlett exhibited a similar wide range of fertility (48% to 84%) in a detailed study (Wasmund, 1990). Haven (1979) pointed out that most Pi ll species are annuals or biennials, with about 10% of the known lyfH species in Ona¬ graceae being perennial. If O. luciae-julianiae proves to be a PrH species, it will represent an¬ other atypical species, as it clearly has a perennial habit from an enlarged root. Oenothera luciae-julia¬ niae does not appear to be closely related to other species of the white-flowered species group of sec¬ tion Hartmannia. It may share a common ancestor with O. tetraptera , an outcrossing pair-forming spe¬ cies that appears to be directly related to the other PTH species in this group, O. kunthiana. Paratyjies (those used to study pollen fertility have % fertile pollen given in square brackets). MEXICO. Chi¬ huahua: ca. 51 ini. S of (Teel, Straw & Forman 1900 (USA) [26%]; Llano Grande, Pennington 181 (TEX) [37%]; Guachochi, 8 km al W de Cabbrachi, Hernandez 8778 (MEXU); Ocampo, Cascada de Basaseachi, Tenorio A' Torres 4510 (MEXU); Urique, Rye el al. 15612 (MEXU) |90%], Tenorio et al. 9 985 (MEXU) [62%]; base of Ml. Mohinora, 8 mi. from Guadalupe y Calvo, Nelson 4848 (US). Duriinpi: Canolas, Bolanos 1565 (MEXU); 55-60 km SW of Durango City on road to I, a Elor, Breedlove 44184 (MO); from the Sierra Madre, W ol Durango, Forrer 12 (UC); ca. 77 rd. km S of Durango by hwy. to La Elor, Worthington 8826 (MO); 48 km WNW of Huejuquilla Id Alto. Jalisco to Canoas, Breedlove & Almeda 59165 (MO); 6 mi. E of Buenos Aires tow. El Salto, Straw & Forman 1795 (BSA) 1 83% |; 31 mi. N of Estaeion Coyotes. Breed¬ love 18741 (MO); 54 mi. N Estaeion Coyotes, just NW ot Guachichilas, Breedlove 18788 (MO) [59%]; 71 mi. W of Durango. E of La Campafia, Oliver 715 (MO); I I km SW of La Ciudad near Buenos Aires. Breedlove 86465 (MO) |27%|; llwy. 40. 11.5 mi. W of La Ciudad, Wagner & Solomon 4808 (MO); llwy. 40, at La Campana, Reveal et al. 2689 (MO) [74%]; llwy. 40. about 81 mi. W of Du¬ rango, Reveal & Atwood 8477 (MO) [39%]; betw. Pinos Altos & Conohefto, llewitt 118 ((-II) [88%]; El Salto, El Tapextle, Tenorio A Romero 81 6 (MEXU); 1 km al SW de El Salto, Tenorio A Romero 927 (MEXU); “Arroyo de El Salto” al N de El Salto, Tenorio A Romero 759 (MEXU); 17 mi. W of El Salto, Waterfall 12698 ((41. US) [35% |; 25 mi. E of El Salto along Mexican Hwy. 40, Breedlove 15758 (MO); 7 km W of Llano Grande, Hendrickson 1704 (BSA) 1 72%]; ca. 120 road mi. NW of Santiago Papa- squiaro, Spellenberg A 'Zimmerman 6688 (MO. NMC); 10 Volume 14, Number 1 2004 Wagner Oenothera Sect. Hartmannia 133 mi. W of El Salto on Mazatlan rd., Straw & (Gregory 1270 (USA); Sierra Madre Occidental, 5.1 rd. mi. by hwy. 40 SW of El Salto at Arroyo de Agua, Worthington H902 (MO); Durango, Ejido Cienega de Eos Caballos, Ortega & Pacheco HI (CIIDIR, IEB. MEXU, US); Ejido, Encina, Ortega 20 (CIIDIR, US); El Mezquital, El Zapote, Soil's 902 (CIIDIR, US); Ea Guajolota, Garcia 260 (CIIDIR. US); Eaguna del Chivo, Acevedo 445 (CIIDIR, IRUG, US); Eas Minas, Solis II (CIIDIR, US); Nombre de Dios, San Jose de Ea Parrilla, Sanchez 455a (US), 544 (CIIDIR, US); Pueblo Nuevo, El Salto, Valenzuela 3—25 (CIIDIR, US); Predio Eas Rayas de la UJED, Garcia & Acevedo 9H0 (US); San Rernardino de Milpillas Chico, Sanchez (INI) 9 (US), 19 (CIIDIR, US); San Francisco de Eajas, I.N.I. 31 (CII¬ DIR, US); Santa Barbara, Garcia 1 137a (US); San Dimas, Eos Aposentos, Garcia et al. 435 (CIIDIR, US). Guana¬ juato: Guanajuato, Rincdn del Toro, Martinez 760 (MEXU 1 2]); Canada de la Virgen, Agua Sabrosa, Martinez 935 (MEXU); 34—35 mi. E of San Euis de la Paz tow. Xichu, Straw & Forman 1474 (MEXU. RSA). Jalisco: 14— 18 km SW of Tequila on Volcan de Tequila, Breedlove 39226 (MO) [47%]; Volcan Tequila, Reveal & Harley 4095 ll. MEXU, MO. NY. TEX) [94%]. Michoacan: betw. Los Cabras & Pueblito, Chilchota to Zaeapu, Sharp 45507 (RSA). Nayarit: 105 km WNW of lluejuquilla El Alto along road to Jesus Maria, Breedlove 6I50H (MO) [94%|. Queretaro: Colon, parte alta del Cerro Zamorano, Rze- dowski 44449 (MICH). Acknowledgments. The holotype and the distri¬ bution of isotypes of Oenothera luciae-julianiae were kindly provided by S. Gonzalez. Thanks are given to Alice Tangerini for tin* preparation of the three excellent plates t licit amply demonstrate the distinctions among these difficult species of Oenot¬ hera sect. Hartmannia. I thank Denise Mix for comments on the manuscript, compilation of the specimens examined, assistance with the loans, and preparation of Figure 1. I appreciate translations of the* diagnosis of O. luciae-julianiae to Latin provid¬ ed bv Dan Nicolson. and the translation of the ab¬ stract into Spanish by Pedro Acevedo. I greatly ap¬ preciate the help of Socorro Gonzalez in obtaining an excellent set of collections of O. luciae-julianiae from the region surrounding Durango, including the type. Finally, I am grateful to Peter Hoch for his excellent comments during the review process, which improved the clarity and accuracy of this paper. Literature Cited Alexander, M. P. 1969. Differential staining of aborted and non-aborted pollen. Stain Technol. 44: 117—122. Gleland, R. E. 1972. Oenothera Cytogenetics and Evolu¬ tion. Academic Press, London. Gregory, I). P. & W. M. Klein. I960. Investigations of mei- otic chromosomes of six genera in the Onagraceae. A I i - so 4: 505-521. Katinas, E.. J. Crisci, W. E. Wagner & P. C. Hoch. In press. Geographical diversification of tribes Epilobieae, Gongylocarpeae, and Onagreae (Onagraceae) in North America. Ann. Missouri Rot. Card. Levin, R. A., W. L. Wagner, P. Hoch, M. Nepokroeff, J. C. Pires, E. A. Zimmer & K. J. Sytsma. 2003. Family-level relationships of Onagraceae based on chloroplast rhc\. and ndhV data. Amer. J. Rot. 90: 107—115. - , - . - . W. J. Hahn, A. Rodriguez, I). A. Baum, L. Katinas, E. A. Zimmer & K. J. Sytsma. 2004. Evolutionary relationships among and within Tribes Onagreae and Epilobieae (Onagraceae). Syst. Rot., in press. Loesener, L. E. T. 1913. Mexikanische und zentralameri- kanische Novitaten. IV. Report. Sp. Nov. 12: 217—244. Munz, P. A. 1932. Studies in Onagraceae VIII. The sub¬ genera Hartmannia and Gauropsis of the genus Oeno¬ thera. The genus Gayophytum. Amer. J. Rot. 19: 755— 778. - . 1965. Onagraceae. N. Amer. FI. II. 5: 1-278. Raven, P. H. 1979. A survey of reproductive biology in Onagraceae. New Zealand J. Rot. 17: 575—593. Rose, J. N. 1905. Studies of Mexican and Central Amer¬ ican plants — No. 4. Contr. U.S. Natl. Herb. 8: 281—339. Wagner, W. E. 1984 [ 1 985 1 . Reconsideration of Oenothera subg. Gauropsis (Onagraceae). Ann. Missouri Rot. Card. 71: 1 1 14-1127. Wasmund, O. 1990. Cytogenetic investigations on Oeno¬ thera nutans (Onagraceae). PI. Syst. Evol. 169: 69-80. Four New Species of Nasa ser. Alatae (Loasaceae) in the Amotape— Huancabamba Zone of Peru Maxim ilian Weigend Inslilut fiir Biologie— Systematische Botanik mid Pflanzengeographie, Freie Universitat Berlin, Altensteinstr. 6, D- 14195 Berlin, Germany, weigend@zedat.fu-berlin.de Abstract. Four new species of Nasa ser. Alatae (Urban & Gilg) Weigend are described from the Amotape— Huancabamba Zone in northern Peru. The new species Nasa stolonifera has extensive un¬ derground stolons and forms clonal stands, whereas N. victorii has angular stems with elevated longi¬ tudinal ridges. New species Nasa pongalamesa and N. lambayequensis are closely allied to N. carnea from northern Peru and N. loxensis front Ecuador. Nasa amaluzensis Weigend, previously only known from tin* type collection in southern Ecuador, is reported as new to the Peruvian flora. The flowers of Nasa ser. Alatae are discussed as funnel-revolver flowers, based on their morphology. A distribution map of the newly described taxa and a key to all known species of Nasa ser. Alatae in Peru are pro¬ vided. Key words: Amotape— Huancabamba Zone, fun¬ nel-revolver flower, Loasaceae, Nasa ser. Alatae, Peru. A revision of Nasa ser. Alatae in Peru was re¬ cently provided together with an overview of its morphology and ecology (Weigend, 2000a). Nasa ser. Alatae was originally described as Loasa subg. Loasa ser. Alatae Urban & Gilg (Urban & Gilg, 1000), but has since been removed to the new ge¬ nus Nasa and includes the type of this genus (Nasa rubrastra (Weigend) Weigend). Nasa ser. Alatae is closely allied to Nasa ser. Grandiflorae, and its pre¬ cise limits are currently under investigation (Wei¬ gend et al.. in prep.). Both groups share more or less campanulate corollas usually with yellow, red, or orange petals and floral scales with distinct api¬ cal wings. The series as currently defined comprises ca. 27 species, which range throughout the northern and central Andes (southern limit in south-central Peru with an outlier in northern Bolivia) and are usually restricted to the moister eastern slopes in the drier central Andes. The geographical range has been recently expanded northward by the description of a new species from Central America (N. panamen- sis Weigend, Weigend, 2002b). The species here described reported from Peru bring the species to¬ tal of Nasa ser. Alatae in the strictest sense (i.e., sensu Urban & Gilg. 1900) up to 17 from 13 in the year 2000, and an original number of only 5 spe¬ cies before the beginning of the current revisionary effort. I'he species totals known for Colombia and Ecuador are 7 and 8. respectively, so that Peru now houses more than twice as many species as each of these other two countries. This has nothing to do with the different sizes of the countries, since Nasa ser. Alatae in Peru is largely restricted to the north¬ ern part and there mostly to the eastern cordilleras, i.e., an area substantially smaller than their range in Ecuador and Colombia. MATERIALS AND METHODS New collections were obtained during field stud¬ ies in Peru in October 2000 and March— April 2001 in addition to the collections cited in Weigend (2000a); vouchers are cited under the respective species below. Pressed specimens, FA A-material. and photographs are available from these collec¬ tions. Duplicates have been sent by Asuncion Cano E. from USM, Segundo Leiva G. (Herbario de La Universidad Antenor Orrega, Trujillo, Peru), and Victor Quipuscoa S. from HUSA (Universidad Na- cional de San Augustin, Arequipa, Peru). Also, two species, N. olmosiana (Killip) Weigend and N. pon¬ galamesa, have been taken into cultivation in Ber¬ lin. Nasa si.it. Aiatae in the Amotapk- Huancabamba Zone The Amotape— Huancabamba Zone has been pos¬ tulated as a particular center of diversity for the Loasaceae (Weigend, 2002a), and this fact is fur¬ ther underscored by the four new species and the new record. All five species here discussed are found in the area called the Amotape— Huancabam¬ ba Zone (Fig. I, Ayers, 1999; Berry, 1982; Young Beynel, 1997) and close apparent distribution gaps ol Nasa ser. Alatae (Weigend. 2000a). The species show the narrow endemieity and morpho- Novon 14: 134-146. 2004. Volume 14, Number 1 2004 Weigend Nasa ser. Alatae from Peru 135 Figure 1. Distribution of the newly described and the newly reported species of Nasa ser. Alulae, bold circles: Nasa amaluzensis (Weigend) Weigend; black stars: Nasa lambayequensis ; empty star: Nasa stolonifera ; inverted black trian¬ gles: Nasa victorii ; inverted empty triangle: Nasa pongalamesa (map modified from Weigend, 2002a). logical divergence that is so typical of many plant species in this geographical region. A total of 14 of the 17 species of Nasa ser. Alatae known from Peru are endemic to the Peruvian part of the Amotape— Huancabamba Zone, another 3 species are found in both the Ecuadorean and the Peruvian part of the Amotape— Huancabamba Zone, and 1 species is endemic to the Ecuadorean part of the Amotape— Huancabamba Zone. This means that more than 50% of all the known species of Nasa ser. Alatae are restricted to that relatively small zone, in spite of the fact that the group is widespread and ranges from central Peru to Panama. Nasa ser. Alatae comprises mostly narrowly en¬ demic species, and all the newly described taxa fall into that category. In some cases apparent narrow endemicity may reflect a lack of collecting, espe¬ cially in the poorly known cloud forests on the east¬ ern slopes of the Andes. This is borne out by the fact that some species previously known from oidy one locality have now been recollected at different places (e.g., N. lenta (Urban & Gilg) Weigend). Nasa ser. Alatae are usually plants from cloud and mistbelt forests, with only very few species ranging either into paramo habitats (e.g., N. loxensis (Kunth) Weigend from Ecuador, the new species N. 136 Novon Figure 2. I\asa pongalamesa Weigend. — A. Young plant showing typical decussate phyllotaxy of series A/otae. — B. Flower, lateral view, showing dense cover with black stinging setae. — C. Flower, male phase, showing numerous stamens in the center of the flower. —1). Ovary with dense cover with black stinging setae. — E. Stinging hairs on back of petal. — F. Stamens and Moral scales in old flower (lemale phase), showing the funnel-shaped scale apices giving access to the nectar. — O. Floral scale, full dorsal view with basal sacs, scale back, and apical wings. — H. Base of floral scale showing dorsal sacs with their oblique folds. — I. Apical part of floral scale showing recurved scale neck and apical wings. — K. Papillose staminodes alter removal of floral scale. — L. Densely setose ovary roof (androeeium, scales, petals, and two sepals removed) surrounded by the white annular nectary. — M. Receptacle as in L, lateral view, showing the elevated ovary roof with black stinging setae, the thick, white annular nectary and the base of the style (Weigend et al. 852C, type material cultivated at Berlin). Volume 14, Number 1 2004 Weigend Nasa ser. Alatae from Peru 137 pongalamesa ) and into dry scree-slope habitats (N. u re nl ivel utina We i gen d) . Floral Morphology The basic structure and terminology of the floral elements of Nasa ser. Alatae have been discussed previously (Weigend, 2000a) and are largely con¬ gruent with the morphology observed in Nasa ser. Grandiflorae. The finer detail, especially petal shape, color, and orientation, size and shape of the scale back, presence or absence of dorsal threads, and die orientation and shape of the apical wings are crucial for taxon delimitation. A detailed over¬ view is here provided on the basis of photographs of cultivated Nasa pongalamesa (Fig. 2). The pen- tamerous flowers have a largely inferior ovary, five calyx lobes, and five membranaceous, shallowly cymbiform petals (Fig. 2B, 2C). The petals are usu¬ ally setose on the back (f ig. 2C, 2K), but stinging hairs are usually densest on the ovary (Fig. 2D). Phe flowers are proterandric and the mature anthers are folded into the center of the flower during the male phase; during the female phase the stigma elongates and is then situated in the center of the flower, while the anthers have by then wilted (Fig. 2F). Idle floral scales seen from above (i.e., in the flower visitor’s view) form a typical “revolver” (Fig. 2F), i.e., each scale apex forms a funnel (Fig. 2H. 21) that guides the beak of visiting hummingbirds to the basal nectar reservoir (Fig. 211). Pile term revolver flower is generally used for flowers where there is more than one access point to the nectariferous part (Endress, 1994: 116). In Nasa ser. Alatae (and most Loasaceae sublam. Loa- soideae) there are five nectaries, each covered by a floral scale and two staminodes, and thus each floral scale has to be probed to extract all the nectar from an individual flower. Each floral scale forms a distally widened funnel providing mechanical guid¬ ance to the nectaries and also ensuring direct con¬ tact of the beak or the head of the visiting hum¬ mingbird with either the anthers or the stigma. This is a very peculiar case of a revolver flower, and I would like to introduce the term funnel-revolver (lower for this particular situation. Phe basal nectar sacs are not the place where the nectar is secreted; they only function as nectar storage containers. Nectar secretion takes place in five basal nectaries (f ig. 2F. 2M), and nectar is secreted between the bases of the two free staminodes (Fig. 2K) into the base of the nectar scale. The ovary roof, i.e., the receptacle within the nectary, is densely covered with stinging hairs (Fig. 2M), presumably to deter nectar theft by small insects, which could otherwise land on the nectar scales, climb onto the recepta¬ cle, and then remove the nectar from the scales without touching either anthers or stigmas. Key to the Taxa of Nasa ser. Alatae in Peru 1. Calyx tube and fruit without stinging hairs . Nasa anderssonii 1'. Calyx tube and fruit usually densely covered with stinging hairs, at least in proximal por¬ tion . 2 2(1). Stem with strongly elevated longitudinal ridges (usually 6); stinging hairs on ovary concentrated at base, distal portion with very few, or without stinging hairs .... Nasa victorii 2'. Stem terete, never with elevated ridges; stinging hairs ± evenly distributed on fruit, usually very numerous . 3 3(2). Leaves pentagonous with acute leaf lobes; stem densely covered with uniseriate glan¬ dular hairs, especially in distal portion, setae 4—5 mm long; petals narrowly oblong and se¬ tose on hack . Nasa amaluzensis 3'. Leaves ovate to widely ovate, il widely ovate then leaf lobes always rounded to acuminate; stem without or w ith few uniseriate glandular hairs, setae typically less than 4 mm long; lamina typically ovate, if subcircular in out¬ line, then petals widely ovate to subcircular and esetulose on hack . 4 4(3). Corolla yellow, leaves dark green with white lines along primary and secondary veins adaxially . N. driesslei 4'. Corolla pink, orange, or red, never yellow, leaves without white line along veins or, it with white veins, then lamina bright green and flowers bright red (only N. tingamarien- sis , occasionally) . 5 5(4). Inflorescence with large, sessile, semi-am- plexieaul bracts; petals very deeply cymbi¬ form and bright red; corolla balloon-shaped, rarely half spreading . N. olmosiana 5'. Inflorescence with petiolate or sessile bracts, these not semi-amplexicaul; petals variable in shape and color, often orange or pink; co¬ rolla campanulate or star-shaped, never bal¬ loon-shaped . 6 6(5). Lamina very densely pubescent, velvety to the touch with numerous stinging hairs be¬ tween the trichomes; vegetative shoots with numerous, mostly spirally inserted leaves (rarely opposite); stiffly erect, sparsely branched shrub; floral scales yellow' with or¬ ange nectar sacs and apically with filaments . N. urentivelutina 6'. Lamina hairy and sometimes densely so, hut never velvety to the touch; leaves never spi¬ rally inserted, opposite, internodes > 3 cm; plant often branched from base; floral scales red or yellow, nectar sacs not contrastingly colored . 7 7(6). (.eaves approximately as long as wide, dark green callus below' insertion of petiole pre¬ sent; petals either very narrow (5—6 X as long as wide) or widely ovate, carnose and completely esetulose on back;; capsule cla- 138 Novon vate, erect on post-anthetieally elongating pedicel . 8 7'. Dark green callus below insertion of petiole absent; petals not narrow (< 4 X as long as w ide); capsule variable in shape, but pedicel never elongating post-anthetieally . 10 8(7). Petals ovate (< 4 X as long as wide), carnose . !\as 6 X as long as wide), membranaceous . 9 9(8). Leaves ovate, up to 90 X 50 mm; found in Piura . Nasa solata 9'. Leaves widely ovate to subcircular, 90—150 mm long and wide; Cajamarca . . . Nasa dillonii 10(7). Corolla brilliant red. inside of petals densely glandular distally; soft-stemmed rain forest herb with widely ovate leaves 100-180 mm long and w ide, found below (2000— )1000 m . N. tingomariensis 10'. Characters not as above, leaves smaller, found only above 2200 m . II 11(10). Petals half-spreading and corolla star¬ shaped; floral scales white or pale yellow 12 11'. Petals erect and corolla campanulate; floral scales white, pale yellow, orange, or deep pink . 13 12(1 1). Sparsely branched, stiffly erect herb with few to many yellow to red setae on stem and leaf veins; petals pink, half-spreading; Cajamar¬ ca: Bambamarca & Cutervo . Nasa carnea 12'. Much-branched stoloniferous herb or sub¬ shrub with numerous yellow to red setae on stem and leaf veins; petals orange; La Lib- ertad: Bolfvar . Nasa stolonifera 13(12). Erect shrubs; capsule turbinate to clavate, 2—2.5 X as long as wide; Cajamarca, Lam- bayeque, Piura . 14 13'. Erect herbs or scandent shrubs; capsule glo¬ bose or shortly turbinate, 1—1.5 X as long as wide; Amazonas, Pasco, Huanueo, San Mar¬ tin . 15 14(13). leaves widely ovate, ca. 1.2— 1.5 X as long as wide with 2 to 3 leaf lobes on each side; stem and pedicels densely glandular; petals orange; Lambayeque: Ferrefiafe . . Nasa lambayeque nsis 14'. Leaves ovate, ca. 2 X as long as wide with 5 to 7 lobes on each side; stem and pedicels sparsely glandular; petals pink; Cajamarca: San Miguel . Nasa pongalamesa 15(13). Erect herb with soft stems (collapsing upon drying); sepals 10—13 X 4.5—7 mm; petals 30-37 mm long, narrowly ovate, apical ly acuminate, deeply cymbiform, membrana¬ ceous; plant very densely covered with setae; Pasco, Junin . N. pascoensis 15'. Subscandent shrub with lignescent stems (not collapsing upon drying); sepals 8—10 X 2-4- mm; petals < 25 mm long, oblong, api¬ cal ly rounded, shallowly cymbiform to ± pla¬ nar; plants sparsely setose to nearly esetu- lose . 16 16(15). Petals widely ovate, dorsally with numerous red setae, membranaceous; Amazonas . . . N. nubicolorum 16'. Petals oblong, apically rounded, without se¬ tae, carnose; Huanueo, San Martin ... A. tenia 1 . Nasa lambayequensis Weigend, sp. nov. TYPE: Peru. Dept. Lambeyeque: Province Ferrenafe, Incahuasi, to Laguna Tenibladera, 3000 m, 13 Sep. 1985, 1. Sagdstegui A., I). Skill man, Mostacero & L Ramirez 12872 (holotype, HUT; isotypes, F, MO). Figures 1, 3. Haec species N. pongalamesae affinis, sed ab ea foliis late ovatis ambobus lateribus lobis 2 vel 3 instructis (nee anguste ovatis ambobus lateribus lobis 5—7 instructis), co¬ rolla aurantiaca (nec rosea) atque caule apicem versus dense (nec parce) glanduloso differ!. Strictly erect shrub up to 1.5 in tall; stein Idled with white pith, to 0.5 cm diatn., densely covered with red-brown setae to 3 mm long and scabrid hairs, young parts of the plant and pedicels very densely covered with uniseriate, gland-tipped, white trichomes consisting of 2 to 5 cells each. Leaves opposite, petioles 30—45 mm long, densely setose; lamina ovate, 50—75 X 50-65 mm, mem¬ branaceous, base shallowly cordate (sinus 4—6 mm deep), apex acuminate, margin lobed with 2 to 3 widely triangular lobes on each side, each 10—15 X 17-20 mm, lobe margin serrate; abaxial surface setose on veins, otherwise covered with scabrid-glo- chidiate hairs (0.7-1 mm), adaxial surface densely setose and densely covered with longer scabrid hairs (ca. 1 mm); venation pinnate with 2 to 3 lat¬ eral veins on each side. Inflorescence a terminal monochasium, rarely diehasium, 10-20 cm long, with 5 to 8 pendent flowers, bracts subsessile, up to 30 X 12 mm; pedicel 20—40 mm long. Calyx very densely setose, tube conical to ovoidal, 5—6 X 5 nun, calyx lobes 5, ovate-acuminate, 12-14 X 5 mm, densely setose and covered with scabrid tri¬ chomes on the outside; corolla widely campanulate; petals obovate, cymbiform, 18—22 X 10 mm, apex rounded to slightly acuminate, abruptly widening ca. 5 mm from base, densely setose and set with scabrid and glochidiate hairs on back, with gland- tipped, 3-celled uniseriate trichomes, orange; nec¬ tar scales 5, with rectangular back, 8X2 mm, base incurved, basally on back with two depressed glo¬ bose sacs 2 mm diam.. pale red. scale neck slightly thickened, recurved and deeply red, without lili- form appendages, laterally protracted into 2 in¬ curved, suberect wings 4 X 1.5 mm; staminodia 2 per scale, 7 mm long, base dilated, 1 mm wide, filiform above, base curved, papillose, pale yellow; stamens numerous, in 5 epipetalous fascicles of 12 to 14 each; filaments 12 mm long, white, anthers 1 X 0.5 mm, black. Capsule clavate, crowned with the persistent calyx lobes, pedicel erect, 25—35 mm long, capsule 20—25 X 10 nun, very densely red setose, covered with very short, white, glochidiate Volume 14, Number 1 2004 Weigend Nasa ser. Alatae from Peru 139 Figure 3. Nasa lambayequensis Weigend. — A. Leaf. — B. Capsule. — C. Sepal. — D. Petal. — E. Floral scale. — E Staminode, lateral view. (Drawn from Sagdstegui el al. 12872, HUT.) and scabrid trichomes and uniseriate, gland-tipped trichomes, opening with 3 apical valves; seeds nu¬ merous, ovoid, testa reticulate. Nasa lambayequensis lias been collected in flow¬ er and fruit in August and September, but is likely to show peak flowering toward the end of the rainy season in late April and May; no detailed field data are so far available. Nasa lambayequensis is evi¬ dently closely allied to N. pongalamesa described below. They share the shrubby, poorly branched habit and grow in similar situations along rocks and dry stone walls in jalca (dry paramo of Peru) hab¬ itats. It differs from N. pongalamesa and closely allied N. carnea in leaf shape (widely ovate with 2 to 3 lateral lobes vs. ovate blades with more than 4 lateral lobes), petal color (orange vs. pink), and its much denser glandular indument. Both N. pon¬ galamesa and N. lambayequensis are segregates of N. carnea (Urban & Gilg) Weigend s.l. (sensu Wei¬ gend, 2000a), and the material here referred to the new species was previously treated under that tax¬ on. The three taxa are allopatric, with both new species found on the western slope of the Andes, while N. carnea is restricted to inner Ande< in val¬ leys in the Central Cordillera (centered around Dept. Cajamarca, Prov. Bambamarca). The northern specimens of what now remains in N. carnea (e.g.. Dept. Cajamarca, Prov. Cutervo) remain problem¬ atical, and field studies will be required to clarify where they actually belong, since the available her¬ barium material is unsatisfactory. Paratypes. PERU. Dept. Lambeyeque: Prov. Eer- renafe, between Incahuasi and Sinchihual, 3000 m, 25 Aug. 2000. A. Sagdstegui A. & M. Zapata 18279 (HAO, HUT. M). 140 Novon 2. INasa pongalamesa Weigend, s[). nov. TYPE: Peru. Dept. Cajamarca: Prov. San Miguel de Pallaques, Road Agua Blanea to Oyotun, pass height “Ponga la Mesa," 3500—3600 m, 14 Oct. 2000 (cultivated in Berlin since Sep. 2002), M. Weigend, //. Fiirlher, N. Dostert & E. Rodriguez R. 2000/752C (holotype, USM; isotypes. BM. BSB, HUT, M. MO. NY. W). Fig¬ ures 1, 2, 4. Haec species l\. carneae affinis, sed ah ea hal)itu fru- ticis rigide erectis (nec herbae arete ramosae), squamis nectariferis roseis (nec luteolis) et corolla campanulata (nec stellata) dilfert. Strictly erect shrub 1-2 m tall; stem filled with white pith, to 1.5 cm diam., densely covered with reddish brownish setae to 3 mm long and scabrid hairs, young parts ol the plant and pedicels with scattered to many uniseriate, gland-tipped, white triehomes consisting ol 2 to 5 cells each. Leaves opposite, petioles 30—55 mm long, densely setose; lamina ovate, 100-170 X 50—70 mm, membrana¬ ceous, base shallowly cordate (sinus 3-5 mm deep), apex acuminate, margin lobed with 5 to 7 widely triangular lobes on each side, each 7—12 X 15—18 mm, lobe margin serrate; abaxial surface very densely setose on veins and covered with glochid- iate hairs (0.5 mm), adaxial surface sparsely setose and densely covered with longer scabrid hairs (ca. I mm); venation pinnate with 3 to 5 lateral veins on each side. Inflorescence a terminal monoehasi- urn, rarely dichasium, 20—50 cm long, with 8 to 12 pendent flowers, bracts petiolate, up to 55 X 18 mm; pedicel ca. 20 mm long, calyx very densely setose, tube conical to ovoidal, 6—7 X 4 mm, calyx lobes 5, ovate-acuminate, 13—15 X 5 mm, densely setose and covered with scabrid triehomes on the outside; corolla widely campanulate; petals ob- ovate, cymbiform, 20—25 X 15 mm, apex rounded to slightly acuminate, abruptly widening ca. 2 mm from base, setose and set with scabrid anti glochid- iate hairs on back, with gland-tipped, 3-celled un¬ iseriate triehomes, pale pink; nectar scales 5, with rectangular back, 6X2 mm, base incurved, basally on back with two depressedly globose sacs 2 mm diam., pale red, scale neck slightly thickened, re¬ curved and deeply red, without filiform appendag¬ es, laterally protracted into two incurved, suberect wings 3 X 2 mm wide; staminodia 2 per scale, 7 mm long, base dilated, I mm wide, filiform above, base curved, papillose, pale pink; stamens numer¬ ous, in 5 epipetalous fascicles of 12 to 14 each; filaments 12 mm long, white, anthers 1 X 0.5 mm, black. Capsule clavale, crowned with the persistent calyx lobes, pedicel erect, 25—35 mm long, capsule 25—30 X 10 mm, very densely black setose and covered with very short, white, gloehidiate tri- chomes, opening with three apical valves. Seeds numerous, ovoid, testa reticulate. Nasa pongalamesa was encountered in fruit in October 2000, i.e., in the middle of the dry season. Like most species of Nasa in the region, it likely flowers March to May at the end of the rainy season. Nasa pongalamesa is closely allied to three other species, N. carnea (Urban & (dig) Weigend, N. urentivelutina Weigend (Weigend, 2000), and N. lamhayequensis (see above). It differs from N. cor¬ nea, e.g., by the stiffly erect habit and perennial growth form with a strongly lignified stem, a very dense cover with dark red-brown stinging hairs (vs. stinging hairs scattered, pale brown), and the pres¬ ence of stinging hairs on the back of the petals. Nasa urentivelutina differs in apricot-colored petals (vs. pink), densely spiral phyllotaxis (vs. opposite leaves), and a much more strongly branched habit (shrubs may be as large as 2 X 1.5 m). Its floral scales have a back that is very strongly narrowed above the nectar sacs and ends in three apical fi¬ liform appendages, whereas the scales of N. pon¬ galamesa are only slightly narrowed above the sacs and have no filiform appendages. The new species is also ecologically clearly dif¬ ferentiated: It grows on an exposed ridge with strong cloud condensation at an elevation of ca. 3500 m between rocks in otherwise grassy puna vegetation. Nasa carnea, on the other hand, grows in shrub forest in inner Andean valleys at ca. 2200—3000 m, and N. urentivelutina is restricted to dry scree slopes in full sun at elevations below 3000 m. Nasa pongalamesa is thus the species of Nasa ser. Alatae that reaches the highest elevation so far known from this group and is found together with a species of the N. cymbopetala group, the typical puna group in the genus. Nasa pongalamesa was found in fruit only in Oc¬ tober 2000 and was then brought into cultivation in the greenhouses of the Institute of Biology in Berlin. The type specimens were prepared from the cultivated material when it came into flower. Nasa pongalamesa is named after the locality of the orig¬ inal collection, “Ponga la Mesa” (lay the table). The epithet is here used as a noun in apposition, with its spelling not changed, sensu Article 23.5. of the ICBN (Greuter et ah, 2000). Paratypes. PERU. Dept. Cajamarca: Prov. San Mi¬ guel de Pallaques, road Agua Blanca to Oyotun, pass height “Ponga la Mesa,” 3500— 36(H) in. 14 Oct. 2000. M. Weigend et ea ovario liasi trichomatibus urentibus instructo (nec omnino tri- chomatibus urentibus carente) et habitu fruticis scanden- lis (nec herbae rigide erectae), ab omnibus speciebus cet¬ eris Loasacearum caule costis longitudinalibus elevatis instructo differt. Erect to ascending shrub 0.6— 2 m tall: stem te¬ rete with typically 6 elevated longitudinal ridges, with white pith, to 1.5 cm diam., with very few reddish brown setae 1—3 mm long and densely cov¬ ered with minute scabrid hairs < 0.2 mm long, eglandular. Leaves opposite, petioles 40—65 mm long, very sparsely setose; lamina widely ovate, 100-150 X 60—90 mm, membranaceous, base sub- truncate (sinus < 2 mm deep), apex long acumi¬ nate. margin lobed with 5 to 7 indistinct, triangular lobes on each side, each up to 5 X 15 mm, lobe margin serrate to serrate-denticulate; abaxial sur¬ face with scattered setae on veins and covered with numerous scabrid hairs (< 0.2 mm), adaxial sur- Volume 14, Number 1 2004 Weigend Nasa ser. Alatae from Peru 143 Figure 5. Nasa stolonifera Weigend. — A. Habit. — B. Sepal. — C. Petal. — I). E. Moral scale, dorsal and lateral view. — F. Staminode, lateral view. (Drawn from Weigend el al. 2000/822, BSB.) 144 Novon Figure 6. A hisa victorii Weigend. — A. Habit. — lb Single flower. — C. Sepal. — I). Petal. — E. Floral scale, lateral view. — F. Staminode, dorsal view. (Drawn from Quipuscoa cl al. 2480, HSR.) face very sparsely setose on veins and densely cov¬ ered with short seabrid hairs (< 0.3 mm); venation pinnate with 3 to 5 lateral veins on each side. In¬ florescence a terminal monochasium 1.3—20 cm long, with up to 6 pendent flowers, bracts petiolate, lamina ovate, up to 55 X 20 mm; pedicel 20-40 mm long; calyx setose only at the base, tithe sub- globose, 5 X 5 nun, calyx lobes 5, narrowly trian¬ gular-ovate, ea. 10 X 3^1 mm, densely setose and densely covered with short seabrid trichomes on the outside; corolla widely campanulate; petals ob- ovate, shallowly cymbiform, 15 X 11 mm, apex Volume 14, Number 1 2004 Weigend Nasa ser. Alatae from Peru 145 rounded, without triangular teelli, esetulose and densely set with short scabrid hairs on hack, eglan- dular, orange-red; nectar scales 5, with rectangular back, 8X3 mm, base incurved, basally on back with 2 indistinct, depressedly globose sacs 3 mm diam., yellow, scale neck not thickened, without ap¬ pendages. laterally protracted into 2 incurved, hor¬ izontal wings 3 X 2 mm; staminodia 2 per scale. 13 mm long, base dilated, 1 mm wide, filiform above, base and tip curved, papillose, white; sta¬ mens numerous, in 5 epipetalous fascicles of 12 to 14 each; filaments 12—15 mm long, white, anthers 2 X 1.5 mm, black. Capsule subglobose, ca. 20 nun diam.. with setae only in proximal part and covered with numerous very short, white, scabrid trichomes, opening with 3 apical valves. Seeds nu¬ merous. ovoidal. testa reticulate. Nasa victorii probably flowers and fruits sporad¬ ically throughout the year, since its habitat does not experience a marked dry season. It has been col¬ lected with flower buds in September and with flow¬ ers and fruits in September. While most species of Nasa are readily recog¬ nized, N. victorii is certainly one of the most readily identifiable taxa. Its stems with longitudinal ridges are unique not only in the species group, but in the entire family Loasaceae. The species is a subscan- dent to seandent shrub in dense, wet cloud forest on the eastern slope of the Andes, a notoriously poorly known region. Like N. nubicolorum from the same general region (Weigend, 2000a) it grows in deep leaf litter and humus in extremely wet habitats. The species is now known from two collections in the department San Martin, but may be more widespread in this inaccessible region. Like most species of the group it is likely to be sensitive to habitat alteration and therefore vanishes rapidly from areas as they become accessible to civilization. Its sparse indument is reminiscent of Nasa an- derssonii, which is known from the Cordillera Golan in Peru and also neighboring Ecuador, but it differs from that taxon, e.g., in stem and (lower morphology, and habit. Nasa victorii is named in honor of Victor Quipuseoa Silvestre (LIUS A. Arequipa. Peru). Paratypes. PERU. Dept. San Martin: Prov. Rioja, road from Pedro Ruiz to Rioja, between La Ksperanza and Nuevo Cajamarca, near km 377, 2000—2300 m, 05°42'09"S, 077°48'27"W, 21 Oct. 2000. M. Weigend el al. 2000/929 (BSR, HUT, M, USM). Nkw Rkcoko 5. Nasa amaluzensis (Weigend) Weigend, Mon- ogr. Syst. Bot. Missouri Bot. Card. 75: 954. 1999. Loasa amaluzensis Weigend, Bot. Jahrb. Syst. 118: 264. 1996. TYPE: Ecuador. Loja: 26.4 km S of Amaluza on road to Zumba, river valley, 2800 m. S. Clemants 2392 (holotype, QGA; isotype, QCNE). Figure I (see also Wei¬ gend. 2000b: fig. I6J-R). Nasa amaluzensis apparently flowers during the wet season (January to April) and probably fruits from April onward, like most species of Nasa from the western part of the Andes. Nasa amaluzensis was described from a single collection from the extreme south of Ecuador in 1996. Material exactly matching the type has now been collected by S. Leiva G. in the extreme north of Peru, so that this species is now known from both countries. Nasa amaluzensis shows clear affinity to Nasa loxensis from Ecuador (it shares horizontally indexed wings on its nectar scales, and cymbiform, setose petals), but differs in distally densely glan¬ dular stems and pentagonous leaves with acute leaf lobes. Its closest geographical neighbors are Nasa solata (leaves ovate, petals linear), N. sagasteguii (similar leaf shape but with ovate leaf lobes, car- nose, esetulose petals), and N. glabra (plant virtu¬ ally esetulose, carnose, esetulose petals). Material examined. ECUADOR. Prov. Loja: 26.4 km S of Amaluza on road to Zumba, river valley, 2800 m, ,S. Clemants 2392 (QGA, QCNE). PERU. Dept. Piura: Prov. Ayavaca, Cerro Aypate, 2650 m, 10 Jan. 2002, .S’. h>iva G. 2632 (BSB, F, HAD. HUT). Acknowledgments. I express my sincere grati¬ tude to Dominik Kneer (Berlin, Germany) for the preparation of the drawings and to Katja Weigend (Berlin) for the preparation of the photographic plate. I also thank E. Rodriguez R. ( 1 1 LIT). S. Leiva G. (HAO), Victor Quipuseoa S. (HUSA), and A. Cano E. (USM) for providing valuable collections, and N. Dostert (Berlin) and H. Forther (both Mu¬ nich) for their assistance during fieldwork. I thank the curators and directors of the following herbaria lor access to and loans of specimens: E, HAO, HUT, M, MO, USM. Part of the studies was supported by the Deutscher Akademiseher Austauschdienst, tin* Deutsche Eorschungsgemeinschaft, and boteon- sult GmbH (Berlin), which are here gratefully ac¬ knowledged. I .iterature Cited Ayers, T. 1999. Biogeography of Lysipomia (Campanula- ceae), a high elevation endemic: An illustration of spe¬ cies richness at the Huancabamba Depression, Peru. Arnaldoa 6: 13—28. Berry, P. E. 1982. The systematics and evolution of Fuch¬ sia sect. Fuchsia (Onagraceae). Ann. Missouri Bot. Card. 69: 1-198. Endress, P. k. 1996. Diversity and Evolutionary Biology 146 Novon of Tropical Flowers, 2nd ed. Cambridge Univ. Press, Cambridge, U.k. Greuter, W., J. McNeill. F. R. Barrie, H. M. Burdet, V. Demoulin, T. S. Filgueiras, 1). H. Nicolson, P. C. Silva, J. F. Skog, P. Trehane, N. J. Turland & 1). L. Hawk- sworth (editors). 2000. International Code of Botanical Nomenclature (Saint Louis Cotie). Regnum Veg. 1 38. Urban. I. & W. Gilg. 1000. Monographia I .oasacearum. Nova Acta Caes. Leop. -Carol. German. Nat. Cur. 76(1): 1-368. Weigend, M. 1990. Loasaeeae. In P. M. Jprgensen & S. Le6n-Ydnez (editors). Catalogue of the Vascular Plants of Ecuador. Monogr. Syst. Bot. Missouri Bot. Card. 75: 537-539, 954-955. - . 2000a. A revision of the Peruvian species of AV«a scr. Alulae (Loasaeeae). Nordic J. Bot. 20: 15—32. - . 2000b. 132. Loasaeeae. In G. Hailing cV L. An- dersson. Flora of Ecuador 64. - . 2002a. Observations on the biogeography of the Amotape— Huancabamba Zone in northern Peru. In: k. Young et al.. Plant Evolution and Endemism in Andean South America. Bot. Rev. (Lancaster) 68: 38—54. - . 2002b. Nasa panamemis, a new species of Nasa (Loasaeeae) from Central America. Bot. Jahrb. Syst. 124: 21 1-216. Young, K. R. & C. Reynel. 1997. Huancabamba Region, Peru and Ecuador. Pp. 465—469 in S. I). Davis, V. II. Heywood, 0. Herrera-MacBryde, J. Villa-Lobos & A. C. Hamilton (editors). Centers of Plant Diversity, a Guide and Strategy for their Conservation, Vol. 3. The Amer¬ icas. 1UCN Publications Unit, Cambridge. Aconitum ouvrardianum var. acutiusculum (Ranunculaceae), a New Combination from Yunnan, China Yang Qiner and Luo Yan Laboratory of Systematic and Evolutionary Botany, Institute of Botany, Chinese Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, People’s Republic of China. qeyang@ns.ibcas.ac.cn; luoyan@ns.ibcas.ac.cn ABSTRACT. Close examination of the type speci¬ mens of Aconitum acutiusculum and of A. acutius¬ culum var. aureopilosum has shown that their inflo¬ rescence rachis and pedicels are all yellowish spreading pubescent, and thus the two taxa show no essential difference from each other. They are merged herein. A new combination, A. ouvrardian¬ um var. acutiusculum (Fletcher & Lauener) Q. E. Yang & Y. Luo, is proposed, considering that A. acutiusculum is most similar to A. ouvrardianum in having leaves 3-sect with ultimate lobules linear or lanceolate-linear, inflorescence rachis and pedicels yellowish spreading puberulent, and the upper se¬ pal navicular-galeate, yet distinguishable by the acute tip of the lateral sepals; they are sympatric in geographical distribution, both occurring in De¬ qen County, northwestern Yunnan. Key words: Aconitum, China, Ranunculaceae, Yunnan. Fletcher and Lauener (1950: 198) described ,4c- onitum acutiusculum based on Forrest 14696 from Raima Shan, Deqen County, northwestern Yunnan, and stated that this species is clearly distinguished from all others in the genus by the acute tip of the lateral sepals, an otherwise unimportant feature in the classification of Aconitum L. because the lateral sepals in this genus are generally subrounded in outline. In the original Latin description, the upper part of the stem and the inflorescence are men¬ tioned as densely and extremely minutely white crispy pubescent (Caulis . . . praesertim superne cum inflorescentia dense et minutissime albo-cris- po-pubescens). Wang (1965) described a new va¬ riety, A. acutiusculum var. aureopilosum, from Yangtsa, Deqen County, and stated that this variety is distinguished from the typical one by having the inflorescence yellowish spreading puberulent. When describing this new variety, he had no op¬ portunity to examine the type specimens of A. acu¬ tiusculum and thus had to base his judgment on Fletcher and Lauener’s description (Wang, pers. comm.). We have checked the holotype and isotype of A. acutiusculum var. aureopilosum at PE and found that the inflorescence rachis and pedicels are indeed densely yellowish spreading puberulent. Pedicel pubescence is one of the most important diagnostic characters in the classification of the ge¬ nus Aconitum. Close examination of the type material of Aco¬ nitum acutiusculum, Forrest 14696, at BM, E, and K has shown that its inflorescence and pedicels, like those of A. acutiusculum var. aureopilosum , are densely yellowish spreading puberulent and that other important characters, such as the division of leaves and the shape of the upper sepal, are by no means different from those of A. acutiusculum var. aureopilosum. Their leaves are 3-sect with ultimate lobules linear or lanceolate-linear, and the upper sepal is navicular-galeate. As this variety shows no essential difference from A. acutiusculum in all the characters, they should be merged. Aconitum ouvrardianum Handel-Mazzetti (1931) is a common species in Deqen, northwestern Yun¬ nan. We have not seen its type. Handel-Mazzetti 9702. but fortunately this species is quite richly represented in the two major Chinese herbaria, PE and KUN, and its identity has long been deter¬ mined with certainty. This species shows no essen¬ tial difference from A. acutiusculum in all the char¬ acters except in the shape of the lateral sepals, with those in A. acutiusculum being acute at the apex while those in A. ouvrardianum are subrounded. In our herbarium observations we found this feature is quite constant in these two entities, but this differ¬ ence is too tenuous to allow treating them as two independent species. In fact, the type specimen of A. acutiusculum var. aureopilosum at PE was once referred to A. ouvrardianum by Chen and Fiu (1941). When Handel-Mazzetti (1931 ) described A. ouvrardianum, he had noticed the similarity of For¬ rest 14696 with his new species but also their dif¬ ferences, particularly in the shape of the lateral se¬ pals. Thus, he did not refer this specimen to any species then known to him. Considering the con¬ sistent difference in the shape of the lateral sepals Novon 14: 147-148. 2004. 148 No von yet the great similarity in other characters between A. ouvrardianum and A. acutiusculum, as well as their sympatric distribution, it seems best to reduce A. acutiusculum as a variety under A. ouvrardian¬ um. The following new combination is thus pro¬ posed. Acoiiitum ouvrardianum Handel-Mazzetti var. aciitiiiseulum (Fletcher & Lauener) Q. E. Yang & Y. Luo, comb. nov. Basionym: Aconi¬ tum acutiusculum Fletcher & Lauener, Notes Roy. Bot. Card. Edinburgh 20: 198. 1950. TYPE: China. Yunnan: Deqen County, Baima Shan, heavy pasture on the margins of pine forests, 12,000 ft., Aug. 1917, G. Forrest 1469 6 (holotype, E; isotypes, K. BM). Aconitum acutiusculum var. aureopilosum W. T. Wang, Acta Phytotax. Sin., addit. 1: 85. 1965. Syn. nov. TV PE: China. Yunnan: Deqen County, Yangtsa, Snow Range, Rusala, mountain grassy slope, 4100 m, Aug. 1937, T. T. Yu 9671 (holotype, PE; isotype, PE). Aconitum ouvrardianum var. acutiusculum is only known from Deqen County, northwestern Yun¬ nan, and seems quite uncommon. Although we had made several botanical expeditions to this area, we tailed to find it. The typical variety ouvrardianum , however, is very common in this area, growing in alpine meadows or bushes at altitudes from 3000 to 4100 m. Acknowledgments. We are grateful to the Keep¬ ers of BM, E, K, and PE for permission to examine specimens or for the loan of material. This work was supported by the National Natural Science Foundation of China (grant no. 30070057) and a Key Project of the Chinese Academy of Sciences (KSCXZ-SW-101A). Literature Cited Chen, F. H. & Y. Liu. 1941. An enumeration of Aconitum collected by T. T. Y u from Yunnan. Bull. Fan Mem. Inst. Biol., Rot. ser. 11: 43-49. Fletcher, H. R. & L. A. Lauener. 1950. Notes on some Chinese species of Aconitum with an enumeration of the Chinese species represented in the Edinburgh Herbar¬ ium. Notes Roy. Bot. Card. Edinburgh 20: 181-206. Handel-Mazzetti, H. 1931. Aconitum. Pp. 282—297 in Symbolae Sinieae, Vol. 7. Julius Springer, Vienna. Wang, W. T. 1965. Notulae de Ranunculaceis sinensibus II. Acta Phytotax. Sin., addit. 1: 49-103. Miscellaneous Notes on Pteridophytes from China and Neighboring Regions (IV) — Validation of Some Combinations in Diplopterygium (Pteridophyta: Gleicheniaceae) Xian-Chun Zhang Institute of Botany, Chinese Academy of Sciences, Beijing 100093, People’s Republic of China, xianchunzhang@yahoo.com.cn Abstract. Some names of Diplopterygium widely used in Chinese literature of pteridophytes are not valid; therefore, formal combinations of five names for Diplopterygium species from China are validat¬ ed. i.e., D. criticum (Ching & P. S. Chiu) Ching ex X. C. Zhang, D. rufum (Ching) Ching ex X. C. Zhang, D. rufopilosum (Ching & P. S. Chiu) Ching ex X. C. Zhang, D. simulans (Ching) Ching ex X. C. Zhang, and I). yunnanense (Ching) tilling ex X. C. Zhang. Diplopterygium rufopilosum is lectotypi- fied. Key words: China, Diplopterygium, Gleicheni¬ aceae, Pteridophyta. Diplopterygium (Diels) Nakai was traditionally treated as a subgenus of Gleichenia Smith (Holttum, 1957, 1959; Tagawa & Iwatsuki. 1979; Tryon & Tryon, 1982), but is now generally accepted as a separate genus (Ching, 1978; Kramer, 1990; Wu & Ching, 1991; 0stergaard Andersen & 011gaard, 2001 ). The Chinese species of Diplopterygium were recorded under the generic name Hicriopteris C. Presl (Ching, 1940; Ching et al., 1959). Holttum (1959) discussed the reason why Hicriopteris was misused in this sense. The type of Hicriopteris was first described as a species of Dicranopteris Bern- hardi, i.e., I). speciosa (C. Presl) Holttum. When Presl (1851) described the genus he compared it with Gleichenia glauca (Thunberg) Hooker, a spe¬ cies of Diplopterygium. Tater authors (Ching, 1940; Copeland, 1947; Ching et ah, 1959) thought he was describing a species of Gleichenia sect. Diplopter¬ ygium and consequently misapplied the generic name Hicriopteris. Hicriopteris is properly a syno¬ nym of Dicranopteris (see Holttum, 1959; 0ster- gaard Andersen & 011gaard, 2001). Three genera of the fern family Gleicheniaceae occur in China: Diplopterygium (Diels) Nakai, Sticherus C. Presl, and Dicranopteris Bernhardi. Plants of Diplopterygium can be easily distin¬ guished from those of Sticherus by their bipinnate or pinnate-pinnatifid branches, and from Dicran¬ opteris by their dormant buds and rhizomes with scales, veinlets of segments once forked, and spo¬ rangia (2)3 to 5(7) per sorus; in Dicranopteris the dormant buds and rhizomes with hairs, veinlets of segments 2 to 4 times forked, and sporangia ca. 8 to 25 per sorus (0stergaard Andersen & 011gaard. 2001) separate it from both Diplopterygium and Sticherus. Diplopterygium (Diels) Nakai, Bull. Nat. Sci. Mus. Tokvo 29: 47. 1950. Gleichenia sect. Di¬ plopterygium Diels, in Engler N Plant I, Nat. Pflanzenfam. 1(4): 353. 1900. Dicranopteris sect. Diplopterygium (Diels) Underwood, Bull. Torrey Bot. Club 34: 251. 1907. Gleichenia subg. Diplopterygium Holttum. Reinwardtia 4: 261. 1957. TYPE: Diplopterygium glaucum (Thunberg ex Houttuyn) Nakai (Polypodium glaucum Thunberg ex Houttuyn). Diplopterygium contains about 25 species, from Asia to Queensland, Australia, and Polynesia (in¬ cluding Hawaii), mainly in tropical Asia, not in Af¬ rica. A single species, D. hancroftii (Hooker) A. R. Smith, occurs in tropical America. Ching et al. (1959) recorded 16 species of Di¬ plopterygium under the generic name Hicriopteris from China and one from northern Vietnam, the region bordering China. Six of these already have been transferred into Diplopterygium, including: Diplopterygium blotiunum (C. Christensen) Nakai, Bull. Nat. Sci. Mus. Tokyo 29: 49. 1950. Diplopterygium cantonense (Ching) Nakai. Bull. Nat. Sci. Mus. Tokyo 29: 49. 1950. Diplopterygium chinense (Rosenstock) De Vol, in H. L. Li et al., El. Taiwan I: 92. 1975, “ chine ns is.” Diplopterygium giguiiteum (Aallich ex Hook¬ er) Nakai, Bull. Nat. Sci. Mus. Tokyo 29: 50. 1950. Diplopterygium glaucum (Thunberg ex Hout- Novon 14: 149-151. 2004. 150 Novon luyn) Nakai, Bull. Nat. Sci. Mus. Tokyo 20: 51. 1950. Diplnplcrygiimi Inrvissimimi (H. Christ) Na¬ kai. Bull. Nat. Sci. Mus. Tokyo 29: 52. 1950. Chiu^ (in Shing, 1982) published numerous combinations for Chinese species of Diploptery- gium, but these were not validated properly ac¬ cording to Article 33.3 of the ICBN (Greuter et al., 2000). Two of these were validated later, i.e., Di- plopterygium maximum Ching & II. S. Rung (in Rung, 1988: 134), and Diplopterygium glaucoides (Ching) Ching ex P. S. Wang & X. Y. Wang (in Wang & Wang, 2001: 279). Some of them are presently regarded as synonyms of various species and there¬ fore need not be transferred into Diplopterygium. The following five combinations for Diploptery¬ gium are formally proposed here in order to solve the nomenclatural problems of these Chinese spe¬ cies and for the revision of fern flora of China. Diplopterygium eriticuni (Ching & P. S. Chiu) Ching ex X. C. Zhang, comb. nov. Basionym: HicriopterLs critica Ching & P. S. Chiu, in S. S. Chien X Chun, FI. Beipubl. Popul. Sin. 2: 348. 1959. Diplopterygium criticum (Ching & P. S. Chiu) Ching, in R. II. Shing, Gloss. Term. Nam. Kerns 50. 1982, nom. inval. TYPE: Chi¬ na. Yunnan: Pingbian, Dawei Shan, R. C. Ching 55 (holotype, PE). Diplopterygium rufuin (Ching) Ching ex X. C. Zhang, comb. nov. Basionym: Hicriopteris rufa Ching, in S. S. Chien & Chun, FI. Reipubl. Popul. Sin. 2: 347. 1959. Diplopterygium ruf- um (Ching) Ching, in R. II. Shing, Gloss. Term. Nam. Ferns 50. 1982. nom. inval. T\ PE: China. Yunnan: Pingbina, Dawei Shan, K. M. Feng 4832 (holotype, PE). The collection number was misprinted as 4823 when the species was published, but on the label of the type in PE the number is 4832. Diploplerygiimi riifopilosmn (Ching & P. S. Chiu) Ching ex X. C. Zhang, comb. nov. Bas¬ ionym: Hicriopteris rufopilosa Ching & P. S. Chiu, in S. S. Chien & Chun, FI. Reipubl. Po¬ ind. Sin. 2: 348. 1959, “ rufo-pilosa Diplop¬ terygium rufopilosum (Ching X P. S. Chiu) Ching, in R. II. Shing, Gloss. Term. Nam. Ferns 50. 1982, " refo-pilosum nom. inval. TYPE: China. Yunnan: Dali. Cang Shan, C. Y. Wu 1 1825 (lectotype, designated here, PE). A second syntype, H. C. Wang 3541 (PE), is of a young sterile plant, but C. Y. Wu 1 1825 is a fertile plant; therefore, it is selected as the lectotype. Diplopterygium siiniilans (Ching) Ching ex X. C. Zhang, comb. nov. Basionym: Hicriopteris si- mulans Ching, Acta Phvtotax. Sin. 8: 133, 101. 1959. Diplopterygium simulans (Ching) Ching, in R. H. Shing, Gloss. Term. Nam. Ferns 50. 1982, nom. inval. TYPE: China. Hainan: C. Wang 35496 (holotype, PE). Diplopterygium yunnanense (Ching) Ching ex X. C. Zhang, comb. nov. Basionym: Hicriopteris yunnanensis Ching, in S. S. Chien & Chun, FI. Reipubl. Popul. Sin. 2: 350. 1959. Diplopter¬ ygium yunnanense (Ching) Ching, in R. II. Shing, Gloss. Term. Nam. Ferns 50. 1982, nom. inval. TYPE: China. Yunnan: Jingdong, Wuliang Shan, W. //. Hsu 6 (holotype, PE). Acknowledgments. I am indebted to Alan Smith for discussion of the generic classification of Di¬ plopterygium and Hicriopteris. Thanks also to Prof. Werner Greuter for nomenclatural comments. Literature Cited Ching, 1C C. 1940. On natural classification of the family “Polypodiaceae.” Sunyatsenia 5: 201—268. - . 1978. The Chinese fern families and genera: Sys¬ tematic arrangement and historical origin. Acta Phyto- tax. Sin. 16: 1-37. - . S. H. I n, Chu 11. Wang & k. H. Shing. 1959. Gleicheniaceae. In: S. S. Chien & W. Y. Chun, Flora Reipuhlicae Popularis Sinicae 2: 1 16—132. Science Press. Beijing. Copeland, E. B. 1947. Genera Filieum. Chronica Botani- ca, Waltham, Massachusetts. Greuter, W., J. McNeill. F. B. Barrie, H. M. Burdet, V. Demoulin, T. S. Filgueiras, I). II. Nicolson, P. C. Silva, J. E. Skog, P. Trehane, N. J. Turland & I). E. Hawk- sworth (editors). 2000. International Code of Botanical Nomenclature (Saint Louis Code). Regnum Veg. 138. Holttum, R. E. 1957. On the taxonomic subdivision of the Gleicheniaceae, with description of new Malaysian spe¬ cies and varieties. Reinwardtia 4: 257—280. - . 1959. Gleicheniaceae. Flora Malesiana ser. 2, 1: 1—36. N.V. Erven P. Noordhoff, Groningen. Kramer, k. U. 1990. Gleicheniaceae. Pp. 145—152 in k. Kubitzki (editor). The Families and Genera of Vascular Plants, Vol. 1. Pteridophytes and Gymnosperms. Vol. editors, k. U. kramer X P. S. Green. Springer- Verlag, Berlin. kung, S. H. 1988. Flora Sichuanica, Tom. 6. Sichuan Sci¬ ence & Technology Publishing House, Chengdu. Ostergaard Andersen, E. & B. Ollgaard. 2001. Gleichen¬ iaceae. In: G. Hailing & L. Andersson (editors). Flora of Ecuador 66: 105—170. Botanical Institute, Goteborg University, Goteborg, Sweden. Presl, C. 1851. Abh. kiinigl. Boh. ties. Wiss., ser. 5, 6: 386. [Epimel. Bot. 26. 1851.] Shing, K. 11. 1982. Glossary and Terms of Ferns. Science Press, Beijing. Volume 14, Number 1 2004 Zhang Pteridophytes from China 151 Tagawa, M. & K. Iwatsuki. 1979. Gleicheniaceae. In: T. Smitinand & K. Larsen, Flora of Thailand 3(1): 50—56. Tryon, 11. M. & A. F. Tryon. 1982. Ferns and Allied Plants with Special lieferenee to Tropical America. Springer, Berlin/Heidelberg/New York. Wang, P. S. & X. Y. Wang. 2(H) I . Pteridophyte Flora of Guizhou. Guizhou Science & Technology Press, Gui- yang. Wu, S. 11. & If. C. Ching. 1991. Fern Families and Genera of China. Science Press, Beijing. Volume 14, Number 1, pp. 1-152 of NOVON was published on 23 March 2004. Volume 14 NOVON Number 2 2004 Novelties and Notes on Miscellaneous Asian Brassicaceae Ihsan A. Al-Shehbaz Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299, U.S.A. ihsan.al-shehbaz@mobot.org Abstract. Draba bartholomewii (Brassicaceae), a new species from Qinghai, China, is described and illustrated. The monotypic genera Coelonema and Nesocrambe are reduced to the synonymy of Draba and Hemicrambe, respectively. The new combina¬ tions Alyssum dahuricum, Draba draboides, Hemi¬ crambe socotrana, Lepidium coronapus, and Pe- gaeophyton purii are proposed. Key words: Alyssum , Asia, Brassicaceae, Coe¬ lonema, Dilophia, Draba, Hemicrambe, Lepidium, Nesocrambe, Pegaeophyton, Ptilotrichum. Delimitation of genera of Brassicaceae (Crucifer- ae) is a well-known problem, and 125 of the 337 genera recognized by Appel and Al-Shehbaz (2003) are monotypic, with about an additional 100 con¬ taining two to four species (oligotypic). Molecular studies (see review by Koch et al., 2003) have clearly demonstrated that the vast majority of monotypic genera are indistinguishable from and nested within larger genera. Upon a critical re-eval¬ uation of morphology, two monotypic genera, Coe¬ lonema Maximowicz and Nesocrambe A. G. Miller, are reduced herein to synonymy of other genera well established on the basis of morphological and molecular grounds (Koch & Al-Shehbaz, 2002; Koch et ah, 2003; Warwick & Black, 1997; Gomez- Campo, 1999). I'he present paper deals with the description of a new species of Draba, as well as nomenclatural adjustments in the genera Alyssum I.., Draba L, Hemicrambe Webb, Lepidium L., and Pegaeophyton Hayek & Handel-Mazzetti. Alyssum versus Ptilotrichum Although most authors place Ptilotrichum C. A. Meyer in the synonymy of Alyssum (e.g., Dudley, 1964; Ball & Dudley, 1993; Zhou et ah, 2001; Ap¬ pel & Al-Shehbaz, 2003), a few others (e.g., Re- benskaya, 1994; Czerepanov, 1995) maintain both genera. As indicated by Zhou et ah (2001), the characters used to distinguish Ptilotrichum from Alyssum (e.g., white vs. yellow flowers, edentate vs. dentate filaments, and 1 -seeded vs. 2- or 3-seeded locules) are unreliable and found within Alyssum excluding Ptilotrichum. Therefore, these alleged generic boundaries are artificial, and the two gen¬ era should be united. All except one of the species originally described in Ptilotrichum have names in Alyssum. The following new combination is needed. Alyssum dahuricum (Peschkova) Al-Shehbaz, comb. nov. Basionym: Ptilotrichum dahuricum Peschkova, Novosti Sist. Vyssh. Bast. 15: 230. 1979. TYPE: Russia. Ghita: Onon River sys¬ tem, mountain at Bukukun River, near Buku- kunskij, 18 July 1913, V. Smirnov 1986 (ho- lotype, LE). Alyssum dahuricum is easily distinguished at an- thesis from the closely related A. tenuifolium Ste¬ phan ex Willdenow by having stramineous stems and by lacking the basal and lowermost cauline leaves. By contrast, A. tenuifolium has purplish or greenish stems and persistent basal rosette and cauline leaves. Draba versus Coelonema The monotypic Coelonema was recognized in various Chinese floras (e.g., Zhou et al., 2001) as endemic and was said to differ from Draba solely by including stoloniferous perennials with flattened Novon 14: 153-157. 2004. 154 Novon bases of staminal filaments. Draba was said to con¬ sist of annuals, biennials, or perennials with slen¬ der or slightly flattened staminal filaments. A crit¬ ical examination at LE of the type of C. draboides Maximowicz, as well as numerous other species of Asian Draba, reveals that these alleged differences are insignificant because several species of Draba (e.g., D. sibirica (Pallas) Thellung) are typically sto- loniferous, and the filament bases vary in this com¬ plex genus of ca. 350 species from slender to dis¬ tinctly flattened. Therefore, the conclusion reached by Appel and Al-Shehbaz (2003) in uniting Coe- lonema with Draba is justified, and the following new combination is needed to accommodate C. dra¬ boides in Draba. Draba draboides (Maximowicz) Al-Shehbaz, comb. nov. Basionym: Coelonema draboides Maximowicz, Bull. Acad. Imp. Sci. Saint-Pe- tersbourg 20: 424. 1880, TYPE: China. [Gan¬ su]: “Terra Tangutorum. Jugum S a fl. Tetung ad declivitates denudates mintium rarius, 30 A pri 1 — 1 2 May 1873,” N. M. Prezewalski 14 (holotype, LE). Draba During a visit to LE in 2002, I examined the four syntypes of Draba eriopoda Turczaninow, a species widely distributed in Bhutan, China, India, Mongolia. Nepal, and Russia. It became immedi¬ ately evident that the treatment of D. eriopoda in Zhou et al. (2001) represented two distinct species, of which one is described below as new. Draba bartbolomewii Al-Shehbaz, sp. nov. TYPE: China. Qinghai: Dari (Darlag) Xian, just N of Manzhang, along Manzhang He, belw. Dari & Banma (Baima), at base of valley sides in loose soil, 4000 m, 33°17'51"N, 100°25'55"E, 12 Aug. 1993, Ho Tingnung, Bruce Bartholomew A- Michael G. Gilbert 1 185 (holotype, MO; isotypes, BM. CAS, E, GH, HNWP, PE). Figure 1. Herba annua. Caules decumbentes, ad basim ramosi, pilis stellatis praediti. Folia basalia nulla; folia caulina 2- 6(— 10), sessilia, nonauriculata, ovata vel elliptica, 1—3.5 X 0.4—1. 5 cm, subtus pilis subsessilibus stellatis 4-radia- tis praedita, supra pilis subsetosis simplicibus et stipitato- furcatis praedita. Racemi 8— 20-flori, rharhidibus rectis. Pedicelli fructiferi horizontales, 5—15 mm longi. Petala Hava, spathulata, 1.2-1. 6 X 0.5— 0.6 mm. Ovarium 14— 20-ovulatum. Fructus oblongi, D — 1 ()(— 1 2) X (2.5— )3— 3.5 mm, compressi, pilis simplicibus et subsessilif)us furcatis praediti. Semina 1—1.3 X 0.7-0.9 mm. Herbs annual, 5-25 cm tall; stems decumbent. branched primarily at or near base, pubescent with subsessile stellate trichomes rarely mixed with sim¬ ple ones, rarely glabrous distally. Basal leaves w ith¬ ered by anthesis; cauline leaves 2 to 6(to 10), ses¬ sile, not auriculate; leaf blade ovate to elliptic, I — 3.5 X 0.4— 1.5 cm, abaxiallv pubescent with subsessile or short-stalked, 4-rayed, stellate tri¬ chomes with simple rays, adaxially with subsetose simple or long-stalked forked trichomes sometimes mixed with fewer, smaller, 4-rayed stellate ones, base cuneate, margin 3- to 5-toothed on each side, apex acute. Racemes 8- to 20-flowered, ebracteate, elongated in fruit; rachis straight; fruiting pedicels 5—15 mm long, horizontal, straight, pubescent all around, or rarely glabrescent. Sepals oblong, 1-1.3 X 0.6-0. 8 mm, erect, abaxially pilose with stellate trichomes, base of lateral pair not saccate, margin membranous; petals yellow, spatulate, 1.2— 1.6 X 0.5— 0.6 mm, apex retuse, obscurely clawed at base; filaments 0.8-1 mm; anthers ovate, 0. 1-0.2 mm; ovules 14 to 20 per ovary. Fruit oblong, 8-10(-12) X (2.5— )3— 3.5 mm, often horizontal, latiseptate, not twisted; valves sparsely puberulent with simple and subsessile forked trichomes, rarely glabrescent, ob¬ scurely veined, base and apex acute; style obsolete, rarely to 0.1 mm; seeds brown, ovate, 1-1.3 X 0.7— 0.9 mm. Flowering and fruiting in August. fhe new species is named after Bruce Barthol¬ omew, one of the collectors of the type collection. Draba bartholomewii is easily distinguished from D. eriopoda by having decumbent stems, horizontal fruiting pedicels pubescent all around or rarely gla¬ brous, pilose sepals with stellate trichomes, small petals 1.2— 1.6 X 0.5— 0.6 mm, oblong, often hori¬ zontal fruits (2.5-)3— 3.5 mm wide, and fruit valves puberulent with simple and subsessile forked tri¬ chomes. By contrast, D. eriopoda has erect stems, divaricate-ascending fruiting pedicels often gla¬ brous adaxially and pubescent abaxially, pilose se¬ pals with simple trichomes, larger petals 2— 3(— 3.5) X (0.5— )0. 8—1 mm, erect or ascending, ovate or ovate-elliptic fruits 1.5-2. 5 mm wide, and glabrous fruit valves. Draba bartholomewii is also related to I), alticola Komarov (Afghanistan, China, Kyrgyzstan. Tajikis¬ tan), which it resembles in flower size and fruit shape. However, the latter has rosulate basal leaves, leafless stems, a (lexuous rachis in fruiting racemes, pilose sepals with simple and forked tri¬ chomes, glabrous fruits 1-2 mm wide, and seeds 0.4— 0.5 mm long. Draba bartholomewii lacks the basal leaf rosette and has 2 to 6(to 10) cauline leaves, a straight rachis in fruiting racemes, pilose Volume 14, Number 2 2004 Al-Shehbaz Asian Brassicaceae 155 Figure 1. Draba bartholomewii Al-Shehbaz. —A. Plant. — B. Trichomes of abaxial leaf surface. — C. Trichomes of adaxial leaf surface. — D. Petal. Scales: A = 1 cm; B. C = 0.2 mm; D = 0.5 mm. Drawn by Al-Shehbaz from the holotype, Ho , Bartholomew & Gilbert I IBS (MO). 156 Novon sepals with stellate trichomes, puberulent fruits (2.5— )3— 3.5 mm wide, and seeds 0.7— 0.9 mm long. Paratypes. CHINA. <^inghai: Nangqen Xian, Xiao- long Gou, W of Nangqen on road to Domba, 32°17'N, 96°16'E, T. /V. Ho, H. Bartholomew, M. Watson & M. G. Gilbert 2822 (CAS, HNWP, MO); Tungjen Xian, T. P. Wang 6186 (PE). Sieliuan: Haitzeshan, //. Smith 1 1598 (MO, TI, UPS). Hemicrambe V ERSUS Nesocrambe Hemicrambe consists of two species, of which one, //. fruticulosa Webb, is endemic to Morocco and the other, H. fruticosa (C. C. Townsend) Gdmez- Campo, is endemic to the island Soqotra (Yemen). The two species are subshrubs disjunetly separated by some 0900 km (Gdmez-Campo, 1977, 1978; Townsend, 1971). The genus is characterized by having yellow or white flowers, petiolate and den¬ tate or lyrate-pinnatipartite leaves, patent median nectaries, reflexed fruiting pedieels. and segment¬ ed, flattened fruits with a 2- to 4-seeded, indehis- cent distal segment and a seedless or 1 -seeded, de¬ hiscent proximal segment (Appel & Al-Shehbaz, 2003). The recently described Nesocrambe (Miller et al., 2002) is indistinguishable from Hemicrambe in al¬ most every morphological aspect. It is said to be a perennial herb with a woody base instead of a sub¬ shrub, but this is an unreliable distinction. Both genera have well-developed, [latent, median nectar glands, a feature unique in the entire tribe Bras- sieeae (Townsend, 1971; Gdmez-Campo, 1977; Miller et al., 2002). Miller et al. (2002) did not include Nesocrambe socotrana A. G. Miller in Hem¬ icrambe because when they compared it with the Soqotran //. fruticosa they found that H. fruticosa has white instead of yellow flowers, dentate instead of lyrate-pinnatisect leaves, and cylindric instead of flattened fruits. The Moroccan //. fruticulosa has both yellow flowers and lyrate-pinnatipartite leaves. The only other difference between the two genera is that in Hemicrambe the distal fruit segment is compressed and the proximal segment is sometimes seedless, whereas in Nesocrambe the distal fruit segment is cylindric and the proximal segment is 1 -seeded. However, such highly artificial separation is good only at the species rather than the generic level. Miller et al. (2002: 62) have indicated that Nesocrambe “is clearly related to Hemicrambe." In my opinion, the remarkable similarities in nectar gland and almost every aspect of the plant (see ta¬ ble 1 in Miller et al., 2002: 63) strongly argue for the maintenance of one genus. Furthermore, there is no need to create a new monotypic genus when its nearest relatives, species of Hemicrambe, are known. Several other genera of the tribe Brassiceae (e.g., Conicya Porta & Rigor ex Roy, Vella L.) show fruit diversity far greater than in Hemicrambe in¬ cluding Nesocrambe (Leadlay & Heywood, 1990; Warwick & Al-Shehbaz, 1998). Miller et al. (2002: 66) stated that “the perennial habit is relatively uncommon in the Brassicaceae.” However, Al-Shehbaz (1984: 349) indicated that “more than 62 percent [of the family] are peren¬ nials” and only about 5% are typically woody. Key to the Species of Hemicrambe la. Leaves crenate-dentate; flowers white; Soqotra . H. fruticosa 1 b. Leaves lyrate-pinnatisect or lyrate-pinnatipartite; flowers yellow. 2a. Perennial herbs woody only at base; fruits cylindric; Soqotra . H. socotrana 2b. Subshrubs; fruits flattened; Morocco . . . . H. fruticulosa Hemicrambe socotrana (A. G. Miller) Al-Sheh¬ baz, comb. nov. Basionym: Nesocrambe soco¬ trana A. G. Miller, in Miller et al., Willden- owia 32: 63. 2002. TYPE: Republic of Yemen. Western Soqotra, Sharahin, 12°31'N, 53°20'E, 450 m, 6 Feb. 2001, A. G. Miller, R. Atkinson, A. W. Kulaidi & N. Taleeb M -2002 1 (holotype, E; isotypes, B not seen, UPS not seen). Lepidium Based on extensive molecular studies (Brtigge- mann, 2000) and critical evaluation ol morphology (Appel & Al-Shehbaz, 2003; Al-Shehbaz et al., 2002), Coronopus Zinn was reduced to the synon¬ ymy of Lepidium. All except one species of Coron¬ opus have correct names in Lepidium. The excep¬ tion is C. squamatus (Forsskdl) Ascherson, a species originally described in Lepidium by Forss- kal (1775). However, the earliest validly published name for the species is Cochlearia coronopus L. (Linnaeus, 1753), and this specific epithet should be used in Lepidium when this genus is united with Coronopus. Therefore, the following new combina¬ tion is needed. Lepidium coronopus (L.) Al-Shehbaz, comb. nov. Basionym: Cochlearia coronopus L., Sp. PI. 2: 648. 1753. TYPE: (lectotype, designated by Jonsell & Jarvis, 2002: 68) Herb. Linn. 826.5 (LINN). Pegaeophyton versus Dilophia Rawat et al. (1996) described the new species Dilophia purii from Garhwal in Northwest India. Volume 14, Number 2 2004 Al-Shehbaz Asian Brassicaceae 157 The excellent illustration immediately suggested a possible affinity to Pegaeophyton. However, the de¬ scription and illustration indicated the presence of glandular trichomes, a feature not known in any species of Pegaeophyton and Dilophia Thomson. When I examined the MO isotype of D. purii, I found none of the trichomes were glandular, and the generic affinity of this plant to Pegaeophyton became abundantly clear. Pegaeophyton differs from Dilophia by having accumbent instead of in¬ cumbent cotyledons, nonfleshy instead of fleshy fu¬ siform roots, obtuse instead of strongly apiculate anthers, and smooth instead of apically cristate fruits. Pegaeophyton purii (1). S. Rawat, L. R. Dangwal & R. D. Gaur) Al-Shehbaz, comb. nov. Ras- ionym: Dilophia purii I). S. Rawat, L. R. Dang¬ wal & R. I). Gaur, J. Bombay Nat. Hist. Soc. 93: 262. 1996. TYPE: India. Northwest Him¬ alaya, Roopknnd, 4850 m, 11 Aug. 1993, D. S. Rawat .s'.//, (holotype, GUH 22,498 not seen; isotype, MO). Except for having flattened instead of terete and 8- instead of 2- to 4-seeded fruits. Pegaeophyton purii is quite similar in all aspects of leaves and flowers to P. nepalense Al-Shehbaz, Arai & H. Ohba, a species originally thought to be endemic to Nepal (Al-Shehbaz et ah, 1998) but later re¬ ported from Bhutan, China, and India (Al-Shehbaz, 2000). The collection of mature fruiting material of P. purii should help in the determination of whether or not the two species are conspecific. Acknowledgments. I am profoundly grateful to Henk van der Werff for correcting the Latin, John H. Wiersma for his note on Lepidium coronopus, Antony G. Miller for allowing me to examine the type collection of Nesocrambe socotrana, Steve Caf- ferty for the typification of Cochlearia coronopus, D. S. Rawat for sending an isotype of Dilophia purii , and Nicholas J. Turland for opinions on various no- menclatural matters. I am deeply indebted to Vic¬ toria C. Hollowell. Neil A. Harriman, and Steve L. O'Kane, Jr., for their editorial advice and review of the manuscript. Literature Cited Al-Shehbaz. I. A. 1984. The tribes of Cruciferae (Brassi¬ caceae) in the southeastern United States. .1. Arnold Ar¬ bor. 65: 343—373. - . 2000. A revision of Pegaeophyton (Brassicaceae). Edinburgh J. Bot. 57: 157-170. - . K. Arai & H. Ohba. 1998. A new Pegaeophyton from Nepal. Novon 8: 327—329. - , K. Mummenhoff & O. Appel. 2002. Cardona, Coronopus, and Stroganowia are united with Lepidium (Brassicaceae). Novon 12: 5—12. Appel, O. & 1. A. Al-Shehbaz. 2003. Cruciferae. In: K. Kubitzki (editor), Families and Genera of Vascular Plants. 5: 75—174. Springer- Verlag, Berlin, Heidelberg. Ball, P. W. & T. R. Dudley. 1993. Alyssum. In T. G. Putin, N. A. Burges, A. (). Chater, J. R. Edmondson, V. 11. Heywood, D. M. Moore, I). H. Valentine, S. M. Walters & 1). A. Webb (editors), FI. Europaea, ed. 2, 1: 359— 369. Cambridge Univ. Press, Cambridge. Briiggemann, H. 2000. Molekulare Systematik und Bio- geographie der Gattung Lepidium L. und verwandter Sippen (Brassicaceae). Ph.D. Thesis, University of Os- nabriick. Czerepanov, S. K. 1995. Vascular Plants of Russia and Adjacent States (the Former USSR). Cambridge Univ. Press, Cambridge. Dudley, T. R. 1964. Synopsis of the genus Alyssum. J. Arnold Arbor. 45: 358—373. ForsskM, P. 1775. Flora aegyptiaco-arabica. Moller, Kjwbenhavn [Copenhagen], Gomez-Campo, C. 1977. Studies on Cruciferae: III. Hem- icramhe townsendii nom. nov. an example of geographic disjunction. Anal. Inst. Bot. Cavanilles 34: 151—155. - . 1978. Hemicrambe fruticosa (Townsend) Gomez- Campo. comb. nov. Uagascalia 7: 189—190. - . 1999. Taxonomy. Pp. 3—32 in C. Gomez-Campo (editor). Biology of Brassica Coenospecies. Elsevier, Amsterdam. Jonsell, B. & C. F. Jarvis. 2002. Lectotypifications ol Lin- naean Names for Flora Nordica (Brassicaceae— Api- aceae). Nordic J. Bot. 22: 67-86. Koch, M. & I. A. Al-Shehbaz. 2002 Molecular data in¬ dicate complex intra- and intercontinental differentia¬ tion of American Draba (Brassicaceae). Ann. Missouri Bot. Card. 89: 88-109. - , - & K. Mummenhoff. 2003. Molecular sys- tematics, evolution, and population biology in the mus¬ tard family (Brassicaceae). Ann. Missouri Bot. Card. 90: 151-171.' Leadlay, E. A. & V. H. Heywood. 1990. The biology and systematics of the genus Coincya Porta & Rigo ex Rouy (Cruciferae). Bot. J. Linn. Soc. 102: 313—398. Linnaeus, C. 1753. Species Plantarurri. Impensis laurentii Slavii, Holmiae [Stockholm]. Miller, A. G.. R. Atkinson, A. W. Kulaidi & N. Taleeb. 2002. Nesocrambe, a new genus of Cruciferae (Brassi¬ caceae) from Soqotra, Yemen. W illdenowia 32: 61-67. Rawat, I). S., L. R. Dangwal & R. 1) Gaur. 1996. A new species of Dilophia Thoms. (Brassicaceae) from Ga- rhwal Himalaya (India). J. Bombay Nat. Hist. Soc. 93: 262—264. Rebenskaya, E. V. 1994. Ptilotrichum C. A. Meyer. Pp. 106—107 in L. I. Malyschev & G. A. Pesc-hova (editors), EL Sibiriae, 7. Nauka, Novosibirsk. Townsend, C. C. 1971. Fabrisinapis fruticosus. Hooker’s Icon. PL 7(4): tab. 3673. Warwick. S. I. & I. A. Al-Shehbaz. 1998. Generic evalu¬ ation of Boleum, Euzomodendron, and Vella (Brassica¬ ceae). Novon 8: 321-325. - & L. D. Black. 1997. Phylogenetic implications of chloroplast DN A restriction site variation in subtribe Raphaninae and Cakilinae (Brassicaceae, tribe Brassi- ceae). Canad. J. Bot. 75: 960—973. Zhou, T. Y., L. L. Lu, G. Yang & 1. A. Al-Shehbaz. 2001. Brassicaceae. Pp. 1—193 m C. V. Wu & P. H. Raven (editors). Flora of China, Vol. 8. Science Press, Beijing, and Missouri Botanical Garden Press, St. Louis. Psychotria carrascoana (Rubiaceae), a New Species from the Carrasco Vegetation of Northeastern Brazil Piero G. Delprete Institute of Systematic Botany, The New York Botanical Garden, Bronx, New York 10458- 5126, U.S.A. pdelprete@nybg.org. Current address: National Herbarium of the Netherlands, Utrecht University Branch, Plant Systematics, Heidelberglaan 2, 3584 CS Utrecht, The Netherlands, p.delprete@bio.uu.nl Elnatan Bezerra de Souza Curso de Biologia, Universidade Estadual Vale do Acarau, Sobral, Ceara, 62040-370, Brazil. ebsouza@nvanet.br Abstract. Psychotria carrascoana (Rubiaceae, Psychotrieae) is here described and illustrated. It is endemic to carrasco vegetation, a closed, shrub¬ by, xerophilous vegetation on quartzitic sand soils, on the plateaus of the Serra da Ibiapaba and Serra do Araripe, at altitudes of 800 to 000 m, state of Ceara, Brazil. Because of its reduced, triangular stipules, leaves drying pale green, commonly soli¬ tary flowers, and dorsally tricostate pyrenes, this species is placed in Psychotria subg. Heteropsycho- tria. Key words: carrasco , Ceara, northeastern Bra¬ zil, Psychotria, Psychotrieae, Rubiaceae, shrub- land. As part of the project Rubiaceae do Estado do Ceara, in northeastern Brazil, fieldwork was con¬ ducted during 1998—2000, which has produced about 400 collections of species of this family. A preliminary checklist of the Rubiaceae of Ceara was recently presented (Delprete et ah, 2001), and an annotated checklist will be published in the near future (Delprete & Souza, in prep.). Two new species of Rubiaceae were recently described from the arid vegetation of this state (Simira gardneriana Barbosa & Peixoto (2000); Mitracarpus longicalyx Souza & Sales (2001)). and the identity of some other taxa collected during this project is currently under study. Among these, a species of Psychotria restricted to carrasco vegetation, and readily distin¬ guishable by its small leaves drying pale green, tri¬ angular stipules, flowers commonly solitary, and dorsally trieostate pyrenes, is described below. Ecological Obskhvations About Carrasco Vegetation The definition of carrasco vegetation lias long been debated, as it is not readily distinguishable from that of caatinga, the widespread xerophilous vegetation of northeastern Brazil. Caatinga is com¬ monly characterized by two vegetational layers: sparse to moderately dense trees and shrubs fre¬ quently armed with thorns and almost completely leafless during the dry season, and the lower laver with the presence of Bromeliaceae and Cactaceae, and annual herbs (Rizzini, 1997). Andrade-Lima (1978) stated that carrasco vegetation is physiog- nomically distinct from caatinga vegetation be¬ cause of the high density of shrubs and treelets, single-layered vegetation, and the almost complete absence of Cactaceae and Bromeliaceae. Rizzini (1997) and Fernandes (1981) recognized carrasco vegetation as sufficiently distinct from caatinga vegetation. Fernandes (1990) and Fernandes and Bezerra (1990) postulated two possible hypotheses about the identity and origin of carrasco vegetation: either as a distinct natural vegetation, or as a result of partial degradation of the cerradao vegetation, which gives the general aspect of a dense capoeira (secondary vegetation that can be originated from the degradation of several possible vegetation types). Cerradao is usually characterized by three vegetational layers: a tree layer 8— 1 2( — 1 8) m tall, with closed canopy and almost completely devoid of epiphytes; an intermediate arbustive layer about 3—5 m tall and with sparse shrubs and rare lianas; and a lower herbaceous layer, composed primarily of Cyperaceae, Poaceae, and Bromeliaceae (ef. Riz¬ zini. 1997). Araujo et al. (1998a, 1998b), in a project focused on the plant community and floristic composition of carrasco, studied three plots in the southern portion of the Planalto da Ibiapaba, near Novo Oriente, Ceara. Brazil. As a result of that study, only 31 out of 102 shrubby and arboreal species were found to Novon 14: 158-162. 2004. Volume 14, Number 2 2004 Delprete & Souza Psychotria carrascoana from Brazil 159 be exclusive to carrasco, while the other species were shared with the contiguous vegetation types. Those data alone confirmed the difficulty of decid¬ ing if carrasco vegetation should be treated as a degraded cerradao or a distinct vegetation type. Af¬ ter a series of additional studies, Araujo and Mar¬ tins (1999) and Araujo et al. (1999) recently con¬ cluded that carrasco vegetation can be defined as a deciduous, dense, single-layered shrubland, with vines, irregular canopy, and sparse trees, with an average annual precipitation of 668—1289 mm, a dry season ranging from 5 to 7 months a year, grow¬ ing on deep, acidic, distrofic quartzitic sands, at 700—900 m elevation. According to those authors, carrasco vegetation occurs within the semi-arid do¬ main of northeastern Brazil, at the Chapada do Ar- aripe, southern Ceara, and Planalto da Ibiapaba, at the border area of the states of Ceara and Piauf. The new species of Psychotria described below is endemic to this vegetation (hence the specific ep¬ ithet), and it grows in the shade of shrubs and tree- lets. Psychotria carrascoana Delprete & E. B. Souza, sp. nov. TYPE: Brazil. Ceara: Planalto da Ibia¬ paba, Mun. Ubajara, Jaburuna Sul. 830 m, 5 Jan. 1995 (fl), F. S. Araujo 1054 (holotype, EAC; isotype, NY). Figure 1. Haec species, suffrutex usque ad 1.3 m altus, quoad inflorescentiam terminalem subsessilem paucifloram P. subtriflorae similis, sed ab ea stipulis triangularibus (nec bilobis), inflorescentia ebracteata (nec bracteis naviculi- formibus subtenta), tubo corollae 1.7— 2.8 mm (nec 6—7 mm) longo atque fructibus rubris (nec cyano-purpureis) diversa. Subshrub 0.4— 1.3 m tall, densely branched from the base; main stems often tortuous; bark thin, creamy white; raphides present in all plant parts. Stipules free at base, triangular. 1-2 X 0.8—2 mm, margin entire or sometimes denticulate, apex acute, glabrous outside, pubescent inside; leaves opposite, sessile to subpetiolate; petioles to 1.5 mm long; blades 9—30 X 2-9 mm, linear, narrowly elliptic to lanceolate, base attenuate, apex acute to obtuse, margins narrowly revolute, chartaceous, glabrous throughout, pale green; midvein prominent below, discolorous; secondary veins 5 to 9 each side, em¬ bedded within the lamina; domatia absent. Inflo¬ rescence uniflorous (rarely biflorous), terminal, on lateral short-shoots of 5 to 7 reduced nodes. Flow¬ ers subsessile to short-pedicellate; pedicels to 1.3 mm long; hypanthium broadly turbinate, ea. 2 X 1.2 mm, glabrous; calyx shallowly cupular; tube 0.2— 0.3 mm long; calyx lobes 5(6), 0.3— 0.5 mm long, deltoid to narrowly triangular, acute; corolla narrowly infundibuliform, 4.5— 6.5 mm long, gla¬ brous outside, with a ring of hairs from stamen in¬ sertion to mouth inside; corolla tube 1.7— 2.8 mm long; corolla lobes 5 (rarely 6), 2. 5-3.8 X 1.4-1. 7 mm, oblong-ovate, acute and slightly thickened at apex, antrorse pubescent at basal portion inside; stamens subsessile, partially exserted; filaments 0.2— 0.5 mm long, glabrous; anthers oblong, 0.7— 0.9 X 0.3— 0.5 mm; disk entire; style terete, 4—6 mm long; style branches spatuiate, 0.5— 0.7 X 0.3— 0.4 mm. Fruits drupaceous, ovoid to ellipsoid, 3—5 X 3—4 mm. red; pyrenes plano-convex, dorsally 3-cos¬ tate; seeds suborbicular, ca. 2.5 X 2 mm, pale brown; exotesta smooth. Distribution, habitat, and phenology. Endemic to the carrasco vegetation of the Planalto da Ibia¬ paba and Chapada do Araripe, slate of Ceara, and probably occurring in the same vegetation type in the contiguous state of Piauf, always observed and collected in the shade of taller shrubs, at 800—900 m elevation. Flowering specimens were collected in January, and fruiting specimens in March and Sep¬ tember. Paratypes. BRAZIL. Ceara: Planalto da Ibiapaba, Mun. Ubajara, Jaburuna Sul. 830 m. 27 Apr. 1994 (st). F. S. Araujo 679 (LAG), 23 May 1994 (st), 768 (EAC); Plan¬ alto da Ibiapaba, Mun. Novo Oriente, Baixa Fria, 16 Feb. 1991 (st). F. .S'. Araujo 278 (EAC); Planalto da Ibiapaba, Mun. Tiangua. 19 Nov. 1994 (st). A. G. Fernandes & F. ,/. A. Matos FAC 21465 (EAC); Planalto da Ibiapaba, BR- 222. 3 km from Tiangud, 3°45'S, 41°01 'W, ca. 800 m, 21 Mar. 2000 (st), Delprete et al. 7196 (EAC. NY. uva [Univ- ersidade Vale do Acarau|), 7197 (EAC, uva), 7198 (EAC, NY, uva), 7199 (EAC, NY, uva), 7203 (EAC, uva); Cha¬ pada do Araripe, Mun. Crato, Sftio Mirindiva, 7°17'S, 39°33'W, 900 m. 29 Mar. 2000 (voting f r). Delprete et id. 7304 (EAC. NY, uva), s.n. (EAC.‘ k. MO, NY. US); Cha¬ pada do Araripe, Mun. Crato, 19 Sep. 2001 (fr), F. S. Cavalcanti tV F. Silveira 843 (EAC, NY, uva). Psychotria L., as traditionally recognized, is one of the largest genera of flowering plants, with about 2000 species worldwide. Several authors have tried to organize this taxonomic conundrum, either by dividing it into several smaller genera (e.g., Bre- mekamp, 1934), or by treating it as a large genus with many sections (e.g., Steyermark, 1972, 1974). Psychotria s.l. has been shown to be paraphyletic in the molecular phytogenies presented by Anders- son and Rova (rp.s l6; 1999), and Nepokroeff el al. (ITS and rbch; 1999). Following ihe molecular data published by these authors and additional morpho¬ logical evidence, the genera IS'otopleura Breme- kamp and Carapichea Aublet have been recently resurrected by Taylor (2001) and Andersson (2002), respectively. Notopleura is easily distinguished, among other morphological characters, by its pseu- 160 Novon Figure 1. Psychotria carrascoana Delprete & E. B. Souza. — A. Branch with immature fruits. — B. Short-shoot with solitary, terminal (lower hud. — C. Flower in anthesis. — 1). Style. — E. Section of open corolla. — F. Mature fruit. A and F from photos of living material; B— E, from Araujo 1054 (types, EAC, NY). Volume 14, Number 2 2004 Delprete & Souza Psychotria carrascoana from Brazil 161 doaxillary inflorescences, and the sheathing stip¬ ules with an interpetiolar appendage inserted below the upper margin (Taylor, 2001); and Cnrapichen by its terminal, condensed inflorescences subtend¬ ed by foliose bracts, pyrenes with germination slits on abaxial ridges, and persistent stipules, withering on the stem (Andersson, 2002; Delprete, 2001, 2003). The rest of the Psychotria complex, cur¬ rently divided into subgenus Psychotria (Steyer- mark, 1972: 444) and subgenus Heteropsychotria Steyermark (1972: 484), remains taxonomically un¬ resolved, and its close relationship with Palicourea Aublet has been addressed (e.g., Muller Argovien- sis, 1881: Nepokroeff et ah, 1999), but not yet re¬ solved (Taylor, 1996. 1997, and work in progress). About 1 I species of Psychotria are known from the state of Ceara (Delprete et al., 2001), and among them Psychotria carrascoana is the species with the smallest leaves, and the only one with sol¬ itary (or rarely paired) flowers. This species has of¬ ten been collected when sterile, mostly because of its minute, usually solitary, greenish white flowers, and therefore difficult to spot during anthesis. Be¬ cause of the reduced, triangular stipules, leaves drying pale green, terminal flowers commonly sol¬ itary (or rarely in pairs), fruits red, and pyrenes dorsally tricostate, Psychotria carrascoana is placed in Psychotria subg. Heteropsychotria. 'I bis species is most similar to Psychotria subtriflora Muller Ar- goviensis (Zappi & Stannard, 1995: 571) because of the pauciflorous, subsessile, terminal inflores¬ cences, but it differs by having stipules triangular (vs. bilobed, lobes narrowly triangular in P. subtri¬ flora). inflorescences not subtended by bracts (vs. subtended by navicular bracts in P. subtriflora), co¬ rolla lube 1.7-2. 8 mm long (vs. 6-7 mm long in P. subtriflora), and red fruits (vs. purplish blue in P. subtriflora). Acknowledgments. We are grateful to Francisca Simdes Cavalcanti, Curator of the Herbario Prisco Bezerra (EAC; Universidade Federal do Ceara, For¬ taleza) and Coordinator of the project Bubiaceas do Ceara. for help with field and herbarium work; to Afranio Fernandes (EAC) for accompanying us dur¬ ing fieldwork in the carrasco vegetation of the Plan- alto da Ibiapaba and Chapada do Araripe; to Fran¬ cisco Jose de Abreu Matos (Laboratorio de Produtos Naturais, Universidade Federal do Ceara, Fortale¬ za) for contributing use of the university vehicle for our expeditions during 1998-2000; to the Conselho Nacional de Pesquisa (CNPq) for granting support and permission for collecting Rubiaceae in Ceara; and to The New York Botanical Garden, the Univ¬ ersidade Federal do Ceara, and the Universidade Estadual Vale do Aearau (uva; Sobral, Ceara) for economic and logistic support. The herbarium of the Universidade Estadual Vale do Aearau (ca. 4000 specimens), the institution of the junior au¬ thor, is not yet included in the Index Herbariorum. We are thankful to Michael Nee (NY) for proof¬ reading a draft of the manuscript, Bruno Manara (VEN) and Roy Gereau (MO) for correcting the Lat¬ in diagnosis, and William Burger (F) and an anon¬ ymous reviewer for helpful comments on the man¬ uscript. Literature Cited Andersson, L. 2002. Re-establishment of Carapichea (Ru¬ biaceae, Psychotrieae). Kew Bull. 57: 363—374. - & J. H. E. Rova. 1999. The rps 16 intron in the phylogeny of the Rubioideae (Rubiaceae). PI. Syst. Evol. 214: 161-186. Andrade-Lima, D. 1978. Vegetagao. Pp. 131—135 in R. C. Lins (editor), Bacia do Parnafba: Aspeetos Fisio- graficos. fnstituto Joaquim Nabuco de Pesquisas So- ciais, Sdrie Estudos e Pesquisas 9. Araujo, F. S. & F. R. Martins. 1999. Fisionomia e organ- izagao da vegetagao do carrasco no planalto da Ibia¬ paba, Estado do Ceard. Acta Bot. Brasil. 13(1): 1—13. - . F. V. S. B. Sampaio, M. A. Figueiredo, M. ,1. N. Rodal & A. G. Fernandes. 1998a. Composigao florfstica da vegetagao de carrasco. Novo Oriente, CE. Rev. Bras- d. Bot. 21: 10.5-116. - . - , M. J. N. Rodal & M. A. Figueiredo. 1998b. Organizagao comunitdria do componente len- hoso de tres areas de carrasco em Novo Oriente — CE. Rev. Brasil. Biol. 58: 85—95. - , - & G. J. Shepherd. 1999. VariagSes es- truturais e florfsticas do carrasco no planalto da Ibia¬ paba, estado do Ceard. Rev. Bras. Biol. 59: 663—678. Barbosa, M. R. de V. & A. I.. Peixoto. 2000. A new spe¬ cies of Simira (Rubiaceae, Rondeletieae) from north¬ eastern Brazil. Novon 10: 110—112. Bremekamp, C. F. B. 1934. Rubiaceae. Pp. 1 13-298 in A. Pulle, Flora of Surinam, Vol. 4. J. H. de Bussy, Am¬ sterdam, The Netherlands. Delprete, P. G. 2001. Notes on some South American spe¬ cies of Psychotria subgenus Heteropsychotria (Rubi¬ aceae), with observations on rubiaceous taxonomic characters. Brittonia 53: 396—404. - . 2003. Carapichea guianensis: The correct name for Psychotria carapichea (Rubiaceae), with a correction on the authority of P. carapichea, and a new combina¬ tion in Carapichea. Brittonia 55: 88-89. - . E. B. Souza & F. Simoes Cavalcanti. 2001. Checklist preliminar das Rubiaceae do Estado do Ceard (Brasil). Poster. 52° Congresso Nacional de Botanica (Joao Pessoa, 22-28 Jul. 2001). Programa e Resumos, p. 309. Grafica UFPB, Joao Pessoa, Paraiba, Brazil. Fernandes, A. 1981. Vegetagao do Piauf. XXXII Congres¬ so Nacional de Botanica. Programa e Resumos, pp. 7— 9. Grafica FUFPI, Teresina, Piaut, Brazil. - . 1990. Temas fitogeogrdfieos. Stylus Comunica- goes, Fortaleza, Ceara, Brazil. - & P. Bezerra. 1990. Estudo fitogeografico do Bras¬ il. Stylus Comunicagoes, Fortaleza, Ceard, Brazil. Muller Argoviensis, J. J. 1881. Rubiaceae, Tribus VI. Psy¬ chotrieae. Pp. 162^158 in C. F. P. von Martius, A. G. 162 Novon Eichler & I. Urban (editors), Flora Brasiliensis, Vol. 6, part 5. Fleisher, Leipzig, Germany. Nepokroeff, M., B. Bremer & K. J. Sytsma. 1999. Reor- ganization of the genus Psychotria and the tribe Psy- ohotrieae (Rubiaceae) inferred from ITS and rbcL se¬ quence data. Syst. Bot. 24: 5—27. Rizzini, C. T. 1997. Tratado de Biogeografia do Brasil: Aspetos Ecoldgicos, Sociologieos e Florfstieos, 2nd Ed. (revised by C. M. Rizzini). Ambito Cultural Edigoes, Rio de Janeiro, Brazil. Souza, E. B. de & M. F Sales. 2001 [2002]. Mitracarpus longicalyx (Rubiaceae, Spermacoceae), a new species from northeastern Brazil. Brittonia 53: 482—486. Steyermark, .1. A. 1972. Psychotria (Rubiaceae). In B. Ma¬ guire, J. J. Wurdack & Collaborators, Botany of the Guayana Highlands, Part IX. Mem. New York Bot. Gard. 23: 406—717. - . 1974. Psychotria (Rubiaceae). In T. Lasser & J. A. Steyermark, Flora de Venezuela 9(3): 1 I 1 1 — 1683. Institute Botanico, Caracas, Venezuela. Taylor, C. M. 1996. Overview of the Psychotrieae (Rubi¬ aceae) in the Neotropics. Opera Bot. Belg. 7: 261—270. - . 1997. Conspectus of the genus Palicourea (Ru¬ biaceae: Psychotrieae) with the description of some new species from Ecuador and Colombia. Ann. Missouri Bot. Gard. 84: 224-262. - . 2001. Overview ol the Neotropical genus Noto- pleura (Rubiaceae, Psychotrieae), with the description of some new species. Ann. Missouri Bot. Gard. 88: 478— 5 1 5. Zappi, I). C. & B. F. Stannard. 1995. Rubiaceae. Pp. 546— 578 in B. F. Stannard (editor), Flora of the Pico das Almas, Chapada Diamantina— Bahia, Brazil. Royal Bo¬ tanic Gardens, Kew. Una Nueva Especie de Tibouchina (Melastomataceae) de Guerrero, Mexico Jesus Ricardo de Santiago Gomez Laboratorio tie Plantas Vasculares, Facultad de Ciencias, UNAM, Cireuilo Exterior, Ciudad Universitaria. Mexico, D.F. 04510. rdesantiago@correo.unam.mx RESUMEN. Se describe e iluslra Tibouchina ara- neicalyx de la vertiente del Oceano Pacifico de la Sierra Madre del Sur en Guerrero, Mexico. Se dis- cuten las posibles relaciones de esta especie con T. rnexicana (D. Don) Cogniaux y otras especies con flores pentameras y estambres fuertemente dimor- ficos, de las que se distingue principalmente por tener ramas agudamente tetragonas, lobulos del ea- liz triangulares mas cortos que el hipantio y petalos con cilios eglandulares. Abstract. Tibouchina araneicalyx is described and illustrated from the Pacific slopes of the Sierra Madre del Sur in Guerrero, Mexico. Its possible relationships with T. rnexicana (D. Don) Cogniaux and another Mexican species of Tibouchina with pentamerous flowers and strongly dimorphic sta¬ mens are discussed; it is principally distinguished from these species by the sharply tetragonal branches, the triangular calyx lobes shorter than the hypanthium, anti the petals with eglandular cil¬ ia. Key words: Guerrero, Melastomataceae, Mexi¬ co, Tibouchina. Tibouchina Aublet, con poco mas de 350 espe¬ cies, es el genero de frutos capsulares con mayor numero de especies de la familia Melastomataceae (Almeda, 1993; Renner, 1993). Se distribuye destle el norte tie Argentina hasta el noroeste tie Mexico, con su maxima diversidad en la euenea del Ama¬ zonas y regiones antlinas adyacentes, con alrededor de 10 especies en las Antiilas y en America Cen¬ tral. En Mexico esta representado por 25 especies de las cuales 21 o 22 son endemicas del pais, para un endemismo tie 84 a 88%, el mas alto dentro de los distintos generos tie Melastomataceae presentes en Mexico, esto de acuerdo con los principales tra- tamientos que han descrito o revisado las distintas especies del genero presentes en Mexico (Cog¬ niaux, 1891; Rrantlegee, 1913, 1914; Standley, 1924; Wilson, 1958; Williams, 1963; Wurdack. 1967; Almeda, 1993; Totlzia, 1999). De las 25 especies antes mencionadas, 9 de ellas, incluyendo la que se describe en este trabajo, presentan dos series de estambres fuertemente di- morfieos, los mayores con las tecas y conectivo ro- sados a rojizo-violaceos mucho mas largos que en los estambres menores que tienen anteras y conec¬ tivo amarillos, constituyendo un grupo claramente diferenciado en el que Tibouchina breedlovei Wur¬ dack. 7. chiapensis Wurdack y 7. roseotincta Totlzia presentan flores estrictamente tetrameras, mientras que T. araneicalyx descrita aquf, T. durangensis Standley, T. galeottiana (Triana) Cogniaux, T. me- xicana (D. Don) Cogniaux, T. rufipilis (Schlechten- dal) Cogniaux y T. thulia Todzia son basicamente pentameras, con algunas flores tetrameras, hexa- meras o muy raramente heptameras en las inflores- cencias fobs. pers.). En los herbarios mexicanos mas importantes Ti¬ bouchina se encuentra representado [tor un numero abundante de colectas. Debido a la compleja ta- xonomfa del genero, a su abundante variacion mor- fologica y a la earencia tie claves y descripciones adecuadas para las especies mexicanas, en nume- rosas ocasiones el material se encuentra mal de- terminado o sin asignacidn de esjtecies. Por tal mo- tivo, aetualmente se realiza una revision de los ejemplares de este genero con el objetivo de ubicar claramente las especies presentes en Mexico. Del material revisado, tanto de los herbarios antes men- cionados como de la amplia coleccidn del proyecto “Flora de Guerrero” de la Facultad de Ciencias de la Universidad Nacional Autonoma de Mexico (UNAM) aetualmente en revision, se detectaron va¬ ries espeefmenes de un taxon procedente de la Si¬ erra Madre del Sur de Guerrero con estambres mar- cadamente dimdrficos. Dado que tiene hipantios con pelos no glandulares, el taxon se encuentra re- lacionatlo con /.’ rnexicana, T. rufipilis y T. thulia, especies de las que se distingue principalmente por tener las ramas agudamente tetragonas. fits lobulos del caliz triangulares mas cortos que el hipantio y los petalos con cilios no glandulares, por lo que se propone en este trabajo como una especie nueva para la ciencia. Novon 14: 163-167. 2004. 164 Novon Tihoiichinn araneicalyx de Santiago, sp. nov. TIPO: Mexico. Guerrero: Mun. Atoyac . Hipantio con tricomas adpresos no glandu¬ lares . T. roseotincta 1 1). Flores basieamente pentdmeras (ocasionalmente hexitmeras o heptameras en T. galeottiana y te- trdmeras, hexameras o heptameras en T. aranei¬ calyx). 4a. Hipantio con pelos glandulares. 5a. Ramas con pelos alargados adpresos o abiertos, hojas basalmente nervadas . . . T. galeottiana 5b. Ramas con pelos cortos adpresos, hojas pi i nervadas . T. durangensis 41). Hipantio con pelos no glandulares. 6a. Lobulos del cdliz mds cortos que el hi¬ pantio, triangulares o lineares, petalos con cilios eglandulares ... 7! araneicalyx 6b. Ldbulos del ealiz iguales o mas largos que el hipantio, generalmente oblanceo¬ lados, petalos con cilios glandulares. 7a. Hipantio con pelos adpresos, tallo con algunos pelos glandulares . . . . T. thulia Volume 14, Number 2 2004 de Santiago Tibouchina araneicalyx de Mexico 167 71). Hipantio con pelos ascendentes o patentes. 8a. Ramas con pelos largos (0.8—3 mm) cafe-rojizos adpresos a patentes, hipantio con pelos rectos subadpresos . . 7. rufipilis 8b. Ramas con pelos blanquecinos adpresos de menos de 2 mm, hipantio con pelos patentes mas o menos flexuosos .... . T. mexicana Pardtipos. MEXICO. Guerrero: Mun. Atoyae de Al¬ varez. 2 km al W de El Molote, 5 die. 2000, de Santiago 868 (MEXU), 5 die. 2000. de Santiago 86 9 (CAS. ENCB. FCMK, MO, UAGC, UAMIZ, XAL); Canada a 0.5 km V de El Molote, 5 die. 2000, de Santiago 888 (ENCB. ECME. MEXU), 18 abr. 2001, de Santiago 902 (MEXU), 21 abr. 2001, de Santiago 911 (FCME, MEXU). 10 die. 2001, de Santiago 939 (CAS, ENCB, FCME, MEXU, MO, UAGC), 10 die. 2001, de Santiago 9 40 (FCME, MEXU), 10 die. 2001, de Santiago 941 (FCME, MEXU); Mun. Petatlan. El Venado, 1 nov. 1991, de Santiago 81 (ENCB, ECME, MEXU. MO); El Venado, 17 nov. 1991. de San¬ tiago 486 (ENCB, ECME, MEXU, MO), 17 nov. 1994, de Santiago 489 (ENCB, FCME, MEXU, MO). Agradecimientos. Agradezco el apoyo prestado, tanto para revisar la coleccion de Melastomataceae del Laboratorio de Plantas Vasculares de la Eacul- tad de Ciencias de la UNAM como para la reali- zacion de coleetas botanicas en el estado de Gue¬ rrero, a la coordinadora del Area de Taxonomfa de dicho laboratorio, la M. en C. Nelly Diego; al Cu- rador del Herbario Naeional de Mexico (MEXU) M. en C. Mario Sousa por el apoyo recibido para la revision de la coleccion de Melastomataceae del herbario; al M. en C. Lucio Lozada por su continuo apoyo en el trabajo de campo y en la redaecion del escrito; al Dr. Jose Luis Villasenor, investigador del Instituto de Biologfa de la UNAM por la revision crftica del manuscrito, y a la dibujante Ana E. Vi- niegra por la elaboracion de la ilustracion que acompana este trabajo. Eileratura Citada Almeda, F. 1993. Melastomataceae. Pp. 1—38 in J. Rze- dowski y G. Calderon (editores). Flora del Bajfo y de regiones adyacentes, fascfculo 10. Instituto de Ecologfa, A. C., Patzeuaro, Miehoacan. Brandegee, T. S. 1913. Plantae Mexicanae Purpusianae, V. Univ. Calif. Publ. Bot. 4: 375—388. - . 1914. Plantae Mexicanae Purpusianae, VI. Univ. Calif. Publ. Bot. 6: 51-77. Cogniaux, C. A. 1891. Melastomataceae (Monographiae Phanerogamarum). Pp. 1-1256 in A. y C. de Candolle (editores), Monogr. Phan. 7. Lozada, I,.. E. Leon, ,1. Rojas y R. de Santiago. 2003. Bosque mesofilo de montana en El Molote. Pp. 1—35 in N. Diego y R. M. Fonseca (editores), Estudios fiorfsticos en Guerrero. Num. 13. Facultad de Ciencias, UNAM. Renner, S. S. 1993. Phylogeny and classification of the Melastomataceae and Memecylaceae. Nordic J. Bot. 13: 519-540. Standley, P. C. 1924. Melastomataceae. In: Trees and shrubs of Mexico. Contr. U.S. Natl. Herb. 23: 1046- 1074. Todzia, C. A. 1999. Ten new species of Tibouchina (Me¬ lastomataceae) from Mexico. Brittonia 51: 255—279. Williams, E. (). 1963. Tropical American plants. V. Fiel- diana, Bot. 29: 543—586. Vi i Ison, P. G. 1958. Contributions to the flora of tropical America LXIII: Plantae Hintonianae XII. Kew Bull. 13: 160-161. Wurdack, J. J. 1967. Certamen Melastomataceis XI. Phytologia 14: 257—274. A New Species of Dyckia (Bromeliaceae) from Rio de Janeiro State, Brazil Rafaela Campostrini Forzza and Bruno Rezende Silva Jardim Botanico do Rio de Janeiro (JBRJ), Rua Pacheco Eeao 915, CEP 22460-030, Rio de Janeiro, Brasil, rafaela@jbrj.gov.br; brsilva@jbrj.gov.br ABSTRACT. A new seaside endemic species, Dy¬ ckia martinellii B. R. Silva & Forzza, from Rio de Janeiro State, Brazil, is described and illustrated. The genus was represented in this state only hy D. pseudococcinea L. B. Smith, from which D. marti¬ nellii can he distinguished by a rosette with a larger diameter; longer and wider leaves hearing straight, patent-antrorse larger spines; basal scape bracts densely arranged, with the others longer than or equaling the internodes; floral bracts castaneous, entire, white-lepidote; and sepals orange with a castaneous apex. Rksumo. Uma nova especie endemiea do litoral, Dyckia martinellii B. R. Silva & Forzza, do estado do Rio de Janeiro, Brasil e descrita e ilustrada. 0 genero estava representado neste estado apenas por I), pseudococcinea L. B. Smith da qual D. marti¬ nellii pode ser diferenciada pela roseta de maior diametro; pelas folhas maiores e mais largas, es- pinhos patente-antrosos maiores; pelas br&cteas basais do escapo congestas as demais maiores ou igualando os entrenos; I trade as florais castanhas, inteiras, alvo-lepidotas e pelas s£palas laranja com dpice castanho. Key words: Brazil, Bromeliaceae, Dyckia, Pit- cairnioideae, Rio de Janeiro state. Dyckia Schultes & Schultes f., with ca. 200 spe¬ cies, is among the largest genera in Bromeliaceae subfamily Pitcairnioideae and the one with greatest diversity in Brazil, where approximately 80% of the species occur (Forzza, 2001). The members of this genus have an exclusively South American distri¬ bution, with diversity centers in southern Brazil within the cerrado and campos rupestres formations (Forzza, 2001). According to a phylogenetic anal¬ ysis based on morphological characters, the genus Dyckia is monophyletic (excluding D. biflora Mez; Forzza, 2004). This is supported by four synapo- morphies: an axial inflorescence; the scape bracts different from leaves; presence of nectaries on the sepals; and a petal -staminal ring (Forzza, 2001). Dyckia martinellii B. R. Silva & Forzza, sp. nov. TYPE: Brazil. Rio de Janeiro: Mun. Paraty, Paratymirim, costoes rochosos entre Paratym- irim e Saco do Mamangud, 5 m elev., 27 Sep. 1990 (fl), G. Martinelli 14413 (holotype, RB). Figure 1. Species nova Dyckiae pseudococcineae affinis, sed ro- sula foliorum circa 60 cm diam., foliis longioribus, latior- ibusque, spinis rectis vel patente-antrorsis et majoribus, bracteis scapalibus infernis foliis similibus, dense dispos- itis, supernis longioribus vel rare internodia aequantibus, bracteis floriferis castaneis, integris et albo-lepidotis, se- palis aurantiaceis apice brunneis differt; D. encholirioidi similis, sed propagulis basalibus brevioribus, rosula fo¬ liorum breviore, foliis brevioribus angustioribusque, flori- bus per anthesim patentibus, bracteis floriferis integris et albo-lepidotis, sepalis albo-lepidotis, aurantiaceis apice brunneis, petalis obtrullatis, aurantiaceis differt. Plant rupicolous, (lowering 70-100 cm high, propagating by stout basal shoots; rosettes ca. 60 cm diam. Leaves suberect-arcuate; sheaths elliptic, 2.5—3 X 5-6 cm, castaneous abaxially, whitish adaxially, margin entire except for a few evanescent spines in the transition to the blade; leaf blades 20—35 X 2. 2-2. 6 cm, lanceolate, apex attenuate, whitish abaxially, light green adaxially, margins spinose, spines straight, patent-antrorse, 3—4 mm long, 1-1.5 cm apart, light green with a castaneous apex. Scape erect, 55—97 cm long, 0.7—10 mm diam., green, sparsely white lepidote; scape bracts the basal ones subfoliaceous, densely arranged, spinulose, 11—12.5 X 0.9— 1.2 mm, erect, charta- ceous, sparsely lepidote; the upper ones exceeding to rarely equaling the internodes, lanceolate, char- taceous, stramineous, glabrous to sparsely lepidote, clasping the scape, apex attenuate, margins spi¬ nulose to inconspicuously serrulate; inflorescence simple, racemose, 30-43 cm long, laxly flowered, 38- to 65-flowered, rachis orange, sparsely lepidote, straight; floral bracts elliptic to elliptic-attenuate, acuminate, exceeding to equaling the middle of the sepals, 4— 12 X 3—6 mm, castaneous, entire, white- lepidote. Mowers patent at anthesis, becoming sub- erect afterward, pedicellate, pedicels cylindrical, 1-2 mm long, orange, sparsely lepidote, reaching 4 Novon 14: 168-170. 2004. Volume 14, Number 2 2004 Forzza & Silva Dyckia martinelli from Brazil 169 Figure 1. Dyckia martinellii B. R. Silva & Forzza. — A. Habit. — B. Leaf. — C. Spine. — 1). Scape base. — E. Inflorescence. — F. Floral bract. — G. Flower. — H. Sepal. — I. Petal and stamens. — J. Gynoecium. — K. Ovule. — L. Capsule. Drawings based on R. C. Forzza el al. 2386 (A— C), G. Martinelli 14413 (D-K), R. R. Silva 961 (L). mm long in fructification; sepals ovate, apex round¬ ed, 7-9 X 4—6 mm, white-lepidote, orange with a castaneous apex; petals orange, obtrullate, gla¬ brous. apex emarginate, erect at antbesis, 9—12 X 9 mm; stamens included; filament connate for 3.5 mm, adnate to petal for 2.5 mm; anthers slightly sagittate at base, attenuate and reflexed at apex, ca. 3 mm long; ovary 7 mm long, ovule alate, ca. 0.7 170 Novon Table 1 . Comparison of Dyckia encholirioides. 1). martinellii, and 1). pseudoococcinea. 1). encholirioides D. martinellii 1). pseudococcinea Vegetative propagation rhizomatous basal shoots basal shoots Rosette diameter (cm) ca. I(M) ca. 60 ca. 40 Foliar blades (cm) 30-100 X 2.5^1 20-35 X 2.2-2 .6 20-30 X 1-1.8 Foliar spines (mm) antrorse, curved, 5-7 patent-antrorse, straight, 3-4 subfoliaceous, densely ar- retrorse, curved, 1.5—2 Basal scape bracts subfoliaceous, densely ar- not foliaceous, sparse, in- ranged, spinulose ranged, spinulose conspicuously serru¬ late Scape bracts longer than to rarely longer than to rarely much shorter than inter- equaling the internodes equaling the internodes nodes Anthesal flower position reflexed patent patent Floral bracts ferrugineous-lanate, serru¬ late white-lepidote, entire reddish, serrulate Sepals ferrugineous-lanate orange with eastaneous apex, white-lepidote reddish, glabrous Petals yellow, unguiculate orange, obtrullate orange, obtrullate mm long. Fruits ca. 15 mm long; seeds alate, ea. 3 mm long. Dyckia martinellii presents morphological affin¬ ities both toward I). pseudococcinea and D. encho- lirioides (Gaudichaud-Beaupr6) Mez (Table 1), these being t lie only three species known from coastal southeastern Brazil. Although the southern states of Rio Grande do Sul. Santa Catarina, and Parana possess many Dy¬ ckia species, including seaside endemics, only one species was known from the southeastern State of Rio de Janeiro, D. pseudococcinea. Sao Paulo, an¬ other southeastern state of Brazil, is also relatively poor in seaside species: only D. encholirioides is known at present, with llha do Cardoso being its northern limit of distribution. The new species was found growing on granite outcrops, from 3 to 10 m above the high tide line, the rocks often being washed by rainwater. By com¬ parison, Dyckia pseudococcinea is a restricted en¬ demic from an area of periodically inundated Eri¬ caceae restinga in the municipality of Marica, Rio de Janeiro State. Hie specimen cited by Smith and Downs (1974) as originating in Atibaia, Sao Paulo, actually belongs to another species, D. tuberosa (Vellozo) Beer. The less selective D. encholirioides occurs on rocks or sand along the coast from Rio Grande do Sul to Sao Paulo, a completely subtrop¬ ical distribution (Smith & Downs, 1974). This new taxon has only been collected from a population on a granite outcrop with an inclination angle of ca. 45°. The outcrop occupies an area of approximately one hectare and is densely vegetated with Vellozia Candida Mikan, Alcantarea edmundoi (Leme) .1. R. Grant, Vriesea neoglutinosa Mez, Cat- tleya intermedia Graham ex Hooker, Cyrtopodium polyphyllum (Vellozo) L. C. Menezes, Cereus, Ti- bouchina, and Pitcairnia. Well represented here, Dyckia martinellii grows mainly in the shade of the shrubs, the deepest shade being the place where the largest specimens were found. This fact may explain its late discovery since it renders the spec¬ imens almost invisible from passing boats, the only means of transportation to the site. Many other sea¬ side outcrops with similar characteristics exist in the region of Paraty and the nearby Ubatuba in Sao Paulo and should be surveyed to establish the spe¬ cies' biological range. Paratypes. BRAZIL. Rio de Janeiro: Mun. Paraty, Paratymirim, costoes rochosos entre Paratymirim e Ma- mangud, 5 m elev.. 30 May 1990 (fr), G. Martinelli 1208V (RB): 23°14'06"S, 44°37'62"W, 17 Nov. 2002, fl cult. Dec. 2002, H. R. Silva 961 (RB), 5 Feb. 2003 (fr), R. C. Forzza 2386. R. R. Silva & E. Borges (MO. RB). Acknowledgments. The authors are grateful to Elton Leme and Claudio Nieoletti de Fraga for cor¬ rections and critical comments, and to Paulo Or- mindo for preparing the illustration. l.iterature Cited Forzza, If. C. 2001. Filogenia da tribo Puyeae Wittm. e revisao taxondmica do genero Encholirium Marl, ex Schult. & Schult. f. (Pitcairnioideae— Bromeliaceae). Tese de Doutorado, Instituto de Biociencias, Universi- dade de Sao Paulo, Sao Paulo. - . 2004 (in press). Revisao taxondmica de Encho¬ lirium Mart, ex Schult. & Schult. f. (Pitcairnioideae— Bromeliaceae). Bol. Bot. Univ. Sao Paulo. Smith, L. B. & R. J. Downs. 1974. Bromeliaceae (Pitcair¬ nioideae). FI. Neotrop. Monogr. 14(1): 1—662. Hafner Press, New York. A New Species of Polyalthia (Annonaceae) from China Hou Xueliang School of Life Sciences, Xiamen University, Xiamen 361005, China, and South China Agriculture University, Guangzhou 510642, China *Zi Shijin South China Institute of Botany, the Chinese Academy of Sciences, Guangzhou 510650, China, lisj@scib.ac.cn Abstract. The new species Polyalthia zhui X. L. flou & S. J. Li, found from Hainan and southern Guangdong in southern China, is described and il¬ lustrated. It has previously been confused with Po¬ lyalthia nemoralis A. DC. or P. littoralis (Blume) Boerlage in China, but is distinguished on the basis of its green fleshy ovate petals, outer petals spread¬ ing ea. 4 X 2.5 mm, inner petals not fully spread¬ ing, somewhat bent to the inside, ea. 5—6 X 4—4.2 mm, 1 mm thick or more, its carpels pilose only in the lower half, and its non-lobed stigma. Key words: Annonaceae, China, Polyalthia. Polyalthia Blume, with 150 to 160 species, is the largest annonaceous genus in Asia (Van Heusden, 1992). Its species are mainly distributed in tropical Asia. New Guinea, Australia, the Solomon Islands, Fiji, Madagascar, and East Africa (Van Heusden. 1992; Verdcourt, 1971) and are especially numerous in Southeast Asia (Sinclair, 1955). There are 32 spe¬ cies in Singapore (Sinclair, 1955). 27 in Vietnam (Ban. 2000), 7 in Sri Lanka (Huber. 1985), and 8 in Burma (Kurz. 1974). The northern distribution edge of Polyalthia is in southern China; there are 17 spe¬ cies in Guangdong, Guangxi, Yunnan, and Hainan Provinces (Tsiang & Li, 1979). The systematics of Annonaceae from China is the first author’s Ph.D. thesis, and the present paper is a part of this study. In the last three years, nearly all Annonaceae herbarium collections in China (about 10,000 sheets) were checked, and several field investigations were carried out concurrently. At the South China Institute of Botany (IBSC), we found that some specimens cited as Polyalthia nemoralis A. DC. by Merrill and Chun (1940) were heterogeneous with the others, i.e.. How 70423 (IBSC). Lau 2843 (IBSC), Liang 66027 (IBSC), Liang 62116 (IBSC), Liang 62794 (IBSC). Liang 63904 (IBSC), Liang 66500 (IBSC). Tso & Chun 44581 (IBSC), Tso & Chun 44717 (IBSC). We tem¬ porarily regarded them as doubtful specimens for lacking flowers, and we did not confirm they are our new species, P zhui, until we had observed the flowers of the Hainan Western Expedition 264 (IBSC). Its outer petals are ovate, 4 X 2.5 mm, its inner petals are ovate, 5—6 X 4-4.2 mm, I mm thick or more, pubescent outside, while P. nemor¬ alis petals are oblong, 8X4 mm, only 0.4 mm Table 1 . Morphologic al comparison of Polyalthia nemoralis and Polyalthia zhui. Polyalthia nemoralis Polylalthia zhui Petals White, di ensely pubescent outer side, eharta- Green, pubescent outer side, fleshy, 1 mm thick ceous. less than 0.4 mm thick, oblong; inner or more, ovate-rounded; inner petals 5—6 mm petals 8 mm long long Stamens Connect i ve apex convex, densely puberulent Connective apex slightly concave, nearly gla- brous Ovaries Densely gray pubescent, stigma bifid Pilose only in the lower half, stigma unlobed Fruit walls Coherent with seeds Separate from seeds heaf blade Secondar y veins incurved; intersecondary veins Secondary' veins straight; intersecondary veins 6 veins absent ; veinlets unbranched; tracheids 0.3— to 8 pairs; veinlets 1- or 2-branched; tra- 0.5 mm cheids less than 0.1 mm Leaf Length : width = 2.5—3 : 1 Length : width = 3-3.6 : 1 * Author for correspondence. Novon 14: 171-175. 2004. 172 Novon Figure la-c. Polyalthia nemoralis (from H. R. Liang 61573 , IBSC). — a. Flower. — b. Leaf venation. — c. Leaf venation magnified, d— f. Polyalthia zhui X. L. Hou & S. J. Li (from the holotype, Hainan Western Expedition 264 , IBSC). — d. Flower. — e. Leaf venation. — f. Leaf venation magnified. Scale bars: a, c, d & f = 1 mm; b & e = 1 cm. Volume 14, Number 2 2004 Hou & Li Polyalthia zhui from China 173 Table 2. Morphological comparison of Polyalthia littoralis and Polyalthia zhui. Polyalthia littoralis Polyalthia zhui Ratio of inner petal length to 2-2.6 : 1 1 .4—1.6 : 1 outer petal length Petal texture thin-membranous fleshy, 1 mm thick or more Ratio of monocarpel length to 2-3.1 : 1 4-5 : 1 the stalk of monocarpel length Length of monocarpel stalk 3—6 mm 3 mm Length ol fruiting peduncles ca. 12 mm 3—1 mm Leaf blade lanceolate, usually 2—3 cm wide oblong-lanceolate or ovate-lan¬ ceolate, usually 3. 5-4. 2 cm wide thick, and pilose outside. Further observation shows that these specimens also differ from P. ne- moralis in carpels, fruit, stamens, and leaf venation (Table 1, Fig. 1). After having compared the specimens with the known species of Polyalthia recorded in various floras and systematic treatments (Blume, 1830; Hooker & Thomson, 1855, 1872; Merrill, 1912, 1923; Sinclair, 1955; Backer & Bakhuizen, 1963; Huber, 1985; Ban, 2000), we found a few species with petals less than 10 mm long. Among these. Figure 2. Holotype of Polyalthia zhui ( Hainan Western Expedition 264. I BSC). only P. littoralis (Blume) Boerlage, whose distri¬ bution extends to Indonesia and Vietnam, is similar to P. zhui. We made sure, however, that P. zhui was distinct from P. littoralis. Polyalthia littoralis petals are thin-membranous, whereas P. zhui petals are thick-fleshy up to 1 mm or more. Furthermore, the two species can be distinguished by their leaves, monocarpels, and fruiting peduncles (Table 2). Po¬ lyalthia littoralis was reported as new to China by Li (1993). According to the specimens cited by Li (1993), however, his P. littoralis was Merrill and Chun’s P. nemoralis, that is, it included P. nemor- alis and P. zhui. Polyalthia zhui X. L. Hou & S. J. Li. sp. nov. TYPE: China. Hainan: Darizhou City, Nada, Lianchangjiaoyuan, 15 May 1954, Hainan Western Expedition 264 (holotype, 1BSC; iso¬ types. IBK. KUN, PE). Figures ID— F, 2. 3. Affinis Polyalthiae nemorali, sed floribus ca. 6 mm diam., petalis exterioribus ovato-rotundis 4 mm longis, 2.5 mm latis, petalis interioribus ovato-rotundis, 5—6 mm lon¬ gis, 4—4.2 mm latis, externis rare pubescentibus, carpellis meris pubescentibus in dimidio inferiore, stigmate non lobato, connectivo parum concavo differt. Shrubs usually 1.5—3 m, rarely up to 5 m tall; bark dark brown; young twigs dark purple, striate, pubescent with appressed gray-yellow hairs, but quickly glabrescent. Leaves coriaceous, oblong- lanceolate or ovate-lanceolate, 9-16(— 19) X 2.5- 4.5(— 5) cm, apex acuminate to obtuse-acuminate, base cuneate, glabrous, dark green above, lucid and pale green beneath, midrib slightly grooved above and obviously convex beneath, secondary veins 8 to 10 pairs, at 60° to 80° with the midrib, interarching 3—5 mm from the margin, reticulations somewhat prominent on both sides, intersecondary veins in 6 to 8 pairs; petioles 3—5 mm long, yel¬ lowish, rugose. Inflorescences extra-axillary with 1 (or 2) flowers; peduncles ca. 1 mm long; pedicels 174 No von F igure .'1. Polyallhia zhui. — a. Abaxial view of stamen. — b. Adaxial view of stamen. — c. Carpel. — d. Longitudinal section of carpel. Scale bar = 1 mm. (From the holotype, Hainan Western Expedition 264 , IBSC.) 1—2 mm long, pubescent with gray-yellow hairs, bract 1. broadly ovate, 1-2 X 2 mm. acuminate. Flower green, ea. 6 mm diam. when mature; sepals connate at base, lobes 3, ovate, 2 X 1 .8 mm, pu¬ bescent outside, glabrous inside, persistent; petals 6 in 2 whorls, fleshy, acute, pubescent outside, gla¬ brous inside, outer petals ovate, flat, spreading, ca. 4 X 2.5 mm, inner petals ovate, flat, not fully spreading, somewhat bent to inside, 5—6 X 4-4.2 mm, ca. 1 mm thick or more; stamens ca. 50 to 100, cuneate, 1—1.3 mm long, ca. 1 mm wide, con¬ nective apex slightly concave, dark purple, gla- brescent; carpels ca. 30. with one basal ovule each. 1.5—2 mm long, pilose with brown-yellow hairs in the lower half; stigma capitate, sessile, puberulent, not lobed. Monocarpels 4 to 7 (to 11), ellipsoid, 12— 15 mm long, 8—10 mm diam.. purple-red when dry, glabrous, densely tiny-dotted, stalks 3 mm long, pubescent; seed I per loeule, ellipsoid, red, 10—12 mm long, 6—8 mm diam., separated from fruit walls, endosperm ruminative. Distribution and habitat. In wet forest at 250— 600 m, Hainan and southern Guangdong. In forest edges, sometimes along small streams. This was collected in flower April through July and in fruit July through February of the following year. Polyalthia zhui is distinguished from the other species of Polyalthia by its oblong-lanceolate or ovate-lanceolate leaf shape, lucid anti pale green beneath and 6 to 8 pairs of intersecondary veins, its relatively short pedicels and small green flowers with fleshy petals, its nearly glabrous and slightly concave connective apices, its capitate stigma, and its seeds not adherent to fruit walls. Eponymy. Polyalthia zhui was named in honor of our master supervisor. Prof. Zhu Changshan, working in the Henan Agriculture University, who shared his rich knowledge of and correct methods for studying plant taxonomy. Paratypes. CHINA. Guangdong: Xuwen Co., Mailao- wo village, 31 Oct. 1951 (fr), Z. S. Zhu I0,'i6 (IBSC); Xuwen Co., Kengzai village, 26 Apr. 1936 (fr), Hainan Southern Expedition 291 (IBK, IBSC, KUN, PE); Xuwen Co., Xuwenshan hill, near Yugonglou, 27 Sep. 1951 (fr), S. H. Chun 7515 (IBSC): Xuwen Co., Guooheng village, 7 Nov. 1957 (fr), Hainan Southern Expedition 152 (IBSC); Xuwen Co., Baishui village, 15 Apr. 1985 (II). Li & Xing 1985 (IBSC). Hainan: Baisha Co., a valley near Hong- rnaodong, 27 Apr. 1936, S. A. Diu 26524 (IBSC. KUN); Baoting Co., Diaoluosban Mountain. Baishuiling. 24 Nov. 1954, Diaoluoshan Expedition 2815 (IBK, IBSC, PE); Cbangjiang Co., Jiaqieshan bill. 16 Oct. 1933 (fr), S. A. Dm 2842 (A. IBSC); Cbangjiang Co., 20 Dec. 1933 (fr), II. II. Liang 66027 (A, IBK. IBSC); Cbangjiang Co., Dec. 1933 (fr). II. R. Liang 60102 ( \. IBK. PE); Cbangjiang Co.. 20 Dec. 1933 (fr), II. R. Liang 66500 (IBK. IBSC, PE); Dongfang Co., Guangba District, Qicha, 17 Nov. 1956, S. II. Chun II 124 (IBSC); Dongfang Co.. Maanling, 6 Nov. 1978. G. A. Eu 1291 (IBSC); Dongfang Co., Jian- fengling, near Sanmojue village, 23 Mar. 1934, 6. A. Dm 2522 (SYS). .S'. A. Dm 4924 (IBSC); Dongfang Co., //. R. Liang 62094 (A, IBK. IBSC); Ledong Co., Taohui village. Taohuiling, S’. A. Dm 27454 (A. IBK. IBSC. KUN, PE); Eingshui Co., Nanwanling, 21 Jan. 1985, C. A. Eu 5551 (IBSC); Sanya City, Nanshanling, 30 Dec. 1932, Tso S: Chun 44717 (A, IBK, IBSC, PE); Sanya City. Nansbanl- ing. 26 Dec. 1932. Tso A- Chun 44581 (A. IBK. IBSC, Volume 14, Number 2 2004 Hou & Li Polyalthia zhui from China 175 PE); Sanya Cily, Xiaobaokang, 16 July 1933 (fr), H. R. Liang 62116 (A. IBK. IBSC. PE); Sanya City, Daposhan hill, 23 Aug. 1933, H. R. Liang 62794 (A, IBK. IBSC); Sanya City, Yanglin, 24 Mar. 1933, F. C. How 70423 (IBSC); Sanya Cily, Ganzhaling, upriver Fotian Reservoir, 30 Oct. 1987, Z. X. Li 3058 (IBSC); Sanya City. Dabao- kang, 16 July 1933, C. Wang 33087 (IBSC, PE); Sanya City, Nanlin, Guikaishan hill, 16 Oct. 1933 (fr), C. Wang 34633 (IBSC). Acknowledgments. The authors thank Li Ping- tao. South China Agriculture University, for criti¬ cally reading the manuscript and giving valuable suggestions. We are likewise grateful to Zhang Dianxiang (IBSC) for helpful comments on the man¬ uscript. We are also grateful to Zeng Feiyan (IBSC), Wang Jinghua (KLIN), Lin Qi and Ban Qin (PE), Li Mingguang (SYS), and Wei Fanan and Yu Xixi (IBK) for offering assistance in consulting speci¬ mens. fhe study was supported by a grant from the Bureau of Environmental Protection of Guangdong Province (970165). Literature Cited Backer, C. A. & K. C. Bakhuizen van den Brink. 1963. Pp. 100—116 in Flora of Java, Vol. 1. Walters Noordhoff N.V., Groningen. Ban. IN. T. 2000. Flora of Vietnam, Vol. 1. Science & Technics Publishing House, Ha Noi. Illume, C. L. 1830. P. 99 in Flora Javae, Vol. 1. Fugduxi Bataviorum. Hooker, J. D. & T. Thomson. 1855. Annonaceae. Pp. 86- 153 in Flora Indica, Vol. 1. W. Pamplin, London. - & - . 1872. Annonaceae. Pp. 45—94 in J. 1). Hooker (editor), The Flora of British India I. Reeve, London. Huber. H. 1985. Annonaceae. Pp. 1—75 in M. I). Dassan- ayake (editor), A Revised Handbook to the Flora of Ceylon, Vol. 5. A. A. Balkema, Rotterdam. Kurz, S. 1974. Annonaceae. Pp. 25—50 in Forest Flora of British Burma. Vol. 1. Bishen Singh Mahendra Pal Singh. Dehra Dun. Li, P T. 1993. Novelties in Annonaceae from Asia. Gui- haia 13(4): 311-315. Merrill E. D. 1912. Annonaceae. Pp. 205—209 in A Flora of Manila. Bureau of Printing, Manila. - . 1923. Annonaceae. Pp. 154—178 in An Enumer¬ ation of Philippine Flowering Plants II. Bureau of Print¬ ing, Manila. - & W. Y. Chun. 1940. FI. Hainan III. Sunyatsenia 5(1-3): 59. Sinclair, J. 1955. A revision of the Malayan Annonaceae. Card. Bull. Straits Settl. 14: 149—516. Tsiang, Y & P. T. Li. 1979. Annonaceae. Pp. 10—175 in Flora Reipublicae Popularis Sinicae 30(2). Science Press, Beijing. Van Heusden, E. C. H. 1992. Flowers of Annonaceae: Morphology, classification, and evolution. Blumea Suppl. 7: 1-218. Verdcourt, B. 1971. Annonaceae. Pp. 1—132 in E. Milne- Redhead & R. M. Polhill (editors), Flora of Tropical Fast Africa. Crown Agents for Oversea Governments and Administrations, London. A New Species of Didymoplexiella (Orchidaceae) from China Jin Xiaohua, Chen Singchi, and Qin Haining Herbarium (PE), Institute of Botany of Chinese Academy of Sciences, Beijing 100093, China, xiaohuajin@mail.kib.ac.cn; hainingqin@ns.ibcas.ac.cn Guanghua /hit Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166, U.S.A. guanghua.zhu@mobot.org Gloria Siu Laiping Kadoorie Farm and Botanic Garden, Hong Kong. siulpcon@kfbg.org.hk Abstract. Didymoplexiella hainanensis X. H. Jin & S. C. Chen, a new speeies of Orchidaceae from Hainan, China, is described and illustrated. The new species is related to I). ornata (Ridley) Garay, from which it differs by the dorsal sepal briefly connate with the petals, the lateral sepals connate Va to Vi at base, an articulate lip, and larger lateral lobes. Key worth: China, Didymoplexiella, Orchidaceae. Didymoplexiella Garay is a leafless orchid genus of ca. seven species (Seidenfaden & Wood, 1992) mainly distributed in Indonesia, with a few species, e.g., D. siamensis (Rolfe ex Downie) Seidenfaden, extending to northern Thailand, China, and Japan. It is similar in habit and floral appearance to Di- dymoplexis Griffith, with which it was considered by some authors as congeneric (Holttum, 1953). Garay (1955) found enough distinction in floral col¬ umn to justify a separation between them. The col¬ umn of Didymoplexiella has a pair of long stelidia on either side of its stigma and lacks a distinct column foot, while that of Didymoplexis lacks stel¬ idia and the column foot is distinct. The new spe¬ cies was collected in late March from Hainan Is¬ land and is no doubt a new member of Didymoplexiella. It is somewhat related to D. or¬ nata (Ridley) Garay from Malaysia and Thailand, from which the new species is distinct by its almost separate dorsal sepal and petals, articulate lip, and much larger lateral lobes. Didymoplexiella hainanensis X. 11. Jin & S. C. Chen, sp. nov. TYPE: China. Hainan: Ledong, Jianfenlin Mt., 17 Mar. 2002 (fl). //. K. Ka¬ doorie Program Team 3087 (holotype, PE; iso- type [spirit], MO). Figure 1. Habitu Didymoplexiellae ornatae subsimilis, sed sepalo dorsali petalis breviter vel vix connato, sepalis lateralibus inter se tantum Va ad Vi parte inferiore connatis, labello multo longiore quam columna prope medium mobili-arti- culato bene differt. Mycoheterotrophic plants; tuber creeping, cylin¬ drical, 5—6 cm long, 3—10 mm diam., wrinkled, nodal; stem erect, slender, terete, 5—15 cm long, brown, nodal, sheathing on the lower half, with in¬ ternodes 2-9 mm long; sheaths 1 .5—2.5 mm long. Inflorescence 7-9 mm long, compactly 2- to 4-flow- ered; bracts triangular-lanceolate, 1.5—2 mm long; petlicel and ovary 5—15 mm long, brown. Flowers ca. 1.5 cm across, white except the lip mid-lobe yellow, dorsal sepal elliptic, 7-10 X 13 mm, briefly or scarcely united at base with petals, 3-nerved, obtuse; petals similar to dorsal sepal. 6—7 X 10 mm, joined with dorsal sepals for Va to Vi of their length; lateral sepals elliptic, 10—11 X 3.5-4 nun, 3-nerved, obtuse, connate at base to each other for Va to Vi of the length; lip 11-12 mm long, consisting of an apical blade 5.5—6 mm long and a basal claw 4.5—5 mm long, with an articulate joint between them; blade tri-lobed, with a basal claw, its upper surface a U-shaped callus covered with small warts; lateral lobes sub-oblong, ca. 5X4 mm with a slightly undulate margin, much larger than the mid¬ lobe; mid-lobe fleshy, ca. 3 X 2 mm at the apex, with stelidia on either side of the stigma; stelidia curved downward, 2.5—3 mm long; pollinia 4, mealy. Capsule not seen. Distribution. China, Hainan Province, Ledong County, Jianfengling Mountains, alt. 700 to 800 m. in humid evergreen forest. Paratype. CHINA. Hainan: Ledong, Jianfengling Mtn., 15 Mar. 2002 (fl), Hong Kong Kadoorie Program Team 3081 (MO. |spirit| KUN). Acknowledgments. We are indebted to the Ka- Novon 14: 176-177. 2004. Volume 14, Number 2 2004 Jin et al. Didymoplexiella (Orchidaceae) 177 Figure 1. Didymoplexiella hainanensis X. H. Jin & S. C. Chen. — A. Habit and inflorescence. — B. Front view of sepals and petals. — C. Front view of lip. — D. Stelids. — E. The articulate claw. ( Based on Hong Kong Kadoorie Program Team 3087, PE.) doorie Farm and Botanic Garden, Hong Kong, for its financial support and to the CITES officials of China for their kind help. We also thank Yinbao Sun for his excellent illustration. Literature Cited Garay, L. A. 1955. Notatio Orchidologica 3. Arq. Jard. Got. Kio. 13: 31-54. Holttum, R. E. 1953. Didymoplexis. P. 107 in R. E. Holt- tum (editor). Flora of Malaya. Vol. 1. Orchids of Malaya. Government Printing Office, Singapore. Seidenfaden, G. & J. J. Wood. 1992. Didymoplexiella. Pp. 127—139 in G. Seidenfaden & J. J. Wood (editors). The Orchids of Peninsular Malaysia and Singapore. Olsen & Olsen, Fredensborg. A New Species of Holcoglossum (Orchidaceae) from China Jin Xiaohua, Chen Singchi, and (Jin Haining Herbarium (PH). Institute of Botany of Chinese Academy of Sciences, Beijing 100093, China, xiaohuajin@mail.kib.ac.cn; hainingqin@ns.ihcas.ac.cn Guanghua Zhu Missouri Botanical Garden, P.0. Box 299. St. Louis, Missouri 63166, U.S.A. guanghua.zhu@mobot.org Gloria Sin Lai ping Kadoorie Farm and Botanic Garden, Hong Kong. siulpcon@klBg.org.hk ABSTRACT. Holcoglossum weixiense X. H. Jin & S. C. Chen, a new species of Orchidaceae from Yunnan, China, is described and illustrated. It re¬ sembles morphologically H. sinicum Christenson and //. flavescens, but it can be distinguished from the former by having a transverse callus on the lip, a papillose spur, and a semi-orbicular mid-lobe, and from the latter by its pending habit, longer and semiterete leaves (more than 10 cm), and the semi- orbicular mid-lobe. Key words: China, Holcoglossum, Orchidaceae. Holcoglossum Schlechter (Orchidaceae) is dis¬ tributed mainly in southwestern China, with a few species, e.g., H. subulifolium (Reichenbach f.) Christenson, extending to Laos, Thailand. Vietnam, and Malaysia. Because of its pollinia, three-lobed lip, and the short bifid rostellum, this genus is sim¬ ilar in floral structure to Aerules, Papilionanthe, and Vanda, from which several species have previously been transferred, e.g., H. rupestre (Handel-Mazzetti) Garay (1972), H. flavescens (Schlechter) Tsi (1982). and H. subulifolium (Reichenbach f.) Christenson (1987). Although Seidenfaden (1988) stated that it was difficult to define Holcoglossum, the new spe¬ cies somewhat coincides with Garay’s (1972: 181) delimitation of the genus: “. . . characterized by the short, footless column with prominent wings. Cli- nandrium deeply cleft in front: rostellum short, bi¬ fid. Pollinia 2, notched, on linear, tapering stipes. Lip sessile with a slender, arcuate spur. Stem rather short, completely enclosed by distichously arranged leaf-sheaths. Leaves articulate, triquetrous or te¬ rete." The new species was collected from the Langcan River in northwestern Yunnan. It is somewhat re¬ lated to Holcoglossum sinicum Christenson and H. flavescens (Schlechter) Tsi, but is distinguished from the former by its semi-orbicular mid-lobe, papillose spur, notable callus on the labellum, and from the latter by its pending habit, longer and semiterete leaves (more than 10 cm), and the semi-orbicular mid-lobe. Holcoglossum weixiense X. H. Jin & S. C. Chen, sp. nov. TYPE: China. Yunnan: Weixi, La¬ ngcan River, 21 May 2002 (fl), //. K. Kadoorie Program Team 3490 (holotype, PE; isotypes [spirit], KUN. MO, PE). Figure 1. Habitu Holcoglosso sinico subsimile, sed calcari intus papillato, inflorescentia caule longiore, labello medio callo nolabili instructo bene differ!. Epiphytic plant, pendulous; root terete, thick, white; stem short, 1—2 cm long, enclosed by the leaf-sheathing bases. Leaves 20—30 X 1.5— 2.5 cm, coriaceous, semiterete, channeled, articulate at base. Inflorescence lateral, erect, 3-4 cm long, 2- to 4-flowered, exceeding the stem, bracts ovate, 0.7 cm long. Flowers spreading, opening fully, white, tinged with pink; pedicel with ovary 2 cm long; dorsal sepal erect, ovate, 1 X 0.7 cm, 3-veined; lateral sepals oblique at base, seculate, 1.5 X 0.9 cm wide, 4-veined; petals spathulate, 1.1 X 0.0 cm, 3-veined; lip adnate to the column foot, fixed; three-lobed; lateral lobes triangular, erect, with purple streaks on the inner surface; mid-lobe semi- orbicular, entire, 1 X 1.2 cm. with a transverse callus at base toward the entrance of the spur; spur funnel-shaped, papillose, 0.7 cm long; column 0.4 cm long, column foot 0.1 cm long; pollinia 2, notched; stipes obcuneate, 3 mm long. Capsule un¬ known. Distribution. China: Yunnan, Weixi, Bilou Snow Mountain, at 2300 to 2900 m. in evergreen forest. Novon 14: 178-179. 2004. Volume 14, Number 2 2004 Jin et al. Holcoglossum (Orchidaceae) 179 Figure 1. Holcoglossum weixiense X. II. Jin & S. C. Chen. — A. Habit and inflorescence. — B. Lateral view of flower. — C. Front view of flower. — L). Lateral view ol lateral lobes. — E. Inner wall of spur. Drawn from the type specimen (Hong Kong Kadoorie Program Team 3490) by Yingbao Sun. Paratypes. CHINA. Yunnan: Langcan River, 20 July 2001, Hong Kong Kadoorie Program Team 1166 (KUN, PE), 21 May 2002 (fl), Hong Kong Kadoorie Program learn 3436 (KUN), 3497 (PE). Acknowledgments. We are indebted to Wen-tasi Wang for his critical reading of the manuscript, to the Kadoorie Farm and Botanic Garden, Hong Kong, for its financial support, and to the CITES officials of China for their kind help. We also thank Yinbao Sun for his excellent illustration. Literature Cited Christenson, E. A. 1987. An intrageneric classification of Holcoglossum Schltr. (Orchidaceae: Sarcanthinae) with a key to the genera of the Aerides-Vanda alliance. Notes Roy. But. Card. Edinburgh 44: 249—256. Garay, L. A. 1972. On the systematics of the monopodial orchids I. But. Mus. Leaf!. 23(4): 149-212. Seidenfaden, G. 1988. Orchid genera in Thailand XIV. Fifty-nine genera. Opera Bot. 95: 304—308. Tsi, Z. H. 1982. A study of the genus Holcoglossum of Orchidaceae. Acta Phytotax. Sin. 20(4): 439—444. New Taxa of Cimicifuga (Ranunculaceae) from Korea and the United Stales Hyun-Woo I^ee El A Division, Korea Environment Institute, Seoul 122-706, Korea. hwlee@kei.re.kr Chong-Wook Park School of Biological Sciences, Seoul National University, Seoul 151-742. Korea. parkc@plaza.snu.ac.kr Abstract. A new species and a new variety of Cimicifuga (Ranunculaceae) are described and il¬ lustrated. Cimicifuga austrokoreana, restricted to south-central Korea, is distinguished from the other species of the genus by its strongly arcuate inflo¬ rescence axis, much shorter pedicels, and three bracteoles, one at the base and the other two at the middle of the pedicel. Cimicifuga data var. alpes- tris, found in southern Oregon in the United States, is distinguished from variety data and the other members of the genus by its sheathing scales sur¬ rounding the lower nodes of the stem. Key words: Cimicifuga, Korea, Ranunculaceae, United States. The genus Cimicifuga Wernischeck (Ranuncu¬ laceae) comprises as many as 22 species that are widely distributed in temperate regions of the Northern Hemisphere, including Europe, Asia, and North America; the center of diversity is in eastern Asia (Tamura, 1966, 1990, 1995; Ramsey, 1965, 1997; Compton et al., 1998). The plants of Cimi¬ cifuga are erect rhizomatous herbs with long-peti- oled, ternately compound leaves, racemose or pa¬ niculate inflorescences bearing many small flowers, and follicular fruits (Tamura, 1966, 1990, 1995; Park & Lee, 1995, 1996; Ramsey, 1965, 1997). Classification of Cimicifuga and its related gen¬ era has been controversial. Compton et al. (1998) merged Cimicifuga and Souliea Franchet with Ac- taea L. mainly on the basis of the analyses of ITS and /rnL-F sequences. However, Wang et al. (1999, 2001) argued that they are independent genera based on characters from morphology, palynology, and cytology. Cimicifuga is distinguished from Ac- taea in having three (rarely one) bracteoles, five sepals, staminode-like petals, one to eight pistils, follicular fruits, seeds with membranous scales on the surface, and more symmetric karyotypes with¬ out telocentric chromosomes. Cimicifuga is also distinguished from Souliea in having inflorescences with numerous flowers, caducous sepals, oblong to ovoid follicles, seeds with membranous scales on the surface, and mostly tricolpate pollen grains. In diis study we follow the generic concept and delim¬ itation of Tamura (1995) and Wang et al. (1999, 2001), in which Cimicifuga was recognized as a distinct genus. During the course of a comprehensive systematic study on the genus Cimicifuga, a new species and a new variety were discovered from Korea and the United States, respectively. We describe these new taxa herein to make their names available for the Flora of Korea and other works in progress. 1. (.imicifiigu austrokoreana H.-W. Lee & C.- W. Park, sp. nov. TYPE: Korea. Chungbuk: Youngdong-gun, Mt. Minjuji-san, Samdo-bong, Minimigol, 700 m, 19 Sep. 2000, H.-W. Ise 1542 (holotype, SNU; isotypes, MO, SNU, TI). Figure 1. Differt a C. simplici inflorescentiis axibus arcuatis, ped- icellis multo brevibus et unibracteolatis basi. Herbs perennial, hermaphroditic, 40—80 cm tall; rhizomes thick, 5—8 cm long, knotted, bearing fi¬ brous roots; stem simple, erect. 1.5-4 mm thick, glabrous to sparsely pubescent with filiform uni¬ cellular trichomes. Leaves cauline, usually 3, al¬ ternate, 1- to 3-ternately compound, long-petioled; terminal leaflet broadly elliptic, 7.0—10.2 X 5.4— 7. 1 cm, 3-lobed, acuminate to cuspidate at tip, cu- ueate at base; both surfaces moderately to densely pubescent along major veins with filiform unicel¬ lular trichomes ca. 0.2 mm long; petiolules 1.5— 2.5 cm long, grooved, sparsely to moderately pubescent with filiform unicellular trichomes; lateral leaflets similar to terminal one, but slightly smaller and inequilateral; petiole 10—20 cm long, grooved, sparsely to moderately pubescent with filiform uni¬ cellular trichomes. Inflorescence a terminal ra¬ ceme, 10—20 cm long, often bearing 1 or 2 short Novon 14: 180-184. 2004. Volume 14, Number 2 2004 Lee & Park New Taxa of Cimicifuga 181 Figure 1. Cimicifuga austrokoreana H.-W. Lee & C.-W. Park. — A. Inflorescence anil upper leaf. — B. Stem with lower leaf. — C. Petals. — 1). Pistils and a pedicel with bracteoles. — E. Follicles. — F. Seed. Scale bars 5 cm (A, B), 5 mm (E), 2 mm (D), 1 mm (C, F). A-D, drawn from the holotype (Lee 1542, SNU), and E, F from the paratype (Oh 1355, SNU), by H.-W. Lee. 182 Novon lateral branches near base, densely pubescent with saccate unicellular trichomes 0.1— 0.2 mm long; in¬ florescence axis distinctly arcuate; pedicels very short, 1.0— 2.5 mm long; bracteoles 3, 1 at base and 2 at middle of pedicel, triangular, 0.4— 0.7 X 0.3— 0.5 mm. Flowers bisexual, actinomorphic, small; sepals 5, petaloid, pale brown, broadly elliptic, concave, 3.5— 4.8 X 2.8— 3.5 mm; petal I. broadly elliptic, 2. 0-4.6 X 1.4— 3.3 mm, bearing 2 small white appendages at tip, short-stipitate, nectarifer¬ ous at base; stamens 18 to 27; filaments filiform, 5—8 mm long, slightly dilated upward; anthers bi¬ locular, broadly oblong, 0.6 X 0.5 mm. basifixed, latrorsely dehiscent; pistils usually 2. rarely 1 or 3, elliptic to broadly elliptic, 1-3 X 0.7-2 mm, stip- itate, sparsely pubescent with saccate unicellular trichomes ca. 0.1 mm long; style 1. slender, unci¬ nate when mature, 0.8-1. 7 mm long; stigma I. mi¬ nute, appressed to style. Follicles oblong, 5.5—9 X 1—6 mm, chartaceous, glabrous or sparsely pubes¬ cent with saccate unicellular trichomes; stipe 4—6 mm long; seeds 1 to 5 per follicle, elliptic, 2.2— 2.6 X 1 mm, winged; wings ca. 1 mm long, chaffy, crenate. Distribution. Restricted to high mountains of south-central Korea; moist, shady places along streamsides and mountain slopes of mixed decid¬ uous forests, alt. 500 to 1000 in. Phenology. Flowering mid-September to Octo¬ ber (pers. obs.). Cimicifuga austrokoreana is closely related to C. simplex (DC.) Turczaninow, but it clearly differs from the latter species by its strongly arcuate inflo¬ rescence axis, much shorter pedicels ca. 1.0— 2.5 mm long, and three bracteoles, one at base and the other two at middle of pedicel (Fig. 1); in particular, the inflorescence axis is distinctly and consistently bent downward from the early stage of its devel¬ opment. Cimicifuga simplex has an erect, straight inflorescence axis, relatively long pedicels ca. 4-8 mm long, and bracteoles borne at the base of the pedicel. In addition, the karyotypic study and allozyme analysis showed that Cimicifuga austrokoreana is genetically distinct from C. simplex (Fee & Park, 1998; Fee et al., 2000). Their karyotypes differ in the position of a secondary constriction on a pair of submetacentric chromosomes (Fee & Park. 1998), and populations of C. austrokoreana are dis¬ tinguished from those of C. simplex by their allelic compositions at four loci ( Fe-3 , Gdh , Lap, anil Pgi- 2) and significantly low genetic identity values (mean = 0.688) (Fee et al., 2000). Paratypes. KOREA. Chiinhuk: Muju-gun, Mt. l)u- kyu-san, Baekryundam, 16 Sep. 1996, Lee 771 (SIN LJ ). Chuiighuk: Youngdong-gun, Mt. Minjuji-san, Samdo- bong, 29 Sep. 1995, lee 747, 748 (SNU), 8 Oct. 1995, lee 775, 776 (SNU). Chiinnain: Gurye-gun, Mt. Cliiri, Piagol, Sunyugyo, under deciduous forest, 24 Sep. 1990, //vim 5012 (SNU); Jilmaejae, 6 July 1995, lee 566, 567 (SNU), 18 Sep. 2(KM). lee 1541 (SNU). 30 Sep. 2. g, h from Lorence & Stone 8380 ; e— f, from Lorence & Stone 8385. Scale bar 10 cm in b, c; 7 mm in d, e; 6 mm in f; 7.5 cm in g; 2.5 cm in h. Volume 14, Number 2 2004 Lorence & Gemmill Pritchardia flynnii from Kaua'i 187 felt-like, pale brown or gray. Inflorescences 1 or occasionally 2 per leaf axil, in flower erect or ar¬ cuate, 58—88 cm long, equaling or generally ex¬ ceeding petiole and reaching Vi (rarely %) length of blade, in fruit 80—100 cm long, erect to arcuate and slightly shorter to slightly longer than the blade; prophyll externally brown or tan lepidote, tomen- tose basally, disintegrating; peduncular bracts 4, overlapping, eventually disintegrating, externally uniformly brown or tan tomentose, denser basally; peduncle 48—60 cm long, 8—1 1 mm diam., terete or compressed, brown lanate-tomentose; panicle branched to second degree, the floriferous portion 1 1-20 cm long, the basal branches 10—13 cm long with 3 or 4 rachillae, the middle branches with a single bifurcate branch of 2 rachillae or less com¬ monly unbranched, apical portion of rachis with 9 to I I unbranched rachillae, these sinuous in flower, 6—11 cm long, at first sparsely to densely lanate- tomentose, in fruit 10—14 cm long, tomentose or glabrescent; floral bracts subulate-filiform, 1 .5—3 mm, brown. Flowers spirally arranged, 8—10 mm long in bud, glabrous, calyx plus pedicel 3—5 mm long, calyx cupular, 1.5— 2.5 mm wide at base, 3—4 mm wide at rim of cup, green when fresh, when dry with prominulous nerves converging in abruptly acuminate teeth 0.3— 0.8 mm long; petals 5.7-7 X 2.8— 3.5 mm, acute to acuminate at apex, yellow when fresh, veins prominulous when dry; staminal cup 3.5—6 mm long, exserted beyond calyx rim. deep orange-yellow when fresh; filaments 1—2 mm long, anthers 3—5 mm long; pistil 5—6.5 mm long, including style 3-4 mm long; style present in fruit, ca. 2 mm long. Fruit ellipsoid or ovoid-ellipsoid, 25—35 X 18—23 mm when fresh, 22—25 X 15—16 mm when dry, often slightly asymmetrical or acen¬ tric, with persistent calyx and corolla at base and accrescent style at apex, smooth, glabrous, shiny dark green, ripening purplish black; seed ovoid- ellipsoid, 14—18 X 9—12 mm. Distribution. Known from south-central to north-central Kaua'i. Populations are known from the Wahiawa Drainage (Wahiawa Stream and Moun¬ tains and Mt. Kahili) in the south, the north fork of the Wailua River (“Blue Hole”), Makaleha Moun¬ tains, Power Line Trail, and Wainiha Valley in the north. Intervening areas likely harbor additional plants, in effect linking these known populations. Habitat. This new species occurs from approx¬ imately 488 to 890 m elevation, usually on mod¬ erate to steep slopes in low-stature lowland wet for¬ est. shrubland, or herbland usually dominated by Metrosideros polymorpha Gaudichaud, M. waiale- alae (Rock) Rock var. waialealae, and Dicranop- teris linearis (Burmann) Underwood. Other associ¬ ates include Antidesma platyphyllum II. Mann var. hillebrandii Pax & K. Hoffmann, Bobea elatior Gaudichaud, Diospyros sandwicensis (A. DC.) Fos- berg, Freycinetia arborea Gaudichaud, Ilex anom- ala Hooker & Arnott, Machaerina angustifolia (Gaudichaud) T. Koyama, Melicope, Syzygium sand¬ wicensis (A. Gray) Niedenzu, and species of Cibo- tium, Cyanea, Cyrtandra, Myrsine, Psychotria, Scaevola, and Tetraplasandra. Weedy alien species invading this habitat include Psidium cattleianum Sabine var. cattleianum and Melastoma candidum D. Don, both of which pose a significant threat to native plant communities in the vicinity (Lorence & Flynn, unpublished), and a variety of alien grass¬ es including Paspalum conjugatum Bergius, Setaria gracilis Kunth, and Schizachyrium condensatum ( Kunth) Nees. Population size and conservation status. This species occurs as scattered individuals and small groves throughout the Wahiawa Drainage area, where the population size is estimated at 250 to 300 individuals. Additional populations are known from: the ridge between Mt. La'auhiha'ihai and Mt. Kahili (6 individuals), the Blue Hole (exact number unknown). Power Line Trail (8), the Makaleha Mts. (84). and Wainiha Valley (exact number unknown), bringing the total to perhaps 350 to 400 plants. Plants of different size classes including occa¬ sional juveniles are represented in some localities, suggesting at least limited regeneration is taking place. However, invasive weedy alien plant species, notably Psidium cattleianum Sabine and Melastoma candidum 1). Don and aggressive grasses, threaten this species (Lorence & Flynn, unpublished). Ad¬ ditional threats are posed by feral animals includ¬ ing feral pigs (Sus scrofa ) and the Polynesian rat (Rattus exulans), a known seed predator that ad¬ versely impacts regeneration of the Hawaiian Prit¬ chardia species (Cuddihy & Stone, 1990). Feral pigs are present in many areas inhabited by this species and may destroy seedlings and increase op¬ portunities for alien plants to invade through deg¬ radation of the habitat. Based on its relatively low population status, presence of threats, and low re¬ generation, we suggest this new species should be considered for threatened status. Etymology. We are pleased to name this new species for one ol its initial collectors, Timothy W. Flynn, in recognition of his extensive botanical col¬ lections and contributions to our knowledge of the Hawaiian flora. Affinities. Among the Kaua'i species, Pritchar¬ dia flynnii resembles P. hardyi Rock, which is also characterized by abaxially densely gray tomentose- 188 Novon lepidote leaves. This latter speeies differs by its stouter trunk 4-8 m tall and 30-45 cm diam., in¬ florescences equal to or often exceeding the leaves with minutely villous rachillae, and long-drooping infructescences with larger fruits 20—35(40) X 16- 25 mm when dry. Pritchardia flynnii is distinguished from its Kaua'i congeners by the following combination of features: short stature with trunk 0.7-7(11) m tall; relatively small crown with only 10 to 26 leaves; leaf blades relatively small, 57-107 cm long, nearly plane to slightly undulate and abaxially densely ap- pressed tomentose-lepidote; relatively small inflo¬ rescences branched to the second degree, reaching Vz (rarely %) length of blade in flower and often reaching the blade apex in fruit; fruits smooth, el¬ lipsoid or ovoid-ellipsoid. Several collections from the Makaleha Mountains (Lorence & Flynn 7424, PTBG; Perlman & Wood 16260) diller in having leaf blades that are abaxi¬ ally sparsely scattered lepidote, but these speci¬ mens are otherwise referable to P. flynnii. The com¬ paratively sparse foliar pubescence may be due to leaf age or weathering, or it may be genetically con¬ trolled. This variant will key out under 8b in the key below. Seven endemic Pritchardia species including P. flynnii occur on Kaua‘i, and two Pacific species are commonly cultivated there (P. pacifica Seemann & H. Wendland and P. thurstonii F. Mueller & Drude). In order to successfully key out these palms com¬ plete material is required including flowers, fruits, and leaves. In addition, the number of leaves per crown, relative lengths and positions of leaves, in¬ florescences, and infructescences should be noted and photographed when making herbarium speci¬ mens. With adequate herbarium material available, the native Kaua‘i species generally can be sepa¬ rated by the following key, which was adapted in part from Gemmill (1668) and Read and Hodel (1660). Measurements are based on dried herbari¬ um specimens. Kky to tiik Endemic Species oe Pritchardia in Kaua'i, Hawai‘1 lu. Inflorescences nearly equal lo or exceeding leaf blades in flower; infructescences usually extend¬ ing well beyond the blade apex and generally pendulous . P. hardyi lb. Inflorescences shorter than petioles or reaching Vz (rarely f4) length of leaf blade; infructescences shorter than or equaling but not exceeding the blade apex, erect, arcuate, or pendulous. 2a. Leaf blades abaxially densely silvery to pale brown or golden, oppressed lepidote-tomen- tose, the scales and hairs matted and ob¬ scuring the blade surface. 3a. Inflorescence rachillae densely flocco- se-lanate at least when young, becoming glabrate with age . P. minor 3b. Inflorescence rachillae glabrous or sparsely to densely velutinous-tomen- tose when young and soon glabrescent. 4a. Inflorescence rachillae glabrous, viscous with varnish-like resinous coating; flowers and buds shiny, viscous-resinous; fruit 33—35 mm long . P. viscosa 4h. Inflorescence rachillae velutinous- tomentose or glabrescent, not vis¬ cous or resinous; flowers and buds dull, not shiny and viscous; fruit 22-25 mm long . P. flynnii 2b. Leaf blades abaxially with scattered to closely arranged scales and hairs, but these not obscuring the blade surface. 5a. Large palms, up to 20 m tall; trunk large and stout, 50 cm or more DBH; crown with more than 40 leaves; leaf blades waxy or glaucous pale green . . . P. waialealeana 5b. Moderate- to small-sized palms, less than 10 m tall; trunk moderate to slen¬ der, much less than 50 cm DBH; crown with less than 30 leaves; leaf blades plain green. 6a. Leaves with petioles 00— 170 cm long; fruit 30—38 X 18 — 21 mm . P. perlmanii 6b. Leaves with petioles (31)35—90 cm long; fruit 18-28 X 12-21 mm. 7a. Abaxial surface of blade with lepidia 0.2— 0.3 mm long; flow¬ ers with calyx 4- pedicel 1.8— 3 mm; petals 3-3.5 mm .... . P. napaliensis 7b. Abaxial surface of blade with lepidia 0.4— 1.5 mm long; flow¬ ers with calyx + pedicel 2.5— 5 mm; petals 4.5—7 mm. 8a. Rachillae glabrous; floral bracts scarious, ovate-tri¬ angular, 0.5— 1.5 mm; ca¬ lyx + pedicel 2.5—3 mm; petals 4.5—5 mm . . P. limahuliensis 8b. Rachillae sparsely to densely velutinous-tomen- tose when young, often glabrescent; floral bracts stiff, subulate-filiform, 1.5—3 mm; calyx + pedi¬ cel 3—5 mm; petals 5.7—7 mm . P. flynnii (Makaleha Mountains collections) Paratypes. HAWAIIAN ISLANDS (U.S.A.). Kaua'i: koloa District, Li hue— Koloa forest reserve, Wahiawa Stream ud, F. M. 1846. Crueiferae. Pp. 105-185 in C. Gay, Historia Ffsica y Polftica de Chile, Vol. 1, Botany. Paris, Santiago. Volume 14, Number 2 2004 O’Kane & Al-Shehbaz Physaria in South America 205 Boelcke, 0. 1967. Crueiferae. Pp. 281-371 in A. L. Ca¬ brera (editor). Flora de la Provincia de Buenos Aires, Vol. 4, pt. 3. Buenos Aires. - & C. Bomanezuk. 1984. Crueiferae. FI. Patago- nica, Vol. 4A: 373-544. INTA, Buenos Aires. Eichler, A. G. 1865. Crueiferae. Pp. 293-338 in C. F. Martius (editor), FI. Brasil, Vol. 13, pt. 1. Miinchen, Wien, Leipzig. Kurtz, F. 1893. Sertum Cordobense — Observaciones Sobre Plantas Nuevas, Baras 6 Dudosas de la Provincia de Cdrdoba. Revista Mus. La Plata 5: 283-303. Manton, I. 1932. Introduction to the general cytology of the Crueiferae. Ann. Bot. II, 46: 509-556. Muschler, B. 1908. Crueiferae andinae. Bot. Jahrb. Syst. 40: 267-277. O’Kane, S. L., Jr. 1999. Lesquerelln navajoensis (Brassi- caceae), a new species of the L. hitchcockii complex from New Mexico. Madrono 46: 88—91. - & 1. A. Al-Shehbaz. 2002. Paysonia, a new genus segregated from Lesquerella (Brassicaceae). Novon 12: 379-381. - , - & N. J. Turland. 1999. (1393) Proposal to conserve the name Lesquerella against Physaria (Cru- ciferae). Taxon 48: 163—164. Payson, E. B. 1921 [issued 1922], A monograph of the genus Lesquerella. Ann. Missouri Bot. Card. 8: 103— 236. Philippi, B. A. 1864. Plantarum novarum Chilensium: Centuriae, inclusis quibusdam Mendocinis et Patago- nicis. Linnaea 33: 1—308. Rollins, R. C. 1939. The cruciferous genus Physaria. Bhodora 41: 392^415. - . 1993. The Crueiferae of Continental North Amer¬ ica. Stanford Univ. Press, Stanford. - . 1995. Two Lesquerellas (Crueiferae) of south central and western Montana. Novon 5: 71—75. - & E. A. Shaw. 1973. The Genus Lesquerella (Cru- ciferae) in North America. Harvard Univ. Press, Cam¬ bridge. - . K. A. Beck & F. E. Caplow. 1996. An undescri¬ bed species of Lesquerella (Crueiferae) from the State of Washington. Bhodora 97: 201—207. Schulz, O. E. 1934. Neue Cruciferen aus Sudamerika. No- tizbl. Bot. Gart. Berlin-Dahlem 12: 39^11. - . 1936. Crueiferae. Pp. 227—658 in A. Engler & H. Harms (editors), Die Naturlichen Pflanzenfamilien, Vol. 17B. Verlag von Wilhelm Engelmann, Leipzig. Watson, S. 1888. Contributions to American botany. XV II. I. Some new species of plants of the United States, with revisions of Lesquerella ( Vesicaria ) and of the North American species of Draba. Proc. Amer. Acad. Arts 23: 249-267. A New Species of Pariana (Poaceae: Bambusoideae: Olyreae) Endemic to the Atlantic Moist Forest in the State of Bahia, Brazil Reyjane Patricia de Oliveira Departamento de Ciencias Biologicas, Programa de Pos-graduagao em Botanica/Herbario, Universidade Estadual de Feira de Santana, BR 116N km 3, 44031-460, Feira de Santana — BA, Brazil, patricia@uefs.br Hilda Maria Longhi-Wagner Departamento de Botanica, Universidade Federal do Rio Grande do Sul. Av. Bento Gongalves, 9500 (43323), 91501-970, Porto Alegre — RS, Brazil, hmlw@vant.com.br Victoria C. Hollowell Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. victoria.hollowell@mobot.org ABSTRACT. A new species of Pariana from the Atlantic forest of the Brazilian state of Bahia is de¬ scribed. Pariana carvalhoi is an herbaceous bam¬ boo (Poaceae: Bambusoideae: Olyreae) that resem¬ bles P. lanceolata hut differs from it by the very short to absent fimbriae at the leaf sheath apex. Both species are endemic to Bahia, Brazil. The new species is only known from the locality of Una, in southern Bahia. It grows in remnants of humid for¬ est and should be considered as critically endan¬ gered. Key words: Bahia, Bambusoideae, Brazil, Oly¬ reae, Pariana, Poaceae. Pariana Aublet includes about 30 species dis¬ tributed in the Amazon Basin from Costa Rica and Trinidad, to Bolivia and Brazil (Judziewicz et al., 1999). It has been included in its own tribe Pari- aneae by several authors (Calderon & Soderstrom, 1980; Clayton & Renvoize, 1986; Hollowed, 1994), but presently it is considered a genus of the tribe Olyreae (Clark et ah, 1995; Hollowed, 1987, 1997; Zhang & Clark. 2000). Pariana , together with Er- emitis Doll, is classified in the subtribe Parianinae (Hollowed. 1987. 1997), which is characterized by the presence of fimbriae at the apex of the leaf sheaths, termed oral setae (Calderon & Soderstrom, 1980; Hollowed. 1987, 1997), and by the spiciform inflorescence with each whorl presenting one fe¬ male central spikelet surrounded by five male spikelets. Kach of these gynecandrous whorls cor¬ responds to a diaspore of the species, shattering as intact spikelet whorls. According to Soderstrom and Calderon (1971, 1974). the showy inflorescences of Pariana species attract insects especially by the easily visible anthers, and these bamboo grasses are probably entomophilous. The pollen grains of some Pariana species differ from the basic pattern found in grasses (anemophily) by showing an are- olate exine, sometimes vestigial columellae, and no distinct annulus (Salgado-Labouriau & Rinaldi, 1990; Salgado-Labouriau et al., 1993); this would offer more friction in wind transport than the smooth surface of other grass pollen grains. The absence of an annulus in Pariana pollen was con¬ firmed, and the developmental detail of its micro¬ spore has been further investigated by Skvarla et al. (2003). According to Clark (1990) the state of Bahia rep¬ resents a significant center of diversity for Neotrop¬ ical bamboos, with many new species being de¬ scribed from this area. The two species of Pariana found in Bahia, P. lanceolata Trinius (cf. Renvoize, 1984) and the new species described here, differ from other species of the genus by the barbate style, the smaller number of pale stamens (2 vs. 6 to 36 or more, with pale to bright yellow anthers, within Pariana), and the much longer pedicels of the male spikelets. Due to this reduced number of stamens, the inflorescences of the new and Bahian species are less conspicuous than those of other Pariana species. The new species epithet, Pariana carvalhoi, hon¬ ors the outstanding botanist Andre Maurfcio de Carvalho (1951—2002), who for many years was the curator of the herbarium at CEPLAC (Cocoa Re¬ search Center in llheus, Bahia) and was a great Novon 14: 206-209. 2004. Volume 14, Number 2 2004 Oliveira et al. Pariana carvalhoi from Brazil 207 Figure 1. A— C, Pariana lanceolata Trinius. — A. Habit. — B. Inflorescence whorl of male and female spikelets, the solitary female spikelet cryptic. — C. Apex of leaf sheath with fimbriae evident and numerous. D— N, Pariana carvalhoi K. P. Oliveira & 11. M. Longhi-Wagner. — D. Habit. — E. Apex of leaf sheath with fimbriae absent or vestigial. — F. Inflorescence whorl. — G. Central female spikelet. — H. Articulate inflorescence rachis. — I. Female anthecium, ventral aspect. — J. Female anthecium, dorsal aspect. — K. Male spikelet with adnate laminar pedicel, dorsal aspect. — L. Male anthecium, ventral aspect. — M. Male anthecium, dorsal aspect. — N. Caryopsis. A— C drawn from IF IF Thomas et al. 9399 (CEPEC); I ) — N drawn from the holotype, A. M. Carvalho et al. 4382 (CEPEC). Illustration prepared by Ivan Farias de Castro and Reyjane Patricia de Oliveira. 208 Novon defender and advocate for the preservation of the Atlantic forest in eastern Brazil. Key to the Species of Pariana in Bahia, Brazil 1. Leaf sheaths with numerous fimbriae at the apex, 0.7—20 mm long; leaf blades lanceolate (7.5— )11- 16(— 21.4) X 0.7— 1.7(— 2.5) cm . . . Pariana lanceolata 1'. Leaf sheaths without fimbriae at the apex, rarely with 1 to 3 vestigial fimbriae, less than 1 mm long; leaf blades ovate-lanceolate, (6— )1 1.5-20 X 2.1— 3.4 cm . Pariana earvalhoi Pariana earvalhoi R. P. Oliveira & H. M. Longhi- Wagner, sp. nov. TYPE. Brazil. Bahia: Una, along road Sao Josd/Una, 10 Feb. 1994, A. M. Carvalho, L. G. Clark, T. S. dos Santos, W. W. Thomas & S. Sant' Ana 4382 (holotype, CE- PEC; isotype, ISC). Figure I . Herba perennis cespitosa; vaginae glabrae rare fimbria- tae; lolia ovato-laneeolata. Spieulae masculae pedicellis longis, pilosis; stamina 2. Plants with monomorphic stems 40—75 cm tall, 2-4 mm diam., glabrous or pilose, bearing 3 to 6 leaves; nodes glabrous, with a small conspicuous scar beneath. Leaf sheaths glabrous or rarely ciliate at the margins; apex usually without oral fimbriae, rarely with 1 to 3 vestigial fimbriae, less than 1 mm long; leaf blades (6-)l 1 .5—20 X 2. 1-3.4 cm, ovate- lanceolate, base symmetrical, attenuate, apex sym¬ metrical, acuminate, both blade surfaces adaxially and abaxially glabrous or ciliate along the adaxial midrib, blade margins scaberulous; pseudopetioles 1- 1.6 mm long; ligules 0.3-0. 5 mm long. Inflores- cence 6.5—8 X 0.5— 0.8 cm, terminal and spiciform, with 6 to 8 spikelet whorls 12.5-14 X 0.5— 0.8 mm; rachis 9-9.5 mm long, scaberulous. Female spike- lets 10-11.2 X 2.6—3 mm, oblong, glabrous; glumes 9-10 mm long, both acuminate or shortly caudate, totally glabrous or with short trichomes at the apex; anthecium 9.5—1 1 X 2—3 mm, oblong- lanceolate, yellowish or dark brown. Male spikelets 2— 2.5 X 1.2— 1.5 mm, oblong-ovate, brown; pedi¬ cels 7—8.2 mm long, laminar, usually pilose; glumes 2—3 mm long, oblong-triangular, pilose; lemmas pu- berulent, 3-nerved; stamens 2. Caryopsis glabrous, cryptic but free within the persistent anthecium, apex with erect stylar remnants, hilum linear. Pariana earvalhoi differs from P. lanceolata es- peeially by its wider leaf blades that are ovate-lan¬ ceolate (vs. lanceolate in P lanceolata ), and by the absence of fimbriae at the apex of the leaf sheaths, that, i( present, are vestigial. This absence of leaf sheath fimbriae was noted by Tutin (1936) for other species of Pariana but only from Amazonia. Both Pariana lanceolata and P. earvalhoi are en¬ demic to the Brazilian state of Bahia, occurring within moist Atlantic forest. According to IUCN cri¬ teria (IUCN, 1994), P. earvalhoi should be consid¬ ered “critically endangered,” with only a few pop¬ ulations known, all oceurring outside preservation areas. Although many field trips were made to dif¬ ferent areas in Bahia, additional populations of the new species have not been found. Paratypes. BRAZIL. Bahia: Una, estrada Una a Sao JosG 18 Feb. 1994, A. M. Carvalho et al. 4395 (CEPEC), 10 Feb. 1994, A. M. Carvalho et al. 4384 (CEPEC); Fa¬ zenda Dendaevea, 30 km from Una, 28 Apr. 1976, C. E. Calderon et al. 2403 (CEPEC). Acknowledgments. The authors thank the Univ- ersidade Estadual de Feira de Santana for making the field trips possible, and the Cocoa Research Center for specimen loans and logistic support for fieldwork. The second author thanks the Brazilian Research Council (CNPq) for a Research Fellow¬ ship. In memoriam, we are grateful to the unfor¬ gettable Dr. Andre Maurfeio de Carvalho for his enthusiastic support. Literature Cited Calderdn, C. E. & T. R. Soderstrom. 1980. The genera of Bambusoideae (Poaceae) of the American continent: Keys and comments. Smithsonian Contr. Bot. 44: 1-27. Clark, L. G. 1990. Diversity and biogeography of Neo¬ tropical bamboos (Poaceae: Bambusoideae). Acta Bol. Brasil. 4: 125-132. - , W. Zhang & J. E. Wendel. 1995. A phylogeny of the grass family (Poaceae) based on ndhV sequence data. Syst. Bot. 20: 436-460. Clayton, W. D. & S. A. Renvoize. 1986. Genera Gramin- um: Grasses of the World. Royal Botanic Gardens, Kew. Hollowell, V. C. 1987. Systematics of the Subtribe Pari- aninae (Poaceae: Bambusoideae: Olyreae). Ph.D. The¬ sis, University of South Carolina, Columbia. - . 1994. Tribus Parianeae. Pp. 216—218 in G. Dav- idse, M. Sousa & A. 0. Chater (editors). Flora Mesoam- ericana, Vol. 6. Alismataceae a Cyperaceae. Universi- dad Nacional Autonoma de Mexico, Mexico, D.F.; Missouri Botanical Garden, St. Louis; The Natural His¬ tory Museum, London. - . 1997. Systematic relationships of Pariana and associated Neotropical taxa. Pp. 45—60 in G. P. Chap¬ man (editor). The Bamboos. Academic Press, London. IUCN (The World Conservation Union). 1994. The World List of Threatened Trees. Appendix 4: 629-643. World Conservation Press, Gland, Switzerland. Judziewicz, E. M., L. G. Clark, X. Londono & M. ,1. Stern. 1999. American Bamboos. Smithsonian Institution Press, Washington, D.C. Renvoize, S. A. 1984. The Grasses of Bahia. Royal Bo¬ tanic Gardens, Kew. Salgado-Labouriau, M. L. & M. Rinaldi. 1990. Palynology of Gramineae of the Venezuelan mountains. Grana 29: 1 19-128. - , S. Nilsson & M. Rinaldi. 1993. Exine sculpture in Pariana pollen (Gramineae). Grana 32: 243-249. Skvarla, J. J., J. R. Rowley, V. C. Hollowell X' W. E. Chis- Volume 14, Number 2 2004 Oliveira et al. Pariana carvalhoi from Brazil 209 soe. 2003. Annulus-pore relationship in Gramineae (Po- aceae) pollen: The pore margin of Parinrui. Amer. J. Bot. 90: 924—930. Soderstrom, T. R. & C. E. Calderon. 1971. Insect polli¬ nation in tropical rain forest grasses. Biotropica 3: 1— 16. - & - . 1974. Primitive forest grasses and evo¬ lution of the Bambusoideae. Biotropica 6: 141—153. Tutin, G. 1936. A revision of the genus Pariana (Grami¬ neae). J. Linn. Soc., Bot. 50: 337—362. Zhang, W. & L. G. Clark. 2000. Phytogeny and classifi¬ cation of the bamboos (Poaceae: Bambusoideae). Pp. 35^12 in S. W. L. Jacobs & J. Everett (editors). Grasses, Systematics and Evolution. CS1R0, Melbourne. [Pro¬ ceedings of the 2nd International Conference on the Comparative Biology of the Monocotyledons, Vol. 2.] New Taxa of Calyptranthes (Myrtaceae) from Colombia Carlos Parra- 0. Instituto de Ciencias Naturales, Universidad Nat ional de Colombia, Apartado Aereo 7495, Bogota, Colombia, caparrao@unal.edu. co Abstract. Two new species (C. buchenavioides, C. estoraquensis) and one new subspecies (C. loz- anoi subsp. tornentosa ) of Calyptranthes (Myrta- ceae) from Colombia are described and illustrated. The taxonomic affinities and the distribution ol the new taxa are discussed. Key words: Calyptranthes , Colombia, Myrta¬ ceae. Calyptranthes Swartz is a genus of ca. 100 spe¬ cies in Myrtaceae, distributed from Florida (United States), the Caribbean, and Mexico to northern Ar¬ gentina and Uruguay (Legrand, 1962; MeV'augh. 1958, 1963; Sanchez-Vindas, 1990). The genus is distinguished by frequently dibrachiate trichomes, inflorescence usually a pair of panicles, calyptrate calyx, 2(-3)-locular ovary, and myrcioid embryo (Landrum & Kawasaki, 1997; Holst, 2002). In a preliminary study of the genus for the Colombian flora, two new species and a new subspecies of Ca¬ lyptranthes lozanoi C. Parra-0. were recognized as undescribed. Calyptranthes buchenavioides C. Parra-0., sp. nov. TYPE: Colombia. Valle del Cauca: “Hoya del rfo Cali, vertiente derecha, entre Puente Sofia y Yanaconas,” 1580-1730 in, 6 Nov. 1944 (fr), J. Cuatrecasas 19725 (holotype, F). Figure 1. Arbor 3-^4 m alta; ramulis juvenilibus subteretibus, par- ee pubescentibus, vetustis teretibus, glabratis. Folia pe- tiolata, petiolo 0.6— 1.6 mm longo; lamina coriacea, supra glabra, subtus glabrata, obovata, 1.3— 2.6 X 0.7-1. 4 cm, apice obtusata, basi attenuata, costa supra impressa, gla¬ bra. subtus convexa, tantum per proximalem unam terliam partem parce pilosa. Inflorescentia et flores non visi. Fruc- tus globosus, diametro 1.0— 1.3 cm. Tree, 3-^4 in tall, almost glabrous; trichomes when present pale gray (except in the embryo); young branches terete or subterete, sparsely pu¬ bescent; old branches terete, glabrous; vegetative buds lanceolate or conical, 0.8— 1.1 mm long, pu¬ bescent. Leaf blades obovate, 1.3— 2.6 X 0.7-1. 4 cm, coriaceous, the upper surface glabrous, with impressed glandular dots, the lower surface gla¬ brous; apex obtuse, sometimes slightly apiculate; Novon 14: 210-215. 2004. base attenuate; midvein convex above, glabrous, convex below, sparsely pilose only in the proximal VS; lateral veins 7 to 11 pairs (almost impercepti¬ ble), sulcate above, impressed below, joining in a straight marginal vein at 0.5 mm from the margin; petiole 0.6— 1.6 mm long, barely rugose, glabrous or sparsely pubescent, slightly canaliculate. Inflo¬ rescence terminal, apparently a panicle, the axes terete or subterete, glabrous; bracts ovate, 0.5 X 0.3— 0.4 mm. pubescent, deciduous; bracteoles un¬ known. Buds and flowers unknown: calyptra (per¬ sistent in the fruit) discoid, ca. 1.4 mm diam., with¬ out prominent apiculum. glabrous. Fruits globose, blackish, 1-1.3 cm diam., exocarp smooth, gla¬ brous; seeds 1, reniform, 1 — 1.1 X 0.9—1 cm, seed coat papyraceous, pale orange, smooth, without glands; embryo myrcioid, lower portion ol the hy- pocotyl and portions of the cotyledons densely pu¬ bescent with long (1.2— 1.5 mm) gray trichomes mixed with short (ca. 0.5 mm) red trichomes. The specific epithet refers to the striking vege¬ tative similarity (profuse ramification, opposite, ob¬ ovate, and coriaceous leaves, and similar size of the leaves) between this new species and some species ol Buchenavia Eichler (Combretaceae), especially with Buchenavia tetraphylla (Aublet) R. A. Howard and Buchenavia parvifolia Ducke. Calyptranthes buchenavioides is known from coastal and premontane forest of the western An¬ dean Cordillera in Colombia. Despite the lack of complete inflorescences of the studied specimens, other characters such as the calyptra (falling as a unit), the circular scar at the top of the fruit, and the myrcioid embryo place it in Calyptranthes. Calyptranthes buchenavioides is somewhat simi¬ lar to Calyptranthes pulchella DC., a species known from Colombia to Brazil. The leaves of the species are similar in shape (obovate blades) and size. Ca¬ lyptranthes buchenavioides differs from Calyptran¬ thes pulchella by the terete or subterete (vs. 2- winged) branchlets, the base of the leaf blades attenuate (vs. acute or cuneate), and the diameter of the calyptra (ca. 1.4 mm vs. 2-3 mm). Paratypes. COLOMBIA. Valle de Cauca: “Punta Arenas, north shore of Buenaventura Bay, near sea level," Volume 14, Number 2 2004 Parra-O. New Taxa of Calyptranthes 211 Figure 1. Calyptranthes buchenavioides C. Parra-0. — A. Vegetative branch. — B. Detail of a partial inflorescence. — C. View of the calyx scar on the Iruit apex, showing the attached calyptra. — D. Fruit. — E. Embryo. (A, C-E, from the holotype, Cuatrecasas 19725; B, Cuatrecasas 38628 (F paratype).) 2 June 1944. E. R. Killip & J. Cuatrecasas 38628 (COL, F). Calyptranthes estoraquensis C. Parra-O., sp. nov. TYPE: Colombia. Norte de Santander: l.a Playa de Helen, “vereda Peritama, Area Nat¬ ural Unica Los Estoraques,” 1650 m, 20 Feb. 2002 (Ir), R. Galindo, J. Penuela & E. Rodri¬ guez 689 (holotype, COL: isotype, MO). Fig¬ ure 2. Arbor 5 m alta; ramulis juvenilibus subteretibus, parce pubescentibus, vetustis teretibus, glabratis. Folia petiola- ta, petiolo 3.8— 5. 5(— 7.6) mm longo; lamina coriacea, supra glabra vel puberula, subtus puhescente, obovata, 5.0— 7.3 X 3.1— 4.7 cm, apice obtusata, basi cuneata, costa supra impressa, tantum per proximalem unam tertiam partem puhescente, subtus convexa, dense puhescente. Inflores- centia paniculata, 4—6 cm longa, dense pubescens; pedi- cello nullo. Flos alabastro obovoideo, 2.3— 2.6 mm longo; calyptra 2.0— 2.2 mm diam.; staminibus (1.6)2. 1—3.1 mm longis; ovario 2-loculari. Fructus globosus, diametro 5.5- 7.9 mm. Tree, 5 m tall; trichomes when present light red¬ dish brown; young branches subterete, subcom¬ pressed, sparsely pubescent, the light reddish 212 Novon Figure 2. Calyptranthes estoraquensis C. Parra-O. — A. Fruiting branch. — B. Detail of the leaf blade pubescence (lower surface). — C. Flower, after anthesis. — D. Fruit. — E. Seed. — F. Longitudinal section of the fruit and seeds showing the embryo. (A, B, D— F, from the holotype Galindo el al. 689; G, Galindo el al. 1168 (COL paratype).) brown trichomes mixed with gray trichomes, the gray ones located mostly in the margins and apex of the branches; old branches terete, glabrous; veg¬ etative buds semi -elliptic, 1.4— 1.5 mm long, pilose. Leaf blades obovate, 5—7.3 X 3.1— 4.7 cm, coria¬ ceous. the upper surface glabrous to puberulous, sometimes with whitish trichomes, with impressed glandular dots, the lower surface pubescent; apex obtuse; base cuneate; midvein sulcate above, pu¬ bescent only in the proximal Yi, convex below, densely pubescent; lateral veins 17 to 35 pairs in addition to weaker intermediates, convex on both surfaces, joining in a straight marginal vein at 1.4 mm from the margin; petiole 3.8-5. 5(— 7.6) mm long, rugose, puberulous with gray or mixed gray and light reddish brown trichomes (petioles of Volume 14, Number 2 2004 Parra-O. New Taxa of Calyptranthes 213 leaves near the inflorescences are densely pubes¬ cent with light reddish brown trichomes), canalic¬ ulate. Inflorescence axillary or terminal, the paired and opposite panicles 4-6 cm long, with 20 to 30 flowers per panicle, the axes subterete or com¬ pressed. densely pubescent; peduncles 2. 7-3.3 cm X 1.7— 2.4 mm; bracts lanceolate, ca. 4.5 X 2 mm, pubescent, truncate in the base, deciduous; brac- teoles (1)2, the larger one al the base of the hy- panthium, ovate, ca. 0.7 X 0.3 mm, pubescent, de¬ ciduous or persistent alter anthesis, the smaller one incompletely surrounding the base of the hypan- thium, squamiform, ca. 0.5 X 0.9 mm, pubescent, persistent after anthesis. Flower buds obovoid, 2.3— 2.6 mm long, pubescent, sessile; ealyptra discoid, 2—2.2 mm diam., without prominent apiculum, pu¬ bescent; petals absent; hypanthium 1.8— 2.1 X 2.6- 2.9 mm, prolonged 0.7— 0.8 mm above the ovary, pubescent outside, glabrous inside; style generally not developed, when occasionally developed, 1 .2 mm long, glabrous; filaments (1. 4)1.9— 2.9 mm; an¬ thers globose, 0.2 mm long, bearing one apical gland; ovary 1 mm diam., 2-locular, 1 ovule per locule. Fruits globose, dark wine-red, 5.5— 7.9 mm diam., exocarp smooth, sparsely to moderately pu¬ bescent; seeds 2. reniform, 4.8-5. 8 X 2.8— 4.2 mm, seed coat papyraceous, orange, smooth, without glands; embryo myrcioid, glabrous. Calyptranthes estoraquensis is only known from the Colombian Natural Area named “Estoraques,” a semidesert landscape (badlands) made of rock formations caused by erosion. Estoraques is located at the eastern Andean Cordillera in Norte de San¬ tander Department, between 1450 and 1900 m. At present, this particular area is included in the Na¬ tional Park System as a protected area, but the pop¬ ulation of the species there seems to be small, “with only a few individuals” (R. Galindo, pers. comm. 2002). Calyptranthes estoraquensis seems to be related to Calyptranthes brasiliensis Sprengel, and espe¬ cially with Calyptranthes brasiliensis var. mutabilis (Berg) Legrand. A specimen of C. brasiliensis var. mutabilis from Brazil, Duarte 9152 (MO), exhibits some degree of similarity w ith the specimens of C. estoraquesis from Colombia, particularly in the veg¬ etative morphology (trichomes light reddish brown in young branches and petioles, and similar size of the petioles). The pubescence of the adaxial leaf blade and the inflorescence and the size of the in¬ florescence are also common characteristics of these two taxa. Nevertheless, Calyptranthes estora¬ quensis differs from Calyptranthes brasiliensis var. mutabilis by its leaf base (cuneate vs. rounded), the laminae (obovate vs. ovate or ovate-oblong) and the size of the fruit (5.5— 7.9 mm vs. 4—4.5 mm). Paratype. COLOMBIA. Norte de Santander: La Playa de Beldn, “vereda Peritama, Area Natural tlnica Ixjs Estoraques,” 1550 m, 26 Feb. 2003 (fl). H. Galindo, J. Pefiuela & E. Rodriguez 1168 (COL). Calyptranthes lozanoi C. Parra-O. subsp. to- mentosa C. Parra-O., subsp. nov. TYPE: Co¬ lombia. Antioquia: Guarne, Laguna de Guar- ne, 20 July 1938 (bud, fr). Bra. Daniel 2258 (holotype, US; isotype, COL). Figure 3. Haec subspecies a Calyptranthe lozanoi subsp. lozanoi alabastris ellipsoideis vel late ellipsoideis (nee obovo- ideis), ramulis, petiolo, foliis (subtus), inflorescentia et al¬ abastris modice vel dense pubescentibus (nee parce pu- bescentibus trichomatibus albis vel glabratis, interdum glabris), atque fructu bullato (nec laevi), exocarpio spar- sim pubescente (nec glabro) differt. Shrub, 2^1 m tall; trichomes when present yel¬ lowish or blackish; young branches subcompressed, sparsely to densely pubescent, sometimes almost blackish; old branches terete, glabrous; vegetative buds lanceolate or elliptic, 2.2^1. 1 mm long, pu¬ bescent. Leal blades obovate, sometimes broadly elliptic, 3.2— 5.2(-6) X 1.6— 3.1(— 4) cm, coriaceous, the upper surface glabrous to sparsely puberulous, with impressed glandular dots, the lower surface sparsely to densely pubescent; apex acute or ob¬ tuse; base cuneate; midvein convex above, puber¬ ulous to sparsely pubescent in the proximal Vs, con¬ vex below, moderately to densely pubescent; lateral veins 15 to 18 pairs, convex on both surfaces; mar¬ ginal veins 2, the inner one 0.9-1. 1 mm from die margin, the outer one ca. 0.4 mm from the margin; petiole 2.6-6.1(— 8) mm long, blackish, lightly ru¬ gose, sparsely to densely pubescent (petioles of leaves near the inflorescences are densely pubes¬ cent with yellowish trichomes), canaliculate. Inflo¬ rescence subterminal, the paired and opposite pan¬ icles 3.5—7 cm long, with 12 to 25 flowers per panicle, the axes subterete or compressed, densely pubescent, sometimes almost blackish, rarely gla¬ brous; peduncles 2.2—4 cm X 1.3— 1.7 mm; bracts dimorphic, either lanceolate, ca. 6X2 mm, pu¬ bescent with yellowish trichomes, deciduous, or ob¬ ovate, 1.2— 1.7 X 0.4— 0.5 cm. persistent; bracteoles 2, opposite at base of hypanthium, pubescent, de¬ ciduous or persistent after anthesis, the larger one lanceolate, 2.8— 3.4 X 1 — 1.2 mm, the smaller one elliptic, 1.7— 2.8 X 0.6— 0.8 mm. Flower buds ellip¬ soid or widely ellipsoid, 4.6— 6.4 mm long, pubes¬ cent, sometimes almost blackish, rarely glabrous, on pedicel 0.8— 1.3 X 0.7— 0.9 mm, densely pubes¬ cent, rarely glabrescent; ealyptra discoid, 3—3.5 214 No von Figure 3. Calyptranthes lozanoi C. Parra-O. subsp. tomentosa C. Parra-O. — A. Fertile branch. — B. Flower bud. — C. Flower, after anthesis. — D. Longitudinal section of hypanthium and ovary. — E. Petal. — F. Fruit. — G. Embryo without seed coat. (A, B, F. G, from the holotype Bro. Daniel 2258 ; G— E, Cabrera 254 (GOl. paratype).) Volume 14, Number 2 2004 Parra-O. New Taxa of Calyptranthes 215 mm diam., weakly to prominently apiculate, gen¬ erally pubescent, rarely glabrous; petals present, attached to the inner portion of the calvptra, 2 or 3 (early deciduous at anthesis), obovate, 2.2 X ].] mm, dorsally densely pubescent with white tri- chomes; hypanthium 2.5—3 X 4. 1-4.7 mm, pro¬ longed 1.3— 1.6 mm above the ovary, pubescent out¬ side, glabrous inside; style 4-5.5 mm long, glabrous; stamens 50-150; filaments 2.2-6 mm; an¬ thers globose, 0.2— 0.3 mm long, without apical glands; ovary 1.5 mm diam., 2-locular. 1 ovule per locule. Fruits globose, light brown, 8 mm diam., exocarp bullate, sparsely pubescent with whitish trichomes; seeds 2, reniform, 5.4—6 X 5 mm, seed coat papyraceous, red, smooth, without glands; em¬ bryo myrcioid, glabrous. The subspecific epithet refers to the yellowish or blackish tomentum of the plant that varies from moderately dense to dense on tw igs, leaves (petiole and lower surface of the blade), inflorescence, and flower buds. Calyptranthes lozanoi subsp. tomentosa is known only from the montane forest of the Cen¬ tral Andes Cordillera in Colombia. In 2001 I described Calyptranthes lozanoi (Par¬ ra-O.. 2001) as a conspicuous species from the Si¬ erra Nevada de Santa Marta (Colombia), an isolated coastal mountain separated from the Andes Cordil¬ lera by plains and semiarid regions. The specimens cited here for the description of the new subspecies are similar to the specimens collected in Sierra Ne¬ vada de Santa Marta by the obovate leaf blade (and also in its size), the size of the panicles, the size of the hypanthium, the pubescence of petals, and the size of the fruit. Also, the vegetative morphology of all the specimens strikingly resemble the general aspect of Myrcianthes O. Berg species that grow in the montane forests of Colombia (Parra-O., 2001). Calyptranthes lozanoi subsp. tomentosa differs from the typical subspecies by the ellipsoid or w idely ellipsoid (vs. obovoid) flower buds, the mod¬ erate to dense yellowish or blackish pubescence (vs. sparse whitish pubescence or glabrescent, to often glabrous) of the twigs, petioles, lower surface of the leaf blade, inflorescence, and flower buds (vs. sparse whitish pubescence or glabrescent, to often glabrous), and the bullate, sparsely pubescent fruits (vs. smooth and glabrous). Paratypes. COLOMBIA. Antioquia: Medellin, “Hoya de la Estacidn Experimental Eorestal ‘Piedras Blancas,’ ” June— Oct. 1957 (H), /. Cabrera 86 (COL, MEDEL), /. Ca¬ brera 254 (COL, MEDEL); Medellm-Guarne, “Parque Eeologico Piedras Blancas, laguna de Guarne,” 75°29'W, 6°18'N, 2350 m, 22 July 1995 (II), F Roldan et al. 2354 (HUA, MO). Acknowledgments. I express my special grati¬ tude to the Missouri Botanical Garden and the Gen- tty Fellowship Program, which supported me as a visiting researcher from June to July 2003. In St. Louis, I’m indebted to Alina Freire Fierro for her help in all the different aspects related to this pro¬ gram. I am grateful to F and US for sending loans of Calyptranthes to COL, and to HUA and MEDEL for providing me access to their collections. Thanks to Henry Arellano, who prepared the illustrations, and to Peter Hoeh (MO) for reviewing an earlier version of the manuscript. Thanks to Roy Gereau (MO) for reviewing the Latin descriptions and di¬ agnoses, and to B. Holst (SEL) and an anonymous reviewer for their helpful comments on the manu¬ script. Also, I thank the Herbario Nacional Gol- ombiano (COL) of the Instituto de Ciencias Natur- ales, Universidad Nacional de Colombia, for its support. Literature Cited Holst. B. 2002. New species and notes on Myrtaceae from northern South America. Selbyana 23: 137-180. Landrum, L. & M. L. Kawasaki. 1997. The genera of Myr¬ taceae in Brazil: An illustrated synoptic treatment and identification keys. Brittonia 49: 508-536. Legrand, 1). 1962. El genero Calyptranthes en el Brasil austral. Lilloa 31: 183—206. McVaugh, 11. 1958. Myrtaceae. In: ,|. F. Maguire (editor). Flora of Peru. Field Mus. Nat. Hist.. Bot. Ser. 13, part 4(2): 569-818. - . 1963. Myrtaceae. Flora of Guatemala. Fieldiana, Bot. part 24, 7(3): 283-405. Parra-O., C. 2001. L ria nueva especie de Calyptranthes Sw. (Myrtaceae) de Colombia. Caldasia 23(2): 435-439. Sanchez- V indas, P. 1990. Myrtaceae. In: A. G6mez-Pompa (editor). Flora de Veracruz. Instituto de Ecologfa, Xal- apa, Veracmz, Mexico 62: 1-146. New Species and Combinations in Astragalus (Leguminosae) from China and the Himalayas Dietrich Podlech Ludwig-Maximilians-University Munich, Department Biologie 1 — Systematic Botany, Menzinger StraBe 67, D-80638, Munich, Germany, podlech@lrz.uni-muenchen.de Lang-Ran Xu Northwestern Institute of Botany, Academia Sinica, Yangling 712100, Shaanxi, People’s Republic of China ABSTRACT. Ten new species of Astragalus are de¬ scribed here. They belong to the following sections: section Alopecuroidei : A. burqinensis ; section Bra- chycarpus : A. barclayanus, A. golubojensis, A. mieherorum, A. nanshanicus, A. pseudojagnobicus, A. sagastaigolensis ; section Hemiphaca : A. nigro- dentatus; section Polycladus: A. tibeticola ; section Skythropos : A. griersonii. Furthermore, six taxa are raised in rank. Key words: Astragalus, Bhutan, China, Legu- minosae, Nepal. In preparing the treatment of Astragalus L. for the English version of the Flora of China, several new species could he detected and furthermore some taxa, formerly described as varieties, were found to be good species. In the present paper the new species are described and the formal combi¬ nations are made. The flora of China is very rich in Astragalus species. In the Flora, 357 species, of which 189 are endemic in China, will he treated. Astragalus L. sect. Alopecuroidei DC., Prodr. 2: 294. 1825. TYPE: Astragalus alopecuroides L. Astragalus burqinensis Podlech & L. K. Xu, sp. nov. TYPE: China. Prov. Xinjiang: Burqin, 600 m, 24 July 1998, L. R. Xu 2029 (holotype, MSB). Differt ah A. alopecuro planta in partibus vegetativis glabra vel sub stipulis tantum sparse pilosa, petiolo 2.5— 7 cm (nec 1—3 cm) longo, foliolis 13— 15-jugis (nec 17— 27-jugis), late ovatis, 12-30 X 10-22 mm, apice rotun- data vel leviter emarginata (nec anguste ovata, (10— )20— 35(_40) x (6—) 1 0— 1 3(— 20) mm, apice obtusa ad subacuta), bracteolis deficientibus, vexillo 27-28 mm lon¬ go, dorso piloso (nec ca. 20 mm longo, glabro), laminis carinae ca. 15 X 8-9 mm (nec 7 X 3.5 mm), leguminibus ca. 14 mm (nec ca. 7 mm) longis, ab A. pseudovulpino planta in partibus vegetativis glabra vel sub stipulis tan¬ tum sparse pilosa, bracteolis deficientibus, laminis carinae 8—9 mm (nec 4—5 mm) latis. Plants more than 30 cm tall (lower parts are missing). Stem erect, finely sulcate, glabrous or just below the stipules and at the base of stipules sparsely hairy. Stipules ca. 15 mm long, narrowly triangular, shortly adnate to the petiole, heaves 20- 30 cm long: petiole 2.5—7 cm long, finely striate- sulcate. Leaflets in 13 to 15 pairs, widely ovate, 12-30 X 10-22 mm, at the apex rounded to slight¬ ly emarginate. Peduncle very short, up to 1 cm long, densely covered with spreading hairs 1—2 mm long. Racemes (in fruit) cylindric, 7-9 cm long, densely many-flowered; axis loosely covered with spreading hairs. Bracts linear-acute, 12-18 mm long, ciliate at the margins with spreading hairs 1.5-2. 5 mm long. Bracteoles absent. Flowers sub- sessile. Calyx 12-19 mm long, campanulate, dense¬ ly covered with ascending to spreading hairs 2—5 mm long; teeth subulate, 5-7 mm long. Petals yel¬ low. Standard 27-28 mm long; blade elliptic, ca. 9 mm wide, emarginate at the apex, at the base sub- abruptly narrowed into the long c law. Wings un¬ known. Keel ca. 27 mm long; blades triangular, with gibbously curved lower edge and ± straight upper edge, obtuse at the apex, ca. 15 X 8—9 mm; auricle acute, ca. 1 mm long, claw ca. 13 mm long. Legumes sessile, oblong, 12-14 mm long, 7-9 mm high and 4-7 mm wide, obtusely carinate ventrally, slightly grooved dorsally, abruptly contracted at the apex into a beak ca. 2 mm long, bilocular; valves netlike rugulose, loosely covered with subappres- sed soft hairs up to 3 mm long. Seeds 3 X 2—3 mm, dark brown, smooth. Astragalus burqinensis differs from A. alopecurus Pallas by having vegetative parts glabrous or sparsely hairy only, by the petiole 2.5— i (vs. 1—3) cm long, the leaflets in 13 to 15 (vs. 17 to 27) pairs, widely ovate, 12-30 X 10-22 mm, rounded or slightly emarginate (vs. narrowly ovate, (10— )20— 35(_40) X (6—) 1 0-13(— 20) mm, obtuse to sub- Novoin 14: 216-226. 2004. Volume 14, Number 2 2004 Podlech & Xu Astragalus from China 217 acute), bracteoles absent, standard 27—28 mm long, hairy on the back-side (vs. ca. 20 mm long, gla¬ brous), limbs of keel ca. 15 X 8-9 mm (vs. 7 X 3.5 mm), legumes ca. 14 (vs. ca. 7) mm long, from A. pseudovulpino N. Ulziykhutag by being in veg¬ etative parts glabrous or only below the stipules sparsely hairy, bracteoles absent and limbs of keel 8—9 (vs. 4—5) mm wide. Astragalus sect. Ammodendron Bunge, Mem. Acad. Imp. Sci. Saint Petersbourg 1 1(16): 128. 1868. TYPE: Astragalus ammodendron Bunge. Astragalus macrostephanus (S. B. Ho) Podlech & L. R. Xu, stat. nov. Based on Astragalus iliensis Bunge var. macrostephanus S. B. Ho, Bull. Bot. Res. North-East. Forest. Inst. 3(1): 68. 1983. TYPE: |China.] Xinjiang: Kazak Zizhizhou, Huocheng Xian, Sandohezi, 540 m, 14 June 1959, A. R. Li & J. N. Zhu 10619 (holotype, PE; isotvpe, WUK). Because of the very short original diagnosis of this taxon a complete description is given here. Astragalus macrostephanus differs from A. iliensis by stems of the current year up to 30 (vs. up to 20) cm long, leaves 8-12 (vs. 3—5) cm long, petiole 4— 5 (vs. 0.5—1) cm long, leaflets in 3 pairs (vs. in 1 or 2 pairs), 12-25 (vs. 20-40(-50)) mm long, pe¬ duncles 6-10 (vs. 2-5) cm long, calyx 5-6 mm long, rather densely covered with medifixed, ap- pressed hairs 0. 3-0.4 mm long (vs. 3-4 mm long, densely covered with partly flexuose, asymmetri¬ cally bifurcate, subappressed to slightly ascending hairs), petals longer, standard ca. 13 (vs. 8—9) mm, wings 12 (vs. 7—8) mm, keel 10 (vs. 6—7) mm long, legumes 10—12 (vs. 4-5) mm long, 4—5 (vs. ca. 2) mm high and wide. Plants suffruticose, ca. 35 cm tall, covered with medifixed, appressed white hairs. Caudex divided, with long subterranean stolons, covered with gray- brownish bark. Stems erect, branched, very densely covered with white hairs 0.3— 0.6 mm long, shoots of the current year up to 30 cm long. Stipules whit¬ ish to pale brownish hyaline, 2—4 mm long, obliquely triangular, acute, adnate to the petiole for ca. 1 mm, the lower ones distinctly vaginate-con- nate behind the stem, the upper ones mostly free, sparsely to loosely hairy, at the margins with strong¬ ly asymmetrically bifurcate to basifixed hairs, sometimes glabrescent with age. Leaves 8-12 cm long; petiole 4-5 cm long, like the rachis stout, at the base up to 1.5 mm thick, hairy like the stem. Leaflets in 3 pairs, narrowly elliptic to narrowly ovate, 12-25 X 3—5 mm, acute at the apex, on upper side loosely covered with hairs 0.6— 0.8 mm long, on underside loosely to rather densely cov¬ ered with somewhat shorter hairs. Peduncle 6—10 cm long, hairy like the stem. Raceme 5—12 cm long, very remotely 10- to 12-flowered. Bracts whit¬ ish to pale brownish hyaline, 1-1.5 mm long, nar¬ rowly triangular to narrowly ovate, sparsely white hairy. Pedicels ca. 1 mm long, hairy. Calyx 5-6 mm long, campanulate, rather densely covered with white hairs 0.3— 0.4 mm long; teeth subulate, ca. 1 mm long. Standard ca. 13 mm long; blade ca. 5 mm wide, widely elliptic, emarginate at the apex, below the middle subabruptly contracted into the widely cuneate claw longer than the blade. Wings ca. 12 mm long; blades narrowly oblong, round- tipped, 6 X 1.5 mm; auricle short, claw ca. 6 mm long. Keel ca. 10 mm long; blades obliquely ellip¬ tic-curved, with widely curved lower edge and con¬ cave upper edge, obtuse at the apex, 4.5 X 2 mm; auricle short, clawT 5.5 mm long. Ovary sessile, ovoid, densely hairy; style glabrous. Legumes ses¬ sile, ovoid, 10—12 mm long, 4—5 mm high and wide, obtusely carinate ventrally, widely grooved dorsally, acuminate at the apex into a curved beak ca. 1 mm long, nearly fully bilocular; valves thin but tough, densely villous with subbasifixed, ± spreading, tangled, fine, soft white hairs 2-3 mm long, mostly sitting on depressed tubercles. Astragalus sect. Brachycarpus Boriss., FI. URSS 12: 877. 1946. TYPE: Astragalus melanos- tachys Bentham (lectotype, designated by Pod¬ lech (1990: 466)). Astragalus barclayanus Podlech, sp. nov. T^ PE: Nepal. Tilicho Pass path, 4820 m. 7 June 1971. C. Barclay & I1. M. Synge 2620 (holo¬ type, K). Differt ab A. chateri planta pilis albis nigris immixtis 0.2— 0.3(— 0.5) mm longis, in inflorescentiis nigris ad 0.7 mm longis (nec pilis mere albis 0.3— 0.8 mm longis) ob- tecta, stipulis 3^4 (nec 4—6) mm longis, breviter connatis vel superioribus liberis (nec ad medium connatis), foliolis supra glabris (nec sparse ad laxe pilosis), bracteis angus- tissime triangularibus, 2.5—3 mm longis (nec ovati-acu- minatis, 3-5 mm longis), calyce 4 (nec 5—6) mm longo, pilis ascendentibus ad patentibus (nec subappressis) ob- tecto, vexillo 5-6 (nec 7—8) mm longo, ovario glabro (nec piloso). Plants 7-12 cm tall, covered with subappressed to slightly ascending hairs 0.2-0.3(— 5) mm long, in the inflorescence up to 0.7 mm long. Caudex with slightly elongated stolons provided with stipule-like bracts. Stems short, 3-5 cm long, prostrate to as¬ cending, in lower part (the stolon part) glabrous, pale yellowish, in upper part (stem of the current year) rather densely covered with ± straight, sub¬ appressed to slightly ascending hairs, at the lowest 218 Novon internode with mostly spreading white hairs, with a few black hairs mixed in. Stipules greenish, 3-4 mm long, the lower ones shortly connate behind tin* stem, the upper ones free. Leaves 2.5^1 cm long; petiole 0.5—1 cm long, like the rachis loosely cov¬ ered with tangled, subappressed to ascending white and also black hairs. Leaflets in 7 to 9 pairs, nar¬ rowly elliptic, 4—7 X ca. 2 nnn, at the apex mostly rounded, on upper side glabrous, on underside rather densely to densely covered with subap¬ pressed hairs up to 0.5 mm long, at the midvein and the tip often with black hairs mixed in. Pe¬ duncle 3—5 cm long, rather densely to densely cov¬ ered with subappressed to ascending hairs, at the base mostly with white, in upper part mostly with predominantly black hairs. Raceme globular to ovoid, densely many-flowered, ca. 1.5 cm long. Bracts membranous, very narrowly triangular, 2.5— 3 mm long, black hairy. Pedicels 0.5—1 mm long, black hairy. Calyx 4 mm long, campanulate, dense¬ ly covered with ascending to spreading, tangled black hairs up to 0.6 mm long; teeth narrowly tri¬ angular, 1 .5—2 mm long, densely black hairy on inner side. Petals pale pink and white. Standard 5- 6 mm long; blade ca. 3.5 mm wide, widely elliptic, shallowly emarginate at the apex, very shortly nar¬ rowed at the base without distinct claw. Wings 4.5 mm long; blades obovate-spathulate, widely round¬ ed at the apex, ca. 3X2 mm; auricle ca. 0.5 mm long, claw 1.5 mm long. Keel 4 mm long; blades oblique-elliptic, with widely curved lower edge and slightly curved upper edge, subacute at the apex, 2.5 X 1.5 mm; auricle indistinct, claws 1.5 mm long, connate for nearly the entire length. Stamen tube obliquely cut at the mouth. Ovary subsessile, ovoid, glabrous; style short. Legume unknown. Astragalus barclayanus differs from A. chateri Vassilczenko by the indumentum of white and black hairs 0.2— 0.3(— 0.5) mm long, in the inflores¬ cence of black hairs up to 0.7 mm long (vs. of only white hairs 0.3— 0.8 mm long), by stipules 3—4 (vs. 4—6) mm long, shortly connate or the upper ones free from each other (vs. all connate for half of the length), leaflets glabrous (vs. sparsely to loosely hairy) on upper side, bracts very narrowly trian¬ gular, 2.5—3 mm long (vs. ovate-acuminate, 3-5 mm long), calyx 4 (vs. 5—6) mm long, covered with ascending to spreading (vs. subappressed) hairs, standard 5—6 (vs. 7— 8) mm long, ovaries glabrous (vs. hairy). Astragalus goluhnjensis Podlech & L. R. Xu, sp. nov. TYPE: [China.] Prov. Xizang [Qinghai ?]: Kam [Qamdo], river Kam, river Goluboj, near Nry-tschgo, 3590 m, 25 July 1900, Ladygin 358 (holotype, LE). Differt ab A. densifloro planta pilis 0.2— 0.3 mm (nee 0.4—0.5 mm) longis obtecta, caulibus ad f8 cm (nec 2- 5(— 7) cm) longis, sparsissime pilis albis nigrisque (nec laxe ad densiuscule pilis albis) obteetis, dentibus calycis anguste triangularibus 1—1.5 mm longis (nec subulatis, 1.5-2 mm longis), petalis in siceo albidis (nec pallide violaceis), vexillo ca. 8 mm longo et 5 mm lato (nec 6—7 mm longo et 3—3.5 mm lato), alarum laminis apice oblique emarginate (nec obtusa), ovario glabro (nec piloso). Plants ca. 25 cm tall, covered with appressed hairs 0.2— 0.3 mm long. Caudex up to 15 mm thick, branched. Stems up to 18 cm long, slender, pros¬ trate to ascending or erect, angular-sulcate, fur¬ nished with scattered appressed white and black hairs. Stipules narrowly triangular, 3—4 mm long, very shortly adnate to the petiole, shortly vaginate- connate behind the stem, with scattered black hairs, at the margins with minute sesssile glands. Leaves 2.5—4 cm long, sessile; rachis distinctly grooved on upper side, loosely covered with ap¬ pressed white hairs. Leaflets in 5 or 6 pairs, nar¬ rowly elliptic, 6—14 X 1—3 mm, obtuse to minutely emarginate at the apex, on upper side glabrous, on underside sparsely to loosely hairy, nearly always with uprolled margins or folded, often curved. Pe¬ duncle 3-9 cm long, sulcate, sparsely but below the raceme more densely white and predominantly black hairy. Raceme capitate to somewhat elongat¬ ed. densely many-flowered. Bracts whitish mem¬ branous, narrowly triangular, 2-3 mm long, with subappressed to ascending predominantly black hairs. Flowers nearly sessile. Calyx ca. 3 mm long, campanulate, rather densely to densely covered with subappressed to slightly ascending predomi¬ nantly black hairs; teeth narrowly triangular, 1 — 1.5 mm long. Petals in dry state whitish. Standard ca. 8 mm long; blade ca. 5 mm wide, widely elliptic, slightly emarginate at the apex, at the base nar¬ rowed without distinct claw. Wings ca. 7 mm long; blades narrowly oblong, obliquely laterally incised at the apex, 4.5 X 1.2 mm; auricle ca. 0.5 mm long, claw 2 mm long. Keel 5 mm long; blades obliquely obovate, with lower edge rectangularly curved in distal part and upper edge straight, subobtuse at the apex, 3 X 1 .5 mm; auricle ca. 0.3 mm long, claw 2 mm long. Stamen tube truncate at the mouth. Ovary shortly stipitate, glabrous. Legume unknown. Astragalus golubojensis differs from A. densiflorus Karelin & Kirilow by hairs 0.2— 0.3 (vs. 0.4— 0.5) mm long, the stems up to 18 (vs. 2— 5(— 7)) cm long, covered very sparsely with white and black hairs (vs. loosely to rather densely with only white hairs), petals whitish (vs. pale violet) when dry, standard ca. 8X5 mm (vs. 6-7 X 3—3.5 mm), limbs of Volume 14, Number 2 2004 Podlech & Xu Astragalus from China 219 wings obliquely emarginate at the tip (vs. obtuse) and the ovaries glabrous (vs. hairy). Astragalus mieheorum Podlech & L. R. Xu. sp. nov. TYPE: China. Prov. Qinghai: Kunlun Shan, Kunlun Shankou, SW of Golmud, 4690 m. 35°41'N, 94°03'E, 2 Aug. 1993, G. & S. Miehe 9392/OH (holotype, MSB). Differt ah A. densifloro caule, rachidibus pedunculisque pilis oblique patentibus ad patentibus (nec appressis) 0.5— 0.8 mm longis (nec 0.4— 0.5 mm longis) obtectis, foliolis subtus pilis ascendentibus (nec appressis) obtectis, calyce 5—6 (nec 3—5) mm longo, vexillo late ovato, 8—9.5 mm longo et 5-6.5 mm lato (nec elliptica, 6—7 mm longa et ad 3.5 mm lata). Plants 7—12 cm tall. Caudex with ± elongated, slender stolons partly subterranean, partly aerial. Stems 1—2 cm long, slender, prostrate to ascending, densely covered with ascending to spreading, white or black and white hairs 0. 5-0.8 mm long. Stipules greenish, narrowly triangular, 3—6 mm long, adnate to the petiole for ca. 1 mm. the lower ones shortly vaginate-connate behind the stem, the upper ones nearly free from each other, sparsely predominantly black hairy, at the base of margins with minute ses¬ sile glands. Leaves 2— 4(— 5.5) cm long; petiole 0.5— 1 cm long, with the rachis loosely to rather densely hairy like the stem. Leaflets in (3)4 to 6 pairs, nar¬ rowly elliptic, 5—14 X 1.5— 4.5 nun. obtuse to acute at the apex, glabrous on upper side, on underside rather densely covered with tangled, ascending white hairs up to 1 mm long. Peduncle 2^4 cm long, densely covered with ascending to spreading, tangled white or black and white, sometimes only black hairs up to 1 mm long. Raceme ovate, 1.5— 3 cm long, densely many flowered. Bracts white- membranous, linear-acute, 4-6 mm long, black hairy. Flowers nearly sessile. Calyx 5—6 mm long, eampanulate, ± densely covered with ascending to nearly spreading, tangled black hairs up to 1 mm long; teeth subulate, ( 1 .5 — )2 — 3.5 mm long. Petals violet, in dry state often pale yellowish brown. Stan¬ dard 8—9.5 mm long; blade w idely ovate, 5-6.5 mm wide, widely retuse to slightly emarginate at the apex, at the base subabruptly narrowed into the cuneate claw'. Wings 6.5— 7(— 8) mm long; blades narrowly oblong, gradually widened toward the truncate-incised apex, 4—4.5 X 2—2.2 mm; auricle minute, claw 2.5—3 mm long. Keel 5.5—6 mm long; blades oblique-obovate, with widelv nearly rectan¬ gular-curved lower edge and nearly straight upper edge, subacute at the apex, ca. 3X2 mm; auricle minute, claws 2.5—3 mm long, connate for nearly the entire length. Stamen tube truncate at the mouth. Ovary sessile to subsessile. widely ovate to nearly globular, white hairy; style thick, short, gla¬ brous. Legume sessile, globose, ca. 4 mm thick, rounded at the apex without beak but with the per¬ sistent glabrous style, bilocular; valves densely cov¬ ered with ascending to spreading, tangled, partly straight partly curly white and black hairs up to 1 mm long. Seeds probably two. Astragalus mieheorum differs from A. densiflorus Karelin & KirilowT by the stem, rachis, and pedun¬ cle covered with ascending to spreading hairs 0.5— 0.8 mm long (vs. by appressed hairs 0.4— 0.5 mm long), by leaflets covered on the underside with as¬ cending (vs. appressed) hairs, by the calyx 5—6 (vs. 3—5) mm long, and by the standard widely ovate, 8-9.5 mm long and 5—6.5 mm wide (vs. elliptic, 6- 7 mm long and up to 3.5 mm wide). Etymology. Named alter Prof. (1. Miehe and Mrs. S. Miehe, University of Gottingen, explorers of the vegetation of Tibet. Paratypes. CHINA. Qinghai: Kunlun Shan, Kunlun Shankou, SVi of Golmud, 4870 m, 35°40'N, 94°03'E. 3 Aug. 1993, G. & S. Miehe 9394/06 (MSB); Kunlun Shan, E of Kunlun Shankou, SVi of Golmud, 4820 in. 35°41'N, 94°14'E, 1 Aug. 1993, G. & S. Miehe 9374/14 (MSB); NE Tibet, Northeastern Plateau along Golmud-Lhasa highway, Kunlun Shankou-Qumar Heyan, 4570 m, 35°30'N, 93°45'E, 3 Aug. 1993, G. & S. Miehe 9403/12 (MSB); between Madoi and Bayanka La. 4450 m. 34°20'N, 97°55'E, 3 Aug. 1998, G. Miehe, ./. Q. Lm & Sonamco 98-35221 (MSB). Astragalus naiislianieus Podlech & L. R. Xu, sp. nov. TYPE: China. Prov. Qinghai: Qinghai Nan Shan, Caka Basin, 36°40'N. 98°45'E. 3520 m, 27 July 1993, G. & S. Miehe 9329/07 (holo¬ type. MSB). Differt ah A longiracemoso stipulis 2.5—3 (nec 3-7) mm longis, foliis 2-4.5 (nec 3-8) cm longis, 4—6 (nec 6-9) jugis, foliolis 4—8 X 1—3 (nec 9—27 X 2—7) mm, pedun- culo mere albi-piloso (nec in parte superiore saepissime mere nigri-piloso), axi inflorescentiae albi-piloso (nec praecipue nigri-piloso), vexillo ca. 7 (nec 8—10) mm longo. Plants tall, up to 30 cm or more, basal parts unknown. Stem up to 30 cm long, slender, loosely to rather densely covered with irregularly spreading white hairs and sometimes below the nodes with few black hairs 0.15—0.3 mm long. Stipules green¬ ish, narrowly triangular, 2.5—3 mm long, free from petiole, loosely to rather densely covered with as¬ cending to nearly spreading white, at the base also black hairs, at the base of margins with minute, sessile glands. Leaves 2—4.5 cm long; petiole 0.3— I cm long, together with the rachis white hairy like the stem. Leaflets in 4 to 6 pairs, narrowly elliptic to narrowly ovate, 4—8 X 1—3 mm, rounded to slightly emarginate at the apex, on upper side gla- 220 Novon brous, on underside rather densely covered vvilh subappressed to ascending hairs up to 0.3 mm long, mostly folded. Peduncles 1-6 cm long, white hairy like the stem. Racemes at anthesis short, densely many-flowered, distinctly elongating in fruit; axis white hairy. Bracts whitish membranous, narrowly triangular, 2-2.5 mm long, sparsely black hairy. Pedicels 0.3-0. 5 mm long, black hairy. Calyx 3 mm long, campanulate, obliquely cut at the mouth, rather densely covered with subappressed, black and white hairs up to 0.5 mm long; teeth narrowly triangular, 1 — 1.5 mm long, densely white hairy on inner side. Petals violet with darker tip of keel. Standard ea. 7 mm long; blade ca. 4.5 mm wide, widely elliptic, emarginate at the apex, gradually shortly narrowed at the base without a distinct claw. Wings ca. 5.5 mm long; blades obovate, widened toward the rounded apex, 3.5 X 2 mm; auricle 0.5 mm long, claw ca. 2 mm long, curved. Keel ca. 4 mm long; blades half-circular, with widely curved lower edge and straight upper edge, subacute at the apex, 2.5 X 1.5 mm; auricle minute, acute, claws ca. 1 .5 mm long, connate for nearly the whole length. Stamen tube ± truncate at the mouth. Ovary sessile, nearly globular, glabrous; style thick, short. Legumes unknown. Astragalus nanshanicus differs from A. longira- cemosus IN. Ulziykhutag by stipules 2.5-3 (vs. 3—7) mm long, leaves 2-4.5 (vs. 3-8) cm long, leaflets in 4 to 6 (vs. 6 to 9) pairs, 4—8 X 1—3 (vs. 9—27 X 2—7) mm, peduncles only white hairy (vs. in up¬ per part mostly merely black hairs), inflorescence axis white (vs. predominantly black) hairy, standard ca. 7 (vs. 8—10) mm long. Astragalus pseiidujagnobieiis Podlech & L. R. Xu, sp. nov. TYPE: China. Kaschgaria, Ugu, Ulug-tuz, valley of river Tscharlym, 28 June 1909, Divanogorskaja 284 (holotype, LE). Differt ah A. jagnobico stipulis petiolo vix adnatis, bas- alibus alte vaginanti-connatis (nec petiolo per 0.5-1 mm adnatis, inter se liberis), foliolis 4—6 (nee 6-9) jugis, vex- illo ca. 10 mm longo et 5 mm lata (nec 5—6 mm longo et 2. 2-2. 5 mm lata), lamina alarum obovata, 4.5-5 X 2 mm (nec anguste oblonga, ca. 3X2 mm). Plants up to 25 cm tall, with appressed to sub¬ appressed hairs 0.1— 0.3 mm long. Caudex up to 15 mm diam., with a pluricipital root-crown. Stems several, prostrate to ascending, slender, striate-sul- cate, sparsely to loosely covered with white, below the nodes also with black hairs. Stipules greenish, ovate to triangular, white and black hairy, nearly free from the petiole, the lower ones highly connate behind the stem, the middle and upper ones free from each other. Leaves 4-7 cm long; petiole 1-2.5 cm long, like the raehis slender, loosely covered with white and interspersed black hairs. Leaflets in 4 to 6 pairs, narrowly elliptic to elliptic, 5-12 X 3—6 mm, rounded to distinctly retuse at the apex, on upper side glabrous, on underside loosely to rather densely appressed white hairy. Peduncles 6— 10 cm long, slender, ascending-curved, striate-sul- cate, loosely covered in lower part with merely white hairs, in upper part also with increasing black hairs. Raceme at first globose-ovoid, soon elongating and up to 4 cm long, many-flowered. Bracts whitish membranous, narrowly triangular, 1.5— 2 mm long, black hair)'. Pedicels ca. 0.5 mm long, densely black hairy. Calyx 3-4 mm long, cam¬ panulate, obliquely cut at the mouth, loosely to rather densely covered with black and few white hairs; teeth narrowly triangular. 1 .2-1.5 mm long. Petals in dry state whitish to slightly pale-violet- suffused, standard sometimes with pale violet lon¬ gitudinal nerves, keel in upper pari mostly violet. Standard ea. 10 mm long; blade slightly obovate, ca. 5 mm wide, slightly emarginate at the apex, gradually narrowed at the base. Wings 7 mm long; blades obovate, widely rounded at the apex, 4.5-5 X 2 mm; auricle short, claw 2.5 mm long. Keel 5 mm long; blades obliquely obovate, lower edge curved in distal part and upper edge ± straight, subobtuse at the apex, 3.5 X 2 mm; auricle indis¬ tinct, c law ca. 1.5 mm long. Ovary ovoid. Legumes unknown. Astragalus pseudojagnobicus differs from A. jag- nobicus Lipsky by the stipules scarcely adnate to the petiole, the lower ones vaginate-connate high up (vs. adnate to the petiole for 0.5—1 mm, free from each other), leaflets in 4 to 6 (vs. 6 to 9) pairs, standard ca. 10 X 5 (vs. 5—6 X 2. 2-2. 5) mm, limbs of wings narrowly obovate, 4.5—5 X 2 mm (vs. nar¬ rowly oblong, ca. 3 X 2 mm). Astragalus sagastaigolensis N. Ulziykhutag ex Podlech & L. R. Xu, sp. nov. TYPE: [China. Prov. Qinghai:] Dzhungaria, Sagastaigol, 2740 m, 5 Sep. 1879, A. Regel (holotype, LE). Differt ab A. imetensi stipulis inter se liberis (nec fo- liorum inferiorum alte vaginati-connatis) foliolis 6-9-juga (nec 5— 6-juga), bracteis 4—5 (nec 3) mm longis, vexillo 8.5— 9.5 (nec 6—7) mm longo, lamina alarum obtriangulari apice distincte emarginata, ca. 2.5 mm lata (nee anguste oblonga apice obtusa, ca. 1.5 mm lata). Plants ca. 15 cm tall. Caudex strong, up to 15 mm thick, with a pluricipital root-crown, with short branches. Stems many, up to 10 cm long, prostrate to ascending or erect, rather densely covered with appressed to subappressed white hairs 0.4—0. 6 mm long, with some black hairs mixed in especially at Volume 14, Number 2 2004 Podlech & Xu Astragalus from China 221 the nodes, glabrescent with age. Stipules whitish to slightly greenish, 4—6 mm long, free from the pet¬ iole and from each other, sparsely to loosely cov¬ ered with suhappressed, predominantly black hairs, at the margins with minute, sessile glands. Leaves 3— 6 cm long, subsessile; rachis hairy like the stem. Leaflets in (5)6 to 9 pairs, narrowly elliptic, 8—15 X 2-5 mm, rounded to narrowly obtuse at the apex, on upper side glabrous, on underside rather dense¬ ly covered with appressed white hairs 0.3— 0.5 mm long. Peduncle 2.5—5 cm long, slightly angular-sul- cate, white hairy like the stem, toward the raceme with increasing black hairs. Racemes 2—3 cm long, densely many-flowered; axis with ascending black hairs. Bracts whitish membranous, narrowly trian¬ gular, acuminate, 4—5 mm long, with subappressed to ascending, predominantly black hairs. Pedicels ca. 1 mm long, recurved, black hairy. Calyx 4^4.5 mm long, campanulate, loosely to rather densely covered with subappressed to ascending black hairs ca. 0.5 mm long; teeth subulate, ca. 2 mm long, black hairy on inner side. Petals whitish when dry. Standard 8.5— 9.5 mm long; blade elliptic, ca. 4 mm wide, rounded at the apex, at the base sub- abruptly narrowed into the short claw. Wings ca. 7 mm long; blades obtriangular, distinctly dilated to the unequally widely emarginate apex, 4.5 X 2.5 mm; auricle ca. 0.3 mm long, claw 2 mm long. Keel 5 mm long; blades oblique-elliptic, with widely rectangular-curved lower edge and slightly curved upper edge, subobtuse at the apex, ca. 3X2 mm; auricle minute, claw 2 mm long. Ovary shortly stip- itate, densely white hairy; style glabrous. Legumes sessile, globose, ca. 3 mm diam., rounded ventrally, deeply grooved dorsally, rounded at the apex, beak¬ less, bilocular; valves pale brownish, rather densely covered with subappressed to slightly ascending while and black hairs up to 0.8 mm long. Seeds one in each locule, 2—2.5 mm long, pale brownish, smooth. Astragalus sagastaigolensis differs from A. ime- tensis Borissova by the stipules all free from each other (vs. those of lower leaves vaginate-connate high up), leaflets in 6 to 9 (vs. 5 or 6) pairs, bracts 4— 5 (vs. 3) mm long, standard 8.5— 9.5 (vs. 6—7) mm long, limbs of wings obtriangular, distinctly emarginate, ca. 2.5 mm wide (vs. narrowly oblong, obtuse, ca. 1.5 mm wide). Astragalus L. sect. Cenantrum Bunge, Mem. Acad. Imp. Sci. Saint Petersburg 11(16): 23. 1868. TYPE: Astragalus frigidus (L.) A. Gray (lectotype, designated by Podlech (1990: 468)). Astragalus purpurinus (Y. C. Ho) Podlech & L. R. Xu, stat. nov. Based on Astragalus mem- branaceus Bunge var. purpurinus Y. C. Ho, Bull. Bot. Lab. North-East. Forest. Inst. 1980(8): 54. 1880. TYPE: [China.] Gansu: Xiaho Xian [Hsiahohsien], 2600 m, 1 July 1937, T. P. Wang 6925 (holotype, WUK). Because of the short original diagnosis of this taxon a complete description is given here. Astragalus purpurinus differs from A. mongholi- cus Bunge by the strong stem up to 8 (vs. 2.5—3) mm diam., shorter (4(-5) vs. 5—9 mm long) and more densely hairy calyx with shorter teeth (0.3-1 vs. 0.5— 1.5 mm long), purple petals and legumes with a stipe only 4—5 (vs. 6—15) mm long. Plants at least 60—70 cm tall, covered with hairs 0.3— 0.5(— 1.5) mm long. Stem erect, at the base up to 8 mm diam., angular-sulcate, glabrous or with scattered spreading hairs up to 1.5 mm long. Stip¬ ules green, very narrowly to narrowly triangular, free, the lower ones 8—10, the upper ones 4—5 mm long, glabrous or ciliate at the margin. Leaves 5-9 cm long, nearly sessile; rachis grooved on upper side, sparsely covered with spreading hairs up to 1 mm long. Leaflets in 3 to 9 pairs, narrowly ovate to narrowly elliptic, rounded to minutely acuminate at the apex, 10—17 X 3—7 mm, on upper side loosely covered with appressed to subappressed hairs 0.3— 0.5 mm long, on underside loosely with appressed to subappressed hairs 0.5—1 mm long. Peduncles 5-13 cm long, sparsely to loosely covered with ap¬ pressed to ± spreading white hairs. Racemes loose¬ ly many-flowered, 4-5 cm long, soon elongating and at least up to 10 cm long; raceme axis black hairy. Bracts green, linear-subulate, 5—8 mm long, ca. 0.5 mm wide, loosely mainly white ciliate. Pedicels 1— 1.5 mm long, later on elongating and up to 3 mm long, with minute black hairs. Bracteoles whitish, linear, 0.5—1 mm long, in upper part of the pedicel. Calyx campanulate, 4(— 5) mm long, slightly gib¬ bous at the base, strongly obliquely cut at the mouth, the tube loosely covered with short black subappressed hairs up to 0.3 mm long and with few long white hairs, at the upper margin and the teeth more densely black hairy; teeth narrowly triangular to triangular, 0.3—1 mm long. Petals lilac or dark purplish at the apex and whitish at the base. Stan¬ dard 13 mm long; blade distinctly recurved, ob- ovate, 7 mm wide, emarginate at the apex, gradu¬ ally and shortly cuneately narrowed at the base. Wings 13 mm long, distinctly bent at the base of the blade; blades narrowly oblong, rounded at the apex, 5 X 1.5 mm; auricle 0.8 mm long, claw 7—8 mm long, straight. Keel 12 mm long; blades strong- 222 No von ly curved-elliptic, with widely and nearly rectan¬ gularly curved lower edge and distinctly concave upper edge, obtuse at the apex, 5X2 mm; auricle ca. 0.3 mm long, claw 7 mm long. Stamen tube obliquely cut at the mouth. Ovary with a stipe ca. 5 mm long, densely white hairy. Legume with a slender stipe 4—5 mm long, narrowly elliptic, acu¬ minate at both ends; valves loosely to densely cov¬ ered with subappressed white and black hairs ca. 0.3 mm long, glabrescent with age and sometimes subglabrous. Ollier specimen seen. CHINA. Gansu: Yuzhong, 2750 m, 8 Aug. 1959, Y. He 5864 (WUK). Astragalus xitaibaieus (k. T. Ku) Podlech & L.- R. Xu. stat. nov. Based on Astragalus mona- delphus Bunge subsp. xitaibaieus K. T. Fu, Aeta Hot. Bor.-Occ. Sin. 1(2): 17. 1981. TYPE: [China.] Shaanxi: Xitaiba Shan, Ding-peng- shan, 3300 m, Fu Kun-tsun 10321 (holotype, WUK). Because of the short original diagnosis of this taxon a complete description is given here. Astragalus xitaibaieus differs from A. monadel- phus by being totally glabrous with the exceptions of the inner side of the calyx teeth and sometimes the bracts (vs. at least hairy on underside of leaf¬ lets, calyx, and legumes) and by the presence of stipel-like entire or fringed, linear to narrowly tri¬ angular emergences 0.5— 2.5 mm long and/or of dis¬ tinctly stalked minute glands at the base of the leaf¬ lets. Plants 12-22 cm tall, totally glabrous with the exception of the inner side ol the calyx teeth and sometimes the bracts. Caudex strong, 15 mm thick, with a pluricipital root-crown. Stems several, as¬ cending to erect, up to 4 mm thick, angular-striate, 10—17 cm long. Stipules greenish membranous with narrowly hyaline margins, ovate, obtuse, 12-15 mm long and up to 10 mm wide, connate at the base, free from the petiole. Leaves 4—8 cm long; petiole 0.7—2 cm long, at the base of the leaflets with sti¬ pel-like entire or fringed, linear to narrowly trian¬ gular emergences 0.5-2. 5 mm long and/or with dis¬ tinctly stalked minute glands. Leaflets in 5 to 7 pairs, ovate to elliptic, 10-18 X 5-9 mm, at the apex rounded and sometimes minutely mucronu- late. Peduncles 4—7.5 cm long. Racemes 8- to 11- fiowered. Bracts narrowly ovate, acute, ca. 10 mm long, sometimes sparsely ciliate at the margins. Pedicels 2—3 mm long. Calyx campanulate, 7.5—9 mm long; teeth subulate, 2.5-3 mm long, hairy on inner side. Petals yellow. Standard ca. 1 1 mm long; blade obovate, 5—6 mm wide, emarginate at the apex, at the base subabruptly contracted into the rather long claw. Wings ca. 1 1 mm long; blades narrowly oblong, obtuse at the apex, ca. 4 X 1.5 mm; auricle short, claw 6.5-7 mm long, keel ca. 10 mm long; blades obliquely elliptic, ca. 3.5 X 2 mm; auricle indistinct, claw ca. 6.5 mm long. Sta¬ men tube closed (monadelphous), slightly obliquely cut at the mouth. Ovary stipitate, hairy. Legumes with a stipe 5—6 mm long, narrowly elliptic, shortly attenuate at both ends, ea. 15 mm long, unilocular; valves hairy. Paratype. CHINA. Shaanxi: Xitaihai Shan. Hun-ren- ping, in summo alpino, 3300 m, 4 July 1959, Yang Jin- xiang 645 (WUK). Astragalus sect. Corethrum Bunge, Mem. Acad. Imp. Sci. Saint Petersbourg 11(16): 98. 1868. T\ PE: Astragalus scoparius C. A. Meyer (lec- totype, designated by Podlech (1990: 471)). Astragalus eliaidamueiisis (S. B. Ho) Podlech & L. R. Xu, stat. nov. Based on Astragalus kro- nenburgii B. Fedtschenko var. ehaidamuensis S. B. Ho, Bull. Bot. Res. North-East. Forest. Inst. 3(1): 42. 1983. TYPE: [China.] Qinghai: Haixi Mongolzu Zangzu, Kazakzu Zizhizhou, Delingha, 3290 m, 21 July 1959. Exped. Gan¬ su et Qinghai Acad. Sin. 80S (holotype, WUk). Because of the very short original diagnosis of this taxon a complete description is given here. Astragalus ehaidamuensis differs from A. kronen- burgii B. Fedtschenko by stipnles triangular, 2 mm long (vs. narrowly triangular, 3-5 mm long), adnate to the petiole for 0.5 (vs. 1—2) mm, densely hairy (vs. subglabrous to loosely hairy), leaflet in up to 4 (vs. in 3 to 6) pairs, peduncles 5-14 (vs. 9-22) cm long, only white (vs. toward the raceme increasingly black) hairy, raceme very short, 3- to 7-flowered (vs. 3-5 cm long, 10- to 16-flowered), bracts 1-2 (vs. 3—4) mm long, calyx 10 (vs. 5-6) mm long, teeth 2.2-3 (vs. 1-2) mm long. Plants ca. 18 cm tall, nearly acaulescent, caes- pitose, covered in vegetative parts with medifixed, appressed white hairs 0.3—1 mm long. Caudex ca. 8 mm diam., at the apex strongly divided w ith short branches, covered with brown bark. Stems branched at the base, parts of the current year, if developed, up to 0.5 cm long, densely hairy. Stip¬ ules ca. 2 mm long, triangular, adnate to the petiole for ca. 0.5 mm, densely hairy. Leaves of sterile shoots 1—2 cm long; petiole 0.5—1 cm long, with 1 or 2 pairs of small leaflets, 2—3 mm long. Leaves of the flowering shoots 3—7 cm long; petiole 1—2 cm long, like the rachis slender, densely hairy. Leaflets in (2)3 to 4 pairs, linear, 10—15 X 1—1.5 mm. Volume 14, Number 2 2004 Podlech & Xu Astragalus from China 223 acute-tipped, on both sides rather densely to dense¬ ly hairy. Peduncles 5-14 cm long, loosely to rather densely white hairy. Racemes short, subumbellate, 3- to 7-How ered. Bracts membranous, narrowly tri¬ angular, 1—2 mm long, sparsely furnished with mostly basifixed white and black hairs. Pedicels ea. 0.5 mm long, white and black hairy. Calyx ca. 10 mm long, tubular, obliquely cut at the mouth, loose¬ ly to rather densely covered with medifixed ap- pressed white hairs 0.5-1 mm long and mostly slightly shorter black hairs; teeth subulate, 2.2-3 mm long. Petals probably yellow. Standard 15-17 mm long, elliptic, 6-7 mm wide, emarginate at the apex, with a very short, indistinct claw. Wings 15 mm long; blades narrowly oblong, at the apex deep¬ ly incised, 7X2 mm; auricle 0.5 mm long, claw 8 mm long. Keel ca. 12 mm long; blades obliquely obovate, with rectangular-curved lower edge, and nearly straight upper edge, subacute at the apex, 5 X 2.5 mm; auricle acute, 0.3 mm long, claw 7 mm long. Stamen tube obliquely cut at the mouth. Ova¬ ry with a stipe ca. 2 mm long, linear, white hairy; style glabrous. Legumes unknown. Astragalus sect. Ilcmipliaea Bunge, Mem. Acad. Imp. Sei. Saint Petersbourg 1.1(16): 20. 1868. TYPE: Astragalus hemiphaca Karelin & Kiri- low. Astragalus nigrodentatus N. Ulziykhutag ex Pod¬ lech & L. R. Xu. sp. nov. TYPE: China. Prov. Xizang: Distr. Kam, Dabo-Shan pass, 18 July 1893, G. N. Potanin (holotype, LE). Differt ab A. alaschano indumento e pil is 0.6— 0.8(— 1) mm (nee 0.1— 0.3 mm) longis consistente, stipulis 6—7 (nec 2-3) mm longis, foliolis anguste ovatis ad anguste ellip- ticis, 5—14 X 2.5—5 mm (nec ellipticis ad obovatis, 3—7 X 2—5 mm), bracteis 4—6 mm (nec 0.8—1 mm) longis, calyce ca. 5 mm (nec 2.5—3 mm) longis. Plants 8-10 cm tall, covered in vegetative parts with flattened, long-acuminate white hairs widest near the base and elsewhere with slender black hairs mixed in. Stems ascending, at the base pale, stolon-like, glabrous, with few7, remote leafless stip¬ ules, in upper part sulcate, loosely to rather densely covered with stiff, tangled, ascending to spreading while, toward the nodes mostly black hairs 0.6— 0.8 nun long. Stipules greenish. 6—7 mm long, narrowly triangular to triangular, free from the petiole, those without developed leaves connate up to the middle, the other ones free from each other, the lower ones glabrous, the upper ones loosely or, especially to¬ ward the margins and tip, densely white and partly also black hairy. Leaves 3—5 cm long; petiole 0.8— 1.5 cm long, distinctly grooved on the upper side, similar to the rachis loosely white hairy like the stem. Leaflets in 6 to 8 pairs, narrowly ovate to narrowly elliptic, 5-14 X 2.5—5 mm, obtuse at the apex, on upper side glabrous but with densely white hairy margins, on underside loosely covered with appressed hairs 0.2— 0.5 mm long. Peduncles 1.5— 3.5 cm long, sulcate, loosely to densely covered with subappressed black hairs 0.3-1 mm long, at the base with some white hairs mixed in. Racemes short, densely many-flowered. Bracts whitish mem¬ branous, narrowly triangular. 4—6 mm long, with predominantly black hairs, at the base with sessile, minute glands. Elowers subsessile. Calyx ca. 5 mm long, campanulate, densely covered with ascending to partly nearly spreading black hairs 0.3— 0.6 mm long; teeth very narrowly triangular, ca. 2 mm long, hairy on inner side. Petals probably violet in life, pale yellowish brown when dry. Standard ca. 9 mm long; blade ca. 4 mm wide, oblong, at the apex widely and deeply incised, at the base cuneately narrowed without distinct claw. Wings ca. 7.5 mm long; blades obtri angular, widest at the widely and deeply incised apex, 4.5 X 2 mm; auricle 0.3— 0.4 mm long, claw 3 mm long, curved. Keel 6.5 mm long; blades oblique-elliptic, with widely curved lower edge and straight upper edge, subacute at the apex, 3.5 X 2 mm; auricle ca. 0.3 mm long, acute, claw ca. 3 mm long. Stamen tube truncate at the mouth. Ovary subsessile, minute, elliptic, glabrous, 2-ovulate. Legumes unknown. Astragalus nigrodentatus differs from A. alas- chanus Bunge by the hairs 0.6— 0.8(-l) (vs. 0.1 — 0.3) mm long, stipules 6-7 (vs. 2—3) mm long, leaf¬ lets narrowly ovate to narrowly elliptic, 5—14 X 2.5— 5 mm (vs. elliptic to obovate, 3—7 X 2—5 mm), bracts 4—6 (vs. 0.8—1) mm long, calyx ca. 5 (vs. 2.5— 3) mm long. This collection was named Astragalus nigroden¬ tatus by Ulziykhutag in the herbarium at St. Pe¬ tersburg (LE) without description. Astragalus sect. Laguropsis Bunge, Mem. Acad. Imp. Sci. Saint Petersbourg 11(16): 137. 1868. TYPE: Astragalus laguroides Pallas (lectotype, designated by Podlech (1990: 479)). Astragalus iiovissiinus Podlech & L. R. Xu. nom. et stat. nov. Based on Astragalus laguroides Pallas var. micranthus S. B. Ho, Bull. Bot. Res. North-East. Forest. Inst. 3(4): 57. 1983 “non A. micranthus Desv., 181 4.” TYPE: |China.| Nei Mongol, Baganmaotasum (patria ignota), 29 May 1931, T. N. Liou 2147 (holotype, WUK). Figure: Ho (1983: 66, fig. 5; as A. la¬ guroides var. micranthus). 224 Novon Because of the short original diagnosis of this taxon a complete description is given here. Plants 8-12 cm tall, acaulescent, with merely white, in vegetative parts distinctly warty, hairs. Caudex slender, simple or with a pluricipital root- crown. Stipules whitish, 3—4 mm long, triangular, nearly free from the petiole, appressed hairy, at the margins with basifixed hairs. Leaves 3-6 cm long; petiole 1-3 cm long, like the rachis slender, loosely to rather densely covered with medifixed, appressed hairs 0.5—1 mm long. Leaflets in 2 to 4 pairs, in the basal leaves often only in 1 or 2 pairs, narrowly elliptic to elliptic, 7-12 X 3-5 mm, acute at the apex, on both sides rather densely to densely cov¬ ered with medifixed, appressed hairs 1.5— 2(— 3) mm long. Peduncles 3.5-6 cm long, slightly angular- sulcate, loosely to rather densely covered w ith med¬ ifixed, appressed hairs 0.5—1 mm long, partly gla- brescent with age. Racemes ovoid, 3—5 cm long, densely many-flowered. Bracts whitish, ca. 2 mm long, narrowly triangular, covered with mostly bas¬ ifixed hairs. Flowers subsessile. Calyx 11-12 mm long, tubular at beginning of anthesis, soon ovoid- inflated, rather densely covered with subbasifixed to basifixed, rigid, spreading, straight white hairs 2-3 mm long; teeth subulate, 3-4 mm long. Stan¬ dard ca. 14 mm long; blade ca. 5 mm wide, elliptic, narrowly triangular toward the slightly emarginate apex, slightly constricted below the middle, at the base obtusely angularly passing into the claw. Wings 13 mm long; blades narrowly oblong, round- tipped, ca. 5 X 1.5 mm; auricle 0.5 mm long, claw 8 mm long. Keel 12 mm long; blades obliquely ob- ovate, with rectangular-curved lower edge and nearly straight upper edge, acute-tipped, 3.5 X 1.5 mm; auricle acute, 0.5 mm long, claw ca. 8 mm long. Stamen tube truncate at the mouth. Ovary ses¬ sile, hairy. Legumes unknown. Astragalus sect. Polyoladus Y. (7 Ho, Bull. Bot. Res. North-East. Forest. Inst. 1(3): 107. 1981. TYPE: Astragalus polycladus Bureau & Fran- chet. The following species was proposed as a new species in Sendtnera 7: 198. 2001, but unfortu¬ nately without a Latin diagnosis. Astragalus tihetieola Podlech & L. R. Xu, sp. nov. TYPE: China. Prov. Xizang: S Tibet, Cha- yul Dzong, Loro Chu, 23 Apr. 1936, F. Ludlow & G. Sherrifj 1326 (holotype, BM). Differt ab A. monbeigii caulibus pilis 0.4— 0.7 mm (nec 0.1— 0.3 mm) longis obtectis, calyce 4—5 mm longo tlen- tibus ejus 1.5—2 mm longis (nec 5—6 mm longo dentibus ejus 2.5—4 mm longis), vexillo ca. 6 mm longo et 5 mm lato, lamina transverse elliptica (nec ca. 10.5 mm longo, lamina late obovate), ovario sessile (nec stipite 1.5—2 mm longo suffulto). Plants 15—25 cm tall. Stems 10-17 cm long, branched at the base, ascending to erect, striate- sulcate, sparsely to loosely covered with appressed to subappressed white, at the nodes also black, hairs 0.4— 0.7 mm long. Stipules green, 6—9 mm long, ovate-acuminate to narrowly triangular, free from the petiole, behind the stem high up vaginate- connate, sparsely to loosely covered with appressed white and sometimes also with some black hairs. Leaves 6—12 cm long; petiole 1—3 cm long, like the rachis loosely covered with appressed white hairs 0.2-0. 5 mm long. Leaflets in 9 to 1 1 pairs, narrowly elliptic to elliptic, 7—17 X 3-6 mm, rounded to slightly emarginate at the apex, on upper side gla¬ brous, on underside sparsely and mainly at the midrib furnished with appressed white hairs 0.5—1 mm long. Peduncles 5-6 cm long, hairy like the stem but in upper part with predominantly blac k hairs. Racemes ovoid, rather loosely 7- to 15-flow- ered. Bracts whitish, linear-acute, 2.5—3 mm long, predominantly black hairy. Pedicels ca. 1 mm long, black hairy. Calyx eampanulate, 4—5 mm long, slightly oblique at the base, slightly obliquely cut at the mouth, rather densely covered with ap¬ pressed black hairs 0.2— 0.3 mm long and with few¬ er, slightly longer white hairs; teeth subulate, 1.5— 2 mm long. Petals mauve with whitish center, keel purplish mauve. Standard ca. 6 mm long; blade 5 mm wide, transverse-elliptic, deeply incised at the apex, at the base subabruptly contracted into the very short claw. Wings ca. 6 mm long; blades ob¬ long, rounded at the apex, 3.5 X 2 mm; auricle wide, ca. 0.5 mm long, claw 2.5 mm long. Keel 5 mm long; blades oblique-elliptic, rounded at the apex, 3X2 mm; claw 2 mm long. Stamen tube truncate at the mouth. Ovary sessile, white-hairy; style glabrous. Legumes unknown. Astragalus tihetieola differs from A. monbeigii N. D. Simpson by stems covered with hairs 0.4— 0.7 (vs. 0.1— 0.3) mm long, calyx 4—5 mm long with teeth 1.5—2 mm long (vs. 5—6 mm long with teeth 2.5— 4 mm long), standard ca. 6 X 5 mm, the limb transversely elliptic (vs. ca. 10.5 mm long, the limb widely obovate), ovaries sessile (vs. with a stipe 1 .5— 2 mm long). Astragalus sect. Skythropos N. D. Simpson, Notes Roy. Bot. Card. Edinburgh 8: 255. 1915. TYPE: Astragalus skythropos Bunge (lectotype, designated by Podlech (1990: 488)). Volume 14, Number 2 2004 Podlech & Xu Astragalus from China 225 Astragalus griersonii Podlech, sp. nov. TYPE: Bhutan, Me La, 27°58'N, 91°37'E, 14,000 ft., 7 Aug. 1933, F. Ludlow & G. Sherrijf 429 fho- lotype, BM). Differt ab A. kongrensi plantis 20—25 (nec 6—14) cm altis, in partibus vegetativis pilis praecipue nigris (nec mere albis) obtectis, calyce ca. 8 (nec 9—12.5) mm longo, campanulati-tubuloso (nec tubuloso), dentibus 3—4 (nec 4—7) mm longis, petalis brevioribus. Plants 20—25 cm tall, with subappressed to ± spreading, thin, long white and black hairs. Caudex with rather long stolons up to 10 cm long, furnished with leafless, pale, scale-like stipules. Stems erect, short, terete to angular-sulcate, loosely to rather densely covered with subappressed to mostly ± spreading, predominantly black hairs 0.3—1 mm long. Stipules 8—12 mm long, dirty whitish to slightly greenish, ovate-acuminate, free from the petiole and from each other, sparsely to loosely cov¬ ered with tangled, subappressed to ascending white and black hairs, sometimes subglabrous. Leaves 9— 16 cm long; petiole 3—5 cm long, like the rachis sulcate. black hairy like the stem. Leaflets in 9 to 1 1 pairs, elliptic to nearly orbicular, 5—12 X 4—9 mm, emarginate at the apex, on upper side sparsely, on underside loosely covered with subappressed white or white and black hairs 0.5—1 mm long. Pe¬ duncle 10—12 cm long, in lower part distinctly, in upper part slightly angular-sulcate, rather densely covered with ascending to spreading blackish hairs 0.5— 1.5 mm long. Racemes short, often globose, ca. 3 cm long, rather densely 9- to 12-flowered; axis black villous. Bracts whitish, narrowly oblong to narrowly elliptic, 5-6 mm long, 1—1.5 mm wide, on outer side rather densely covered with tangled black hairs up to 1.5 mm long, on inner side (upper side) sparsely hairy. Pedicels 1 .5—2 mm long, very densely black hairy. Bracteoles absent. Calyx ca. 8 mm long, campanulate-tubular, truncate at the mouth, rather densely to densely covered with sub¬ appressed to ascending black hairs 0.3-1 mm long; teeth narrowly triangular, 3—4 mm long, sparsely hairy on inner side. Petals purple to bluish white. Standard ca. 16 mm long; blade ca. 7 mm wide, widely rectangular, distinctly emarginate at the wide apex, at the base with a cuneate claw of equal length. Wings ca. 15 mm long; blades narrowly ob¬ long, rounded at the apex, 8X3 mm; auricle nar¬ row, 2.5—3 mm long, claw ca. 7 mm long. Keel ca. 16 mm long; blades oblique-obovate, with lower edge rectangularly curved in distal part and upper edge nearly straight, subobtuse at the apex, ca. 8.5 X 4 mm; auricle ca. 1 mm long, claw ca. 8 mm long. Staminal tube ± truncate at the mouth. Ovary with a stipe ca. 4 mm long, densely covered with subappressed white and predominantly black hairs ca. 1 mm long; style thickish, hairy in lower half. Legumes unknown. Astragalus griersonii differs from A. kongrensis Baker by being 20—25 (vs. 6—14) cm tall, in veg¬ etative parts with predominantly black (vs. merely white) hairs, calyx ca. 8 (vs. 9-12.5) mm long, with teeth 3—4 (vs. 4—7) mm long, standard ca. 16 (vs. 16-21) mm long. Astragalus sect. Trachycercis Bunge, Mem. Acad. Imp. Sci. Saint Petersbourg I 1(16); 1 14. 1868. TYPE: Astragalus humilis M. Bieber- stein (lectotype, designated by Podlech (1990: 490)). Astragalus lang-ranii Podlech, nom. et slat. nov. Based on Astragalus salsugineus Karelin & Kirilow var. multijugus S. B. Ho, Bull. Bot. Res. North-East. Forest. Inst. 3(1): 52. 1983, non Astragalus multijugus DC. TY PL: [China.] Ningxia: Yinchuan, prope viam ferream, 1200 m, 14 May 1959, Y. C. Ho 2135 (holotype, WUK; isotvpe, PE). Because of the short original diagnosis of this taxon a complete description is given here. Astragalus lang-ranii is near to A. alaschanensis H. C. Fu, but differs by smaller leaflets (3-4 X 1.3 mm vs. 5—15 X 3-7 mm), shorter bracts (4—6 mm vs. 10—15 mm long), shorter calyx (12—13 mm vs. 15—18 mm long; teeth 2.5—4 mm vs. 5—9 mm long). Plants ca. 10 cm tall, shortly caulescent, densely covered with symmetrically to asymmetrically bi¬ furcate, spreading white hairs, the branches up to 1 mm long. Caudex slender, simple or shortly branched. Stems up to 5 cm long, densely tomen- tose. Stipules whitish, adnate to the petiole for 2— 4 mm, the lower triangular, 3—4 mm long, the upper narrowly triangular-acuminate 5-7 mm long, all densely hairy. Leaves 8—12 cm long; petiole 3—4 cm long, like the rachis slender, flexuous or curved, rather densely hairy. Leaflets in 1 1 to 13 pairs, el¬ liptic, 3-6 X 1.5—3 mm, rounded to acute at the apex, on both sides covered with tangled, ± spreading hairs. Peduncles 1-1.5 cm long, hairy like the stem. Raceme loosely 4- to 6-flowered. Bracts whitish. 4—6 mm long, ovate-acuminate, loosely white-hairy, partly w ith basifixed hairs. Ped¬ icels 2—3 mm long, hairy. Calyx 12-13 mm long, tubular, obliquely truncate at the mouth, densely covered with short to long spreading, tangled hairs; teeth subulate, 2.5—4 mm long. Petals pale purple. Standard 20—22 mm long; blade ca. 7 mm wide, narrowly ovate, ± rounded at the apex, very shortly 226 Novon narrowed at the base. Wings ca. 19 mm long; blades narrowly obovate, obliquely slightly incised at the apex, 10 X 3 mm; auricle 1 mm long, claw 9 mm long. Keel 15 mm long; blades obliquely el¬ liptic-triangular, with widely curved lower edge and nearly straight upper edge, obtuse at the apex, 6 X 3 mm; auricle short, claw 9 mm long. Etymology. Named after Prof. Xu Lang-Ran, co-worker on the Flora of China , who has recog¬ nized the distinctness of this species. The hyphen in the Latin name lang-ranii must be retained be¬ cause two forenames are combined. Literature Cited Ho, S. B. 1983. Trascursores Florae Astragalorum Sinen- sium. Bull. Bot. lies. North-East. Forest. Inst. 3(4): 51 — 70. Podlech, I). 1990. Die Typifizierung der altweltlielien Sek- tionen der Gattung Astragalus L. (Leguminosae). Mitt. Bot. Staatss. Mtlnehen 29: 461—494. A New Species of Ledermanniella (Podostemaceae) from Cameroon John ./. Schenk Department of Botany and Plant Pathology, Oregon State University, 2082 Cordley Hall, Corvallis, Oregon 97331-2902, U.S.A. Author for correspondence; present address: School of Biological Sciences, Washington State University, Pullman, Washington 99164-4236, U.S.A. john_sch@mail.wsu.edu Duncan W. Thomas Department of Forest Science, Oregon State University, Corvallis, Oregon 97331-2902, U.S.A. DuncanWT@aol.com Abstract. Ledermanniella prasina J. Schenk & I). W. Thomas in the family Podostemaceae, sub¬ genus Ledermanniella, is proposed as a new spe¬ cies. It is known only from its type, collected at a single waterfall in Cameroon. The ecology, mor¬ phology. and variation among individuals are dis¬ cussed. The new species is compared to the mor¬ phologically similar L. bifurcata, L. bowlingii, L. guineensis, L. jaegeri, L. minutissima, and L. mon- andra. Illustrations and a key to the monostaminate species in the subgenus are provided. Key words: aquatic plants, Cameroon, Leder¬ manniella, narrow endemic, Podostemaceae. With its center of diversity in Cameroon, Leder¬ manniella Engler is the second largest genus in Podostemaceae, consisting of about 44 species, which are all endemic to tropical Africa (Cook, 1996; Gusset. 1983, 1984). As defined by Cusset (1974), Ledermanniella is distinguished from other genera in the family through the combination of the following characters: the presence of alternate or whorled leaves, solitary or occasionally clustered flowers inverted within a closed spathellum, and an ovoid to ellipsoidal capsule with eight ribs. This broader classification ignored pollen and stamen number traits previously used and resulted in the inclusion of Inversodicraeia Engler and Monati- driella Engler within Ledermanniella (Cusset. 1974). Subsequently, Cusset divided the genus into two subgenera, Phyllosoma and Ledermanniella, distinguished by the presence or absence of scale¬ like leaves, respectively (Cusset. 1983, 1984). The new taxon is clearly a member of Podoste¬ maceae due to its semi-aquatic habitat, thallus-like base (root), the presence of spathella, flowers that consist of two tepals, a syncarpous superior gynoe- cium with two styles, a single anther, and capsular fruit. The new taxon is placed in Ledermanniella based on its alternate leaves, solitary to clustered flowers each inverted within a closed spathellum, two filamentous tepals, flowers borne along shoots above a thalloid-root, and ellipsoidal capsules. It is placed in the subgenus Ledermanniella based on the absence of scale-like leaves and the presence of elongated leaves. The new species is known only from the type collection, which consists of approx¬ imately six pressed individuals along with 25 stems and shoots (which were separated from each other and/or other individuals at the time of collection), and two individuals along with 13 stems and shoots preserved in alcohol. Both collections have buds, flowers, and fruits in all stages of development. Taxonomy Ledermanniella prasina J. Schenk & 1). W. Thomas, sp. nov. TYPE: Cameroon. Southwest Province: 14 km NE of Mundemba, base of a large waterfall in the Mana River, between the villages of Miangwe 2 and Meta, 300 m, 5°02.4'N, 9°01.0'E, 01 Dec. 1998. Duncan Thomas 11550 (holotype, MO; isotypes, K. LBV, MO (preserved), NY, OSC, P, SC A, WAG, YA). Figure 1. Speciebus monostaminiferis subgeneris Ledermanniel- lae similis sed distinguibilis seminibus smaragdinis et flo- ribus (tepalis anthera gynoeeioque includentibus) majori- bus. Semi-aquatic herbs from the spray-zone of a wa¬ terfall; thalloid-root disk-shaped, holdfast attaching plant to rock substratum; plants erect, 9—13 cm tall; main stem arising from thalloid-root, 1.0—32.0 mm long, 1. 0-2.0 mm wide, erect; vegetative shoots arising from main stems, filiform, dichotomously branched, up to 7 cm long; fertile shoots arising from main stem, filiform, dichotomously branched, bearing reproductive structures; leaves up to 4 cm Novon 14: 227-232. 2004. 228 Novon Figure 1. Ledermanniella prasina J. Schenk & 1). W. Thomas, based on I). W. Thomas 11550. — A. Detailed habit of a single main stem showing the structure of the main stem (X'), fertile shoots (X"), vegetative shoots (Xv), leaves (L), and a thalloid-root (R; note that the apical portion and other mains stems are omitted for clarity). — B. A segment of the primary and fertile shoots showing the details of a flowering cluster, unruptured spathella (Sp) with leaves subtending spathella; also seen are stipules (St) located on the intrapeliolar scales (Is). — C. Ruptured spathella, with emerging, inverted flower (left) and fully emerged flower (right), along with tepals (P; only one of two shown). long, 0.1— 0.5 mm wide, dichotomously branched, solitary to verticillate below spathella, basal portion scale (intrapeliolar scale) with stipules, apical por¬ tion elongating from center of scale, filiform, gla¬ brous; spathella erect, 3.0— 8.0 mm long before de¬ hiscence, 1.0— 2.2 mm wide, ovate at beginning of development, then elongating to clavate, glabrous, solitary or in clusters (1—6) in the axils or along fertile shoots, apex globose to acute, dehiscence mostly irregular, occasionally bilobed to trilobed; (lowers solitary within spathella, zygomorphic, in¬ verted prior to the opening of the spathella, upright after elongation, pedicel up to 13 mm long, tepals 2, acicular, 0.4 — 1 0.7 mm long, gynoecia 1.5( — 1 .7) mm long, 0.7— 0.9 mm wide, attached to a gyno- phore, unilocular, styles 2, linear. 0.6— 0.8 mm long, stigmas bilobed; stamens 1 (very rarely 2); fila¬ ments ( = andropodium) 1.1— 1.3 mm long at spa- Volume 14, Number 2 2004 Schenk & Thomas Ledermanniella prasina from Cameroon 229 thellum dehiscence, 5.0— 5.3 mm long after anthe- sis, alternate to the tepals; anthers basifixed, 2.0 mm long, 0.5— 0.8 mm wide, longitudinally dehis¬ cent; pollen white, in monads; capsules elliptic, 2.2— 2.7(— 3) mm long, 1.0— 1.2 mm wide, bivalved, one valve persistent after dehiscence, the other ca¬ ducous, ribs 8 per capsule, with 3 ribs per valve and 2 as the marginal sutures; placentas green; seeds bright emerald green, 0.3-0. 4 mm long, 0.125—0.15 mm wide, ca. 25 to 37 per capsule. Ecology Like many Podostemaceae, species of the genus Ledermanniella are sometimes narrowly endemic to a single geographical area (Cook. 1996). From our current knowledge, L. prasina occurs only at a sin¬ gle site, and is therefore narrowly endemic to the Mana River valley in the Rumpi Hills of Cameroon. On the basis of this information, it should be red- data listed as an endangered species according to the criteria established by the International Union for the Conservation of Nature (IUCN). The new species occupies a very unusual and specific hab¬ itat. The waterfall in the Mana River consists of two stages with a total height of approximately 100 m. At the bottom, there is a large, deep pool and a shelf of exposed gneiss bedrock, about 200 m2 in extent; L. prasina occurs only on this shelf. The shelf is above the observed high-water level in the river, but its largely unvegetated state suggests pe¬ riodic scouring, probably during occasional flash floods. The area has a very wet but strongly two- seasonal climate with a total annual rainfall ex¬ ceeding 5000 mm. There is a short dry season De¬ cember to March, with three months averaging less than 100 mm of rain. The first part of the wet sea¬ son is typified by thunderstorms of varying fre¬ quency and duration, sometimes with dry periods of several weeks’ duration. The wet season peaks in August and September, when monthly rainfall may exceed 1000 mm (Chuyong et a!., 2004). This wet, seasonal climate, combined with the rugged, rocky topography of the Rumpi Hills, generates the type of river that is ideal for Podostemaceae, with a rocky bed and large seasonal variation in flow. With the exception of L. prasina, all Podostemaceae reported from Cameroon follow similar phenological patterns in response to this climate. The plants are obligate hydrophytes and grow submerged in fast¬ flowing water during the wet season, attached to the rocks by thalloid-roots. When the water level in the rivers subsides at the beginning of the dry season, the exposed plants flower and fruit rapidly, after which at least the aerial parts die (Cusset, 1987; Thomas, pers. obs.). Several species of Podoste¬ maceae exhibiting this phenology have been col¬ lected from the Mana River, including Lederman¬ niella ledermannii (Engler) C. Cusset, Macropodiella pellucida (Engler) C. Cusset, and Saxicolella flabellata (G. Taylor) C. Cusset (Cusset, 1987). Ledermanniella prasina exhibits a very different habit and phenology, which is unique for the family in Cameroon. It does not grow submerged as most podostemads do, but instead has erect aerial stems and shoots that are kept moist throughout the grow¬ ing period by spray generated from the waterfall. The spray is continuous during the wetter months with high water flow in the river, and it is this spray that supports the growth of the new species. The aerial shoots are intolerant of desiccation and with¬ er rapidly when removed from the moist environ¬ ment. In the dry season, the waterfall is present but much reduced in volume, and little spray reaches the rock shelf that supports the L. prasina popu¬ lation. During this period, the dry rocks appear to be largely devoid of vegetation and we do not yet know if L. prasina is annual, or if it passes the dry season as a dormant thalloid-root. Instead of having the short flowering period typical of Podostemaceae when the water level drops, the flowering and fruit¬ ing of L. prasina are prolonged and take place dur¬ ing the latter part of the growing season, when its fertile branches are produced. During this same pe¬ riod, the other Podostemaceae in the river are sub¬ merged and growing vegetatively, but they do not flower. For several reasons, we do not believe that Led- ermanniella prasina is a terrestrial form of one ol the many other Podostemaceae that grow in the area. No other species of subgenus Ledermanniella have been collected from the Mana River. In ad¬ dition, the L. prasina population is limited to a rock shelf, at least a meter above the high-water level, and does not extend down to the water, suggesting that it is intolerant of extended periods of submer¬ sion. Finally, some of the traits that separate the new species from similar species are structural or reproductive, and likely to be independent of hab¬ itat-related morphological plasticity. Morphology Podostemads are difficult to describe in terms of standard angiosperm morphology due to their great¬ ly reduced vegetative anatomy and unusual mor¬ phology. Distinguishing vegetative organs using cell and tissue orientation is an extremely difficult if not an impossible task (Rutishauser, 1995; Ameka et 230 Novon al., 2003). The literature contains descriptive in¬ consistencies among the interpreted structures, demonstrating a general discordance of terminology and homology assessment among biologists study¬ ing Podostemaceae (Rutishauser, 1995; Sehgal et al., 2002). We use the terms “root,” “main stems,” “fertile shoots,” “vegetative shoots,” and “leaves” (Fig. I) simply to describe this species; however, homology of such organs remains uncertain (Rutis¬ hauser, i995, 1997). The root structure is represented by an undefined thalloid tissue with a holdfast (= hapteron) located underneath. The function of the holdfast is to sta¬ bilize the plant by attaching it to the substrate. Bot¬ anists working with Podostemaceae use the term holdfast or hapteron, borrowed from algae termi¬ nology, to describe the superficially similar struc¬ ture (Taylor, 1954; Hammond, 1937; Philbrick, 1984). The shape of the thalloid-root is described as being disk-like. However, the variability in thal¬ loid-root shape is not fully understood in this spe¬ cies. The term “thalloid-root” is used to describe the undifferentiated tissue of which the main stem arises and that binds the plant to the substrate. There is no implication of homology to the familiar idea of a root since the structure has been sug¬ gested to be derived from a stem, a coalesced stem and root, or simply a modified root (Sehgal et al., 2002; Rutishauser, 1995). This term conforms to current literature and the root-shoot (CRS) model as the structure located underneath the main stem. However, much anatomical work needs to done on this plant as well as other podostemads before more concise determinations of homology can be made. Approximately four to five main stems develop directly from the thalloid-root for each individual (Fig. 1A). The vegetative shoots and fertile shoots are borne on the main stem. The majority of the flowers are borne on the fertile shoots; however, some parts of the thalloid-root contain a shorter main stem of which there are no apparent vegeta¬ tive or fertile shoots, but do have spathella sub¬ tended by leaves. It is not known if these stems elongate later in development and give rise to the vegetative and fertile shoots or remain as shorter dimorphic stems. This trait is not found in all in¬ dividuals. Two types of shoots are borne from the main stem, fertile shoots and vegetative shoots. The veg¬ etative shoots branch dichotomously and appear leaf-like. There are two to three vegetative shoots borne on the main stem, before the fertile shoots arise (Fig. 1 A). Located where the vegetative shoots diverge from the main stem, ligule-like structures are found that encircle it (Fig. 1A, B). The fertile shoots tend to be thicker than the vegetative shoots where they are borne on the main stem (Fig. 1A, B) . They also differ by having reproductive struc¬ tures along the branches and in the axils of di¬ chotomous branches (F ig. I A, B). Leaves are located below the spathella (F ig. 1 A— C) . The basal portions of the leaves are scale-like, with an elongating thread-like leaf that dichoto¬ mously branches, similar in appearance to the veg¬ etative stems. Stipules are located along the side of the scale. Ameka et al. (2003) referred to the basal scale as an intrapetiolar scale, which can also be rudimentary lateral stipules. We apply the term “leaves” to remain consistent with literature de¬ scribing similar species (Ameka et al., 2003; Ru¬ tishauser, 1997; Hall, 1971; Taylor, 1954). The spathella are quite variable in this species. The shape can vary from ovate to elliptic to clavate as they mature. It is also found that the spathella on the distal parts of the shoots are more ovate than those located toward the base. There is further var¬ iation in the spathellum margin following dehis¬ cence. Most spathella have irregular margins fol¬ lowing dehiscence; some, however, develop bilobed or trilobed margins. The number of stamens is a variable character in this taxon as well as in other select species of Podostemaceae (Cusset, 1974). Two out of approx¬ imately 75 shoots have a single flower (each indi¬ vidual has approximately four to five shoots with an average of ca. 20 (lowers per shoot) that contains two anthers attached to an andropodium (specimen housed at OSC). However, variability in anther number is rare enough that we continue to describe this species as monostaminate. Furthermore, a sin¬ gle individual was found containing two ovaries at¬ tached to a single gynophore (specimen housed at OSC). From our understanding of the group, this is a novel attribute for Ledermanniella. We could not find any literature documenting this mutation, sug¬ gesting its anomaly, and we therefore did not in¬ clude it in the description. The specific epithet “prasina” refers to the em¬ erald green seeds that were removed from dehisced capsules. Less mature seeds seen inside undehis¬ ced capsules are also green. Detailed comparisons were made between the new species and similar described species. The species selected for comparison included all mon¬ ostaminate species from the subgenus, plus one species with two stamens. Published descriptions were used for the comparisons in all but two cases, where we needed to examine herbarium material. Comparisons were made using herbarium mate¬ rial with the two monostaminate species, Leder- Volume 14, Number 2 2004 Schenk & Thomas Ledermanniella prasina from Cameroon 231 manniella bowlingii (J. B. Hall) C. Cusset from Ghana and L guineensis C. Cussel from Guinea. These two speeies were examined because they re¬ semble the general habit of L. prasina: a single sta¬ men, elongated shoots, intrapetiolar scale (L. bow¬ lingii only), and linear leaf or leaf-like structures. The herbarium material along with written de¬ scriptions demonstrate that Ledermanniella bowlin¬ gii differs qualitatively by having papillate spa- thella, pollen in dyads, inflorescence a cyme, and brown to reddish brown seeds, along with quanti¬ tative traits such as longer shoots, shorter tepals, smaller anthers, and smaller gynoecium (Hall, 1971; Cusset, 1984; Ameka et al., 2003). This was determined after examining the isotype (P) and a photograph of the holotype (K) along with descrip¬ tions and illustrations. Ledermanniella guineensis is distinguished from L. prasina by having a total length of one meter, inflorescence a cyme, a bilobed spathellum, the lack of leaves directly subtending the spathellum, smaller overall flower size, and brown to reddish brown seeds (Hall, 1971; Cusset, 1984). The holotype (P), along with four additional collections of Ledermanniella guineensis, was used for this comparison along with written descriptions and illustrations. We did not use herbarium material to compare the three remaining monostaminate species lecler- manniella jaegeri G. Cusset, L. minutissima C. Cus¬ set, both from Sierra Leone, and L. monandra C. Cusset from Cameroon, since the published de¬ scriptions of these species show they are very dif¬ ferent. Both L. minutissima and L. jaegeri differ from L. prasina by containing leaves that subtend the spathella which lack intrapetiolar scales and dichotomous branching. Ledermanniella minutissi¬ ma is further separated from L. prasina by pos¬ sessing spathella that always arise from the thalloid or from a small shoot that is up to 3 mm long, and are always solitary (Cusset, 1984). ledermanniella jaegeri has the additional character traits of leaves borne along the entire shoot instead of solely under the spathellum, as well as a more elliptic ovary (Cusset, 1984). Ledermanniella monandra appears to have an intrapetiolar scale; however, the leaves are borne on the entire length of the fertile shoot and are triangular in shape in contrast to the elon¬ gating leaves of L. prasina (Cusset, 1974, 1984). The only bistaminate species investigated was Ledermanniella bifurcata (Engler) C. Cusset from Cameroon, Gabon, and Congo. Although it does have two stamens, it shares similar morphological traits, such as irregular spathella dehiscence, pol¬ len in monads, and intrapetiolar scales, which are characters we felt needed further comparisons. Be¬ sides the anther number, L. bifurcata differs by having an ovate spathellum (L. prasina also has an ovate spathellum, though it elongates to become clavate as it matures), a fertile shoot that widens at nodes (vs. no widening at nodes in L. prasina ), leaves subtending the spathellum for the entire length of the shoot, shorter tepal length, smaller closed spathella. longer spathella pedicels, smaller anther size, and smaller gynoecium (Cusset, 1984, 1987; Hall. 1971). Key to the Monostaminate Species of Ledermanniella subg. Ledermanniella la. Leaves present under spathella, intrapetiolar scale absent, leaves do not branch dichotomous- ly. 2a. Plants without elongated shoots (0—3 mm); leaves located only under spathella . . . . L. minutissima C. Cusset 2b. Shoots greater than 3 mm, leaves located along entire shoot . L jaegeri C. Cusset lb. Leaves either absent under spathella, or if pre¬ sent, with dichotomous branching and intrapetio¬ lar scale. 3a. Leaves triangular, 1-2 mm long . . L. monandra C. Cusset 3b. Leaves linear, greater than 2 mm long. 4a. Spathella papillate, pollen in dyads . . . /,. bowlingii (J. B. Hall) C. Cusset 4b. Spathella glabrous, pollen in monads. 5a. Stems up to I m long, trailing in moving water; leaves absent below spathella; spathella arranged in a cyme; spathella bilobed at dehis¬ cence; anthers 1 mm long; tepals 0.3— 0.4 mm long; seeds brown/red . L. guineensis C. Cusset 5b. Plants 9—13 cm, erect on wet rock; leaves located below spathella; spa¬ thella solitary or in cluster; spa¬ thella irregular to bilobed at dehis¬ cence; anthers 2 mm long; tepals 0.4— 0.7 mm long; seeds emerald green . L. prasina J. Schenk & 1). W. Thomas Acknowledgments. The authors are indebted to Ffena Schlachter for illustrations, and to Kenton L. Chambers for the Latin description. We thank Aar¬ on Liston for reviewing drafts of this manuscript, and two anonymous reviewers who provided useful comments. Thanks are also due to the curators of K and P for loaned specimens, as well as to Rich¬ ard R. Halse (OSC) for obtaining and housing the loans. Funds for the fieldwork were provided by the International Cooperative Biodiversity Groups (ICBC). Literature Cited Ameka, G. K., G. C. Clerk, E. Pfeifer & R. Rutishauser. 2003. Developmental morphology of ledermanniella 232 Novon bowlingii (Podostemaceae) from Ghana. PI. Syst. Evol. 237: 165-183. Chuyong, G. B., R. Condit, 1). Kenfack, E. Losos, VI. Sa- inge, N. C. Songwe & D. W. Thomas. 2004. Korup Eor- est Dynamics Plot, Cameroon. Chapter 29 in E. Losos & E. G. Leigh, Jr. (editors). Tropical Forest Diversity and Dynamism: Findings from a Large-Scale Plot Net¬ work. Univ. Chicago Press, Chicago. Cook, C. I). K. 1996. The Aquatic Plant Book, 2nd Rev. Ed. SPB Academic Publishing, The Hague. Cusset, C. 1974. Contribution it lYtude des Podostema¬ ceae: 4. Les genres Ledermanniella, Monandriella et Inversodicraeia. Adansonia, ser. 2, 14: 271-275. - . 1983. Contribution it lY'tude des Podostemaceae. 7. Ledermanniella Engl., sous-genre Phyilosoma, C. Cusset. Bull. Mus. Natl. Hist. Nat.. B. Adansonia 5: 361-390. - . 1984. Contribution i lYtude des Podostemaceae: 8. Ledermanniella Engl, sous-genre Ledermanniella. Bull. Mus. Natl. Hist. Nat., B, Adansonia 3: 249—278. - . 1987. Podostemaceae and Tristichaceae. Pp. 51— 99 in B. Satabif4 & P. H. Moral (editors), Flore du Came¬ roon Vol. 30. Ministfere de I'Enseignement Suptirieur et de la Recherche Scientifique, Yaounde. Hall, J. B. 1971. New Podostemaceae from Ghana with notes on related species. Kew Bull. 26: 125-136. Hammond, B. L. 1937. Development of Podostemum cer- atophyllum. Bull. Torr. Bot. Club 64: 17—36. Philbrick, C. T. 1984. Aspects of floral biology, breeding system, and seed and seedling biology in Podostemum ceratophyllum (Podostemaceae). Syst. Bot. 9: 166—174. Rntishauser, R. 1995. Developmental patterns of leaves in Podostemaceae compared with more typical flowering plants: Saltational evolution and fuzzy morphology. Can- ad. J. Bot. 73: 1305-1317. - . 1997. Structural and developmental diversity in Podostemaceae (river-weeds). Aquat. Bot. 57: 29—70. Sehgal, A., M. Sethi & H. Y. Mohan Ram. 2002. Origin, structure, and interpretation of the thallus in Hydrob- ryopsis sessilis (Podostemaceae). Ini. J. PL Sei. 163: 891-905. Taylor, G. 1954. Podostemaceae. Pp. 122—127 in J. Hutchinson & J. M. Dalziel (editors), Flora of West Tropical Africa, vol. 1, 2nd ed., revised by R. W. J. Keay. Crown Agents, London. A New Species of Minasia (Asteraceae, Vernonieae) from the Planalto de Diamantina, Minas Gerais, Brazil Joao Semir Departamento de Botanica, Universidade Estadual de Campinas, C. P. 6109, 13083-970, Campinas SP. Brazil, semir@unicamp.br FI avia F. Jesus Departamento de Genetica, Universidade Estadual de Campinas, C. P. 6109, 13083-970, Campinas SP, Brazil, f931719@dac.un iearnp.br ABSTRACT. A new species of Minasia (Astera- eeae, Vernonieae) was found in the “campus rupes- tres" of the Planalto de Diamantina, Minas Gerais, Brazil. Minasia lewinsohnii is characterized by its overall small size, linear to elliptical subfleshy leaves, sessile to pedunculate capitula, and aehenes with indumentum throughout the surface. So far, it is known from only one population. Key words: Asteraceae, Brazil, Minasia, Ver¬ nonieae. Minasia H. Robinson (Asteraceae, Vernonieae) was recently established by Robinson (1992). who created the genus based on two species previously described as belonging to Vernonia Schreber and established another new species. According to Rob¬ inson (1992, 1995, 1996, 1999), the genus cur¬ rently includes five species: M. alpestris (Gardner) H. Robinson, M. scapigera H. Robinson. M. perei- rae H. Rol >inson, M. splettiae H. Robinson, and M. cabralensis H. Robinson. Minasia species are endemic to the campos ru- pestres of the Kspinhago Range in Minas Gerais, Brazil. More specifically, they occur in the high- altitude areas ranging from the Serra do Cipd to the Planalto de Diamantina, as well as in the isolated western extension of the campos rupestres in Minas Gerais, the Serra do Gabral. The campos rupestres have an immense diversity of species, many of which are endemic (Giulietti & Pirani, 1988). They are considered to present the greatest degree of en¬ demism among Brazilian vegetation types (July, 1970). Therefore, finding a new species in these environments is not surprising. Recent collecting trips were conducted by fac¬ ulty of the Universidade Estadual de Campinas for taxonomic, genetic, and ecological studies on the interactions between Asteraceae and phytophagous insects. During those trips, a new, seemingly micro¬ endemic species of Minasia was found in the Plan¬ alto de Diamantina. This region seems to be the one with the greatest diversity of Minasia species, with four of the five previously described species occurring there (M. alpestris, M. scapigera, M. per- eirae, and M. splettiae), in addition to the new spe¬ cies proposed here. Minasia lewdnsoliiiii .). Semir & F. F. Jesus, sp. nov. TYPE: Brazil. Minas Gerais: Diamantina, Estrada Guinda-Sao Joao da Chapada, campo arenoso com cascalho quartzftico, 18°09.59'S, 43°43.00'W. 7 Sep. 1996 (fl, fr), T. M. Lewin- sohn, P. I. Prado, A. J. Santos 6696 (EF.C). of the adaxial face, margin barbellate, purple on the middle, pale on the margins. Flowers 21 to 43 per capitulum; corolla 7.1-10.6 mm long, lavender to magenta, densely glandular-verrucous on the lobes, sparsely so on the tube, tomentose on the external apex of the lobes, glabrous below; lobes 2.5— 3.2 X 0.6-0. 8 mm, lanceolate; anthers 2. 9-4. 2 mm long, tails ea. 0.7 mm long, apical appendages Volume 14, Number 2 2004 Semir & Jesus Minasia lewinsohnii from Brazil 235 0.6— 1.0 X 0.2— 0.3 mm, triangular to sub-lanceo¬ late, apex acute; style ca. 12 mm long. Achene 2.2— 3.5 mm long, 0.8— 1.5 mm diam., ribbed, ca. 8 ribs, sericeous to setuliferous indumentum, dense to sub-glabrescent throughout the surface, sometimes denser at the base, glandular surface; pappus bris¬ tles 0.4— 7.5 mm tall, in indistinguishable series, the most external ones gradually shorter, lavender to magenta, barbellate. The species is comparable in general size — both of rosette and inflorescence — to Minasia pereirae and M. splettiae. In inflorescence height it is also comparable to M. cabralensis ; however, the rosette leaves are shorter in M. lewinsohnii (up to 9 cm long) than in M. cabralensis (10-16 cm long), giving M. lewinsohnii the appearance of an overall smaller plant. Minasia lewinsohnii is much smaller than M. alpestris and M. scapigera, both of which can be up to 1 m tall. In addition to the difference in general size, the leaves in M. lewinsohnii are linear to el¬ liptical, not oblaneeolate as in M. alpestris , and the inflorescence is not congested like the one in M. scapigera. The capitula in this new species may be pedun¬ culate, as they generally are in Minasia cabralensis, but very rarely in the other species of the genus. However, the peduncles are much shorter in M. lewinsohnii than in M. cabralensis (up to 1.5 cm vs. up to 35 cm long). Also, the achene in M. lewin¬ sohnii has indumentum throughout the surface, whereas in M. cabralensis the achene is distally gla¬ brous (Robinson, 1996). Furthermore, the leaves are shorter and more fleshy in M. lewinsohnii than in M. cabralensis. To date, M. cabralensis seems to be restricted to the Serra do Cabral while M. lew¬ insohnii has been found only in the Planalto de Diamantina. Minasia lewinsohnii differs from M. splettiae in having wider and thicker leaves and from M. pereirae in having linear to elliptic rather than oblaneeolate leaves. To date, Minasia lewinsohnii is known only from one population, in the Planalto de Diamantina, oc¬ curring on sand. The species is named in honor of Thomas Lewinsohn (Departamento de Zoologia, Universidade Estadual de Campinas), one of the first collectors of this population. Paratypes. BRAZIL. Minas Gerais: Diamantina, Es¬ trada Guinda-S. Joao da Chapada, I8°09.58'S, 43°42.96'W, 23 June 2000 (fl, fr). K. S. Yotoko et al. KSY- 00/112 (BllCB, UEC), 29 Apr. 1997, T. M. Lewinsohn et al. in PIC97032 (MBM, SP, UEC). Acknowledgments. We thank Thomas M. Lew¬ insohn, Paulo I. Prado, and Vera N. Solferini for collections and support in the study of this genus; Angela B. Martins for the Latin diagnosis; Angela M. F. Pacheco for the drawings; Christina Muirhead for help with the English text; Fabio A. Vitta and Mariana E. Mansanares for support in the Depar¬ tamento de Botanica (UEC); Karla S. Yotoko and Aluana G. Abreu for the collections in 2000; Har¬ old Robinson (US), John Pruski (MO), and Victoria C. Hollowell (MO) for critical reviews of the man¬ uscript; CNPq and FAPESP (98/05085-2) for finan¬ cial support. This study is part of the program BI- OTA/FAPESP, The Virtual Institute of Biodiversity (www.biota.org.br). Literature Cited Giulietti, A. M. & J. R. Pirani. 1988. Patterns of geograph¬ ic distribution of some plant species from the Espinhago Range, Minas Gerais and Bahia, Brazil. Pp. 39-69 in W. R. Heyer & P. E. Vanzolini (editors), Proceedings of a Workshop on Neotropical Distribution Patterns. Aca¬ demia Brasileira de Ciencias, Rio de Janeiro. Joly, A. B. 1970. Conhega a Vegetagao Brasileira. EDUSP & Polfgono, Sao Paulo. Robinson, H. 1992. Notes on Lychnophorinae from Minas Gerais, Brazil, a synopsis of Lychnophoriopsis Schultz- Bip., and the new genera Anteremanthus and Minasia (Vemonieae: Asteraceae). Proc. Biol. Soc. Wash. 105: 640-652. - . 1995. New combinations and new species in American Vernonieae (Asteraceae). Phytologia 78: 384— 399. - . 1996. A new species of Minasia from the Serra do Cabral, Minas Gerais, Brazil (Vernonieae; Astera¬ ceae). Phytologia 80: 350—35 1 . - . 1999. Generic and subtribal classification of American Vernonieae. Smithsonian Contr. Bot. 89. Eugenia myrciariifolia (Myrtaceae), a New Species from Parana, Brazil Lucia Helena Soares-Silva Herbario, Departamento de Botanica UnB, C.P. 04457 — 70919-970 — Brasilia, I)F, Brazil. lsoares@unb.br Marcos Sobral Departamento de Botanica UFMG, Av. Antonio Carlos 6627, — 31270-110 — Belo Horizonte, MG, Brazil, sobral@mono.icb.ufmg.br ABSTRACT. Eugenia myrciariifolia, a new species from the southern Brazilian state of Parana, is de¬ scribed and illustrated. This species is related to Eugenia oeidocarpa, from which it is set apart by the presence of cataphylls on the twigs, elliptic to lanceolate leaves, smaller or absent pedicels, and axillary, pilose flowers. RESUMO. Eugenia myrciariifolia, uma nova es- pdeie do estado do Parana, Brasil, e descrita e il- ustrada. A nova cspccie e proxima a Eugenia oei¬ docarpa, da qual se distingue pela presen^a de catafilos nos ramos, folhas elfpticas a lanceoladas, pedicelos menores ou ausentes e flores axi lares e pilosas. Key words: Brazil, Eugenia, Myrtaceae. Eugenia L. is one of the largest genera of the Myrtaceae, with about 350 American species (Lan¬ drum & Kawasaki, 1997). The Brazilian species have remained largely unstudied since the para¬ mount treatment of Berg (1857—1859), except for local floristic studies (e.g., Kiaerskou, 1893; Le- grand & Klein, 1969; Kawasaki, 1989; Mattos, 1984; Barroso & Peron, 1994; Marchiori & Sobral, 1997; Sobral, 2003). While studying the family Myrtaceae in the state of Parana, a small-leaved tree of the forests from the northern part of the state came to our attention and is here proposed as new. Eugenia my reiariifoliu Soares-Silva & Sobral, sp. nov. TYPE: Brazil. Parana: Sapopema, Salto das Orqufdeas, margem direita do rio Lageado, 8 Dec. 1995, /.. Soares-Silva 563 (holotype, UB; isotype, FUEL). Figure I. Species E. oeidocarpae proxima, a qua ramulis cata- phyllorum instructa, foliis ellipticis vel lanceolatis (non obovatis), pedicellis usque 2 mm longis (non 2—5 mm lon- gis) et floribus axillaribus ovariis pubescentibus reeedit. Novon 14: 236-238. 2004. Small tree up to 5 m high; twigs red-brown or grayish, exfoliating, the younger ones somewhat an¬ gled, brown and puberulent with white trichomes 0. 1—0.3 nun long, bearing at their bases spathulate to linear cataphylls 2.5— 5.5 X 1 mm, these invo¬ lute, with cilia to 0.1 nun. Leaves chartaceous, with scattered translucid punctations visible to the eye, these more evident abaxially, reticulate, slightly discolored, glabrous except for occasional scattered white trichomes to 0.1 mm long along the adaxial central nerve and 0. 1-0.3 mm long at the base of leaves; blades elliptic to elliptic-lanceolate, 1—3 X 0.5—2 cm, the length/width ratio 2. 2-2. 8:1; apex obtuse to acute; base acute to attenuate; central nerve plane to slightly prominent adaxially and ev¬ idently raised abaxially; secondary nerves 6 to 10 pairs, evident on both faces and diverging at angles of 50°— 75°; marginal vein 0.6— 0.8 mm from margin, the margin itself with a yellowish ridge to 0.1 mm wide; petioles plane adaxially, 1-3 X 0.4-0. 6 mm, with trichomes like those on the branches. Flowers axillary and solitary; pedicels 1—2 X 0.2-0. 5 mm or absent; bracteoles persisting after anthesis, ovate to rounded ovate, 0.5—1 X 0.7— 1.2 mm, ciliate; flower buds globose, 3—4 mm diam.; calyx lobes hemispherical, unequal in size, the external ones 1-1.3 X 1.5-2 mm, the internal ones 1.6—2 X 2- 2.5 mm, sparsely puberulent without and sericeous within, with trichomes about 0.1 mm, the margins sometimes with red cilia to 0.4 mm; petals orbicular and concave, 2.5—3 mm diam., glabrous and ciliate; staminal ring sericeous, 1.5 mm diam.; stamens ca. 100. 4—5 mm, the anthers globose, 0.4— 0.5 mm, with one apical gland; style glabrous, 4—6 mm; ova¬ ry externally pubescent or sericeous, about 1 mm long, bilocular, with 7 to 8 ovules per locule on a central placenta. Fruits not seen. Habitat and distribution. Eugenia myrciariifol¬ ia is a small tree from seasonal forests of central Volume 14, Number 2 2004 Soares-Silva & Sobral Eugenia myrciariifolia from Brazil 237 Figure 1. Eugenia myrciariifolia Soares-Silva & Sobral. — A. Habit. — B. Cataphylls at base of twigs. — C. Detail of adaxial side of leaf. — I). Detail of inflorescence. — E. Flower bud, with bracteoles detached. — F. Flower after polli¬ nation, with bracteoles detached. — G. Stamen with apical gland. From Soares-Silva 570. 238 Novon and western Parana; until now it has been collected in only four municipalities. Phenology. Blooming specimens were collected from October to December; fruiting specimens are still unknown. Etymology. The specific epithet is allusive to the resemblance of this species with some of the genus Myrciaria, especially Myrciaria tenella (DC.) 0. Berg, with which it can be easily confused when sterile due to its small leaves and dense foliation; curiously, one of the vernacular names that we have registered for this species, eambuf, is also largely applied in southern Brazil lor species of Myrciaria. Vernacular names. Cambuf, pitangao (from Lin¬ de man & Haas 2892). Affinities. This species is related to Eugenia oeidocarpa 0. Berg, another southern Brazilian spe¬ cies growing from Parana to Rio Grande do Sul (Sobral, 2003). The following key presents the char¬ acters that set them apart. Key kok Distinguishing Eugenia myrciariifolia and Eu¬ genia OEIDOCARPA IN SOUTHERN BRAZIL la. Twigs without linear cataphylls at base; leaves obovate with revolute margins; petioles to 4 X 0.8 mm; the inflorescences ramiflorous, 2- to 6- florous, the axis to 2 X 0.5 mm; pedicels to 5 X 0.5 mm; flowers glabrous; larger calyx lobes to 3 X 3 mm . Eugenia oeidocarpa lb. Twigs with linear cataphylls at base; leaves el¬ liptic-lanceolate; petioles to 2 X 0.6 mm; pedi¬ cels to 2 X 0.5 mm; flowers pilose, solitary and axillary; larger calyx lobes to 2 X 2 mm . . Eugenia myrciariifolia Additionally, this species shows close resem¬ blance to the protologue of Eugenia coaetanea (). Berg (Berg. 1857-1859), another ramiflorous spe¬ cies distinguished from E. myrciariifolia and E. oei- docarpa by pilose young leaves, linear bracteoles to 4 mm, and flower buds to 8 mm; nevertheless, comparison with this species is problematic since the type was destroyed at B and no isotypes are known. Moreover, although said to be collected in the state of Rio Grande do Sul, no further collec¬ tions of E. coaetanea are known, notwithstanding extensive surveys in the myrtaceous flora of this state (Mattos, 1984; Marchiori & Sobral, 1997; So¬ bral, 2003). Paratypes. BRAZIL. Parana: Aranha, Lindeman & Hass .3432 (MBM, U); Campo Novo, ca. 50 km de Lar- anjeiras do Sul. Lindeman <£ Hass 2892 (MBM. U, LIB); Parque Nacional do Iguayu [municipality of C(iu Azul, not registered in label), 100 m W of rio Floriano, Lindeman & Hass 3544 (MBM. 0); Sapopema, Salto das Orqufdeas, margem direita do rio Lageado, Soares-Silva 441 (FUEL, UB), 9 Dec. 1995, Soares-Silva 570 (FUEL, UB), Chagas e Silva & Soares-Silva 2056 (FUEL, UB), Silva 2127 (FUEL, UB), Medri 744 (BHCB, FUEL), Pavdo et al. s.n. (FUEL 27498, MO). Acknowledgments. We acknowledge Claudenir S. Cares (UB) for his fine drawing of Eugenia myr¬ ciariifolia, the curators of FUEL and MBM for their kind collaboration, and Victoria Hollowed (MO) and an anonymous reviewer for kind and valuable suggestions for the improvement of this paper. Literature Cited Barroso, G. M. & M. V. Peron. 1994. Myrtaceae. In: M. P. M. Lima & R. Guedes-Bruni (org.), Reserva Ecold- gica de Maea6 de Cima, Nova Friburgo, Rio de Janeiro. Aspectos Florfsticos das Esp^cies Vasculares, Vol. 1. Jardim Botiinico do Rio de Janeiro, Rio de Janeiro. Berg, 0. 1857-1859. Myrtaceae. In: K. Martius (editor). Flora Brasil iensis 14(1): 1—656. Kawasaki, M. L. 1989. Flora da serra do Cip6, Minas Gerais; Myrtaceae. Bol. Hot. Univ. Sao Paulo II: 121 — 1 70. Kiaerskou, H. 1893. Enumeratio myrtacearum brasilien- sium. In: E. Warming (editor), Symbolarum ad floram Brasiliae Centralis cognoscendam 39: 1-199. Landrum. L. R. & M. L. Kawasaki. 1997. The genera of Myrtaceae in Brazil: An illustrated synoptic treatment and identification keys. Brittonia 49: 508-536. Legrand, C. I). & R. M. Klein. 1969. Miildceas: Eugenia. Pp. 45—216 in R. Reitz (org.), Flora llustrada Catari- nense, Itajaf. Marchiori, ,1. N. C. — 0.8 mm. Plants subshrubs, woody at base; trichomes den¬ dritic throughout, 0.1— 0.2 mm long; stems 10—17 cm long, erect to ascending, few from base, many- branched above, sparsely pubescent below, gla¬ brous and glaucous above. Leaves all cauline, nar¬ rowly lanceolate, 4-13 X 1-2 mm, fleshy, becoming leathery upon drying, moderately pubes¬ cent, gradually becoming sparsely pubescent up¬ ward, base minutely auriculate, margin entire, apex acute, auricles to I X 0.5 mm. Raceme ebracteate, 6- to 10-flowered; fruiting pedicels 3—5 mm long. Volume 14, Number 3 2004 Al-Shehbaz Synopsis of Neuontobotrys 255 256 Novon glabrous, strongly reflexed or recurved and forming half a loop. Sepals narrowly oblong, 3—3.5 X 0.5— 0.7 mm, moderately pubescent; petals yellow, soon turning reddish with age, linear-oblanceolate, 4.5- 5 X 0.5— 0.7 mm, sparsely pubescent outside; fila¬ ments 2.5—3 mm long; anthers oblong-linear, 0.9— I mm long; ovules 12 to 14 per locule. Fruits oblong-linear, 6-8 X ca. 1 mm, not torulose; valves thick, leathery, with a prominent midvein; gyno- phore 0. 1—0.2 mm long; style slender 0.8—1 mm long; seeds ovoid, brown, uniseriate, 0.9-1 X 0.6— 0.8 mm. Neuontobotrys elloanensis, which is known only from the type gathering, is easily distinguished from its nearest relative, N. deserticola, by having entire, narrowly lanceolate leaves to 2 mm wide and small¬ er petals (4.5—5 mm long) sparsely pubescent out¬ side. Hy contrast, N. deserticola has dentate, broad¬ ly ovate to ovate-lanceolate leaves 4 — 1 0( — 20) mm wide and larger petals (6-8 mm long) glabrous out¬ side. 5. Neuontobotrys deserticola (Philippi) Al- Shehbaz, comb. nov. Basionym: Cardamine de¬ serticola Philippi, Anal. Univ. Chile 81: 86. 1892. TYPE: Chile. “Deserto Atacama ad fon- tem Acerillos,” Oct. 1877, 0. Villanueva 1606 (holotype, SGO-63889). Sisymbrium philippianum I. M. Johnston, Rev. Chil. Hist. Nat. 33: 26. 1929. Syn. nov. Based on Sisymbrium amplexicaule Philippi, FI. Atac. 8: 10. I860, non A. Cray, U.S. Explor. Exped. Wilkes, Bot. XV. 61. 1854. TYPE: Chile. Atacama: Pajonal, |23°46'S, 3170 m], R. A. Philippi s.n. (lectotype, designated here, SCO-49243). Distribution. Argentina (Prov. Catamarca, Ju- juy, and Salta), Bolivia (Depto. Potosf), and Chile (Regidn II and III). Representative specimens. ARGENTINA. Catamarca: Antofagasta de la Sierra, bos Nacimientos, Cabrera et al. 31862 (MO). Jujuy: Susquesientre Cauchori y Catua, Ca¬ brera el al. 22344 (SI). Salta: Salar de Gallego, Camino a buea-huasi, Cabrera 8882 (F, GH). B01.1VIA. Potosi: Prov. Nor bipes, Chiguana, Asplund 3031 (S, UPS). CHILE. Region II (Antofagasta): San Pedro, near river Loa, Zollner 8842 (MO, US). Region III (Atacama): Depto. Copiapd, Cord. Rio Figueroa, Cl. Paredones, Wer- dermann 96 8 (B, CAS, F, GH, MO, UC). Although the type of Neuontobotrys deserticola consists of three racemes and lacks the lower parts of the plant, it is indistinguishable in every aspect of (lower, fruit, and indumentum from plants of Sis¬ ymbrium amplexicaule Philippi. In fart, Schulz (1924) was the first to point out this similarity but failed to unite the two species. Munoz-Pizarro (1960) cited two sheets, SGO- 49242 and SGO-49243, as the type of Sisymbrium amplexicaule. The latter specimen, which is the most complete and consists of four plants all col¬ lected from one locality, is designated herein as the lectotype. The other sheet, SCO-49242, has five fragments each collected from a different place, and it is impossible to assign any of these fragments to a given locality. As delimited hy Knntze (1898), Hesperis intri¬ catissima (Philippi) Knntze included two species that differ substantially in every morphological as¬ pect except for fruit size and orientation. In order to simplify the complicated nomenclature of these two species, they are designated herein as “sp. A” and “sp. B.” One species (sp. A), which is the bas¬ ionym of //. intricatissima, is a glabrous annual originally described as Draba intricatissima Philip¬ pi and now recognized as Eremodraba intricatissi¬ ma (Philippi) O. E. Schulz (Schulz, 1924; Al-Sheh- baz, 1990). I he second of Kuntze’s (1898) species (sp. B) is a pubescent perennial or subshrub cor¬ responding to Sisymbrium amplexicaule Philippi. Schulz (1924) mishandled the nomenclature of Kuntzes (1898) //. intricatissima in four ways. First, he excluded the material based on sp. A to Eremodraba, but maintained the epithet “'intricatis¬ sima" in Sisymbrium for sp. B. Second, he called the species Hesperis intricatissima Knntze, instead of //. intricatissima (Philippi) Knntze, even though Knntze (1898) cited the basionym I). intricatissima and did not treat the plant as a new species. Third, Schulz (1924, 1928, 1932) overlooked Beiche’s (1896) earlier transfer of I). intricatissima (sp. A) to Sisymbrium. Finally, although Johnston (1929) correctly interpreted the nomenclature of sp. B and proposed the name S. philippianum I. M. Johnston to replace the later homonym S. amplexicaule Phi¬ lippi, Schulz (1932) continued to use S. intricatis- sirnum for sp. B and placed S. philippianum in its synonymy. Acknowledgments. I am most grateful to Melica Munoz-Schick for her help with the Philippi types at SGO, Carmen Ulloa for translating the abstract into Spanish. Robert Vogt lor sending the images of Neuontobotrys berningeri , Anthony R. Brach for providing needed literature, and the directors and curators of the herbaria cited. I am most thankful to Steve L. O’Kane Jr.. Victoria C. Hollowed, Amy McPherson, Diana Gunter, and an anonymous re¬ viewer for their valuable comments on the manu¬ script. Literature Cited Al-Shehbaz, I. A. 1990. The South American Eremodraba (Brassicaoeae). Ann. Missouri Bot. Gard. 77: 602—604. Volume 14, Number 3 2004 Al-Shehbaz Synopsis of Neuontobotrys 257 Appel, 0. & I. A. Al-Shehbaz. 2003. Crueiferae. Pp. 75— 174 in k. Kubitzki (editor). Families and Genera of Vascular Plants 5. Springer- Verlag, Berlin, Heidelberg. Baehni, C. & J. F. Macbride. 1937. Remarques sur les Gruciferae-Sisymbrieae. Candollea 7: 291-296. Boelcke, 0. 1961. Nota sobre Sisymbrium robustum Chod. et Wilcz. Bol. Soc. Argent. But. 9: 387—392. Chodat, R. & F. Wilczek. 1902. Contributions & la (lore de la Republique Argentine. Bull. Herb. Boissier, Ser. 2. 2: 281-296. Greuter, W., J. McNeill, F. R. Barrie, H. M. Burdet, V. Demoulin, T. S. Filgueiras, D. H. Nicolson, P. C. Silva, J. E. Skog, P. Trehane, N. J. Turland & I). L. Hawk- sworth (editors). 2000. International Code of Botanical Nomenclature (Saint Louis Code). Regnum Veg. 138. Johnston, I. M. 1929. A new Chilean plant and some no- menclatorial changes. Rev. Chilena Hist. Nat. 33: 25— 27. Kuntze, 0. 1898. Revisio Genera Plantarum. Vol. 3, part 2. Arthur Felix, Leipzig. Macbride, J. F. 1938. Crueiferae. Field Mus. Nat. Hist., Bot. Ser. 13: 937-983. Munoz-Pizarro, C. 1960. Las Especies de Plantas Descri- tas por R. A. Philippi en el Siglo XIX. Ediciones de la Universidad de Chile, Santiago. Reiche, C. 1896. Flora de Chile, Vol. 1. Imprenta Cer¬ vantes, Santiago. Romanczuk, M. C. 1981. Novedades en el genero Sisym¬ brium (Crueiferae) y sinopsis de las especies Patagdni- cas. Parodiana 1(1): 1—29. - . 1982. El genero Sisymbrium (Crueiferae) en la Argentina. Darwiniana 24: 75—156. Schulz, 0. E. 1924. Crueiferae— Sisymbrieae. Pp. 1—388 in A. Engler (editor), Pflanzenreich IV 105(Heft 86). Verlag von Wilhelm Engelmann, Leipzig. - . 1928. Beilrage zur Kenntnis der Flora von Chile. Notizbl. Bot. Gart. Berlin-Dahlem 10: 44)0-472. - . 1932. Crueiferae variae. Notizbl. Bot. Gart. Ber¬ lin-Dahlem 11: 389-392. - . 1936. Crueiferae. Pp. 227—658 in A. Engler & H. Harms (editors). Die Natiirlichen Pflanzenfamilien, Ed. 2. 17B. Verlag von Wilhelm Engelmann, Leipzig. Stafleu, F. A. & R. S. Cowan. 1976. Taxonomic Literature, Vol. 1: A-G. Bohn, Scheltema & Holkema, Utrecht. Torrey, J. & A. Gray. 1838. A Flora of North America, Vol. 1, part 1. Wiley & Putnam, New York. Warwick, S. I. & I. A. Al-Shehbaz. 2003. Nomenclatural notes on Sisymbrium (Brassicaceae). Novon 13: 265— 267. - . - , R. A. Price & C. Sauder. 2002. Phylog- eny of Sisymbrium (Brassicaceae) based on FI’S se¬ quences of nuclear ribosomal DNA. Canad. J. Bot. 80: 1002-1017. A Synopsis of t lie South American Weberbauera (Brassicaceae) Ihsan A. Al-Shehbaz Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. ihsan.al-shehbaz@mobot.org ABSTRACT. Twenty-two species are recognized in Weberbauera (Brassicaceae), and a key to distin¬ guish them is given. Weberbauera dillonii, W. sca- brifolia, and W. violacea arc described and illus¬ trated. and their distinguishing characters from nearest relatives are discussed. The new combina¬ tions W. peruviana and W. lechleri are proposed. Sisymbrium petraeum and .S’, fastigiatum are re¬ duced to synonymy of W. lechleri. Resumen. Se reconocen veintidos especies en Weberbauera (Brassicaceae), y se presenta una clave para distinguirlas. Se describen e ilustran We¬ berbauera dillonii, W. scabrifolia y W. violacea y se discuten los caracteres que las distinguen de otras especies cercanas. Se proponen nuevas combina- ciones de W. peruviana y II' lechleri. Los nombres Sisymbrium petraeum y S. fastigiatum se reducen a sindnimos de W. lechleri. Key words: Brassicaceae, Sisymbrium, South America, Stenodraba, Weberbauera. Weberbauera 0. E. Schulz (Brassicaceae), a ge¬ nus widely distributed in Peru, Bolivia, Argentina, and Chile (Appel & Al-Shehbaz, 2003), was re¬ cently revised and its boundaries critically delim¬ ited (Al-Shehbaz, 1990a, 1990b). The present pa¬ per continues this revision, recognizing 22 species in Weberbauera, including three new species and two new combinations. The two names transferred herein to Weberbauera, lb peruviana (DC.) Al-Sheh¬ baz and W. lechleri (Fournier) Al-Shehbaz, were originally described in Sisymbrium L. Sisymbrium was believed to have one of its major centers of diversity in South America (Schulz, 1924; Al-Shehbaz, 1988), but that opinion has been shown to be incorrect (Warwick et al., 2002), and the genus is considered to be restricted primarily to the Old World, with only one species, S. linifol- ium Nuttall, native to the New World (western North America). Ravenna (1981) and Boelcke and Romanczuk (1984) placed in Stenodraba 0. E. Schulz some species recognized here as Weber¬ bauera. Schulz (1924, 1936) separated Stenodraba from Weberbauera solely on the absence instead of the presence of bracts. This is an unreliable feature for the delimitation of genera in Brassicaceae, and numerous other genera (e.g.. Cardamine I... Draba L., Rorippa Scopoli, Sisymbrium ) have species with or without bracts (Appel & Al-Shehbaz, 2003). As indicated by Al-Shehbaz (1990a), Stenodraba docs not merit recognition. In the enumeration below, the information pro¬ vided includes the bibliographic citation of each species and its type collections. If the species is known from collections other than the type, one representative collection for every major locale is cited. For the full synonymy and typification of spe¬ cies 2, 3, 7, 8, 10, 12, 18—21, see Al-Shehbaz (1990a). For a detailed description of Weberbauera and full account of its generic synonyms and their types, see Al-Shehbaz (1990a). Weberbauera Gilg & Muschler, Bot. Jahrb. Syst. 42: 481. 1909. TYPE: Weberbauera densiflora (Muschler) Gilg & Muschler = Weberbauera sputhulifolia (A. Gray) (). E. Schulz. Novon 14: 258-268. 2004. Volume 14, Number 3 2004 Al-Shehbaz South American Weberbauera 259 Key to the Species of Weberbavera la. Plant, including fruit, densely covered with malpighiaceous trichomes; petals often pubescenl . . 21. IP. trichocarpa lb. Plants glabrous or with simple and/or stalked forked trichomes, rarely with malpighiaceous trichomes and then only on basal leaves; fruit glabrous or rarely sparsely pubescent with simple trichomes; petals glabrous. 2a. Inflorescence bracteate throughout, rarely uppermost flowers ebracteate. 3a. Petals violet. 6.5— 7.5 X 3-3.5 mm . 22. W. violacea 3b. Petals white, 2-3.5(-5) X 0.7-2(-2.5) mm. 4a. Plants densely scabrous with appressed antrorse trichomes; basal leaves entire, densely scabrous adaxially, glabrous abaxially . 16. IP scabrifolia 4b. Plants glabrous or pilose with spreading trichomes; at least some leaves dentate, repand, lyrate, or pinnatifid, all glabrous or pubescent but never scabrous. 5a. Basal leaves not ciliale at base, sometimes glabrous or with forked trichomes . . IB. IP spathulifolia 5b. Basal leaves conspicuously ciliate (at least basally) or densely covered with simple trichomes. 6a. Fruit apex attenuate; style 1-3 mm long; petals 3.5-5 mm long ... 7. IP herzogii 6b. Fruit apex obtuse; style to 0.2 mm long; petals 2.5-3.5 mm long. 7a. Basal and lowermost cauline leaves lyrate-pinnatifid, ciliate; stems and leaves glabrous; cauline leaves and bracts entire or repand; seeds 3 to 5 per locule . 1 - 4P bracteata 7b. Basal and lowermost cauline leaves denticulate or dentate, not ciliate; stems and leaves densely pilose or hirsute; cauline leaves and bracts coarsely dentate; seeds 6 to 15 per locule. 8a. Cauline leaves and bracts sessile; stems retrorsely pilose; fruits gla¬ brous; pedicels pilose; petals 2—2.5 mm wide . 6. IP dillonii 8b. Cauline leaves and bracts petiolate; stems hirsute; fruits hirsute; ped¬ icels glabrous; petals 0.7—1 mm wide . 14. W. peruviana 2b. Inflorescences ebracteate, rarely lowermost few flowers bracteate. 9a. Sepals persistent into fruit; fruil sparsely pubescent with retrorse simple trichomes . . 15. IP retropilosa 9b. Sepals soon caducous; fruit glabrous. 10a. Petals yellow; fruit septum perforate; leaves adaxially densely sericeous .... 13. IP perforata 10b. Petals white; fruit septum complete; leaves glabrous or differently pubescent. 11a. Basal leaves numerous; petioles persistent, straw-colored, 3—3.5 cm long, ca. as long as blade, overlapping and forming a dense crown . 5. IP densifolia 1 lb. Basal leaves few to several; petioles soon caducous or if persistent, then not straw- colored, less than 3 cm long, shorter than blades, not forming a dense crown. 12a. Stems several-branched above; plants with some dendritic trichomes at least on stems and sepals. 13a. Basal leaves ciliate with simple trichomes, pubescent with dendritic tri¬ chomes . 12. IP! parvifolia 13b. Basal leaves not ciliate, exclusively with malpighiaceous trichomes . . 10. IP lechleri 12h. Stems simple above; plants glabrous or with simple and/or forked trichomes. 14a. Basal and cauline leaves not ciliate, sometimes glabrous or with some branched trichomes. 15a. Leaves fleshy, glabrous; petals 6.5-8 mm long; caudex thick, 1.5-2 cm diam . 1 < . IP smithii 15b. Leaves not fleshy, often pubescent or rarely glabrous; petals 2—3.5 (^4) mm long; caudex slender, almost always less than 1 cm diam. 16a. Basal leaves filiform to narrowly linear; cauline leaves coarsely dentate-serrate; trichomes minute, 0.03— 0.1 (-0.1 5) mm long. . . . . 11. IP! minutipila 16b. Basal leaves oblong, ovate, lanceolate, or spatulate, very rarely linear; cauline leaves entire to repand or dentate; trichomes coarser, (0.2-)0.4-0.7(-l.l) mm long . 18. IP spathulifolia 14b. Basal and cauline leaves ciliate with simple trichomes. 17a. Basal leaves pinnatifid; inflorescence 1- to 3-flowered; petals less than 2 mm long; stems less than 2 cm tall . 4. IP cymosa 17b. Basal leaves entire, rarely dentate or sinuate; inflorescence (3- or)4- to 10(to 15)-flowered; petals 2-5.5 mm long; stems more than 2 cm tall. 18a. Basal leaves semiterete, thick, linear . 20. IP suffruticosa 18b. Basal leaves flat, thin, oblong to oblaneeolate or spatulate, rare¬ ly linear-lanceolate. 19a. Style obsolete or rarely to 0.6 mm long in fruit. 260 Novon 20a. Fruil torulose; infructescence lax racemes; fruiting pedicel slender, divaricate, 4— 8(-l2) mm long; basal leaves entire, to 1.5 mm wide . 9. IF lagunae 20b. Fruit smooth; infructescence usually dense, sub- umbellate; iruiting pedicels stout, subappressed, 1.5-4.5(— 7) mm long; basal leaves usually dentate, 2 — 4.5( — 6) mm wide . 3. IF colchaguensis 19b. Style 1-3 mm long in fruit, if shorter then fruits conspic¬ uously flattened. 21a. Leaves abaxially with trichomes shorter than those adaxially or on margin; fruit torulose; stems usually leafless; petioles of basal leaves stout, swollen . . 8. IF imbricatifolia 21b. Leaves abaxially glabrous; fruit smooth; stems I - to 4-leaved; petioles of basal leaves slender, not swol¬ len. 22a. Basal leaves entire; petals (3.5— )4— 5 mm long; style (0.8-)1.5-2 mm long in fruit . . 19. IF stenophylla 22b. Basal leaves dentate; petals 2.5— 3.5 mm long; style 0.5— 0.9(— 1.1) mm long in fruit . . 2. IF chillanensis 1. Weberbauera bracteata ((). E. Schulz) ,). F. Macbride, Candollea 5: 356. 1934. Pelagatia bracteata 0. E. Schulz, Pflanzenreich IV. l()5(Heft 86): 192. 1924. TYPE: IVru. An- cash: Pallasca, Cordillera of Pelagatos, 4600 m, 23 J an. 1920, A. Weberbauer 7234 (holo- type, B; isotypes, F, G). Distribution. Known oidy from the type collection. 2. Weberbauera chillauensis (Philippi) Al- Shehbaz, J. Arnold Arbor. 71: 244. 1990. Dra- ba chillanensis Philippi, Anal. Univ. Chile 2: 377. 1862. TYPE: Chile. [Regibn 1 X ] Termas de Chilian, s.d., Philippi s.n. (holotype, SCO). Distribution. Argentina (Prov. Mendoza) and Chile (Region VIII and IX). Representative specimens. ARGENTINA. Mendoza: Malargiie, Banos del Azufre, 19 Jan. 1941, Castellanos s.n. (BA). CHILE. Region VIII: Volean Peteroa, Werder- mann 6 04 (B, BM, E, F, G, GH, K. M, MO, NY, S, U, UC, /). Region IX: Nuble, Cordillera de Chilian, Jaffuel 3722 (GH). 3. Weberbauera colrhugurusis (Barneoud) Al- Shehbaz, J. Arnold Arbor. 71: 241. 1990. Car- damine colchaguensis Barneoud, in Gay, FI. Chile 1: 1 15. 1846. TYPE: Chile. [Region VII] Colchagua, Cordillera del Cajon del Azufre, cerca de volean de Talcaregue, 8000-9000 ft. [2438—2743 rn|. s.d., C. Gay 171 (holotype, P; isotype, G). Distribution. Argentina (Prov. Mendoza, Neu- qubn, and Rfo Negro) and Chile (Region IV, Me- tropolitana, VII, VIII, X, XI, and XII). Representative specimens. ARGENTINA. Mendoza: Laguna Atuel, Bticher, Hjerting & Rahn 1976 (C. MO). Neuquen: Ixjs Lagos, Filo Machete al Co. Rothleugal, Dierre 929 (LIL). Rfo Negro: Cordon del Rfo Colorado, Cerro Gorra. 19 Feb. 1940, Moreau s.n. (BA). CHILE. Region IV: Choapa, La Vega Redonda, E of La Vega Es- condida, Morrison 16995 (1)S, G, K, MO, S, UC). Region Metropolitana: Rio Blancos, Frodin 580 (UPS). Region VII: Colehaqua, Las Damas, Philippi 9 ll> (SCO). Region VIII : Curicb, El Valle de los Ciegos, near volcano of Pe¬ teroa, Bridges 1120 (BM, E). Region X: Cautfn, Villa r- riea, Neger s.n. (M). Region XI: Osorno, Paso Puyehue, Sparre & Constance 10812 (UC). Region XII: Cerro Agu- do, 50°49'S, 72°57'W, Arroyo X Squeo 870017 (C0NC). 4. Weberbauera cymosa Al-Shehbaz, J. Arnold Arbor. 71: 239. 1990. TYPE: Bolivia. La Paz: Murillo, near Palca, base of Illimani, 4800— 5000 m. 25 Feb. 1979,4. Ceballos, A. Chapin , ./. Fernandez- Casas & E. Valddz-bermeijo 543 (holotype, C). Distribution. Endemic to Bolivia (Depto. La Paz). Representative specimens. BOLIVIA. La Paz: Murillo, La Paz-EI Alto-Valle de Zongo, 16°17'S, 68°7'W, Reek- 21752 (LPB, MO). l he species is known thus far only from the two collections cited above. 5. Weberbauera densifolia Al-Shehbaz, ,). Ar¬ nold Arbor. 71: 229. 1990. TYPE: Argentina. Prov. Catamarca: Depto. Santa Marfa, Sierra del Aconquija, 4600 in. 20 Feb. 1925. 5. Ven¬ turi 6610 (holotype, US). Distribution. Argentina (Prov. Catamarca) and Bolivia. Volume 14, Number 3 2004 Al-Shehbaz South American Weberbauera 261 Representative specimens. ARGENTINA. Catamarca: Catamarca, Rodriguez 1360 (MO). BOLIVIA. Without lo¬ cality, Navarro Sanchez 1147 (MO). The species is known thus far only from the three collections cited above. 6. Weberbauera dillonii Al-Shehbaz, sp. nov. TYPE: Peru. La Libertad: Bolivar, Ascenso Nevado de Cajamarquilla, 3000 m, 7°08'S, 77°42'W, 10 Nov. 2001, /. Sanchez V, M. Dil¬ lon & G. Iberico 1 1 172 (holotype, MO; isotypes not seen, CPUN, F). Figure 1. Herba perennis retrorse pilosa, 15—25 cm alta. Folia basalia rosulata, petiolata, oblonga vel oblanceolata, 2—6 X 0.7— 1.5 cm, denticulata, retrorse pilosa; folia caulina sessilia, oblonga vel ovata. grosse dentata. Racemi 20-50- flori, omnino bracteati; pedicelli fructiferi 4.5— 6.5 mm longi, sigmoidei vel recurvati, retrorse pilosi. Sepala ob¬ longa, 2.5—3 X 0.7—1 mm; petala alba, late obovata, 3— 3.5 X 2—2.5 mm. Fructus lineares vel oblongo-lineares, 8—12 X 1.5—2 mm, glabri, curvati; stylus 0.1— 0.2 mm longus; semina oblonga, 1—1.3 X 0.6— 0.7 mm; cotyle- dones incumbentes. Perennial herbs, retrorsely pilose throughout ex¬ cept for petals and fruits; caudex with petiolar re¬ mains of previous years; trichomes simple, retrorse; stems 15—25 cm long, erect or ascending, densely pilose with appressed retrorse trichomes, usually a few from the base, simple or branched above, densely pilose with appressed retrorse trichomes. Basal leaves rosulate; petiole 1—3 cm long, pubes¬ cent as stems; leaf blade oblong to oblanceolate, 2— 6 X 0.7— 1.5 cm, densely and retrorsely pilose on both sides, base cuneate to attenuate, margin den¬ ticulate, apex subacute; middle cauline leaves ses¬ sile, oblong to ovate, 1—2 cm X 4—8 mm, base cu¬ neate, margin coarsely dentate, apex acute, pubescent as basal leaves. Raceme 20- to 50-flow- ered, bracteate throughout, dense, slightly elongat¬ ed in fruit; bracts sessile, cuneate to subauriculate at base, similar to cauline leaves but progressively smaller upward; fruiting pedicels 4.5— 6.5 mm long, sigmoid and slightly recurved, retrorsely pubescent. Sepals oblong, 2.5—3 X 0.7—1 mm, not saccate at base, sparsely pubescent below apex, ascending; petals white, broadly obovate, 3—3.5 X 2-2.5 mm, not clawed, apex rounded; filaments 2-2.5 mm long; anthers oblong, ca. 0.5 mm long. Fruits linear to oblong-linear, 8-12 X 1.5—2 mm, glabrous, curved, slightly flattened at a right angle to the sep¬ tum; valves smooth, not veined; septum complete; style 0.1— 0.2 mm long; stigma entire; seeds 8 to 10 per locule, oblong, brown, 1—1.3 X 0.6— 0.7 mm; funieles slender along entire length; cotyledons in¬ cumbent. Weberbauera dillonii , which is known only from the type collections, is named in honor of Michael Dillon, one of the collectors of the type material who has done extensive fieldwork in Peru. It be¬ longs to a group of four other species with fully bracteate racemes. It is most closely related to W. peruviana , from which it is easily distinguished by having sessile cauline leaves and bracts, retrorsely pilose stems and leaves, glabrous fruits, pilose fruiting pedicels, and broadly obovate petals 2-2.5 mm wide. Weberbauera peruviana has petiolate cau¬ line leaves and bracts, hirsute stems and leaves, sparsely to densely hirsute fruits, glabrous fruiting pedicels, and narrowly spatulate petals 0.7—1 mm wide. The new species resembles (in its small petals < 4 mm long, minute styles to 0.2 mm long, and bracteate racemes) Weberbauera bracteata (0. F. Schulz) J. F. Macbride, a species known only from the type collection made by August Weberbauer on January 1920 from Depto. Ancash and which has not yet been re-collected. From this, W. dillonii is distinguished by having densely and retrorsely pi¬ lose stems and leaves, denticulate and non-ciliate basal leaves, coarsely dentate cauline leaves and bracts, broadly obovate petals 2-2.5 mm wide, sig¬ moid and slightly recurved fruiting pedicels, linear to oblong-linear and curved fruits, veinless valves, and eight to ten seeds per locule. By contrast, W. bracteata has glabrous stems and leaves, lyrate-pin- natifid and ciliate basal and lowermost cauline leaves, entire or repand cauline leaves and bracts, spatulate petals ca. I mm wide, ascending fruiting pedicels subappressed at base to rachis, oblong and straight fruits, prominently veined valves, and three to five seeds per locule. From the other two Peruvian species of Weber¬ bauera with fully bracteate racemes, W. spathuli- folia (A. Gray) 0. E. Schulz and W. herzogii (O. E. Schulz) Al-Shehbaz, W. dillonii is separated by hav¬ ing stems and leaves densely and retrorsely pilose (instead of glabrous or sparsely pubescent with spreading trichomes) and funieles slender along the entire length (instead of distinctly flattened at base). From W. spathulifolia it is also separated by having exclusively simple instead of forked and submalpighiaceous trichomes mixed with simple ones, and from W. herzogii it differs by the minute style 0. 1-0.2 mm long instead of (0.8— )1.2— 2.5 mm long. 7. Weberbauera herzogii (0. E. Schulz) Al- Shehbaz, J. Arnold Arbor. 71: 236. 1990. Sar- codraba herzogii O. E. Schulz, Notizbl. Bot. Gart. Berlin-Dahlem 10: 563. 1929. TYPE: Bolivia Felsplatten der Hiigel iiber dem Titi- 262 Novon Figure 1. Weberbauera dillonii Al-Shehbaz. — A. Plant with remains of infructescence (left) of previous season. — I?. Sepal. — C. Petal. — I). Bract, fruiting pedicel, and fruit. Drawn from the MO holotype (/. Sdnchez V., M. Dillon & G. Iberico 1 1 172) by Al-Shehbaz. Seale: A = 1 cm, B— I) = 1 mm. Volume 14, Number 3 2004 Al-Shehbaz South American Weberbauera 263 cacasee bei Guagqui, ca. 3900 m, s.d., T. Her¬ zog 2510 (holotype, B; isotypes, G. S, Z). Distribution. Argentina (Prov. Jnjny), Bolivia (Depto. La Paz and Potosf), and Peru (Depto. Puno). Representative specimens. ARGENTINA. Jujuy: Hu- mahuaca, Esquinas l>lancas, between Tres Cruces and Hu- mahuaca, Ruthsatz 13/18 (GH). BOLIVIA. I,a Paz: I11- gavei, Guaqui, Asplund 2226 (S, UPS). Potosf: Frias, ca. 25 kin N of Yocalla towards Ventanilla, Wood 11743 (K, MO). PERU. Puno: Puno, Shepard 38 (GH, US). 8. Weberbauera imbricatifolia (Barneoud) Al- Shehbaz. J. Arnold Arbor. 71: 247. 1990. Dra- ba imbricatifolia Barneoud, in Gay, FI. Chile I: 158. 1846. TYPE: Chile. [Region IV] Cor¬ dillera de Coquimbo, 12.000 ft. [3658 m], C. Gay s.n. (holotype, P; isotype. B). Distribution. Argentina (Prov. San Juan) and Chile (Region IV, V, and VI). Representative specimens. ARGENTINA. San Juan: Dept. Iglesia, 31 km W of Arrequintin, 15 Feb. 1985, Hunziberg & Gamenro s.n. (OS). CHILE. Region IV: Co¬ quimbo, Cordillera de Combarbala, Laguna Tibia, 31°17'S, 70°45'W, Jib's 4864 (CONC, M). Region V: Pe- torca, 5 km S of Junta de Piuquenes, Ri'o Sobrante, Mor¬ rison 17290 (l)S, k. UC). Region VI: Santiago, Andes of San Jos6, Gay 1028 (SGO). 9. Weberbauera laguuae (0. E. Schulz) Al- Shehbaz. J. Arnold Arbor. 71: 246. 1990. Sten- odraba suffruticosa (Barneoud) 0. E. Schulz var. lagunae 0. E. Schulz, Notizbl. Bot. Gart. Berlin-Dahlem 10: 469. 1928. TYPE. Chile. Vallenar, Cordillera Laguna Chica, ca. 4000 in, Jan. 1924, E. Werdermann 262 (holotype, B; isotypes, BM, CONC, E, E, G, GH, k, MO, UC). Distribution. Endemic to Chile (Region III). Representative specimens. CHILE. Region III: vicin¬ ity of Laguna Valeriano, Johnston 6 065 (CONC, GH, US). 10. Weberbauera leehleri (Fournier) Al-Sheh¬ baz, comb. nov. Basionvm: Sisymbrium leehleri Fournier, Rech. Crucif. 129. 1865. TYPE: Chile. Terra Pehuenchorum, Dec. 1854, W. Lechler 3080 (holotype, P; isotypes, G[2], K, P[ 2 1). Sisymbrium petraeum Philippi, Linnaea 28: 668. 1856, non S. petraeum (L.) Delarbre, FI. Auv. ed. 2: 349. 1800. Syn. nov. Stenodraba glareosa Ravenna, Nor¬ dic J. Bot. 1: 141. 1981. TYPE: Chile. [Prov. Nuble] “in Andibus prope oppidum Chilian,” R. A. Philippi s.n. (holotype, SGO 49254). Sisymbrium fastigiatum Philippi, Anal. Univ. Chile 41: 670. 1872. Syn. nov. TYPE: Chile. [Prov. Santiago|: Mina Cristo, valley of Maipo, B. Ddvilla s.n. (lecto- type, designated by Munoz-Schick (1973: 30), SGO 45138). Distribution. Argentina (Prov. Neuqufm and Rio Negro) and Chile (Region IX). Representative specimens. ARGENTINA. Neuquen: Cerro Colohuincu 1, Comber 876 E (K). Rfo Negro: Bar- iloche, lago Moreno Este, Hosseus 107 (RAF). CHILE. Re¬ gion IX: Cordillera of Chilian, 1856—1857, P. Germain s.n. (G, K, W). Munoz-Schick (1973) designated the above lec- totype of Sisymbrium fastigiatum, but Ravenna (1981) overlooked that lectotypification and erro¬ neously designated another one of Davilla’s collec¬ tions (SGO 47168) as the lectotype. Ravenna (1981) treated S. petraeum Philippi, S. leehleri, S. caespitosum Philippi, and S. fastigiatum as four in¬ dependent species that he placed in the genus Stenodraba 0. E. Schulz, but the alleged differenc¬ es given by him are trivial and quantitative in nature. All are indistinguishable in every aspect, especially in their trichomes, which are mal- pighiaceous on the basal leaves and dendritic on the stems, cauline leaves, and sepals. The density of trichomes is quite variable in Weberbauera lech- leri, and the holotype has malpighiaceous trichomes restricted to the margins of basal leaves, whereas the isotypes have such trichomes on both leal sur¬ faces. As indicated above, Stenodraba does not merit recognition independent of Weberbauera. Schulz (1924) first placed Weberbauera leehleri in Heterothrix (B. L. Robinson) Rydberg and later (Schulz, 1936) in Pennellia Nieuwland, but he used the illegitimate later homonym Sisymbrium pe¬ traeum Philippi as the basionym for his combina¬ tions. As indicated by Al-Shehbaz (1990c) and Rol¬ lins (1980), the generic name Heterothrix is an illegitimate later homonym for a genus in the Apo- cynaceae, and the correct name for the mustard genus is Pennellia. Pennellia differs from Weber¬ bauera by having a cup-shaped calyx with purple sepals about as long as the petals. Weberbauera has an oblong calyx with green sepals distinctly shorter than the petals. 1 1. Weberbauera niinutipila Al-Shehbaz, J. Ar¬ nold Arbor. 71: 231. 1990. TYPE: Peru. Puno: San Roman, Puno— Arequipa road at km 1 12.8, ca. 4 km (air) E of Tinocopalca (km 119.5), ca. 10 km W of road turnoff to Sta. Lucia (at km 101), ca. 4000 m, 12 Jan. 1963, //. //. & C. M. litis with D. & V. Ugent 1455 (holotype, GH: isotype, WIS). 264 Novon Distribution. Bolivia (Depto. La Paz) anti Peru (l)epto. Arequipa. Puno). Representative specimens. BOLIVIA. La Paz: Valle Chuquiaguillo, Asplund 1888 (S, UPS). PERU. Arequipa: Arequipa, S of Sta. Lucfa on roarl from Puno to Arequipa, ca. 4—5 km E of Sta. Lucfa, litis & llgent 1415 (Vi IS). 12. Weberbauera parvifolia (Philippi) Al-Sheh- baz, J. Arnold Arbor. 71: 248. 1990. Sisym¬ brium parvifolium Philippi, Linnaea 28: 667. 1856. TYPE: Chile. Cordillera de Linares, s.d., Germain s.n. (holotype, SCO). Distribution. Argentina (Prov. Neuquen) and Chile (Region Metropolitana, VII, and VIII). Representative specimens. ARGENTINA. Neuquen: Cordillera del Viento, cruzada de Trieao, Malal al Cajon de Butald, Boelcke et al. 11565 (BAA, SI). CHILE. Re¬ gion Metropolitana: Monumento Natural el Morado, Price 1409 (MO). Region VII: Colchagua, San Fernando, Trrrnas El Falco, Montero 6043 (SI). Region VIII: Cu¬ bed, Termas del Flaco (Andes de Colchagua), Aravena 33349 (G, GH, MO). 13. Weberbauera perforata Al-Shehbaz, Ann. Missouri Rot. Card. 77: 841. 1990. TYPE: Peru. Cuzco: Nevado Auzangate |as Ausanga- te], 71°24'W, 13°38'S, 4800 m. 1 1 May 1957, R. Hirsch PI 255 (holotype, GH). Distribution. Known only from the type collec¬ tion. 14. Weberbauera peruviana (DC.) Al-Shehbaz, comb. nov. Basionym: Sisymbrium peruvianum DC., Syst. Nat. 2: 477. 1821. TYPE: Peru. H. Ruiz & ./. A. Pavdn s.n. (holotype, BM; iso¬ types, B, G-DC). Distribution. Argentina (Prov. Catamarca, Ju- juy, and Tucuman). Bolivia (Depto. Chuquisaca, Cochabamba, La Paz. Oruro, and Potosf), and Peru (Depto. Ayacucho, Cuzco, Huancavelica, Lima, Puno, and Tacna). Representative specimens. ARGENTINA. Catamarca: Cerro Overo, Rriicher 95017 (M). Jiijuy: Humahuaca, Mina Aguilar, 14 km N of Molina de Minera Aguilar. Hun- ziker et al. 10568 (SI). Tucuman: Tafi, cumbres Calcha- (|ufes. Co. Bayo. 20‘1.1'S. 65°42'W, Gomez-Sosa & Miil- gura 204 (M(), SI). BOLIVIA. Clniqnisaca: Zudanez, Cordillera de los Sombreros, along Ida and Azurduy road. Wood A- Carretero 16149 (k. MO). Cochabamba: (,)uilla- collo, camino Sipe Sipe a Kami, cuenca del valle de Co¬ chabamba. Beck et id. 18064 (MO). La Paz: 47.2 km NF of Penas on road to Valle Hiehucota, Solomon 4957 (GH. MO). Oruro: Sajama, Curahura de Carangas, 10 km SW on backload, 68°25'W, 17°50'S, Johns 83-39 (F. I PH. MO). Potosf: 10 km S of Potosf, ILc.sV 6359 (GH, MO). PERU. Ayacucho: Luncanas, Pampa Galeras, Tovar 6704 (MO). Cuzco: Urubamba, Chichero, Cuper, Qoriwayra- china, 72°0'W, 13°25'S, C. & E. Franquemonl 314 (F). Huancavelica: Morococha, Grant 7573 (A, F). Lima: Huarochiri, laguna de Tuctucocha, Cerrate 1866 (GH). Puno: Macusani, road from Ollachea to Macusani, 70°30'W, 13°53'S, M. & K. Weigend 2000/120 (MO). Tac- na: Tarata, Corillera del Barroso, Torre et al. 2135 (MO). As a result of its exclusion from Sisymbrium (Warwick et al., 2002) and the discovery herein of its nearest relative, Weberbauera dillonii , S. peru¬ vianum is appropriately transferred to Weberbauera after being recognized in Sisymbrium since its dis¬ covery more than 180 years ago. In every aspect of morphology (see discussion under W. dillonii ), the species is at home in Weberbauera. 15. Weberbauera relropilosa Al-Shehbaz, J. Arnold Arbor. 71: 234. 1990. TYPE: Bolivia. Depto. La Paz: Prov. Pacajes, Charana, 4000 m, 2 Mar. 1921, Asplund s.n. (holotype, US). Distribution. Endemic to Bolivia (Depto. La Paz). Representative specimens. BOLIVIA. La Paz: Pacajes, Charana, Asplund 2700 (S, UPS). The species is known thus far only from the two collections above. 16, Weberbauera seabrifolia Al-Shehbaz, sp. nov. TYPE: Peru. Iluanuco: Dos de Mayo, road from Pachas to Plata, 4030 m, 9°42'S, 76°47'W. flat puna, 18 Mar. 2001. M. Weigend, K. Weigend. M. Hinder & E. Rodriguez 5220 (holotype, M; isotype, MO). Figure 2. Herba perennis, 3-6 cm alta. Folia basalia petiolata, oblonga, 7-20 X 3—9 mm, integra, superne antrorse sca- bra, inferne glabra; folia eaulina sessilia, oblaneeolata vel linearia, integra. Racemi 10— 27-flori, omnino bracteati; pedicelli fructiferi 4-7 mm longi, recti, pubescentes. Se- pala oblonga, 2. 5- .3 X ca. I mm; petala alba, spathulata, 3.5—4 X ca. 1.5 mm. Fructus lineares vel oblongo-line- ares, 5—9 X ca. 1 .5 mm, glabri, curvati: stylus ca. 1 mm longus; semina oblonga, ca. 1 X 0.5 mm; cotyledones in- cumbentes. Perennial herbs; caudex slender, with leaf re¬ mains of previous years; trichomes simple, op¬ pressed, antrorse; stems 2 to 5 from base, un¬ branched, 3—6 cm long, decumbent or ascending, densely pubescent with oppressed, antrorse tri¬ chomes to 0.6 mm long and to 0.04 mm wide at base. Basal leaves rosulate; petiole 4—10 mm long, ciliate; leal blade oblong, 7—20 X 3—9 mm, abax- ially glabrous, adaxially scabrous with antrorse, ap- pressed trichomes to 0.4 mm long and to 0.12 mm wide at base, base cuneate, margin entire, apex ob¬ tuse; cauline leaves sessile or nearly so, oblanceo- Volume 14, Number 3 2004 Al-Shehbaz South American Weberbauera 265 figure 2. Weberbauera scabrifolia Al-Shehbaz. — A. Plant. — B. Trichomes of adaxial surface of basal leaves. — C. Trichomes of stems and pedicels. — D. Petal. — E. Fruit and fruiting pedicel. — F. Seed and funicle. Drawn from the M holotype (M. Weigend, K. Weigend, M. Hinder & E. Rodriguez 5220) by Al-Shehbaz. Scale: A = 1 cm. B, C = 0.4 mm, D— F = 1 mm. 266 No von late to linear, 5—10 X 1—4 mm, base attenuate, margin entire, upper ones and bracts ciliate and glabrous except midvein, trichomes longer and nar¬ rower than those of basal leaves. Raceme 10- to 27-flowered, bracteate throughout, dense, slightly elongated in fruit; bracts sessile, similar to cauline leaves but progressively smaller upward; fruiting pedicels 4—7 mm long, straight, suberect to as¬ cending, antrorsely pubescent. Sepals oblong, 2.5— 3 X ca. 1 mm, not saccate at base, sparsely [al¬ bescent, ascending; petals white, spatulate, 3. 5-^4 X ca. 1 .5 mm, apex rounded, claw' ca. 1.5 mm long; filaments ca. 2.5 mm long; anthers oblong, ca. 0.5 mm long. Fruits linear to oblong-linear, 5-9 X ca. 1 .5 mm, glabrous, curved, terete; valves smooth, with a distinct midvein; septum complete; style ca. 1 mm long; stigma entire; seeds 5 to 12 per locule, oblong, brown, ca. I X 0.5 mm; funicles thickened at base; cotyledons incumbent. Weberbauera scabrifolia, which is known only from the type gathering, is easily separated from all species of the genus by its appressed, antrorse, sim¬ ple trichomes much thicker and shorter on the bas¬ al leaves than on the rest of the plant and by having basal leaves glabrous abaxially and densely sca¬ brous adaxiallv. 17. Weberbauera smilliii Al-Shehbaz, J. Arnold Arbor. 71: 233. 1990. TYPE: Peru. Ancash: Yungay, Huascaran National Park, Llanganuco Sector, Quebrada Ancosh at Portachuelo, 77°35'W, 9°03'S, 31 Dec. 1984, D. /V. Smith & h. Godwin BB94 (holotype, MO). Distribution. Endemic to Peru (Depto. Ancash). Representative specimen. PERU. Ancash: Yungay, Huascaran National Park, Llanganuco Sector, Quebrada Ancosh at Portachuelo, Smith I I298A (MO). The species is known thus far only from the two collections above. 18. Weberbauera spatbulifolia (A. Gray) 0. E. Schulz, Pflanzenreich IV. 105(1 left 86): 193. 1924. Basionym: Sisymbrium spathulifolium A. Gray, U.S. Expl. Exped. Phan. 15(1): 60. 1854. TYPE: Peru. [Junfn]: Orbajillo, W ilkes Expedition, Anonymous s.n. (holotype, US; iso¬ type, NY). Distribution. Argentina (Prov. Catamarca, Jujuy, and La Rioja), Bolivia (Depto. La Paz, Oruro, and Potosf), and Peru (Depto. Ancash. Huancavelica. Ju¬ nto, Moquegua, Pasco, Puno, and Santa Rosa). Representative specimens. ARGENTINA. Catamarca: Rfo Potrero, Sleumer 1905 (8). jujuy: Hurnahuaca, Mina Aguilar, Sleumer 5402 (LIE). La Rioja: Sierra Famatina, Cueva de Pdrez, Hieronymus A ; Niederlein 376 (B). BO¬ LIVIA. La Paz: Larecaja, vicinity of Combaya, Mandon 914 (BM, G, P, k). Oruro: Abaroa, Challepata, Asplund 3293 (UPS). Potosf: Frias, Cerro Potosf, Petersen & Hjert- ing 1030 (C, LIL). PERU. Ancash: Carhuds, lluascardn National Park, Quebrada Los Cedros, Smith , Valencia & Minaya 9924 (F, GH. MO). Huancavelica: Viseo, Mac- bride & Featherstone 590 (F, G, NY). Moquegua: Mo¬ quegua, above Torata, Weberbauer 7471 (BM, F, G, US). Pasco: Cerro, Cerro de Pasco, Macbride 3065 (CAS, F, US). Puno: Carabaya, Antapampa, Vargas 6837 (F). San¬ ta Rosa: Cuzco, Stafford 512 (BM. k). 19. Weberbauera stenopbylla (Leybold) Al- Shehbaz, J. Arnold Arbor. 71: 245. 1990. Dra- ba stenophylla Leybold, Anal. Univ. Chile 16: 679. 1859. T\ PE: Chile. Cord. Santiago, Cerro Colorado, Mapocho Valley, 6000—7000 ft. [1829—2134 m], s.d., Leybold s.n. (lectotype, designated here, the plate accompanying the original publication). Distribution. Endemic to Chile (Region VIM). Representative specimens. CHILE: Region VIII: El Valle de los Ciegos, near volcano of Petreroa, Bridges 1121 (BM, E, GH, k). As indicated by Al-Shehbaz (1990a), no type material of this species has been found, and the well-illustrated plate accompanying the detailed original description of Draba stenophylla is desig¬ nated herein as the lectotype. 20. Weberbauera suffruticosa (Barneoud) Al- Shehbaz, J. Arnold Arbor. 71: 247. 1990. Dra¬ ba suffruticosa Barneoud, in Cay, FI. Chile 1: 157. 1846. TYPE: Chile. Cordillera Ovalle, 12,000 ft. [3658 m], C. Gay s.n. (holotype, P; isotype, B). Distribution. Endemic to Chile (Region IV). Representative specimen. CHILE. Region IV: Quebra¬ da Larga [30°44'S, 70°23'W], Jiles 3408 (CONC). 21. Weberbauera tricliocarpa (Muschler) J. F. Macbride, Candollea 5: 356. 1934. Basionym: Eudema trichocarpum Muschler, Bot. Jarhb. Syst. 40: 276. 1908. TYPE: Peru. Above Lima, near Alpamia [Alpamina], 4500 m, 2 Mar. 1904, A. Weberbauer 5119 (holotype, B). Distribution. Endemic to Peru (Depto. Ancash, Lima, and Pasco). Representative specimens. PERU. Ancash: Pallasca, Conchucds, Weberbauer 7229 (B. F, GH). Lima: Rfo Blan¬ co, Macbride 2990 (F, GH. MO, NY. US). Pasco: Cerro de Pasco, Macbride 3073 (CAS, F, GH, MO, NY, US). 22. Weberbauera violacea Al-Shehbaz, sp. nov. Volume 14, Number 3 2004 Al-Shehbaz South American Weberbauera 267 Figure 3. Weberbauera violacea Al-Shehbaz. — A. Plant. — B. Bract. — C. Sepal. — D. Petal. — E. Young fruit. Drawn from the MO holotvpe (A. Sagdstegui A. et al. 11175) by Al-Shehbaz. Scale: A = 1 cm. B-E = 1 mm. TYPE: Peru. Cajamarca: Cajabamba, Caja- bamba-Luchubamba, jalca vegetation, 3800 m, 1 7 Nov. 1983, A. Sagdstegui A. et al. 11175 (holotype, MO; isolype, HUT). Figure 3. Herba perennis, 4—15 cm alta. Eolia basalia rosulata, petiolata, spathulata, 1. 5-3.5 X 1-2.2 cm, integra vel re¬ panda; folia caulina sessilia, oblonga vel oblanceolata, in¬ tegra vel repanda, basi ciliata. Racemi 20— 35-flori, om- nino bracteati; pedicelli Horiferi 7-10 mm longi, recti. 268 Novon subappressi, glabri. Sepala oblonga, 4—5 X 1.5—2 mm; petala violacea, late obovata, 6.5— 7.5 X 3—3.5 mm; stylus 1 .5—2 mm longus. Perennial herbs, subglabrous or sparsely pubes¬ cent along petiole ami at base of cauline leaves; trichomes simple, spreading, to 1.5 mm long; eau- dex thick, simple; stems 4—15 cm long, decumbent or ascending, unbranched, glabrous. Basal leaves rosulate; petiole 0.5-3. 5 cm long, ciliate; leaf blade spatulate, 1.5— 3.5 X 1-2.2 cm, glabrous, base eu- neate to attenuate, margin entire or repand, apex rounded; cauline leaves sessile, oblong, 6—20 X I - 5 mm, base cuneate, margin entire or repand, apex obtuse, sparsely ciliate at base. Raceme 20- to 35- flowered, bracteate throughout, dense; racliis straight, glabrous; bracts sessile, similar to cauline leaves but progressively smaller upward; flowering pedicels 7—10 mm long, erect to ascending, straight, subappressed, glabrous. Sepals oblong, 4— 5 X 1 .5—2 mm, not saccate at base, glabrous or with a few trichomes below apex, ascending; petals violet, broadly obovate, 6.5— 7.5 X 3—3.5 mm, not clawed, apex rounded; filaments 2—3 mm long; an¬ thers oblong, 0.6— 0.8 mm long; style 1.5-2 mm long. Immature fruits linear, ca. 8X1 mm; valves glabrous, with a conspicuous midvein; seeds not seen. Weberbauera violacea, which is known only from the type collection, is readily distinguished from all other species of Weberbauera by having violet, large petals 6.5— 7.5 mm long and racemes bracteate throughout. All other species of the genus have white or rarely yellow (If; perforata) flowers almost always less than 5 mm long. The only exception is W. smithii, a species with ebracteate racemes and white flowers 6.5—8 mm long. Petal size and color easily separate the new species from those with ful¬ ly bracteate racemes (see lead 2a in the key above). Weberbauera violacea is most closely related to W. herzogii and W. scabrifolia. All three differ from the remaining species of Weberbauera by a combi¬ nation of fully bracteate racemes, attenuate, slender styles 1-3 mm long, and receptacle considerably wider than the pedicel. Acknowledgments. I am much indebted to Olga Martha Montiel for translating the summary to Spanish, to Henk van der Werff for correcting the Latin, to Michael Dillon for sending his Peruvian collections of mustards for my study, and to Susan- ne Renner for sending a loan of South American mustards and for allowing a plant of the type col¬ lection of Weberbauera scabrifolia he deposited at MO. 1 am grateful to Steve L. O' Kane Jr., Neil A. Harriman, and Victoria C. Hollowell for valuable comments on the manuscript. I thank the directors and curators of the herbaria cited above. I .iterature Cited Al-Shehbaz, I. A. 1988. The genera of Sisymbrieae (Cru- eiferae; Brassicaceae) in the southeastern United States. .1. Arnold Arbor. 69: 213—237. - . 1990a. A revision of Weberbauera (Brassicaceae). J. Arnold Arbor. 71: 221-250. - . 1990b. Weberbauera perforata (Brassicaceae), a new species from Peru. Ann. Missouri Bot. Card. 77: 481-482. - . 1990c. New or noteworthy species in the South American genera Mancoa, Pennellia, and Sisymbrium. Harvard Pap. Bot. 2: 1 1—16. Appel, 0. & I. A. Al-Shehbaz. 2003. Cruciferae. Pp. 75— 174 in: K. Kubitzki (editor). Families and Genera of Vascular Plants 5. Springer- Verlag, Berlin, Heidelberg. Boelcke, O. & C. Uomanczuk. 1984. Cmciferae. FI. Pa- tagonica, Vol. 4A: 373—544. INTA, Buenos Aires. Munoz-Schick. M. 1973. Complemento de “Las especies de plantas deseritas por H. A. Philippi durante del siglo XIX.” Anal. Univ. Chile 128: 5-69. Ravenna, P. 1981. Taxonomic notes on certain Chilean Cruciferae. Nordic J. Bot. 1: 140—142. Rollins, R. C. 1980. The genus Pennellia in North Amer¬ ica. Contr. Gray Herb. 210: 5-21. Schulz, (). F. 1924. Cruciferae— Sisymbrieae. Pp. 1-388 in: A. Engler (editor). Das Pflanzenreich IV. 105(Heft 86). Verlag von Wilhelm Engelmann, Leipzig. - . 1936. Cruciferae. Pp. 227—658 in: A. Engler & H. Harms (editors). Die natiirlichen Pflanzenfamilien, ed. 2. 17B. Verlag von Wilhelm Engelmann, Leipzig. Warwick, S. I., 1. A. Al-Shehbaz, R. A. Price & C. Sander. 2002. Phylogeny of Sisymbrium (Brassicaceae) based on ITS sequences of nuclear ribosomal DNA. Canad. J. Bot. 80: 1002-1017. Transfer of Arabidopsis gamosepala and Torularia brachycarpa to Braya (Brassicaceae) Ihsan A. Al-Shehbaz Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. ihsan.al-shehbaz@mobot.org Suzanne I. Warwick Eastern Cereal and Oilseed Research Centre, Agriculture and Agri-Food Canada, Central Experimental Farm, Ottawa, Ontario K1A 0C6, Canada. warwicks@agr.gc.ca Abstract. The new combinations Braya gamo¬ sepala and B. brachycarpa (Brassicaceae) are pro¬ posed. The distinguishing characters, especially of habit and fruit and trichome morphology, separating Arabidopsis, Braya, anti Neotorularia (formerly To¬ rularia) are discussed. Key words: Arabidopsis, Brassicaceae, Braya, Neotorularia. Arabidopsis gamosepala Hedge (Brassicaceae), which is endemic to Afghanistan (Hedge, 1968), was transferred to Neotorularia Hedge & J. Leonard by Al-Shehbaz and O’Kane (1997) and has been maintained in that genus to the present. It was ex¬ cluded from Arabidopsis Heynhold by these authors because it has pubescent, torulose fruits, and den- dritieally branched trichomes. As delimited by O’Kane and Al-Shehbaz (1997), Al-Shehbaz et al. (1999), and Al-Shehbaz and O’Kane (2002), Ara¬ bidopsis lias glabrous, non-torulose fruits and a mixture of simple, forked, or rarely stellate stalked trichomes. Torularia brachycarpa Vassilczenko, originally collected only from Tajikistan (Vassilczenko, 1939), is now documented as widespread in four provinces in China (Zhou el ah, 2001). It was transferred to Neotorularia (Leonard, 1986) because Torularia (Cosson) 0. E. Schulz is an illegitimate later hom¬ onym of the red algae Torularia Bonnemaison. Recent phylogenetic studies, based on ITS se¬ quences of nuclear ribosomal DNA and frnL intron of chloroplast DNA (Warwick et ah, 2004), on Braya Sterenberg & Hoppe and Neotorularia (dear¬ ly showed that Arabidopsis gamosepala, Neotoru¬ laria brachycarpa (Vassilczenko) Hedge & Leonard, and B. humilis (C. A. Meyer) B. L. Robinson (N. humilis (C. A. Meyer) Hedge & Leonard) are well nested within Braya and are most closely related to species of this genus with linear fruits, including B. alpina Sterenberg & Hoppe, the type species of Braya. These four species differ from all members of Neotorularia, including the generic type N. to- rulosa (Desfontaines) Hedge & Leonard, by being perennials with a well-defined basal rosette (instead of annuals with no basal rosette) and by having at least basally bracteate instead of ebracteate ra¬ cemes. On the basis of molecular and morphologi¬ cal data, Arabidopsis gamosepala and Neotorularia brachycarpa are best accommodated in Braya, and their transfer to this genus is herein effected. Braya brachycarpa (Vassilczenko) Al-Shehbaz & Warwick, comb. nov. Basionym: Torularia bra¬ chycarpa Vassilczenko, in Komarov, Fh URSS 8: 635. 1939. TYPE: Tajikistan. Pamir: near Ak-baital, confluence of Murgab, ca. 3900 m. 7 July 1901. M. 1. Alexeenko 2391 (holotype, LE). Braya brachycarpa is easily distinguished from the remaining species of Braya by having racemes bracteate throughout or rarely only along the low¬ ermost part, oblong to linear fruits widest basally and 3— 10(— 15) mm long, and seeds sub-biseriate only at the basal portion of the fruit. Braya gamosepala (Hedge) Al-Shehbaz & War¬ wick, comb. nov. Basionym: Arabidopsis ga¬ mosepala Hedge, in K. II. Rechinger, Flora Iranica 57: 334. 1968. TYPE: Afghanistan. Munjan: above Anjuman valley, near Anju- man, 3100 m, 14 Aug. 1965, I). Podlech 12379 (holotype, M: isotypes, E. M). The occurrence of gamosepaly, although rather rare in the Brassicaceae (e.g., Brayopsis gamose¬ pala Al-Shehbaz, Desideria mirabilis Pampanini, Sisymbrium gamosepalum Hedge), is not consid¬ ered to be a useful character at the generic rank (Al-Shehbaz, 2001), but readily distinguishes B. Novon 14: 269-270. 2004. 270 Novon gamosepala from t lie remaining species of the ge¬ nus. Acknowledgments. We are grateful to Thomas G. hammers and Victoria C. Hollowed for their crit¬ ical comments on the manuscript. I .iterature Cited Al-Shehbaz, I. A. 2001. A review of gamosepaly in the Brassicaceae and a revision of Desideria, with a critical evaluation of related genera. Ann. Missouri Bot. Card. 87: 549-563. - & S. L. O’Kane Jr. 1997. Arabidopsis gamosepala and A. tuemurnica belong to Neotorularia (Brassica- ceae). Novon 7: 93—94. - & - . 2002. Taxonomy and phylogeny of 4r- abidopsis (Brassicaceae). (22 August, 2002). In: C. R. Somerville & E. M. Meyerow it/, (editors). The Arabidop¬ sis Book, American Society of Plant Biologists, Rock¬ ville, Maryland, doi/10. 1 199/tab. 0001 (http:// www.aspb.org/publications/arabidopsis/). - . - & R. A. Price. 1999. Generic placement of species excluded from Arabidopsis. Novon 9: 296— 307. Hedge. I. C. 1968. Sisymbrieae. Pp. 309—342 in: K. 11. Rechinger (editor), Flora Iranica 57. Akademisehe Druck-u.Verlagsanstalt, Graz, Austria. Leonard, J. 1986. Neotorularia Hedge and J. Leonard nom g^n^rique nouveau de Cruciferae. Bull. Jard. Bot. Nat. Belg. 56: 389-395. O’Kane, S. I... Jr. & I. A. Al-Shchbaz. 1997. A synopsis of Arabidopsis. Novon 7: 323—327. Vassilczenko, I. T. 1939. Sisymbrium. Pp. 59—69, 635— 636 in: V. E. Komarov (editor), Flora URSS, Vol. 8. Akademii Nauk SSSR, Moscow and Leningrad. Warwick, S. I., I. A. Al-Shehbaz, C. Sander. J. G. Harris & M. Koch. 2004. Phylogeny of Braya and Neotorularia based on ITS sequences of nuclear ribosomal l)NA and trn I. intron of chloroplast I)NA. Canad. J. Bot. 82: 376— 392. Zhou, T. Y., I,. L. Lu, G. Yang & I. A. Al-Shehbaz. 2001. Brassicaceae (Cruciferae). Pp. 1—193 in Z. Y. Wu & P. H. Raven (editors), Flora of China, Vol. 8. Science Press, Beijing, and Missouri Botanical Garden Press, St. Louis. Shangrilaia (Brassicaceae), a New Genus from China Ihsan A. Al-Shehbaz Missouri Botanical Garden, P.0. Box 299, St. Louis, Missouri 63166-0299, U.S.A. ihsan.al-shehbaz@mohot.org Jipei Yue and Hang Sun Herbarium, Kunming Institute of Botany, Chinese Academy of Sciences, Heilongtan, Kunming, Yunnan 650204, People’s Republic of China, yuejipei@mail.kib.ac.cn; hsun@mail.kib.ac.cn ABSTRACT. The new genus and species Shangri¬ laia nana (Brassicaceae) are described and illus¬ trated, and their relationships to distinguishing characters from Braya and Baimashania are dis¬ cussed. Key words: Baimashania, Brassicaceae, Braya, China, Shangrilaia, Yunnan. Although a comprehensive account of die Bras¬ sicaceae for the Flora of China has recently been published (Zhou et al., 2001), many new taxa and new nomenclatural adjustments have since been added for China, especially those that are based on plants from Sichuan, Tibet (Xizang), Yunnan, and Xinjiang provinces or autonomous regions (Al- Shehbaz, 2002a, 2002b, 2002c, 2002d, 2003, 2004a, 2004b; Al-Shehbaz & Koch, 2003; Al- Shehbaz & Warwick, 2004; Al-Shehbaz et ah, 2002a, 2002b). Such additions clearly indicate that our knowledge of the Chinese mustards, especially from the provinces and regions above, is far from being complete, anil further fieldwork is likely to bring additional novelties. A case in point is the present paper, which describes a new genus and species, Shangrilaia nana, from Yunnan Province. Shangrilaia Al-Shehbaz, J. P. Yue & H. Sun, gen. nov. TYPE: Shangrilaia nana Al-Shehbaz, J. P. Yue & H. Sun. Herba nana, perennis, scaposa, pulvinata. Folia acicu- lari-linearia, ciliata, integra, persistentia, dense imbricata, basi complanata, pilis simplicibus instructa. Flores soli- tarii. Sepala ovata, nonsaccata. Petala alba, spathulata. Glandulae nectariferae confluentes. Ovula 6—12. Fructus ovoidei, teretes, breviter stipitati, valvis pubescentibus coriaceis; stylus tenuis, ad 1 mm longus; stigma integrum. Semina uniseriata, ovoidea; cotyledones incumbentes. Dwarf, perennial, scapose, pulvinate herbs, with simple or few-branched caudex covered with leaves of previous years. Trichomes simple. Stems densely leafy, internodes obsolete. Leaves acicular-linear, thick, densely overlapping, ciliate, persistent, with flattened, triangular base. Flowers solitary, termi¬ nating stem. Sepals ovate, deciduous, erect, not saccate at base. Petals white, spatulate, obtuse, clawed. Stamens 6, tetradynamous; filaments slen¬ der at base; anthers ovate, not apiculate at apex. Nectar glands confluent, subtending bases of all stamens; median glands present. Ovules 6 to 12 per ovary. Fruit dehiscent silicle, ovoid, terete, short stipitate; valves leathery, with a distinct midvein at least basally, pubescent; replum flattened at least basally; septum perforate, veinless; style slender, to 1 mm long; stigma capitate, entire. Seeds uniser- iate, wingless, ovoid, not mucilaginous when wet¬ ted; cotyledons incumbent. Initial attempts to identify the plants on which Shangrilaia is based, using Zhou et al. (2001), Ap¬ pel and Al-Shehbaz (2003), or the interactive keys ((http://flora.huh. harvard. edu:8080/actkey/index.jsp)) to the Brassicaceae genera of the world or that of the flora of China, yielded unsatisfactory results. Those keys inadequately led to either Baimashania Al-Shehbaz or Braya Sternberg & Hoppe, and the characters of the new genus did not match those of these two genera. It became abundantly clear that a new genus is needed to accommodate them. Shangrilaia, which is named after the city Shangri- La (formerly Zhong Dian), differs from both Bai¬ mashania and Braya in having exclusively simple instead of a mixture of simple and branched tri¬ chomes, perforate instead of complete septa, and acicular-linear instead of distinctly flattened leaves. It al so differs from Baimashania in having incum¬ bent instead of aceumbent cotyledons, ovoid and terete instead of linear and latiseptate (flattened parallel to the septum) fruits, thick leathery instead of papery valves, veinless instead of veined septum, and flattened instead of rounded replum. From Novon 14: 271-274. 2004. 272 Novon l*'igure I. Shangrilaia nana Al-Shehbaz, J. P. Yue & H. Sun. — A. Plant. — B. Leaf. — C. Portion of leaf showing abaxial surface. — I). Sepal. — -E. Petal. — F. Fruit and pedicel with valve removed to show seed and perforated septum. — G. Fruit valve. — H. Section of seed showing incumbent cotyledons. Scale: A = 1 cm; B, I)— G = I mm; C = 0.2 mm. II = 0.5 mm. Drawn by Al-Shehbaz from the holotype (Yue 0366). Volume 14, Number 3 2004 Al-Shehbaz et al. Shangrilaia from China 273 Braya, it is also readily distinguished by having solitary flowers at the end of the stem instead of flowers in racemes or corymbs on a well-developed peduncle, as well as in having confluent instead of four separate nectar glands. One species assigned to Braya by Zhou et al. (2001), B. forrestii W. W. Smith, was described as having exclusively simple trichomes. However, mo¬ lecular studies by Warwick et al. (2004) clearly demonstrated that this species is quite distinct from the rest of Braya and perhaps should be excluded from that genus. Shangrilaia nana Al-Shehbaz, .). P. Vue & H. Sun, sp. nov. TYPE: China. Yunnan: Shi-ka Shan, near Shangri-La, scree slope, sandy area, 27°47'N, 99°35'E, 4200 m, 5 Aug. 2003, Ji- pei Yue 0366 (holotype, MO; isotype, KUN). Figure 1. Herba pulvinata 1—2.5 cm alta, caudice simplici vel pauciramoso. Pili 0.2-0. 5 mm longi. Folia aciculari-li- nearia, 2—6 X 0.2— 0.4 mm, dense imbricata, sparse cil- iata, basi triangulari, 1-1.5 X 1-1.5 mm. Flores solitarii. Pedicelli fructiferi 0.5—2 mm longi, crassi. Sepala ovata, glabra, 2.5—3 X 1.5—2 mm. Petala alba, spathulata, 4—5 X 1.5— 2.5 mm, unguibus 2—3 mm longis. Ovula 6—12. Fructus ovoidei, teretes, breviter stipitati, valvis 3—3.5 X 2—2.5 mm, pubescentibus coriaeeis, septis ad maturitatem perforatis; stylus 0.4—1 mm longus. Semina ovoidea, 1 — 1 .2 X 0. 6-0.8 mm. Pulvinate herbs 1-2.5 cm tail, caudex simple or few-branched. Trichomes simple, 0.2— 0.5 mm long. Leaves acicular-l inear, 2—6 X 0.2— 0.4 mm, densely overlapping, with a prominent midvein abaxially; base flattened, subtriangular, 1—1.5 X 1-1.5 mm, margin entire, sparsely ciliate to subsetose at least basally or up to the middle, apex obtuse, often with 1 to 3 trichomes. Flowers solitary, in leaf axils, 1 to 4 terminating the stem. Fruiting pedicels stout, somewhat angled, 0.5—2 mm long, glabrous or sparsely pubescent with straight, horizontal tri¬ chomes, distinctly expanded at receptacle. Sepals ovate, glabrous, 2.5—3 X 1 .5—2 mm, with prominent veins. Petals white, spatulate, 4—5 X 1.5— 2.5 mm, obtuse; claw 2—3 mm long, slender at base. F ila¬ ments of median stamens 2—2.5 mm long, those of lateral stamens 1 — 1.2 mm long; anthers ovate, 0.5— 0.6 mm long. Ovules 6 to 12 per ovary. Fruit ovoid, terete; valves leathery, 3—3.5 X 2—2.5 mm, with Hat margin and prominent midvein at least basally, pu- berulent with simple trichomes; septum initially complete, becoming perforate at seed's maturity; gynophore stout, slightly wider than pedicel, 0.2— 0.7 mm long; style slender. 0.4—1 mm long. Seeds ovoid, 1-1.2 X 0.6— 0.8 mm. Fruiting August— Sep- tember. Habitat. Scree slopes on sandy soil; 4180— 4200 m. Yunnan. Both the type collection and the paratype have ample fruits, and the floral description is based on two flowers found on two plants. Collection of flow¬ ering material should be done in late June or early July. There is some variation in the pubescence of leaves, and most plants examined have trichomes at the leaf base and usually one or a few at the leaf apex. Although there are up to six ovules in each locule, usually one or two develop into mature seeds. The septum is complete during early fruit development, and fruits with mature seeds always have perforate septa. Paratypes. CHINA. Yunnan: Shika Shan, Zhong Dian, 27°47'N, 99°35'E, 4180 m, 27 Sep. 2001, J. P Yue 054 (KUN, MO). Acknowledgments. We are most grateful to Henk van der Werff for correcting the Latin and Victoria C. Hollowell for the editorial advice. We thank Fernand Jacquemoud and an anonymous re¬ viewer for their valuable comments. We are much indebted to the Yunnan Natural Science Founda¬ tion (grants-in-aid 2002C0060M), the National Nat¬ ural Science Foundation of China (grants 30270121 and 40332021), and the Innovation Program of the Chinese Academy of Science (grant KSCX2-1-09), all of which supported fieldwork and research of one of us (J. P. Yue). Literature Cited Al-Shehbaz, 1. A. 2002a. New species of Alyssum, Aphrag- mus, Arabis and Sinosophiopsis (Brassieaceae) from Chi¬ na and India. Novon 12: 309—313. - . 2002b. Six new species of Draba (Brassieaceae) from the Himalayas. Novon 12: 314—318. - . 2002c. New combinations in Brassieaceae (Cru- ciferae): Draba serpens is a Hemilophia and D. william- sii is a Lepidostemon (Brassieaceae). Edinburgh J. Bot. 59: 443-450. - . 2002d. Noccaea nepalensis, a new species from Nepal, and four new combinations in Noccaea (Brassi- caceae). Adansonia 24: 89—91. - . 2003. Aphragmus bouffordii, a new species from Tibet and a synopsis of Aphragmus (Brassieaceae). Har¬ vard Pap. Bot. 8: 25—27. - . 2004a. Novelties and notes on miscellaneous Asian Brassieaceae. Novon 14: 153—157. - . 2004b. Two new species of Draba (Brassieaceae): I). mieheorum from Tibet and D. sagasteguii from Peru. Novon 14: 249-252. - & M. Koch. 2003. Drabopsis is united with Draba (Brassieaceae). Novon 13: 173—174. - & S. 1. Warwick. 2004. Transfer of Arabidopsis gamosepala and Torularia brachycarpa to Braya (Bras- sicaceae). Novon 14: 269—270. - . K. Mummenhoff & 0. Appel. 2002a. Cardaria , Coronopus, and Stroganowia are united with Lepidium (Brassieaceae). Novon 12: 5—11. 274 Novon - , B. Bartholomew, A. Abbas & A. Tumur. 2(H)2b. New or noteworthy species of Brassicaceae for the Flora of China. Harvard Bap. Bot. 7(1): 23—24. Appel, 0. & I. A. Al-Shelibaz. 2003. Cruciferae. Bp. 75- 174 in: K. Kubitzki (editor). Families and Genera of Vascular Plants 5. Springer- Verlag, Berlin, Heidelberg. Warwick, S. I., I. A. Al-Shehbaz, C. Sauder, J. G. Harris & M. Koch. 2004. Bhylogeny of Bray a and A 'eotorularia (Brassicaceae) based on nuclear ribosomal ITS and chloroplast trn I . intron sequences. Canad. J. Bot. 82: 376-392. Zhou, T. Y., L. L. Lu, G. Yang & 1. A. Al-Shehbaz. 2001. Brassicaceae. Bp. 1-193 in: Z. Y. Wu & B. H. Raven (editors), F lora of China, Vol. 8. Science Cress, Beijing, and Missouri Botanical Garden Cress, St. Louis. New Combinations in Silphium (Asteraceae: Heliantheae) Jennifer A. Clevinger Department of Biology, James Madison University, Harrisonburg, Virginia 22807, U.S.A. clevinja@jmu.edu Abstract. As part of a revision of Silphium (As¬ teraceae: Heliantheae) for the Flora of North Amer¬ ica (FNA), four new combinations are proposed: S. asteriscus var. simpsonii (Greene) J. A. Clevinger, S. asteriscus var. trifoliatum (L.) J. A. Clevinger, S. asteriscus var. latifolium (A. Gray) J. A. Clevinger, and .S', radula var. gracile (A. Gray) J. A. Clevinger. In addition, Silphium confertifolium Small is re¬ duced to synonymy of Silphium asteriscus var. la¬ tifolium, and Silphium simpsonii var. wrighl ii L M. Perry is reduced to synonymy of Silphium radula Nuttall. Silphium radula var. gracile is lectotypi- fied. Key words: Asteraceae, Compositae, Helian¬ theae, North America, Silphium. As part of a revision of Silphium L. (Asteraceae: Heliantheae) in Clevinger’s (1999) dissertation, 7000 herbarium specimens from 16 herbaria were examined and populations in 22 states were visited. A molecular phylogenetic study of Silphium ex¬ plored relationships in Silphium resulting in the recognition of two monophyletic sections: sect. Sil¬ phium anil sect. Composita (Clevinger & Panero, 2000). As a result of these studies and subsequent eolleeting, four nomenclatural changes in Silphium sect. Silphium are proposed. Three species names are reduced to varietal rank, and one varietal name is transferred to a different species. These nomen¬ clatural changes pave the way for the taxonomic treatment of Silphium that is currently being [ire- pared for the Flora of North America. Silphium asteriscus 1 .. The most current circumscription of Silphium as¬ teriscus can be found in Cronquist (1980). His con¬ cept of Silphium asteriscus includes three varieties: var. asteriscus, var. angustatum A. Gray, and var. scabrum Nuttall. Phis taxonomic concept should be expanded to include S. dentatum Elliott, S. trifol¬ iatum L., and S. confertifolium Small, all of which Cronquist recognized as species. It should also in¬ clude S. simpsonii Greene, which Cronquist placed in synonymy of S. gracile A. Gray. These taxa all share the characteristics of having a variable num¬ ber of ray flowers ranging from 8 to 21 and lance¬ olate to ovate leaves. Cronquist used phyllotaxy and presence or absence of pubescence on stems, leaves, and paleae to distinguish between these species. However, my field observations reveal var¬ iability in these characters within populations found where the geographical ranges of these taxa overlap. This variability is symptomatic of intro- gression between these taxa. Therefore, because of the introgression, they should be recognized at the varietal level rather than as separate species. Silphium dentatum already has varietal status as S. asteriscus var. dentatum (Elliott) Chapman and will be recognized as such (Chapman, 1860). Sil¬ phium trifoliatum var. trifoliatum, S. trifoliatum var. latifolium A. Gray, and S. simpsonii should he transferred herein to S. asteriscus. Silphium confer¬ tifolium will be placed in synonymy with Silphium asteriscus var. latifolium. Silphium asteriscus L., Sp. El. 2: 920. 1753. TYPE: “Asteriscus coronae solis flore et facie” in Dillenius, Hurt. Eltham., 42, t. 37, f. 42. 1732 (lectotype, designated by Perry (1937: 292)). Silphium asteriscus var. simpsonii (Greene) J. A. Clevinger, comb. nov. Basionym: Silphium simpsonii Greene, Pittonia 4: 45. 1899. TYPE: U.S.A. Florida: Palma Sola, in damp ground in pine barrens, 8 July 1890, J. //. Simpson 81 (holotype, US). Silphium asteriscus var. simpsonii is most closely allied to S. asteriscus var. dentatum and S. asteriscus var. angustatum. They share the characteristic of stipitate glands on the paleae. Stipitate glands are absent in other varieties of Silphium asteriscus but are also found in Silphium laciniatum L., Silphium perplexum J. Allison, and Silphium glutinosum J. Allison. Additionally, Silphium asteriscus var. simp¬ sonii, variety dentatum, and variety angustatum, along with variety asteriscus and variety scabrum, share the characteristics of predominantly alternate phyllotaxy and scabrous to hispid leaves. This is in contrast to the predominantly opposite or whorled phyllotaxy and glabrous leaves of variety trifoliatum Novon 14: 275-277. 2004. 276 Novon and variety latifolium. Silphium asteriscus var. simpsonii can be distinguished from its closest al¬ lies, variety dentatum anil variety angustatum, by its height (0.2— 1.1 in vs. 0.5— 1.5 m) and persistent basal rosette. Silphium asteriscus var. simpsonii is restricted to the southeastern U.S. and is primarily found in Florida. Perry (1937) divided Silphium simpsonii into two varieties. Silphium simpsonii var. wrightii Perry, found in the southwestern U.S., should not be included here. Rather, because of its large num¬ ber of ray flowers (20 to 30+), I reduce Silphium simpsonii var. wrightii to synonymy of ,S. radula var. radula. During the summer of 2002, Silphium as¬ teriscus var. simpsonii was collected in Florida and added to the Clevinger and Panero (2000) molec¬ ular data set for phylogenetic analysis. In the re¬ sulting phylogenetic tree, Silphium asteriscus var. simpsonii was allied with Silphium asteriscus rather than Silphium radula. Silpliimn asteriscus var. trifoliatiiin (L.) J. A. Clevinger, comb. nov. Basionym: Silphium tri- foliatum L., Sp. PI. 2: 920. 1753. Silphium ternifolium Michaux, FI. Bor. Amer. (Mi- chaux) 2: 146. 1803. TYPE: “Habitat in Vir¬ ginia” (lectotype, designated by Beveal in Jarvis & Turland (1998: 367), LINN micro¬ fiche # 1032.8, photos, F. GH). Fieldwork shows that Silphium asteriscus var. tri- foliatum is most closely allied to Silphium asteris¬ cus var. latifolium. They share the characteristics of glabrous leaves, peduncles, and stems in contrast to all other varieties of Silphium asteriscus. Sil¬ phium asteriscus var. trifoliatum is easily distin¬ guished in the northeastern U.S. because of its tri¬ foliate leaf arrangement. However, these same populations also contain individuals with alternate, opposite, or 4-foliate leaf arrangements. Along the southern edge of the varietal range, where it over¬ laps with S. asteriscus var. asteriscus and S. aster¬ iscus var. latifolium. these alternate, opposite, and 4-foliate leaf arrangements become more predomi¬ nant. Silpliiuiu asteriscus var. lutil'oliuiu (A. (fray) J. A. Clevinger, comb. nov. Basionym: Silphium trifoliatum L. var. latifolium A. Gray, Syn. FI. N. Amer. 1: 241. 1884. TYPE: U.S. A. Ala¬ bama: July 1840, S. B. Buckley s.n. (holotype, GH). Silphium confertifolium Small. FI. S.E. U.S. 1243, 1340. 1903. Syn. nov. TYPE: U.S. A. Alabama: Choctoaw Co., Cocoa, 13—15 Oct. 1896, C. Schuchert s.n. (ho¬ lotype. NY; isotypes, NY. US). Like the other varieties of Silphium asteriscus, variety latifolium has ray flowers that number 8 to 21. This variety is distinguished by its predomi¬ nantly opposite phyllotaxy and glabrous leaves and stems. In my fieldwork, I found that when Silphium asteriscus var. latifolium populations come in con¬ tact with variety asteriscus there is a mix of glabrous and pubescent plants. When its populations come in contact with variety trifoliatum, there is a mix of opposite and trifoliate leaf arrangements on the plants. Because of this introgression that I observed in the held, variety latifolium is placed in S. aster¬ iscus. I conclude that Silphium confertifolium repre¬ sents an environmental variant of S. trifoliatum var. latifolium and should not retain its status as a dis¬ tinct species. This is based upon examination of herbarium specimens and visits to sites in Dallas Co., Alabama, which is the locality of historic S. confertifolium collections {Godfrey 6 9684, Jones 752, NCU). Cronquist (1980) distinguished these two species on the basis of height and leaf distri¬ bution. Silphium confertifolium ranges in height from 0.4 to I m with basally disposed leaves, while Silphium trifoliatum var. latifolium ranges in height from 1 to 2 m with evenly distributed leaves. In the field I observed a continuum of plants within pop¬ ulations that varied in stature and leaf distribution according to proximity to chalk outcroppings. The plants closest to the chalk outcroppings were al¬ ways the shortest in height with reduced leaves, and the ones furthest were tallest in height with evenly distributed leaves. The recognition of Silphium asteriscus var. trifol¬ iatum and variety latifolium is in agreement with the dissertation work of W. B. Weber (1968). Weber conducted extensive hybridization studies and con¬ cluded that S. asteriscus and S. trifoliatum are not reproductively isolated. Weber also concluded that phyllotaxy was variable between these two species. Weber and his adviser, T. B. Fisher, proposed in Weber’s dissertation that S. trifoliatum var. trifol¬ iatum and S. trifoliatum var. latifolium should be reduced to varietal rank within S. asteriscus. How¬ ever, these changes were never formally published. Silphium radula Nuttai.l Silpliiuiu radula Nuttall, Trans. Amer. Philos. Soc., n.s. 7: 354. 1841. TYPE: U.S. A. Plains of Arkansas, T. Nuttall s.n. (holotype, BM). Silphium simpsonii Greene var. wrightii L. M. Perry, Hho- dora 39: 288. 1037. Syn. nov. TYPE: U.S. A. Texas: 1848. C. Wright s.n. (holotype, (41). Silphium simpsonii var. wrightii is reduced here- Volume 14, Number 3 2004 Clevinger New Combinations in Silphium 277 in to synonymy of Silphium radula based on the examination of the type specimen at GH. Gandhi and Thomas (1989) recognized both S. radula and S. simpsonii var. wrightii and stated that they differ only in the number of capitula and leaf texture. I have observed that these differences are quite var¬ iable, and these two entities should not be sepa¬ rated. Silpliiiiin radula var. graeile (A. Gray) J. A. Clev¬ inger, comb, et slat. nov. Basionym: Silphium graeile A. Gray, Proc. Amer. Acad. Arts 8: 653. 1873. TYPE: U.S.A. Texas: Harris Co., Houston, 1842, F. ./. Lindheimer s.n. (lecto- type, selected here, GH; isotype, (ill). Silphium radula var. graeile is recognized by a persistent basal rosette and ray flowers numbering 12 to 18. In contrast, Silphium radula var. radula has a caducous basal rosette and ray flowers num¬ bering 20 to 30. Perry (1937) and others have sug¬ gested that the southwestern U.S. Silphium radula var. graeile is allied to the southeastern U.S. Sil¬ phium asteriscus var. simpsonii. These two varieties resemble one another in their persistent basal ro¬ settes and reduced stature. However, Silphium rad¬ ula var. graeile and variety radula have pubescent phyllaries and are found in Arkansas, Louisiana, Oklahoma, and Texas, while Silphium asteriscus var. simpsonii has glabrous phyllaries and is found in Alabama, Florida, Georgia, Mississippi, and South Carolina. Silphium radula var. graeile (as Sil¬ phium graeile) was included in the molecular study of Clevinger and Panero (2000). On the basis of ITS and ETS sequence data, this variety was dem¬ onstrated to be sister to S. radula var. radula. Asa Gray cited two collections, those of Hall and of Lindheimer, in his description of Silphium grae¬ ile. He did not designate either as the holotype; therefore, they are syntypes. According to Gray, the Lindheimer specimens have larger leaves and more pubescence than the Hall specimens. Both descrip¬ tions fit Gray’s concept of S. graeile and my concept of S. radula var. graeile. I have seen the two Lind¬ heimer collections at GH. The status of the Hall collections is uncertain; I have seen two 1872 col¬ lections at F and NY labeled //«// 321 (Gray does not cite a number in his description) but none at GH. Given this uncertainty, I have chosen the Lind¬ heimer collection at GH to serve as the lectotype. Acknowledgments. I am grateful to Jose L. Pa¬ nero, Beryl Simpson, Billie Turner, Doug Goldman, Curtis Clevinger, James Allison, Conley K. Mc¬ Mullen, Lowell and Mary Amiek, and the curators of the following herbaria for the loan of specimens: F, FLAS, GH, IA, LL, LSI), MICH, MO, MSC, NCU, NY, OS, TENN, TEX, US, and USCH. This paper represents a portion of a doctoral dissertation by the author submitted to the Department of Bot¬ any at the University of Texas at Austin. Support for some aspects of this study was provided by grants from the American Society of Plant Taxon¬ omists and the Virginia Academy of Science. Literature Cited Chapman, A. W. I860. Flora of the Southern United States: Containing an Abridged Description of the Flow¬ ering Plants and Ferns of Tennessee, North and South Carolina, Georgia, Alabama, Mississippi and Florida: Arranged According to the Natural System. Ivison, Phinney, New York. Clevinger, J. A. 1999. Systematies of Silphium and Its Subtribe Engelmaniinae (Asteraceae: Heliantheae). Ph.D. Dissertation, The University of Texas at Austin. - & J. L. Panero. 2000. Phylogenetic analysis of Silphium and subtribe Engelmanniinae (Asteraceae: Heliantheae) based in ITS and ETS sequence data. Amer. J. Bot. 87: 565-572. Cronquist, A. 1980. Vascular Flora of the Southeastern United States, Vol. I: Asteraceae. Univ. North Carolina Press, Chapel Mill. Gandhi, K. N. & R. I). Thomas. 1989. Asteraceae of Lou¬ isiana. Sida Bot. Misc. 4. Jarvis, C. E. & N. Turland. 1998. Typification ol Linnaean specific and varietal names in the Compositae (Astera¬ ceae). Taxon 47: 347—370. Perry, L. M. 1937. Notes on Silphium. Rhodora 39: 281 — 297. Weber, W. R. 1968. Biosystematic Studies in the Genus Silphium L. (Compositae): Investigations in the Selected Intraspecific Taxa of Silphium asteriscus L. Ph.D. Dis¬ sertation, The Ohio State University. Phylloscirpus (Cyperaceae) Revisited Sandra Dhooge and Paid Goetghebeur Ghent University, Department of Biology, Research Group Spermatophytes, K.I Ledeganckstraat 35, B-9000 Gent, Belgium. Sandra.Dhooge@UGent.be; Paul.Goetghebeur@UGent.be Abstract. Two species of Scirpus are transferred to Phylloscirpus: Scirpus deserticola Philippi and Scirpus boliviensis Barros. Phylloscirpus deserticola has all the typical characteristics of the genus Phyl¬ loscirpus, such as eligulate leaves, a terminal head of spikelets. and scabrid hypogynous bristles. The allocation of Scirpus boliviensis to Phylloscirpus is morphologically less obvious due to its reduced structures, but is well supported by a phylogenetic analysis based on plastid rbcl . anil OtjL-F sequence data. Phylloscirpus deserticola is lectotypified be¬ cause the holotype consists of a mixed collection. Key words: Andes, Cyperaceae, Phylloscirpus. Phylloscirpus C. B. Clarke (1908) is a genus with a single species, Phylloscirpus acaulis (Philippi) Goetghebeur & I). A. Simpson (1990), distributed in Andean regions of South America. Clarke (1908) described the genus based on the presence of a dense head of spikelets subtended by bracts, each spikelet having spirally arranged glumes and bi¬ sexual (lowers comprising five or six scabrid, hy¬ pogynous bristles. During our ongoing taxonomic study of Scirpus s.l. in the Andes, it became clear that one of the species, Scirpus deserticola Philippi, clearly has the diagnostic features of Phylloscirpus , justifying a new combination within the genus. Another and very small species, Scirpus boliviensis Barros, also has characters in common with the species of Phyl¬ loscirpus, such as eligulate leaves and numerous red tannin idioblasts in the glumes. Although Clarke did not mention these characters in his orig¬ inal description, our observations and a molecular analysis discussed below strongly indicate that Scirpus boliviensis Barros should be included in the genus Phylloscirpus. However, as Scirpus boliviensis Barros has one terminal spikelet (opposed to a head of spikelets) and no bristle hairs, it is necessary to redefine Phylloscirpus, to give the genus a broader circumscription. This proposal to make two new combinations in Phylloscirpus is well supported by our phylogenetic analysis based on rbc L and traL-F sequencing data of several Andean Scirpus species. Material and Methods MORPHOLOGICAL DATA Morphological data were obtained from the study of herbarium material with a stereomicroscope. Herbarium material from the following herbaria was studied: AAU, B, BM, C, GENT. GH, GOET, ISC, K, NY, P, OCA. SGO, SI, U. and US. MOLECULAR DATA The plant material used for DNA extraction and its provenance are listed in Table 1. Total DNA was isolated using the modified CTAB method of Doyle and Doyle (1987). Amplification was done in over¬ lapping pieces using the IF, 1024F. 724R. 1024R, and I460R primers for the rbc\ . gene (Muasya et ah, 1998) and the c, d, e, and f primers for /raL-E (Taberlet et ah. 1991). The amplified products were cleaned using the QIAquiek kit according to the manufacturer’s protocols (Qiagen, Ltd.), and se¬ quenced using standard dideoxy methods and run on an ABI 377 automated sequencer (all according to the manufacturer's protocols. Applied Biosys¬ tems, Inc.). Phe sequences were edited in Sequence Navigator, assembled in Autoassembler (Applied Biosystems, Inc.), and aligned by eye in a sequence matrix of Scirpus s.l. (Muasya et ah, 2000). PH'1 LOGENETIC AN At A SIS The molecular data set was analyzed using par¬ simony algorithms of PAUP* 4.0b8 (Swofford, 2002). A heuristic search of 1000 replicates with the TBR (tree-bisection-reconnection) swapping al¬ gorithm and random addition was conducted, re¬ taining five trees at each step. Internal support was estimated with 1000 bootstrap replicates. For each bootstrap replicate, 100 heuristic searches with random sequence addition were conducted. Taxonomy Phylloscirpus deserticola (Philippi) Dhooge & Goetghebeur, comb. nov. Basionym: Scirpus de¬ serticola Philippi, FI. Atacam.: 53. 1860. TYPE: Chile. Agua de profetas in deserto Atacamensi, Jan. 1854, R. A. Philippi s.n. (lectotype, select¬ ed here. SGO 046286 A). Figure 1A, I). Nov ON 14: 278-284. 2004. Volume 14, Number 3 2004 Dhooge & Goetghebeur Phylloscirpus (Cyperaceae) 279 Scirpus semisubterraneus Boeckeler, Eirinaea 36: 495. 1870. Scirpus deserticola Philippi var. semisubterra¬ neus (Boeckeler) Barros, Anal. Mus. Argent. Cienc. Nat. 38: 156. 1935. TYPE: Peru. W. Lechler n° 1977 (holotype, not seen; isotypes, k, P). Caespitose, small herbs, 1.8— 12 cm tall; rhi¬ zomes creeping, branching; culms 0.6-10 cm X 0.5— 1.3 mm, terete, glabrous, sometimes with little red spots (tannin idioblasts), covered with old brown leaf sheaths. Leaves shorter than to as long as the culm; leaf sheaths 2.5— 6.5 mm long, often with red spots; leaf blade 0.3— 1.9 cm X 0.4—0.85 mm, scabrous toward the leaf tip; leaf tip muticous, often orange; ligule absent. Inflorescence a single spikelet or a terminal, dense head composed of 2 to 6 spikelets (Fig. I A), 4—7.2 X 1.5—8 mm, red- brown to dark brown; when 2 or more spikelets are present, each lateral spikelet subtended by a bract and a small, membranous prophyll; bracts 3.4— 6.5 X 2-4 mm, red-brown with many tannin idioblasts and a conspicuous, swollen, pale midrib, apiculate, as long as to slightly shorter than the inflorescence. All glumes fertile, 2.7-4.7 X 1.5-1. 9 mm, ovate, red-brown with many red spots (tannin idioblasts) and a conspicuous, swollen, pale midrib; flowers bisexual; stamens 3; ovary 3-carpellate; style Figure 1. A and D. Phylloscirpus deserticola (Philippi) Dhooge & Goetghebeur ( Philippi s.n., lectotype, SGO). - — A. Inflorescence. — D. Fruit with bristle hairs. — B. Inflorescence of Phylloscirpus acaulis (Philippi) Goetghebeur & D. A. Simpson ( Ruthsatz 8765, GENT). C and E. Phylloscirpus boliviensis (Barros) Dhooge & Goetghebeur ( Petersen & Hjerting 1048b, holotype, G). — C. Inflorescence. — E. Eruil. 280 No von °b _c vO »o vO m o m , _ i I— H CO On c i— H o r- t- 1 - r- r- i - co CO r- l - :: X U- m in m in IO m m m m m IO m io c Ov O' O' On O' & ON O'. On O' On O' U* =b i-i c ^ O —I f35 CM CM LS CO co 2 » - r- t - r- o »o 10 CM ro rt* CM CM CM co ro ro h- h- h- r- io io m -I- -i- 3 o 00 CM CM CM ro ro ro co r - Q >H -2 C CO _ C -C ^ 03 bfi co — ^ < a * >-* >< ^ & r- cm £ vO O O 2 rt CM O ^ <1^ ro co CO t- r ^ io co O' O' lO lO I - I - < C J J= JS c 5 h® ^ © t ^ (/) -S -i- W (/) ^ N _c Oh ca 00 00 cc X 3 X jz X X CS < U — / «< < g 8 8 8 8 3 O' O § O' O'. 3 cm o Uu r >n «< CM ' — 1 I—* ' — 1 ' — 1 CM CM r— 1 H E— I a; O) K. ’ cO "3 "3 ' 3 "3 ' 3 "3 "3 "3 "3 rr Os 7) C/> (/) £jg III oc H / ._. w Ch 'C 5 fc hi'2 £- = t 73 — 73 r\i 73 <-o ^ oc d •-j ^ -C ' — '3 ^ ~ hfi ■? hfi hr >- hfi v— ^ S C 30 •g bp :S "S 5 i? •■S ^ 3 b£ . 5 G‘ kO ^ -w.’ O .VD 5 -c SL u £.2.2 3d ^ I- SC c* — §3 c* | s J ri 2 a; a!> io ^ 5- rfi ~ -s =? -o Uj u. it:' -2 -5' i*; i 3 S .6" v g 5 a£J O 3^ -3 O bQ O ^ § Table 1. Continued. Volume 14, Number 3 2004 Dhooge & Goetghebeur Phylloscirpus (Cyperaceae) 281 =3 = ^ 2 < -C cn CC U) ”3“ 1 — — I I — CO CO CO — uO LO 00 CO O', t- vO O' O' O' O' CM CM CM (N 2 •— < cn < ca C/i CC CO ^ *0 «> CO CC CM CO ON ON LO LO r- r^ i.O io CC f_' w w z s 5 UC H, o. 'cj< iP CM co co NO CM C /J 7] C/J N 1 ° « K « 3 § ^ I .*5C £ -5 ' t: §- . _ JH -§ 9 .5 I .« O in do •£ 3 W £ il ~ N ' u I o e & °y John Manning. Scale bar 10 mm for entire plant, capsules, floral bracts, and side view of flower; detail of the perianth base, entire gynoecium and stamens of dissected flower, 3X scale bar, seed approximately 6X scale bar. Volume 14, Number 3 2004 Goldblatt & Manning African Iridaceae 291 breve quam exteriore, floribus hypocrateriformibus acti- nomorphis pallide ad intense carneis tepalis purpureis ad basem nigris leviter dulce odoris, tubo perianthii pauce curvato cylindrico infime 2—3 mm abrupte expanse in eu- pulam ca. 3 mm profundam, tepalis obovatis patentibus subaequalibus (14— )18— 22 X (f> — ) 10 — 12 mm, filamentis conniventis 5—6 mm longis ereetis nigris, antheris diver- gentibus 6-7 mm longis nigris, polline flavo, ovario ovo- ideo ca. 2 mm longo, stylo erecto furcato ad basem an- therarum ramis 2—3 mm longis nigris, capsulis ovoideis 9—11 X 5 mm, seminibus subglobosis vel angulalis 1.2— 1.5 mm diam. leviter rugosis. Plants 20—60 cm high; conns globose, 15—20 mm diam., the tunics of medium to coarsely tex¬ tured fibers; cataphylls membranous, the upper one reaching just above ground level and flushed red¬ dish brown. leaves 3, all basal, the lower two nar¬ rowly to broadly lanceolate, (3— )5— 10 mm wide, usually about half as long as the stem, the margins and midribs moderately thickened, the uppermost leal entirely sheathing, reaching at least to the mid¬ dle or upper third of the stem. Stem erect, simple or 1- to 3-branched, 0.5—1 mm diam. at the base of the spike, branches held 45°— 90° to the main axis; spike almost straight, the main spike usually 3- to 6-flowered, the lateral spikes I - to 4-flowered; bracts membranous and translucent, the outer 8—10 mm long, with three light purple veins, lightly three-dentate, the inner as long as to slightly longer than the outer, forked at the apex, with two purple veins. Flowers salver-shaped, actinomorphie, pale to deep pink, the tepals each purple to blackish toward the base, lightly sweet scented; perianth tube slightly curved and cylindric for 2-3 mm, wid¬ ening abruptly into a cup ca. 3 mm deep; tepals obovate, spreading at nearly right angles to the tube, subequal, (14— ) 18— 22 X (6—) 10— 12 mm, the inner slightly wider than the outer; filaments in¬ serted at the mouth of the narrow part of the tube, blocking the mouth, connivent, 5—6 mm long, straight and erect, black; anthers divergent, 6—7 mm long (before anthesis), black, the pollen yellow; ovary ovoid, ca. 2 mm long; style straight and erect, dividing opposite the base of the anthers, the branches arching outward, 2—3 mm long, black. Capsules ovoid, 9—1 1 X 5 mm; seeds subglobose or angled by pressure, 1 .2-1.5 mm diam., yellow- brown, lightly wrinkled with surface cells collicu- late. Phenology. Flowering early to mid September, probably also in late August. Etymology. From the Latin superbus , “superb, excellent,” referring to the large, attractive, deep pink and purple-black flowers. Distribution and biology. Ixia superba has a re¬ corded range of just a few acres at the foot of the Langeberg in the Little Karoo east of Montagu. Plants grow on loamy sandstone ground that grades to a light clay at the interface of Table Mountain Sandstone and Bokkeveld Systems on the farm Kli- pheuwel. We collected plants in bloom there in ear¬ ly September 2002 close to the end of their flow¬ ering season. Associated plants were Babiana patula N. E. Brown, Moraea gawleri Sprengel, Wat¬ son ia laccata (Jacquin) Ker-Gawler (Iridaceae), and the shrub Elytropappus rhinocerotis (L. f.) Lessing. The flowers evidently have a bimodal pollination strategy that uses both hopbine scarab beetles (Scarabaeidae: Hopliini) and worker honey bees. Apis mellifiera (and possibly female bees of other genera). The flowers have the typical attributes of the hopbine beetle pollination system, viz. a bright¬ ly colored perianth with dark markings, darkly pig¬ mented stamens, relatively long anthers, and short style branches. The closed perianth tube and ab¬ sence of nectar are also features associated with this pollination system in Ixia (Goldblatt et ah, 2000a). Presence of a sweet floral odor is, however, an attribute of bee pollination and bees do visit the flowers, which evidently last three days, but close at night. Both hopbine beetles and honey bees were found visiting the flowers, the latter actively col¬ lecting pollen. Early in the morning hopbine bee¬ tles were found in half-closed flowers in which they had evidently spent the night. In one other collec¬ tion of /. superba we have found ( Steiner 3239, NBG), field notes indicate that the flowers were vis¬ ited by the hopbine beetle Peritrichia sp. Relationships. Ixia superba is readily confused with a second Little Karoo species, Ixia gloriosa G. J. Lewis (1962). Allhough both have deep pink flow¬ ers with a large blackish glossy central eye and blackish filaments and anthers, there are several im¬ portant floral differences between them. The darker red-purple flowers of /. gloriosa have relatively short, slightly spreading filaments 3—5 mm long, and an¬ thers, said by de Vos (1999b) to be (>-1 mm long and connivent at the tips. Plants studied by us alive ( Goldblatt , Manning & Porter 121 79, MO, NBG), however, had anthers ca. 4.5 mm long diverging throughout, thus not apically connivent. The style normally divides at the mouth of the tube, thus op¬ posite the base of the filaments, and the style branches are ca. 4 mm long. The perianth tube is 4—7 mm long and widens abruptly into a small cup little more than 1 mm deep. Fresh flowers of I. glo¬ riosa have a characteristic umbonate appearance, the result of the tepals being raised above the rim of the perianth cup before spreading. This is not apparent in pressed specimens or closed flowers. In contrast, /. superba has the perianth tube slender below for 3 292 Novon mm, widening abruptly into a cup ea. 3 mm deep and the tepals spread uniformly from the rim of the tube. The filaments are 5—6 mm long, and the di¬ vergent anthers are ea. 6 mm long. The style is en¬ closed above by the closely connivent filaments and divides at or shortly above the top of the filaments into shorter style branches 2—3 mm long. In addition, /. gloriosa typically has linear leaves 15—55 X 1—3 mm and a long slender spike of 8 to 10 flowers on the main axis, whereas /. superba has shorter, lan¬ ceolate leaves 10—30 X (3— )5— 10 mm and a shorter, crowded spike of 3 to 6 flowers. While the bracts of the two species are similar, those of /. gloriosa have more pronounced veins t lint when dry remain dark purple, while the veins in the bracts of /. superba fade and almost disappear. There are several other locally endemic species of Ixia in the Breede River valley that resemble one another closely in their deep pink flowers with a dark center and blackish filaments and anthers (Lewis, 1962). This flower type is adapted for pol¬ lination by hopbine beetles, a pollination system that seems to favor local differentiation among plant species, e.g., Aristea subg. Pseudaristea (Goldblatt & Manning, 1997) and Sparaxis (Goldblatt et al., 2000b). The species of Ixia in the Breede River valley with flowers adapted for this pollination sys¬ tem form a geographic al series, replacing one an¬ other across relatively short distances. Moving down the river valley from its head in the northwest, the series comprises /. vinacea G. J. Lewis (Tul- bagh), /. rouxii G. J. Lewis (Wolseley), /. mostertii M. P. de Vos (Romans River to Worcester), /. van- zyliae L. Bolus (Worcester to Roberston), /. atran- dra Goldblatt & J. C. Manning (south of Worcester to Stettyn), /. superba (Montagu), and /. gloriosa (Barrydale). Among this alliance, /. superba is read¬ ily distinguished by the perianth tube that forms a wide cup ca. 3 mm long in die upper half and the style that divides at or slightly above the base of the filaments, thus close to the base of the anthers, rather than at the* mouth of the perianth tube and we'll below the base of the anthers. Paratypes. SOUTH AFRICA. Western Cape: 3320 (Montagu), Farm Rietvlei No. 1, ca. 10.5 km SF of Mon¬ tagu, scattered on S-facing rocky slope (CC), ca. 370 m, 5 Sep. 1997, Steiner 32.39 (NBC); Scheepers Rust, 20 mi. K of Montagu, 22 Aug. 1936, Martley s.n. (BOL 26930). Tritoniu delpierrei M. P. de Vos, J. S. African But. 49; 403. 1983. Tritoniu marlothii subsp. del¬ pierrei (M. P. de Vos) M. P. de Vos, FI. S. Africa 7(2,1): 121. 1999. TYPE: South Africa. North¬ ern Cape: Richtersveld, top of Helskloof, M. P. de Vos 2462 (type, STL). We do not accept de Vos’s (1999a) reduction of the South African Tritonia delpierrei to subspecific rank in T. marlothii M. P de Vos. We have seen and examined both plants in the wild, and while they are evidently sister laxa, isolated geographi¬ cally and phylogenetieally within Tritonia (de Vos, 1983), there is little overlap in their critical taxo¬ nomic features. We prefer to maintain them as sep¬ arate species differing in floral coloration and di¬ mensions that apparently indicate different primary pollinators. Tritonia delpierrei has lightly sweet- scented, pale yellow flowers, the lower tepals with cream markings, a perianth tube 12—22 mm long, and the dorsal tepal Fi — 1 2 mm long. In contrast, T. marlothii has odorless flowers, a pale to deep pur¬ ple perianth (not lilac as indicated by de Vos) with the lower tepals yellow with purple tips and a me¬ dian dark violet spot, and new collections (e.g., Goldblatt & Porter 11757, MO, NBG) show the perianth tube is 28— 42 mm long and the dorsal tepal (8-) 10- 1 2 mm long. The flowers of T. mar¬ lothii show the stereotyped adaptations for polli¬ nation by the long-proboscid fly Prosoeca peringue- yi (Manning & Goldblatt, 1996), and we have captured this fly while visiting and evidently pol¬ linating the flowers (unpublished obs.). The shorter perianth tube of T. delpierrei suggests pollination by a long-tongued anthophorine bee. The ranges of the two taxa largely overlap. Tritonia marlothii oc¬ curs in the southern Richtersveld of Namaqualand at Brakfontein, southwest of Eksteenfontein, and along the eastern foothills of the Stinkfontein Mountains north of Eksteenfontein, whereas T. del¬ pierrei extends from near Eksteenfontein in the south to Helskloof in the north, a distance of some 60 km that includes a large part of the range of T. marlothii. While T. delpierrei seems to favor sandy slopes, we have only seen T. marlothii in rocky ground, an apparent habitat difference also consis¬ tent with separate species status. Thus despite their evident immediate relation¬ ship, we see no reason to consider Tritonia delpier¬ rei as a subspecies of 7. marlothii. Their geographic ranges partly overlap while their habitats differ, and their critical morphological features show no evi¬ dence of convergence. ImnoiDKAK Ferraria kamiesbergensis M. P. de Vos, J. S. Af¬ rican Bot. 45: 362. 1979. TYPE: South Africa. Northern Cape: Rondefontein, I Sep. 1976, E. G. II. Oliver 5970 (lectotype, here designated, lower left plant on sheet, STE). Described by M. P. de Vos in 1979, Ferraria ka- Volume 14, Number 3 2004 Goldblatt & Manning African Iridaceae 293 miesbergensis is represented by four elements: t he illustration accompanying the protologue (de Vos, 1979: 363), and three plants mounted on a single sheet, Oliver 5970. These were evidently all col¬ lected on the same date at Rondefontein in the Ka- miesberg of central Namaqualand. We have visited the type locality (or as close to it as is possible) and found a small, uniform population of plants with small pale yellow flowers, stems 12—15 cm high, branching if at all. only from near the base, with distinctive dark green leaves, and subequal inflorescence spathes ( Goldblatt cfe Porter 12223, MO. NBG). Most significantly, the lower leaves all curve inward in the same direction. Only one specimen on the type sheet matches these plants, and it also has the paired inflores¬ cence spathes almost equal in length with the inner sheathing only below. The other two plants are tall¬ er (one is ca. 30 cm tall), have narrower, evidently flat leaves, produce branches well above ground level, and the outer inflorescence spathes are about half as long as the inner and are largely sheathing. The plant illustrated in the protologue corresponds to the larger of these two taller specimens, and both appear to be a second species, possibly what is currently called Ferraria divaricata Sweet, either its subspecies aiirea M. R de Vos or subspecies arenosa M. R. de Vos. The flower illustrated is prob¬ ably F. kamiesmontana (and is certainly not F. di¬ varicate z), while the smaller plant on the lower left of the type sheet alone matches plants in the wild and is the lectotype. The plants found at Rondefontein are matched at other sites in the southern Kamiesberg on the farm Karas, to the north, immediately adjacent to Rondefontein. Elsewhere on the Kamiesberg, the only other pale yellow-flowered species of the genus is the florally nearly identical Ferraria macrochla- rnys (treated by de Vos as F. uncinata subsp. ma- crochlamys, see below), which has quite different leaves. The range of Ferraria kamiesbergensis according to de Vos is from Springbok (Thompson 1301 , NBG) in the north to Wallekraal in the southwest ( Comp¬ ton 5405 , BOL), and extending east as far as the Calvinia district (e.g., de Eos 2347, NBG). We have examined all the specimens on which de Vos based her assessment of the range of the species and con¬ sider them to correspond with what she called F. divaricata. None of tin* specimens she cited have the distinctive incurved leaves and subequal spathes of the southern Kamiesberg populations (flower color can no longer be determined). As cir¬ cumscribed here, F. kamiesbergensis is restricted to the southern Kamiesberg at elevations above 1000 m where plants matching the lectotype occur. Ferraria macroehlamys (Baker) Goldblatt & J. C. Manning, comb. nov. Basionym: Lapeirousia macroehlamys Baker, J. Bot. new ser. 5: 338. 1876. Ferraria uncinata subsp. macroehlamys (Baker) M. P. de Vos. J. S. African Bot. 45: 369. 1979. TYPE: South Africa. Namaqua¬ land, without precise locality or date. Herb. Forsyth s.n. [probably collected by J. Niven in 1899] (type. K). In her revision of Ferraria (Iridoideae) de Vos (1979) treated a series of central and northern Na¬ maqualand populations (Northern Gape, South Af¬ rica) with pale yellow flowers as F. uncinata subsp. macroehlamys, while she regarded Western Cape populations with deep blue to violet flowers as F. uncinata Sweet subsp. uncinata. De Vos believed that subspecies macroehlamys merited subspecific status because it differed only in having flowers with a differently colored perianth. The two taxa share with F. brevifolia G. J. Lewis and F. kamies¬ bergensis M. P. de Vos a distinctive flower type in the genus in which the tepal claws are suberect and collectively form a narrow floral cup that contains relatively dilute nectar. Both these latter species also have predominantly pale yellow tepals virtually the same shade and size as those of subspecies macroehlamys. De Vos regarded F. macroehlamys and F. uncinata as sharing similar leaf blades with the margins thickened and the edges crisped and often undulate. In her key (1979: 329) to the spe¬ cies, F. uncinata (including subsp. macroehlamys) is distinguished by “margins of at least some of the foliage leaves crisped.” De Vos commented of sub¬ species macroehlamys and subspecies uncinata that they “differ only in flower coloring and geo¬ graphic distribution.” Closer examination of spec¬ imens she included in subspecies macroehlamys, however, shows that the thickened, hyaline part of the leaf margins is crenate to lightly serrate and sometimes shortly velvety, but the leaf edges them¬ selves are only occasionally lightly crisped. In Fer¬ raria uncinata the thickened part of the leaf margin may be entire, or sometimes lightly serrated or cris- pulate, and not velvety but rather evidently smooth (but under 10X magnification papillate), while the edges of at least some of the leaves are crisped and usually undulate. Thus, examination of the leaf margins alone can usually distinguish the two taxa. In addition, the flowers of F. uncinata have outer tepals (18— )30— 35 mm long, with claws 9—1 1 mm long, thus usually slightly less than one-third the 294 Novon length of the entire tepal. In F. uncinata subsp. macrochlamys the outer tepals are 24—28 mm long and the claws 12.5— 11 mm long, thus half as long as the entire tepal. We believe the differences are sufficient that Ferraria uncinata subsp. macrochlamys should be treated as a separate species. In fact, we are not convinced that it is most closely related to F. un¬ cinata. The central Namaqualand F kamiesmon- tana has flowers virtually identical to those of F uncinata subsp. macrochlamys in size, proportion, and orientation, as well as color, and can be distin¬ guished from the subspecies solely by a few dark spots at the bases of the outer tepal limbs. Leaf margins of F. kamiesmontana are, however, entire, smooth, and barely thickened. Growth form of the two is similar: plants of both taxa are low growing and have a few subequal branches near the stem apex, whereas F. uncinata subsp. uncinata often branches above the base, is frequently taller, and the stems emerge from the leaf sheaths. Relation¬ ships of F uncinata subsp. macrochlamys are by no means obvious, and it is most useful to treat it as a separate species, pending phylogenetic anal¬ ysis of Ferraria using molecular techniques. Regarding the type collection of Ferraria ma¬ crochlamys, the specimen bears the annotation “Forsyth 1835” suggesting a gathering by a Mr. Forsyth. No collector of this name in known in southern Africa, and the name Forsyth, listed by Gunn and Codd (1981) in their compilation of plant collecting in southern Africa, is without initials, dates of birth and death, and lacking additional information. We now believe the annotation refers to William Forsyth, son of W. F. Forsyth of the Chel¬ sea Physic Garden, who maintained a herbarium collection. William Forsyth died in July 1835 and his library, presumably containing herbarium spec¬ imens, was auctioned in November of that year (C. Nelson, pers. comm.). Indications in the accounts of Bahiana spiralis Baker, based on another Forsyth specimen (Baker, 1892, 1896), have led us to con¬ clude that the Forsyth herbarium was purchased by Wiliam Banks, then Keeper of the Kew Herbarium, for the collection there. The most likely collector ol the type of B. spiralis and Ferraria macrochlamys is the British botanist James Niven, who collected in Namaqualand in 1799 and later, and whose specimens are now widely dispersed (Gunn & Codd, 1981). The type citation above reflects this new information. Moraea simplex Goldblatt & J. C. Manning, sp. nov. TYPE: South Africa. Western Cape: 21 km N of Piketberg, 18 Sep. 2002, F. Goldblatt & /,. J. Porter 12231 (holotype, NBG; isotypes, MO, PRE). Figure 2. Plantae 28— M) cm altae, cormo ovoideo ca. 10 mm dia- metro tunicis reticulatis fibrosis brunneis obtecto, foliis productis 1 canaliculatis 2-3 mm latis, caule perramoso, spatha interne inflorescentiis 18—22 mm longis, floribus flavo-cremeis, tepalorum limbis leviter reflexis unguibus ascendentibus oa. 2 mm longis, limbis externis 18—20 X 7.5— 8.5 mm intends ca. 18X7 mm, filamentis liberis ca. 5 mm longis inferne in colunmam gracilem conniventibus superne per 1 mm divergentibus, antheris albis ca. 3 mm longis, stylo ramis filiformibus indivisis ca. 4 mm longis inter filamenta arcuatis, ovario exserto anguste ovato ca. 3 mm longo. Plants 28^40 cm high; corm ovoid, ca. 10 mm diani., the tunics of coarse, light brown fibers, with a neck of fairly fine fibers surrounding the base of the stem; cataphylls papery, the uppermost reaching shortly above the ground, becoming dry and light brown by flowering time, then often irregularly bro¬ ken. Foliage leaf solitary, inserted on the first aerial node, the blade channeled, exceeding the stem, arching outward or trailing distally, 2-3 mm wide. Stem erect, lightly flexuose, sticky below the nodes for half the length of an internode, bearing a sheathing bract-like leaf at each of the upper nodes, these 24—35 mm long, green, becoming dry and light brown above, the apices acute, bearing two or three short branches of one internode each at the upper three or four nodes, branches at the lower nodes sometimes two internodes long and also branched, the branches erect and parallel to the stem below, flexed outward above the sheathing leaf. Inflorescence a rhipidium, terminal on the branches, several-flowered; spathes subequal, green, with dry, brown tips, 18-22 mm long, the outer slightly shorter than the inner. Flowers uni¬ formly pale creamy yellow, the outer tepals each with a pale yellow mark at the base of the limb, unscented, the tepal claws ascending, forming a cup enclosing the base of the filaments; tepals clawed, the outer slightly larger than the inner, the claws ascending, ca. 2 mm long, the limbs ovate, obtuse, initially spreading horizontally, later dip¬ ping up to 30° below the horizontal, the outer 18— 20 X 7.5— 8.5 mm, the inner ca. 18 X 7 mm; fil¬ aments ca. 5 mm long, free but conuivent below and forming a slender column, diverging in the up¬ per I mm; anthers ca. 3 mm long, oblong, ap- pressed to one another, white, the pollen white; ova¬ ry exserted, narrowly ovoid, ca. 3 mm long; style dividing shortly below the base of the anthers into 3 filiform branches, each stigmatic apieally and ex¬ tended between the filaments, ca. 4 mm long. Cap¬ sules and seeds unknown. Volume 14, Number 3 2004 Goldblatt & Manning African Iridaceae 295 Figure 2. Moraea simplex Goldblatt & J. C. Manning. Drawn from living plants from the type population, Goldblatt & Porter 12231, by John Manning. Scale bar 10 mm for entire plant, side view of stamens and style branches approx¬ imately 3X scale bar. 296 Novon Phenology. Flowering in mid to late September, probably into mid October. Etymology. From the Latin simplex, “un¬ adorned, plain.” referring both to the pale-colored (lowers without markings, the spreading subequal tepals, and the filiform style branches lacking style crests. Distribution anil biology. Moraea simplex is known oidy from the relatively moist clay flats at the eastern foot of the Piketberg Mountains. Nu¬ merous plants were found at the beginning of tbeir (lowering or still in bud on September 18 at tbe single known site for the species, which had been burned the previous summer. We infer that (low¬ ering continues until at least early October. The flowers each last a single afternoon, opening at about 14:00H and wilting shortly after nightfall. Relationships. The seventh member of the tax- onomically isolated subgenus Visciramosa (Gold- blatt, 1986; Goldhlatt Manning, 2000b), Moraea simplex has the typical attributes of the subgenus, including sticky internodes, multiple leaves, branched stems, relatively short inflorescence spathes, and free, connivent filaments. Plants are readily mistaken (or the common and widespread M. inconspicua Goldhlatt in the vegetative state as both species share narrow leaves, short, subequal rhipidial spathes, and relatively small flowers. However, M. inconspicua usually has two foliage leaves, unequal tepals, the outer larger than the inner, the tepal limbs strongly reflexed, and rela¬ tively broad, flattened style branches with paired erect crests, and the anthers appressed to the op¬ posed style branches, the latter condition typical of the genus. In contrast, M. simplex has solitary fo¬ liage leaves, the inner and outer tepals nearly equal in size and spreading to slightly dipping below the horizontal, and simple, filiform stvle branches that extend between the filaments. That the species belongs in Moraea cannot be doubted despite the unusual filiform style branches that extend between the filaments, for the resem¬ blance to other species of subgenus Visciramosa is striking. It has firm, channeled leaves, stieky nodes and internodes, and the distinctive lateral branch¬ ing pattern in which the branch axis is parallel to the main stem below, before arching outward almost horizontally above the sheathing bract, but with the rhipidia held erect, and the subequal, obtuse rhip¬ idial spathes of the subgenus. Only the flowers are discordant, not only with the subgenus, but with Moraea in general. Reduced style branches, not or hardly wider than the anthers they subtend, are rel¬ atively common in Moraea anil were the hallmark of the genus Homeria (Goldhlatt, 1981), now in¬ cluded in Moraea (Goldhlatt. 1998; Goldhlatt et ah, 2002). Even in Homeria, however, the style branch¬ es have forked apices and lie opposite the stamens and appressed to the anthers. Completely undivid¬ ed style branches, not opposed to the stamens, were the feature that distinguished Roggeveldia (Gold¬ hlatt, 1979). The two species of Roggeveldia are now also included in Moraea (Goldhlatt, 1998) and are most probably related to Moraea crispa Thun- berg and its immediate allies (Goldhlatt. 1986). Thus the filiform, undivided style branches held alternate to the stamens are a parallel development in a second lineage of Moraea. Discovery and de¬ scription of Moraea simplex brings the number of species in the genus to 197 (Goldhlatt & Manning, 2002). In Moraea tbe following combinations at subspe¬ cies rank were inadvertently not made when spe¬ cies of Hexaglottis and Homeria were transferred to that genus (Goldhlatt, 1998). Moraea bulbillifera subsp. anomala (Goldhlatt) Goldhlatt, comb. nov. Basionym: Homeria bul¬ billifera subsp. anomala Goldhlatt, Ann. Mis¬ souri Hot. Gard. 68: 458. 1981. TYPE: South Africa. Western Cape: Witsands near Ossekop, P. Goldhlatt 4885 (holotype, MO; isotypes, BR, K. NBG. PRE, WAG). Representative specimen. SOUTH AFRICA. Western Cape: 3321 (Ladismith) Cloete's Pass to Wagenbooms (DDK Sep. 1976, Goldhlatt 4157 ( K. MO. PRE, S. WAG). Moraea lewisiae subsp. secunda (Goldhlatt) Goldhlatt, comb. nov. Basionym: Hexaglottis lewisiae subsp. secunda Goldhlatt, Ann. Mis¬ souri Bot. Gard. 74: 558. 1987. TYPE: South Africa. Northern Cape: near the top of Spek- takel Pass, P. Goldhlatt 6678 (holotype, PRE; isotypes, K, MO, NBG, S, US, WAG). Representative specimen. SOUTH AFRICA. Northern Cape: 3119 (Calvinia) Nieuwoudtville, karroid hills NE of Klipkoppies (AC). Sep. 1981, Goldhlatt 7078 (MO). Moraea virgata subsp. karooica (Goldhlatt) Goldhlatt, comb. nov. Basionym: Hexaglottis virgata subsp. karooica Goldblatt, Ann. Mis¬ souri Bot. Gard. 74: 568. 1987. TYPE: South Africa. Northern Cape: 22 km W of Middelpos, D. Snijman 765 (holotype, NBG; isotypes, K, MO, PRE). Representative specimen. SOUTH AFRICA. Northern Cape: 3120 (Williston) Roggeveld Escarpment, 71 mi. SE of Calvinia on Middelpos road via Blomfontein (CC), Sep. 1981, Griddhlatt 4621 (MO. NBG). Volume 14, Number 3 2004 Goldblatt & Manning African Iridaceae 297 Nivenioideae Aristea torulosa Klatt, Abh. Naturf. Ges. Halle 15: 382. 1882. TYPE: South Africa. Eastern Cape: Transkei, betw. Gekau & Bashee, Dec. 1832. J. F. I)rege 4558 (lectotype, effectively designated by Weimarck (1949: 27). B not seen, photo, K; isotype, P). Aristea torulosa var. monostachya Baker, FI. Cap. 6: 49. 1896. TYPK: South Africa. Natal, without precise locality or date, Buchanan s.n. (type, K). Aristea woodii N. E. Brown, Kew Bull. 1931: 192. 1931. Syn. nov. TYPE: South Africa. KwaZulu-Natal, In- anda, J. M. Wood 757 (type, K). Aristea torulosa has been overlooked in recent times, and Vincent (1985) excluded the species from his account of Aristea in eastern southern Af¬ rica because he considered that no specimens could be assigned to it. J. G. Baker (1896) and Weimarck (1940), however, had no misgivings about its identity and status, and Baker (1896) ac¬ tually regarded what was later described as A. woo¬ dii as A. torulosa var. monostachya Baker. Aristea woodii and A. torulosa have identical derived, rel¬ atively large rhipidial spathes and bracts that are dark green below, aging reddish brown, that grade unevenly into broad translucent margins, darker along the veins thus giving them a feathered ap¬ pearance. The pale edges of the spathes and bracts become torn and irregularly fringed with age. The only significant difference between plants assigned to Aristea woodii and the type of A. toru¬ losa is the crowded flowering axis of the latter, which has the lower nodes bearing short lateral branches. Typical A. woodii has a more slender flowering stem usually bearing only sessile flower clusters. Plants readily referable to ,4. woodii may, however, have one or even two of the lowermost nodes bearing stalked flower clusters; thus, in ef¬ fect, the flowering stem is branched. The type of a second species, regarded as a synonym of A. toru¬ losa by Weimarck (1940) but excluded by Vincent (1985), A. congesta, described by N. E. Brown in 1931 and based on plants from the Transkei. South Africa, is just such a plant, and we see no reason to distinguish it from A. woodii. The more congested floral axis of A. congesta in turn, is little different from that of A. torulosa. Plants similar to the types of A. congesta and A. torulosa are matched by Krauss s.n. (K) from Port Natal and Rehmann s.n. (K) from Woodbush in Limpopo Province. The Krauss collection was referred by Weimarck to A. torulosa, while the Rehmann collection was not cit¬ ed by either Weimarck (1940) or Vincent (1985). More extensive collections now available, at NBG, MO, and PRE, suggest that all three named species represent single taxon. The earliest name for this widespread species that extends from Tanzania and eastern Angola (Goldblatt, 1993, 1996) in the north to eastern South Africa in the south thus becomes A. torulosa. Acknowledgments. We thank Ingrid Nanni and Lendon Porter for their assistance and companion¬ ship in the field. Support for this study by grants 7103-01 and 7316-02 from the National Geograph¬ ic Society is gratefully acknowledged. Collecting permits were provided by the Nature Conservation authorities of Northern Cape and Western Gape Provinces, South Africa. I .ilerature Cited Baker, J. O. 1892. Handbook of the Irideae. George Bell & Sons, London. - . 1896. Irideae. Pp. 7-71 in W. T. Thiselton-Dyer (editor). Flora Capensis 6. Ashford, Kent. Brown, N. E. 1931. XXIX — Notes upon some South Af¬ rican plants. Kew Bull. Misc. Inform. 1931: 191 — 197. tie Vos, M. P. 1979. The African genus Ferraria. ,1. S. African Bot. 45: 295—375. - . 1983. The African genus Tritonia Ker-Gawler: Part 2. Sections Subcallosae anti Montbretia. J. S. Af¬ rican Bot. 49: 347—422. - . 1988. Three new species of Ixia I,. (Iridaceae) from the Cape Province. S. African J. Bot. 54: 596— 602. - . 1999a. Tritonia (Iridaceae). In G. Germishuizen (editor), Flora of Southern Africa 7(2: 1): 89—128. Na¬ tional Botanical Institute, Pretoria. - . 1999b. Ixia (Iridaceae). In G. Germishuizen (ed¬ itor), Flora of Southern Africa 7(2: 1): 3—87. National Botanical Institute, Pretoria. Geerinck, 1). 2001. Nouvelle espeee: Gladiolus goldblat- tianus Geerinck ainsi que les deux combinations nou- velles: G. erectiflorus var. verdickii , G. gregarius var. an- gustifolius pour la famille des Iridaceae en Afrique centrale. Taxonomania 4: 2—3. Goldblatt, P. 1979. Roggeveldia, a new genus of southern African Iridaceae— Irideae. Ann. Missouri Bot. Card. 66: 839-844. - . 1981. Systematics and biology of Homeria (Iri¬ daceae). Ann. Missouri Bot. Gard. 68: 413—503. - . 1986. fhe Moraeas of Southern Africa. Ann. Kir- stenboseh Bot. Gard. 14: 1—224. - . 1990. Phylogeny and classification of Iridaceae. Ann. Missouri Bot. Gard. 77: 607-627. - . 1993. Iridaceae. In G. V. Pope (editor), Flora Zambesiaca 12(4): 1—106. Flora Zambesiaca Managing Committee, London. - . 1996. Gladiolus in Tropical Africa. Timber Press, Portland, Oregon. - . 1998. Reduction of Barnardiella, Galaxia, Gyn- andriris, Hexaglottis, Homeria, and Roggeveldia in j Wo¬ men (Iridaceae: Irideae). Novon 8: 371—377. - & J. C. Manning. 1997. New species of Aristea (Iridaceae) from South Africa and notes on the taxono¬ my and pollination biology of section Pseudaristea. No¬ von 7: 137—144. 298 Novon - & - . 20(H). Cape Plants: A conspectus of the vascular plants ol the Cape Region ol South Africa. Strelitzia 7. National Botanical Institute of South Africa, Cape Town. - & - . 2002. Notes and new species of Mo- raea (Iridaceae: Iridoideae) from the southern African winter-rainfall zone. Novon 12: 352—359. - . P. Bernhardt & J. C. Manning. 2000a. Adaptive radiation of pollination mechanisms in the African ge¬ nus Ixia (Iridaceae: Crocoideae). Ann. Missouri Bot. Card. 89: 564-577. - , J. C. Manning & Peter Bernhardt. 2000b. Adap¬ tive radiation of pollination mechanisms in Sparaxis (Ir¬ idaceae: Ixioideae). Adansonia s6r. 3, 22: 57—70. - , V. Savolainen, 0. Porteous, I. Sostaric, M. Powell, G. Reeves, J. C. Manning. T. G. Barraclough & M. W. Chase. 2002. Radiation in the Cape flora and the phy- logeny of peacock irises Moraea (Iridaceae) based on four plastid I)NA regions. Molec. Phvlogenet. Evol. 25: 341-360. Gunn, M. & L. E. Codd. 1981. Botanical Exploration of Southern Africa. Balkema, Cape Town. Lewis, G. J. 1962. South African Iridaceae. The genus Ixia. J. S. African Rot. 28: 45-195. Manning, J. C. & P. Goldblatt. 1996. The Prosoeca per- ingueyi (Diptera: Nemestrinidae) pollination guild in southern Africa: Long-tongued flies and their tubular flowers. Ann. Missouri Bot. Card. 83: 67-86. Vincent, L. P. 1). 1985. A partial revision of the genus Aristea (Iridaceae) in South Africa, Swaziland, Lesotho, Transkei and Ciskei. S. African J. Bot. 51: 209-252. Weimarck, H. 1940. Monograph of the genus Aristea. Acta Univ. Lund (Lunds Univ. Arsskr.) n. s. 36: 1—140. Tradescantia boliviana (Commelinaceae), a New Combination for an Overlooked South American Species Jason R. Grant Laboratoire tie botanique evolutive, Institut de botanique, Faeulte des Sciences, Universite de Neuchatel, rue £mile-Argand 1 1, Case Postale 2, 2007 Neuchatel, Switzerland. jason.grant@unine.ch Abstract. the new combination Tradescantia boliviana (Hasskarl) J. R. Grant is made in recog¬ nition that it is a distinct species widespread in Peru, Bolivia, Paraguay, and Argentina, here sep¬ arated by morphological characters and distribution from its previous inclusion within T. ambigua Mar¬ tins ex J. H. Schultes, which is restricted to north¬ eastern Brazil. Key words: Argentina, Bolivia, Brazil, Mandon- ia, Paraguay, Peru, Tradescantia. While researching the Tradescantia sect. Man- donia complex, problems in species limits and no¬ menclature were discovered in the non-Mexican species. A new combination is proposed as Trad¬ escantia boliviana (Hasskarl) J. R. Grant in recog¬ nition that the material from Peru, Bolivia, Para¬ guay, and Argentina represents a distinct species. This species is here separated from synonymy un¬ der T. ambigua Martius ex J. H. Schultes, which is restricted to northeastern Brazil. Ten species comprise Tradescantia sect. Man- donia D. H unt. The group has a center of diversity in Mexico with six species: T. burchii I). Hunt [found in Jalisco], T. crassifolia Cavanilles (includ¬ ing T. acaulis Martens & Galeotti and T. iridescens Bindley) [northern Mexico to Guatemala], T. exal- tata D. Hunt |()axaca], T. llamasii Matuda [Est. Mexico], T. peninsularis Brandegee [Baja Califor¬ nia], and T. tepoxtlana Matuda [Morelos] (Hunt. 1980, 1986). Four non-Mexican species range from Guatemala to South America: T. ambigua Martius [distributed in northeastern Brazil], T. boliviana (Hasskarl) J. R. Grant [Argentina, Bolivia. Para¬ guay, Peru], T. petricola J. R. Grant [Costa Rica, Venezuela] (Grant, 2000), and T. velutina Kunth & C. 1). Bouehe [Guatemala, Honduras, and Nicara- gua]. While researching Tradescantia petricola , addi¬ tional specimens were located from Venezuela that matched this previously envisioned Costa Rican endemic (Grant, 2000). While further searching among South American indets and among T. am¬ bigua in many herbaria (COL, G, Gil. IAN. INPA, L, MG, MO, NY, P, R, RB, SP, SPF, and US), it became evident based on plant size, leaf disposi¬ tion, sepals, and distribution that the specimens from Peru, Bolivia, Paraguay, and Argentina were clearly different from those from Brazil. While it was initially thought that these represented a new species, the name Mandonia boliviana was found to apply. Type material of Mandonia boliviana was located at Leiden (L, 1 sheet), Paris (P, 3 sheets), and Geneva (G, 2 sheets), and its lectotypification is proposed below. Examination of this type mate¬ rial, and comparison to recently collected speci¬ mens, clarifies its identity as distinct from T. am¬ bigua. Since the combination into Tradescantia has not been made, the new combination is proposed here. Tradescantia ambigua is restricted to the caatin- ga of northeastern Brazil, while Tradescantia boli¬ viana is restricted to the Gran Chaco of Argentina, Bolivia, Paraguay, and Peru (Fig. 1). Ti •adeseantia boliviana (Hasskarl) J. R. Grant, comb. nov. Basionym: Mandonia boliviana Hasskarl, Flora 54: 260. 1871. Skofitzia boli¬ viana (Hasskarl) Hasskarl & Kanitz, Oesterr. Bot. Z. 22: 147. 1872. TYPE: Bolivia. Lare- caja: “Viciniis Sorata montis colles Ullontigi ad scopulorum radicem in regione temperata,” 2700 m, Feb.— Apr. 1858, G. Mandon 1239 (lectotype, designated here, L [Herb. J. K. Hasskarl]; isotypes, G [2], k not seen. P. P [originally from ‘Herb. E. Cossen'], P [origi¬ nally from ‘Herbier E. Drake’)). Specimens examined. ARGENTINA. Cordoba: [entre Panolma y la Villa del Transito (P)], [zwischen Panolma unter der Villa del Transito und der Westseite der Tierra Achala de Cordoba (US)], 23 Feb. 1876, Hieronymus s.n. (P, US); Rodado, Stuckert 10938 (G); Sierra Cleica de Cor¬ doba, Stuckert 13530 (G), lessen 450 (GH); Punilla, Cap- ilia del Monte, Cuezzo & Balegno 2144 (G. US). Salta: M. Sala, Venturi 8180 (GH, US); Cerro San Bernardo, Sal¬ ta, W [no collector?] 6151 (GH); Qrdn, Pintascayo, Pierotti 6558 (G); Cachi, Quebrada de Escoipe, Ruta 30, 2 km Novon 14: 299-301. 2004. 300 Novon Figure 1. Distribution ol Tradescantia boliviano in open circles, and T. ambigna in solid squares. antes de FI Maray, Novara 6067 (G); Rosario de Lerma, Quebrada del Rio Blanco, 2—3 km al SW (aguas arriba) del rio Toro, Novara 7603 (G [3]). Tueunian: Capital, Dunaquito, Venturi 1660 (GH, US), Venturi 1660a ((ill. US); Tufi del Valle, Venturi 2021 ((ill. US). BOLIVIA. Chuquisaea: Tomina, Weddell 3330 (P). Cochabamba: near Taquina, above Cochabamba, Everdam 25231 (G); Cochabamba, Julio 11.174 (NY, US), Cardenas 5560 (US); Camino a Saeabe, cerca a Cochabamba, Ochoa 670 (NY); Cercado de Cochabamba, la Colina de San Pedro, Stein- bach s.ri. (US); Campero, Mique, Steinbach 755 (GH, NY. US); Cerro San Pedro, behind Cochabamba, Cutler & Car¬ denas 3830 (elow 500 m, and is possibly amphibious. (See S. latifolius for further discussion of habitat.) The holotype of Syngonanthus dichroanthus at NY is a mixed sheet including an unidentified ro- sulate species of Eriocaulon of about the same stat¬ ure. The isotype sheet at P is a mixture with the same Eriocaulon as well as with Eleocharis minima Kunth (Cyperaceae). both sheets were distributed as S. simplex (Miquel) Ruhland. This inconspicuous mud-dwelling species is known so far only from mixed collections with other diminutive species. Its relatives include several other small rosulale annuals with pale glabrous ca- pitula, including Syngonanthus gracilis (Bongard) Ruhland, S. biformis (N. E. Brown) Gleason, 5. sim¬ plex (Miquel) Ruhland, and S. latifolius (Moldenke) Hensold. Syngonanthus dichroanthus is distin¬ guished from these in [tart by its delicate habit, membranous threadlike leaves, and involucral bracts exceeding the capitulum. From the closely sympatric and common taxon S. gracilis var. aureus Ruhland, it can usually be distinguished by the lack of glandular hairs on the peduncle sheaths. However, its most striking feature is the strong con¬ trast between the copper-colored staminate flowers and the white pistillate flowers, an unusual char¬ acteristic shared only with S. latifolius. Strong differentiation in the color of the stami¬ nate and pistillate (lowers occurs to my knowledge in only a few divergent species of Syngonanthus. These include the dimorphic-stemmed aquatic S. spongiosus Hensold, with staminate flowers deep yellow-brown, and the rosette species S. angolensis Hess of Africa and S. goyazensis (Kornicke) Ruh¬ land of Brazil, in both of which the staminate ca¬ lyces are deeply pigmented only below the lobes. The specimen from Manaus ( Liitzelburg 21962 p.p.) lacks staminate flowers, which makes positive identification difficult, but is provisionally placed here due to the glabrous flowers, involucral bracts glabrous and surpassing the capitulum (which dis¬ tinguish it from Syngonanthus biformis and S. sim¬ plex), the membranous leaves, and lack of glandular hairs. Paratype. BOLIVIA. Santa Cruz: Prov. Velasco, Es- taci6n Flor de Oro, margen del Rfo It6nez (Guapor6), fron- tera con Rondonia, ea. 20 km N del Serranfa de Huan- chaca, ea. 85 km E del Rfo Paragua, 15 May 1991 (fl), M. Pena et al. 62 p.p. (F, mixed collection with Syngon¬ anthus gracilis var. aureus Ruhland). Additional specimen examined. BRAZIL. Amazonas: Manaus, Villa Municipal, 28 Aug. 1916 (9), Liitzelburg 21962 p.p. (LL = fragment ex NY, mixed collection with Utricularia and sterile graminoid, probably Cyperaceae). Syngonanthus latifolius (Moldenke) Hensold, comb. nov. Basionym: Syngonanthus gracilis var. latifolius Moldenke, Phytologia 21: 418. 1971. TYPE: Brazil. Mato Grosso: Serra do Roncador, gallery margin, ca. 86 km N of Xav- antina, 550 m, 31 May 1966 (fl, fr), H. S. Ir¬ win, J. W. Crear Jr., R. Souza, l\. Reis dos San¬ tos 16349 (holotype, NY). Figure 2G— J. Plants rosulate annuals; roots white, spongy, branched, the larger 0.5— 0.7 mm diam. Leaves re¬ curved, ± prostrate, ca 1.0-1. 2 cm long, 1.0-1. 5 mm wide, ligulate, subacute, the tip minutely re¬ curved and keeled; 3- to 5-veined, the midvein vis¬ ible adaxially as a white line, at least in the basal half; pubescent abaxially with short appressed mal- pighian hairs, and adaxially with long filamentous arachnoid hairs, both surfaces glabrate. Inflores¬ cences numerous, ca. 25 to 50, of mixed ages, some¬ times with new inflorescences initiating outside the older ones; peduncle sheaths 10—17 X 0.6— 0.8 mm diam., the lamina ca. 2 mm long, acute, callous- tipped, slightly recurved; arachnoid pubescent as the leaves, also with scattered upright gland-tipped hairs; white-costate only near the base; peduncles 6—15 cm long, (0.25— )0.3— 0.4 mm wide, 3-costate; the intercostae green and arachnoid pubescent, broad near the apex, narrowing to thin furrows be¬ low; the costae stramineous, glabrous, shiny, with a few gland-tipped hairs when young; capitula 4—5.5 mm diam., ovoid in bud, depressed-hemispheric to globose at maturity; involucral bracts hyaline, gla¬ brous, lustrous, the outer obovate, rounded-apieu- late at apex, pale, shredding with age, the inner oblanceolate-obtuse to linear-acute and gold with a brown tint; shorter than the pistillate flowers; re¬ ceptacle densely white-villous, trichomes about equaling the staminate flowers; receptacular bracts absent. Flowers 3-merous, the staminate and pistil¬ late about evenly mixed and strongly dimorphic, the staminate flowers pigmented and much shorter than the pistillate, the pistillate surpassing the in¬ volucral bracts by 0.6— 0.7 mm, abscising after seed set; staminate flowers : pedicels ca. 0.15—0.25 mm; calyx coppery brown, zygomorphic, the segments ca. 1.0 X 1.0 mm, broadly obovate-spalulate, rounded, slightly cucullate, fused ca. Vi — % of their length, sparingly pubescent to glabrate on the ab- axial face; corolla borne on stipe ca. 0.5 X 0.1 mm; the corolla itself ca. 0.35 mm long, membranous, the lobes involute; filaments ca. 0.35 mm, adnate to corolla; anthers while, 0.2 mm long; pistillodes ca. 0.25 mm long, infundibular, the rim papillate; pistillate flowers: pedicels ca. 0.5 mm long; sepals oblong-elliptic, acute to broadly acuminate, navic¬ ular and slightly sclerified along midvein at base. Volume 14, Number 3 2004 Hensold South American Eriocaulaceae 307 Figure 2. A— F. Syngonanthus dichroanthus Hensold. — A. Habit. — B. Capitulum. — C. Starninate flower. — I). Corolla of starninate flower. — E. Pistillate flower. — F. Gynoecium. G— J. Syngonanthus latifolius (Mold.) Hensold. — G. Sta- minate flower. — H. Pistillate flower. — I. Gynoecium. — J. Habit. Vouchers: A— F Pires & Black 2,165', ( i — J Irwin et al. 16349. 308 Novon 1. 7-2.0 X 1.0 mm. white-hyaline, tinged brown at apex, glabrous; corolla short-stipitate, the petals fused at middle margins, free at base and apex, linear-oblanceolate, acute to acuminate, involute after anthesis, ca. 1.1— 1.3 X 0.25 mm, membra¬ nous, glabrous at maturity; style ca. 0.6 mm long, style branches ca. 0.4 mm long, appendages small and filamentous. Seeds red-brown, ca. 0.4 X 0.2 mm, with raised white longitudinal ribs after wet¬ ting. the epidermis waxy (under SEM). Ecology and distribution. Known from a single collection in marshy campo at 550 m, at the margin of gallery forest in cerrado of northeastern Mato Grosso, Brazil, ca. 52°W, 14°S. Syngonanthus latifolius was originally described as a variety of .S', gracilis (Bongard) Ruhland, dis¬ tinguished from other varieties by the leaves wider and more closely appressed to the ground. The marked size and color dimorphism of the flowers were not noted in the original description, however, and together with the leaf characters, support rec¬ ognition of the taxon as a distinct species. The closest affinity of Syngonanthus latifolius is probably with S. dichroanthus Hensold, which has a nearly identical floral structure and color dimor¬ phism, but flowers subequal in size, and membra¬ nous filiform leaves. In S. latifolius , the sepals and pedicels of the pistillate flowers are nearly twice as long as those of the staminate flowers, such that the small rounded golden-brown staminate flowers are dwarfed by the white acute pistillate flowers with which they are intermixed. Similar differentiation in flower size and a similar aspect to the capitulum are matched only in the species .S’, bijorrnis (N. E. Brown) Gleason, but the latter is distinguished by its uniformly white flowers, pubescent involucres, densely pubescent petals in the pistillate flowers, and prominent style appendages. Affinity is also suggested with Syngonanthus gracilis var. aureus, with which S. latifolius shares the annual habit, nearly glabrous sepals and petals, reduced filiform style appendages, and the waxy coating observed on the seeds under SEM (Hen- sold, unpublished data). Both Syngonanthus latifolius and S. dichroanthus have been collected together with S. gracilis var. aureus in wetlands (veredas) associated with cer¬ rado. Eiten (1975) described in detail the abruptly delineated “seasonally marshy grassy campos" or “hillside campos,” which separate gallery forests from uplantl cerrado in the Brazilian Serra do Ron- eador and elsewhere. Created by rainy season drainage, these campos have gleyed soils, espe¬ cially in their lower, wetter portions, where Erio- caulaceae are reported as a typical element. Syn¬ gonanthus dichroanthus may occur on wetter sites than S. latifolius. since both the Rfo Itenez site and the veredao in Maranhao are inundated for longer periods than the valleys of the Serra do Roneador (Eiten, 1975), but more collections are needed to confirm tbis. Additional specimen examined. BRAZIL. Mato 6 ros¬ so: Serra do Roneador, gallery margin, ca. 86 km N of Xavantina, 550 m, 31 May 1966, II. S. Irwin et al. 16346 p.p. (till) [mixed sheet with X. gracilis var. aureus Ruh¬ land, probably from the same gathering as Irwin et al. 16349. the type]. Acknowledgments. Many thanks to Creg Muell¬ er and the Botany Department at the Field Museum for providing research space; to Peter Jprgensen, editor of the Catdlogo de las Plantas Vasculares de Bolivia, for help and encouragement; to Luzmila Arroyo and Mike Net* for crawling specimens at USZ and NY; and to the curators of GH, LL, MO, NY. and P for loans of material. I .iterature Cited Eiten, G. 1975. The vegetation of the Serra do Roneador. Biotropica 7: 112—135. Giulietti, A. M. 1978. Modificayoes taxonomicas no genero Eriocaulon I.. Bol. But. Univ. Sao Paulo 6: 39-47. - . 1984. Estudos Taxonomicos no Genero Leiothrix Buhl. (Eriocaulaceae). Tese de Livre-Docencia, Univ- ersidade de Sao Paulo, Sao Paulo, Brazil. Killeen. T. J., E. Garefa & S. G. Beck (editors). 1993. Guta de Arboles de Bolivia. Missouri Botanical Garden, St. Louis, and Herbario Nacional de Bolivia, La Paz. Ruhland, W. 1903. Eriocaulaceae. In: A. Engler (editor), Das Pflanzenreieh IV. 30: 1—294. Wilhelm Engelmann, Leipzig. Stiitzel. T. 1988. Untersuchungen zur Wurzelanatomie der Eriocaulaceen. Flora 180: 223—239. - . 1998. Eriocaulaceae. Pp. 197-207 in K. kubitz- ki (editor). The Families and Genera of Vascular Plants, Vol. IV. Flowering Plants. Monocotyledons. Alismatanae and Commelinanae (except Gramineae). Springer- Ver- lag. Berlin. Heidelberg. Tomlinson, P. B. 1969. Anatomy of the Monocotyledons III. t iommel inales— Zingiberales. Clarendon Press, Ox¬ ford. Zhang, Z. 1999. Monographic der Gattung Eriocaulon in Ostasien. l)iss. Bot. 313: i— vii, 1—289. Subtribes of Principally North American Genera of Cichorieae (Compositae) Joongku Lee 1,2 and Bruce G. Baldwin'-' 'Jepson Herbarium and Department of Integrative Biology, 1001 Valley Life Sciences Building #2465, University of California, Berkeley, California 94720-2465, U.S.A. “’Current address: Korea Research Institute of Bioscience and Biotechnology, 52 Oun-dong, Yuseong-ku, Daejeon 305-333, Korea ’Author for correspondence: bbaldwin@uclink4.berkeley.edu ABSTRACT. A new subtribal classification (or principally North American genera of Cichorieae allows for a practical taxonomy that reflects recent phylogenetic hypotheses. We propose the newT sub- tribes Lygodesmiinae (for Chaeladelpha, Lygodes - mia sensu Tomb, and Shinnersoseris), Pinaropap- pinae (for Marshal Ijohnstonia and Pinaropappns), Pyrrhopappinae (for Picrosia and Pyrrhopappus), and three monogeneric subtribes (Glyptopleurinae ( Glyptopleura ), Krigiinae ( Krigia ), and Phalacroser- idinae ( Phalacroseris )) for taxa previously treated in Malacothrieinae, Microseridinae, and Stephanom- eriinae. We recognize novel circumscriptions for Malacothrieinae ( Anisocoma , Atrichoseris, Calyco- seris, and Malacothrix ), Microseridinae (Agoseris, Microseris, Nothocalais, Stebbinsoseris, and Uropap- pus ), and Stephanomeriinae ( Munzothamnus , Pleia- canthus, Prenanthella, Rafinesquia, and Stephan- omeria). The revised classification provides a more precise taxonomic representation of relationships among genera, insofar as robust resolution of mono- phyletic groups currently allows. Key words: Cichorieae, Lactueeae, Malacothri¬ einae, Microseridinae, Stephanomeriinae, sub¬ tribes, taxonomy. Phylogenetic analyses of 18S— 26S nuclear rl)NA external and internal transcribed spacer sequences (Lee et al., 2002, 2003) and chloroplast DNA re¬ striction sites (Jansen et al.. 1991; Whitton et al., 1995) yielded evidence for new hypotheses of re¬ lationship among all genera of Cichorieae (Lactu- ceae) that have natural centers of diversity in North America (plus Picrosia, endemic to South America). On the basis of molecular findings, Lee et al. (2003) concluded that taxa previously treated in subtribes Malacothrieinae, Microseridinae, and Stephano¬ meriinae (e.g., Stebbins, 1953; Bremer, 1993, 1994) or in Jeffrey’s (1966) “Microseris subgroup” and “Stephanomeria subgroup” represent a single major radiation, with a large western North Amer¬ ican component. As previously treated, each of those subtribes and subgroups is evidently para- phyletie or polyphyletic. To better reflect phyloge¬ netic relationships, we propose revised circum¬ scriptions for Malacothrieinae, Microseridinae, and Stephanomeriinae and recognize six new subtribes. Malacothrieinae K. Bremer, Novon 3: 329. 1993. TYPE: Malacothrix DC. Annual or perennial herbs. Leaf blades linear, elliptic, lanceolate, oblanceolate, obovate, or spat- ulate; margins entire, denticulate, lobed, or pinnat- ifid. Capitulescences corymbiform or capitula borne singly (peduncles scapiform). Receptacles not pa- leate, bristly or not bristly. Involucres cylindrical to obconic or campanulate. Phyllaries in 3 to 6 series, unequal (1 or 2 subequal series in Atrichoseris). f lorets 15 to 250 per capitulum. Corollas yellow or white (often with abaxial reddish stripes). Cypselae columnar to fusiform, 5- to 15-ribbed or -veined (5 usually prominent), beaked (Calycoseris) or not beaked. Pappi 0 ( Atrichoseris ) or o( 15 to 40 sub¬ equal, barbellate or, rarely, plumose (Anisocoma), persistent or often (all or mostly) caducous or frag¬ ile bristles and sometimes an outer crown of minute teeth. Pollen echinate (Tomb et al., 1974). Chro¬ mosome number 2 n — 14, 18, 20. Included genera: Anisocoma Torrey & A. Cray, Atrichoseris A. Cray, Calycoseris A. Cray, Malaco¬ thrix DC. Microseridinae Stebbins ex Solbrig, Taxon 12: 234. 1963. TYPE: Microseris I). Don. Annual, biennial, or perennial herbs. Leaf blades linear, oblong, lanceolate, oblanceolate, or spatu- late; margins entire, denticulate, sinuate, lobed, or pinnatifid. Capitulescences branched or, often, ca¬ pitula borne singly (peduncles scapiform). Recep¬ tacles not paleate, not bristly. Involucres subey- lindrical to obconic or campanulate. Phyllaries in Novon 14: 309-313. 2004. 310 No von 2+ series, subequal or unequal, the outer some¬ times forming ealyculi. Florets 5 to 300+ . Corollas yellow, orange, or white (sometimes with abaxial reddish stripes). Cypselae columnar or fusiform, ± 10-ribbed, beaked (Agoseris) or not. Pappi of (0)2 to 90 equal or unequal, smooth to barbellate (rarely plumose) bristles or bristle-tipped scales, persistent or caducous. Pollen echinolophate (Feuer & Tomb, 1977). Chromosome number 2n = 18, 27 (triploid), or 36. Included genera: Agoseris Kafinesque, Microseris 1). Don, Nothocalais (A. Gray) Greene, Stebbinso- seris K. L Chambers, Uropappus Nuttall. Steplianomeriinae Stebbins ex Solbrig, Taxon 12: 235. 1963. TYPE: Stephanomeria Nuttall. Annual or perennial herbs, or shrubs. Leal blades linear, oblong, lanceolate, oblanceolate, ob- ovate, or scale-like; margins entire, sinuate-den¬ tate, lobed, or pinnatifid. Capitulescences corym- biform to thyrsoid-paniculiform. Receptacles not paleate, not bristly. Involucres narrowly cylindrical or obconic. Phyllaries usually in 2 series, unequal (outer forming ealyculi), sometimes graduated (Ste- pharwmeria cichoriacea). Florets 3 to 15 (to 30 in Rafinesquia) per capitulum. Corollas pink, purple, or white. Cypselae clavate, columnar, or fusiform, smooth, sometimes grooved or ^ 5-ribbed, beaked (Rafinesquia) or not. Pappi ol 5 to 20+ equal or unequal ( Prenanthella ), plumose {Rafinesquia, Ste¬ phanomeria), or barbellate, persistent, or deciduous {Munzothamnus) bristles. Pollen eehinate (Tomb ct al., 1974). Chromosome number 2 n = 14 {Pre¬ nanthella), 16, 24 (triploid), or 32. Included genera: Munzothamnus P. II. Raven, Pleiacanthus Rydberg, Prenanthella Rydberg, Raf¬ inesquia Nuttall, Stephanomeria Nuttall. Glyptoplcurinue Joongku Lee & B. G. Baldwin, subtribus nov. TYPE: Glyptopleura I). C. Ea¬ ton. Herbae annuae. Laminae foliorum spatulatae vel oblon- gae, profunde lobatae vel pinnatifidae; margines albi, sca- riosi, dentati vel setoso-ciliati. Capitula portata singulatim vel capitulescentiae parcc ramosae. Receptacula epaleata, non setosa. Involucra cylindracea vel urceolata, ad bases bracteis spatulatis marginihus albis, scariosis, dentatis. Phyllaria 2-seriata, ± aequalia. Flosculi 7—16. Corollae cremeae vel luteolae (in sicco subroseae). Cypselae cla- vatae, tuberculatae, valde costatae, rostratae. Pappi con- slati ex setis 100+, aequalibus, barbellatis, extimis de- ciduis. Pollina echinata. 2 n = 18. Annual herbs. Leaf blades spatulate to oblong, deeply lobed to pinnatifid; margins white, searious, dentate or setose-ciliate. Capitula borne singly or capitulescences sparingly branched. Receptacles not paleate, not bristly. Involucres cylindrical to ur- ceolate, al base with spatulate bracts with margins white, searious, toothed. Phyllaries 2-seriate, ± equal. Florets 7 to 16 per capitulum. Corollas cream-colored to light yellow (when dry, pink). Cypselae clavate, tuberculate, strongly ribbed, beaked. Pappi ol 100+ equal, barbellate bristles, the outermost deciduous. Pollen eehinate (Tomb et al., 1974). Chromosome number 2n = 18. Included genus: Glyptopleura I). C. Eaton. ki -igiiuae Joongku Lee & B. G. Baldwin, subtribus nov. TYPE: Krigia Schreber. Ilerbae annuae vel perennes. Laminae foliorum line- ares, oblongae, lanceolatae, vel oblanceolatae; margines integri, lobati, vel pinnatifidi. Capitula portata singulatim vel capitulescentiae cymosae vel interdum superne sub- umbellatae. Receptacula epaleata. Involucra turbinata, campanulata, vel urceolata. Phyllaria 1— 2 + -seriata, ae¬ qualia. Flosculi 4—30. Corollae llavae vel aurantiacae. Cypselae columnares vel ovoideae, 10-20-nervatae vel - costatae, non rostratae. Pappi null i vel ex squamis 5 + constati (interdum fascientibus coronis minutis), pler- umque cingentibus setis 5 + , inaequalibus, spiculatis. Pollina echinolophata. 2n = 8. 10. 12, 18. 20. 30, vel 60. Plants annual or perennial herbs. Leaf blades linear, oblong, lanceolate, or oblanceolate; margins entire, lobed, or pinnatifid. Capitula borne singly or capitulescences eymiform or subumbelliform. Receptacles not paleate. Involucres turbinate, cam- panulate, or urceolate. Phyllaries I - or 2 + -seriate, equal. Florets 4 to 30 per capitulum. Corollas yel¬ low or orange. Cypselae columnar or ovoid, 10- to 20-nerved or -ribbed, not beaked. Pappi none or of 5+ scales (sometimes forming minute crowns), usu¬ ally surrounding 5 + . unequal, spiculate bristles. Pollen echinolophate (Feuer & Tomb, 1977). Chro¬ mosome number 2 n = 8, 10, 12. 18, 20, 30, or 60. Included genus: Krigia Schreber. Lygodesiniiiiac Joongku Lee & B. G. Baldwin, subtribus nov. TYPE: Lygodesmia I). Don. Herbae annuae ( Shinnersoseride ) vel perennes. Laminae foliorum lineares vel lanceo-lineares (vel squamiformes distaliter); margines plerumque integri, interdum lobati. Capitula raro portata singulatim, capitulescentiae pler¬ umque ramosissimae. Receptacula epaleata. Involucra cy¬ lindracea vel subcylindracea. Phyllaria 2-seriata, inae- qualia (extimis fascientibus calyculis). Flosculi 5—12. Corollae subrosaceae, purpureae, vel albae. Cypselae col¬ umnares vel fusiformes, laeves, strialae, rugosae, vel cos¬ tatae, non rostratae. Pappi ex setis 24—250, aequalis, bar¬ bellatis (el squamis 5, subulatis in Chaetadelpha), constati omnibus persistentibus. Pollina echinata vel echinolopha¬ ta {Lygodesmia). 2 n — 12 (Shinnersoseride) vel HI, vel 27 (triploideis). Volume 14, Number 3 2004 Lee & Baldwin North America Genera of Cichorieae 311 Annual (, Shinnersoseris ) or perennial herbs. Leaf blades linear or lance-linear (or scale-like distally); margins usually entire, sometimes lobed. Capitula rarely borne singly, capitulescences usually much- branched. Receptacles not paleate. Involucres cy¬ lindrical or subcylindrical. Phyllaries 2-seriate. un¬ equal (the outer forming calyculi). Florets 5 to 12 per capitulum. Corollas pink, purple, or white. Cyp- selae columnar or fusiform, smooth, striate, rugose, or ribbed, not beaked. Pappi of 24 to 250 equal, barbellate bristles (and 5, subulate scales in Chae- tadelpha), all persistent. Pollen echinate or eehin- olophate ( Lygodesmia ) (Tomb et al., 1974). Chro¬ mosome number 2n — 12 ( Shinnersoseris ) or 18, or 27 (triploids). Included genera: Chaetadelpha A. Gray, Lygo¬ desmia I). Don (sensu Tomb), Shinnersoseris Tomb. Phalacroseridinae Joongku Lee & B. G. Baldwin, subtribus nov. TYPE: Phalacroseris A. Gray. Herbae perennes. Laminae foliorum lineares, lanceo- lineares, vel oblanceolatae; margines integri. Capitula por- tata singulatim (pedunculis scapiformibus); receptacula epaleata; involucra campanulata; phyllaria 2+-seriata, ae- qualia. Flosculi 13 + , corollae Havae. Cypselae fusiformes, laeves, non rostratae; pappi nulli. Pollina ecliinolophata. 2 n = 18. Perennial herbs. Leaf blades linear, lance-linear, or oblanceolate; margins entire. Capitula borne sin¬ gly (peduncles scapiform). Receptacles not paleate. Involucres campanulate. Phyllaries 2 + -seriate, equal. Florets 13+ per capitulum. Corollas yellow. Cypselae fusiform, smooth, not beaked. Pappi none. Pollen eehinolophate (Feuer & Tomb, 1977). Chro¬ mosome number 2 n = 18. Included genus: Phalacroseris A. Gray. I'inaropappinae Joongku Lee & B. G. Baldwin, subtribus nov. TYPE: Pinaropappus Lessing. Herbae perennes vel frutices rosulatae ( Marshalljohn - stonia). Laminae foliorum lineares (vel squamiformes dis- taliter), ellipticae, vel rhombeae; margines integri, dentati, vel lobati. Capitula portata singulatim. Receptacula pa- leata (. Pinaropappo ) vel epaleata. Involucra obconica vel campanulata. Phyllaria 3 + -seriata, inaequalia. Flosculi 10—18. Corollae subrosaceae, purpureae, vel albae. Cyp¬ selae suleis 5, angustis, aequaliter dispositis, columnares vel fusiformes, non rostratae (distaliter attenuatae in Pi¬ naropappo p.p.). Pappi ex setis 40-60, aequalis, barbel- latis, persistentibus constati. Pollina echinata. 2 n = 18 vel 36. Perennial herbs or rosette shrubs ( Marshalljohn - stonia). Leaf blades linear (or scale-like distally), elliptic, or rhombic; margins entire, dentate, or lobed. Capitula borne singly. Receptacles paleate ( Pinaropappus ) or not paleate. Involucres obconic or campanulate. Phyllaries 3 + -seriate, unequal. Florets 10 to 18 per capitulum. Corollas pink, pur¬ ple, or white. Cypselae columnar or fusiform, with 5 narrow, equally spaced furrows, not beaked (dis¬ tally attenuate in Pinaropappus pro parte). Pappi of 40 to 60 equal, barbellate bristles, persistent. Pol¬ len echinate (Tomb et al., 1974). Chromosome number 2 n — 18 or 36. Included genera: Marshalljohnstonia Henrick- son, Pinaropappus Lessing. Pyrrhopappinae Joongku Lee & B. G. Baldwin, subtribus uov. TYPE: Pyrrhopappus DC. Herbae annuae vel perennes. Laminae foliorum ellip¬ ticae, lanceolatae, vel oblanceolatae; margines integri vel pinnatifidi. Capitula portata singulatim vel capitulescen- tiae laxe corymbiformes. Involucra cylindracea vel anguste campanulata. Phyllaria 2-seriata, inaequalia (extimis fas- cientibus calyculis). Flosculi 30—150+. Corollae flavae, purpureae, vel albae. Cypselae fusiformes, 5— 12-coslatae, rostris fragilibus. Pappi ex setis 119—155, aequalis, bar- bellatis (cum pilis minutis, reflexis subtentis in Pyrrho- pappo) constati, persistentibus. Pollina echinata ( Picrosia ) vel ecliinolophata (Pyrrhopappo). 2 n - 12, 14, vel 24. Annual or perennial herbs. Leaf blades elliptic, lanceolate, or oblanceolate; margins entire or pin- natifid. Capitula borne singly or capitulescences loosely corymbiform. Receptacles not paleate. In¬ volucres cylindrical to narrowly campanulate. Phyl¬ laries 2-seriate, unequal (the outer forming calyc¬ uli). Florets 30 to 150+ per capitulum. Corollas yellow, purple, or white. Cypselae fusiform, 5- to 12-ribbed, beaks fragile. Pappi of 119 to 155 equal, barbellate bristles (subtended by minute, reflexed hairs in Pyrrhopappus), persistent. Pollen echinate (Picrosia) or eehinolophate ( Pyrrhopappus ) (Feuer 6 Tomb, 1977). Chromosome number 2n =12, 14, or 24. Included genera: Picrosia I). Don, Pyrrhopappus DC. Kky to Principally North American Subtribes of Cichorieae la. Capitulescences monocephalic (capitula borne singly); cypselae not ribbed or furrowed; pappi absent .... . Phalacroseridinae lb. Capitulescences corymbiform to thyrsoid-paniculiform or subumbelliform, or if monocephalic, then evpselae ribbed or furrowed and pappi present (rarely absent in Microseris douglasii). 2a. Cypselae beaked. 3a. Some or all pappus bristles caducous or fragile. 312 Novon 4a. Capitulescences stipitate-glandular (gland heads ± disk-shaped); pappus hrislles falling together . Malacothricinae p.p. ( Calycoseris ) 4b. Capitulescences not stipitate-glandular; outer pappus bristles caducous or fragile, the inner persistent . Glyptopleurinae 3b. Pappus bristles persistent. 3a. Pappus bristles plumose . Stephanomeriinae p.p. (Rafinesquia) 5b. Pappus bristles barbellate or smooth. 6a. Capitulescences monocephalic; corollas yellow or orange; beaks shorter than to much longer than bodies of cypselae; apices of beaks glabrous . . Microseridinae p.p. (Agoseris) 6b. Capitulescences usually loosely corymbiform, sometimes monocephalic; corollas yel¬ low, purple, or white; beaks much longer than bodies of cypselae; apices of beaks glabrous or villous (if capitulescences monocephalic and corollas yellow, then apices of beaks villous) . Pyrrhopappinae 2b. Cypselae not beaked (apieally attenuate in Pinaropappus and Uropappus). 7a. Corollas usually yellow or orange (if white, then pappi of scales, scales and bristles, or bristle- tipped scales); pollen echinolophate. 8a. Phyllaries in 1 or 2 series, subequal, the outer not forming calyculi; pappi of scales or scales and bristles . Krigiinae 8b. Phyllaries in 2+ series, subequal or unequal, the outer sometimes forming calyculi; pappi of bristle-tipped scales or bristles . Microseridinae 7b. Corollas often pink or purple (if yellow or white, then pappi absent (Atrichoseris) or of only bristles); pollen usually eehinate (echinolophate in Lygodesmia). 9a. Receptacles not paleate, often bristly; florets 15 to 250 per capitulum; corollas usually yellow, sometimes white; pappus bristles often all or mostly caducous or readily falling . . . Malacothricinae 9b. Receptacles usually not paleate, rarely paleate (. Pinaropappus ), not bristly; florets 3 to 18 per capitulum; corollas usually pink or purple, sometimes white; pappus bristles usually persistent. I()a. Capitulescences monocephalic; phyllaries in 3+ series, unequal . Pinaropappinae l()b. Capitulescences usually corymbiform to thyrsoid-paniculiform, rarely monocephalic; phyllaries usually in 2 unequal series, the outer forming a calyculus. I la. Plants non-thorny herbaceous perennials or opposite-leaved annuals ( Skinner - soseris); pappus bristles barbellate; pollen usually echinolophate, rarely eehinate (Chuetadelpha, Shinnersoseris) . Lygodesmiinae lib. Plants thorny and non-thorny herbaceous perennials, alternate-leaved annuals, or shrubs; pappus bristles usually plumose (if barbellate and plants herbaceous perennials, then stems thorny (. Pleiacanthiis )); pollen eehinate Stephanomeriinae An expanded phylogenetic study of Malcothri- einae is under way to resolve the memberships of lineages within the non-monophyletic Malacothrix, prior to generic redel imitation. We arc also studying relationships of Thamnoseris (endemic to Islas Des- venturadas, Chile), which was included by Stebbins (195.3) in Stephanomeriinae. Data from additional gene regions are needed to gain refined under¬ standing of the relationships among the subtribes of New World Cichorieae treated here. Acknowledgments. We thank Theodore M. Bar¬ kley, Kenton L. Chambers, Victoria C. Hollowell, David J. Keil, John F. Pruski, and John L Strother for helpful comments on the manuscript and JLS and Alan R. Smith for assistance with the Latin diagnoses; Molly Forbes (University ol California Botanical Carden), Jon Rebman (San Diego Natural History Museum), and JLS (University Herbarium) for providing plant material, loans, or access to specimens. We especially thank L. I). Gottlieb for collaborating on the molecular systematic studies that underlie the taxonomic changes proposed here¬ in. Phis work was supported by a postdoctoral fel¬ lowship (to JL) from the Korea Science and Engi¬ neering Foundation and a grant (to JL) from the Lawrence R. Heckard Endowment Fund of the Jep- son Herbarium, University of California, Berkeley. Literature Cited Bremer, K. 1993. New subtribes of the I .acluceae (Aster- aceae). Novon 3: 328—330. - . 1994. Asteraeeae: Cladistics and Classification. Timber Press, Portland. Feuer, S. & A. S. Tomb. 1977. Pollen morphology and detailed structure of Compositae, tribe Cichorieae, 2; Subtribe Microseridinae. Amer. J. But. 64: 230—245. Jansen, It. k., R. S. Wallace, k.-J. kim & k. L. Cham¬ bers. 1991. Systematic implications of ehloroplast I)NA variation in the subtribe Microseridinae (Asteraeeae: Lactuceae). Amer. J. Rot. 78: 1015—1027. Jeffrey, ( 1. 1966. Notes on Compositae: I. The Cichorieae in east tropical Africa, kew Bull. 18: 427—486. Lee, J., B. C. Baldwin & L. I). Gottlieb. 2002. Phylogeny of Stephanameria and related genera (Compositae— Lac¬ tuceae) based on analysis of 18S— 26S nuclear rl)NA ITS and FI'S sequences. Amer. J. Bot. 159: I0(t-I0R. - , - & - . 2003. Phylogenetic relation¬ ships among the primarily North American genera of Volume 14, Number 3 2004 Lee & Baldwin North America Genera of Cichorieae 313 Cichorieae (Compositae) based on analysis of 18S-26S nuclear rl)NA ITS and ETS sequences. Syst. Mot. 28: 6 1 6-626. Stebbins, G. L. 1953. A new classification of the tribe Cichorieae, family Compositae. Madrono 12: 65—81. Tomb. A. S.. D. A. Larson & .1. J. Skvarla. 1974. Pollen morphology and detailed structure of family Composi¬ tae, tribe Cichorieae: I. Subtribe Stephanomeriinae. Amor. J. Bot. 61: 486-498. Whitton, J.. R. S. Wallace & R. k. Jansen. 1995. Phylo¬ genetic relationships and patterns of character change in the tribe Lactuceae (Asteraceae) based on chloroplast DNA restriction site variation. Canad. J. Bot. 73: 1058- 1073. A New Combination in Matelea (Apocynaceae— Asclepiadoideae) Sigrid Liede and IJlrich Meve Department of Plant Systematics, University of Bayreuth, Universitatsstr. 30, D-95440 Bayreuth, Germany, sigrid.liede@uni-bayreuth.de; ulrich.meve@uni-bayreuth.de ABSTRACT. Chthamalia cuyabensis is transferred to Matelea cuyabensis and is lectotypified. Key words: Apocynaceae, Asclepiadoideae, Chthamalia, Matelea. In the course of compiling the Asclepiadoideae for the Prodromus Florae Matogrossensis (ed. B. Dubs, Switzerland), a species attributed to the ge¬ nus Chthamalia Decaisne (Apocynaceae— Asclepia¬ doideae; tribe Aselepiadeae, subtribe Gonolobinae) turned up. The genus Chthamalia, comprising eight species, was sunk by Woodson (1941) under Ma¬ telea Aublet, and Woodson (1941) combined the four North and Central American species, including the type species, C. nummularia Decaisne, under Matelea. The four Brazilian species were not treat¬ ed by Woodson (1941). The genus Matelea, with some 280 species, is one of the least known genera of Asclepiadoideae, and opinions diverge widely concerning the ques¬ tion whether Matelea is one large, variable genus, or whether several monophyletic genera are hidden within Matelea, or whether Matelea even has to be expanded to comprise the ca. 100 Gonolobus Mi- chaux species as well. No molecular phylogeny is yet available, and only some, mostly geographically restricted, revisions have been carried out in the last decades (e.g., Stevens, 1985, 1988, 2001). In Brazil, from where ca. 70 species ol Matelea s.l. are recorded, none of the four Chthamalia species has been mentioned in the literature lately. Nev¬ ertheless, Rapini (2001) cited Chthamalia as a syn¬ onym under Matelea. and there is no apparent mor¬ phological reason to reach a different conclusion. As the name Chthamalia has ceased to exist with the transfer of the type species to Matelea by Wood- son (1941), it is necessary to combine Chthamalia cuyabensis, the only species recorded from Mato Grosso, to Matelea cuyabensis in order to have a valid name available for the Prodromus Florae Ma¬ togrossensis, which is about to be published. As there is, to our knowledge, no ongoing research on Matelea s.l. in Brazil, and as a new name would need to be found ( Chthamalia not being available anymore) if new results were to require the split of the species from Matelea, we consider it best prac¬ tice to transfer the species in question to Matelea, while leaving the remaining three Brazilian species, lor which no immediate need for a name is known, untouched for luture research. Chthamalia cuya¬ bensis was first described by Malme (1927), who considered it closely related to C. majori E. Four¬ nier from Sao Paulo, from which it is said to differ by smaller flowers and the form of the leaves. Both characters may prove to be unreliable if more ma¬ terial comes to light, and it cannot be excluded that C. cuyabensis might constitute a subspecies, a va¬ riety, or even only a synonym of C. majori. How¬ ever, currently available data do not permit such conclusions. Matelea cuyabensis (Malme) Liede & Meve, comb. nov. Basionym: Chthamalia cuyabensis Malme, Ark. Bot. 21A(12): 22. t. 1, f. 2. 1927. TYPE: Brazil. Mato Grosso: Cuyabd, 12 Nov. 1902, G. (). A. N. Malme / / 2562 (lectotype, designated here, G not seen). The protologue is accompanied by an illustration of the habit, flower, and pollinarium. Literature Cited Malme, G. 0. A. N. 1927. Asclepiadaceae mattogrossen- sis. Ark. Bot. 21 A: 1-27. Rapini, A. 2001. Asclepiadoideae (Apocynaceae) da cad- eia do Espinhago de Minas Gerais, Brasil. Bob Bot. Univ. Sao Paulo 19: 55-169. Stevens, W. I). 1985. Asclepiadaceae. Pp. 228—241 in J. Rzedowski & G. C. de Rzedowski (editors), Flora Fa- nerogdmica del Valle de Mexico, Vol. 2. Instituto de Fcologfa, Mexico, 1). F. - . 1988. A synopsis of Matelea subg. Dictyanthus (Apocynaceae: Asclepiadoideae). Ann. Missouri Bot. Gard. 75: 1533-1564. - . 2001. Asclepiadaceae. Pp. 234—270 in W. 1). Stevens, C. Ulloa Ulloa, A. Pool & 0. M. Montiel (ed¬ itors), Flora de Nicaragua, Tomo I. Monogr. Syst. Bot. Missouri Bot. Gard. 85. Woodson, R. E. 1941. The North American Asclepiada¬ ceae. Ann. Missouri Bot. Gard. 28: 193-244. Novon 14: 314. 2004. Three New Speeies of Celastraceae (Hippocrateoideae) from Southeastern Brazil, and a New Combination in Peritassa Julio Antonio Lombardi Departamento de Botanica, Instituto de Ciencias Biologieas, Universidade Federal de Minas Gerais, Av. Antonio Carlos 6627, 31270-110, Belo Horizonte, Minas Gerais, Brazil. cissus@mono.icb.ufmg.br Abstract. The new Peritassa sadleri and Peri¬ tassa longifolia are distinguished by their habit, leaf characteristics, color of flowers, shape of pet¬ als, number of ovules, size of inflorescences, and size of fruits. The new species Salacia nemorosa is distinguished from other southeastern Brazilian species by inflorescences, disc details, and size anti shape of flowers and fruits. The new combination in Peritassa is also proposed: Peritassa glabra (A. C. Smith) Lombardi. KKSUMO. As novas especies Peritassa sadleri e Peritassa longifolia sao distintas pelos habitos, morfologia das folhas, cor das flores, forma das pe- talas, numero de ovulos e tamanho das infloreseen- cias e tlos frutos. Salacia nemorosa difere das out- ras congeneres do sudeste do Brasil pelas inflorescencias, detalhes do disco e tamanho e for¬ ma das flores e dos frutos. Tambem uma nova com- binayao em Peritassa e proposta: Peritassa glabra (A. C. Smith) Lombardi. Key words: Brazil, Celastraceae, H'PI >ocrateo- ideae, Peritassa , Salacia. I he former family Hippocrateaceae is today in¬ cluded in the Celastraceae as a subfamily, which includes 5 tribes, 24 genera, and ca. 357 described species (Hal 1^, 1990; Mennega, 1997) distributed worldwide in tropical and subtropical areas. The delimitations of the Neotropical genera are notoriously controversial, ranging from 2, Salacia and Hippocratea (Peyritsch, 1878), to 17 by Miers’s (1872) account. In this work I followed Smith (1940), who recognized three genera segregated from Salacia L., Cheiloclinium Miers, Peritassa Miers, and Tontelea Aublet, included in the tribe Salacieae with the African Salacighia Loesener and Thyrsosalacia Loesener (Mennega, 1997). The genus Peritassa is restricted to the Neotrop¬ ics with 14 species (Lombardi & Temponi, 1999; Mennega, 1997). Peritassa was described by Miers (1872) and segregated from Salacia by its anthers with longitudinal dehiscence with a prominent con¬ nective and a tubular nectariferous disc. Salacia is a genus of ca. 200 species (Mennega, 1997), distributed worldwide in the tropics and subtropics. Smith (1940) recognized 29 species from the Neotropics. The last comprehensive taxo¬ nomic study of the genus, although unpublished, is Hedin (1999), who recognized 38 species in the Neotropics, including 3 new laxa from Costa Rica, Ecuador, and Peru. The probable paraphyly of Salacia s. str. has been pointed out in recent molecular studies (Sim¬ mons et ah, 2001a, b), but before a more extensive analysis of worldwide representative species on both morphological and molecular aspects, it is premature to propose any generic realignments. While working on a taxonomic study of Brazilian species of Hippocrateoideae, as part of a study of the Neotropical species, I discovered three new species from the Espirito Santo Atlantic ridge (Ser- ra do Mar): two in Peritassa and one in Salacia. These newly described species occur in an area with few forest fragments, remnants from a vege¬ tation continuous in the past hut almost totally de¬ pleted now in Espirito Santo. Peritassa sadleri Lombardi, sp. nov. TYPE: Bra¬ zil. Espfrito Santo: Itarana, Alto Jatiboca, Faz. Frederico Sadler, 20°00'48.6"S, 40°54'39.9"W, 786 m, 26 Feb. 2003, ,/. A. Ixtmbardi 5205. A. Salino, R. C. Mota & L. G. Temponi (holotype, BHCB; isotypes, MBM, MBML, MO, NY, SPE). Figure 1. Peritassae mexiae disco brevi-tubulari et antherarum dehiscentia transversali-obliqua affinis, sod habitu, flomm colore, petalis orbicularibus, ovulorum numero et fruc- tuum forma et dimensione differt. Etiam Peritassae lon- gifoliae hie descriptae affinis, sed habitu, pedicelli di¬ mensione et fructuum forma et dimensione differt. Small tree, 4—5 m, glabrous, branches terete, lenticellate, sulcate, smooth, with seasonal growth marked by scale-like cataphylls. Leaves opposite; stipules 0.4— 0.8 mm long, caducous, triangular. No VON 14: 315-321. 2004. 316 No von Figure 1. Peritassa sadleri Lombardi. — A. Lateral view of flower. — If. Branch with leaves and inflorescences. — C. Lateral and adaxial views of stamens. — D. Lateral view of middle section of open flower showing disc, two stamens, and pistil. Drawn from the holotype, Lombardi el at. 5205 (BHCB). apiculate, minutely erase; petioles 0.4(— 0.8) cm, canaliculate; Blades 5.9—7.2(14.9) X 1.4— 2. 8(4.1) cm, elliptic, chartaceous, drying brawn, the base euneate, the apex acute or acuminate, tip cleft or apiculate, the margin entire and thickened, cream- colored when dry, venation brochidodromous with veins slightly prominulous on both sides. Inflores¬ cence thyrsoid-paniculate, 0.9— 1.4 X 1—1.5 cm, with 28 to 49 (lowers, axillary, sometimes at leafless nodes; bracts 0.7-1 .2 mm long, triangular, the mar¬ gin minutely erase, scarious; peduncle 0.1 — 0.3( — 0.7) cm long, terete, in old leafy nodes covered by tri¬ angular scales, on new shoots sometimes at leafless nodes and naked, then minutely lenticellate-punc- tulate; inflorescence branches opposite or sub¬ opposite, terete; pedicels (1.2— ) 1.7 mm long, terete, minutely punetulate, articulate at base; bracteoles 2, opposite at base of pedicel, triangular, 0.4 mm Volume 14, Number 3 2004 Lombardi Celastraceae from Southeastern Brazil 317 long. Flowers green, 3.7—4 mm wide at anthesis; sepals 5, 1.2— 1.3 X 1—1.1 mm. elliptic, unequal, ± carnose, the margin minutely erose; petals 5, 1.5— 1.8 X 1.6— 1.7 mm, subcircular, ± carnose, spreading at anthesis, the margin minutely erose; disc annular, short-tubular, ea. 0.4 mm high, 0.2 mm thick, carnose, free from reproductive parts, the margin entire; stamens 3, ca. 0.8 mm long, in open flower exserted and reflexed, the filaments flattened, the connective adaxially thickened, not surpassing the thecae, the anthers ca. 0.17 X 0.42 mm, oblong, slightly emarginate at base, basifixed. dehiscence extrorse by almost transverse apical slits; pistil 0.6 X 0.63 mm. conic, ovary slightly 3- lobed, 3-locular, with 2 subapical ovules in each locule, style ca. 0.16 mm long after anthesis, cen¬ tral, terete, stigmas obscure. Mature drupes 4—6.8 X 2.6 X 2. 1 cm, ellipsoid, the epicarp crustaceous, smooth, drying light brown-pruinose, minutely cream-colored punctuate; seeds not seen. Phenology. Collected with flowers and very young fruits in February; collected with mature fruits in October. Distribution and habitat. Peritassa sadleri oc¬ curs at altitudes of 700—800 m in the rain forests of Espfrito Santo, on the southeastern Atlantic coast of Brazil. This area has many small properties with old coffee plantations and pastures; each of these maintains small portions of forest in varied degrees ol preservation in order to fulfill federal government legislation. However protected by environmental laws, the conservation of this species is not sure while it is known only from outside of official con¬ servation sites. Etymology. The specific epithet of this species is an homage to the Brazilian-Pomeranian Sadler family who has preserved an apparently almost in¬ tact tract of forest at the intersection of many family members’ properties. This new species resembles Peritassa mexiae A. C. Smith by the disc, stamens, and pistil, but can be distinguished by the habit (small tree vs. liana), color of dried leaves (brown vs. cinereous), smaller inflorescences (0.9-1. 4 X 1-1.5 cm vs. (0.8— )2. 3— 5.0 X (0.8— )2.8— 7. 1 cm), color of flowers (green vs. yellow), shape of petals (± circular vs. elliptic), number of ovules in each locule (2 vs. 1), and big¬ ger fruits (4—6.8 X 2.6 X 2.1 cm vs. 2.4— 2.7 X 1.6— 2.2 X f.l cm). Peritassa sadleri is also very similar to Peritassa longifolia here described: the flowers are almost identical, but the two species differ in habit (small tree vs. liana), size of the ped¬ icels (1.2— 1.7 mm vs. 2.9 mm), and by different fruits (mature 4—6.8 X 2.6 X 2.1 cm, ellipsoid vs. immature 3.4— 4.8 X 2. 8-4. 2 cm, subpyriform). Paratypes. BRAZIL. Espfrito Santo: Brejetuba [in sched. ‘Brejatuba’], Rancho Dantas, 22 Oct. 2000, G. Hatschbach, M. Hatschbach & J. M. Silva 71523 (BHCB 1 2 J, MBM); Itarana, Jatiboca, Alto de Jatiboca, 3 km da Vila Jatiboca, Fazenda da familia Seidler [sic], 19°51'S, 40°52'W, 14 Feb. 1999. E. M. NicLughadha 204 , A. Mel- lo-Silva. B. L Stannard & M. C. Ass is (BHCB, K not seen, MB ML, SPF). Peritassa longifolia Lombardi, sp. nov. TYPE: Brazil. Espfrito Santo: Santa Teresa, Nova Lombardia, Bes. Biol. Augusto Ruschi, 750 m, 28 Nov. 2001, L. Kollmann 5055 , E. Hansen & W. Pizziolo (holotype, MBML; isotype, BHCB). Figure 2. Peritassae mexiae disco brevi-tubulari et antherarum dehiscentia transversali-obliqua affinis, sed foliomm for¬ ma, fforum colore, petalis orbicularibus, ovulorum numero et fructuum dimensione differt. Ftiam Peritassae sadleri hie descriptae affinis, sed habitu, pedicelli dimensione et fructuum forma et dimensione differt. Lianas, glabrous, branches terete, lenticellate, sulcate, smooth. Leaves opposite; stipules not seen, caducous; petioles 0.7— 0.9 cm, canaliculate; blades 7.5—17.7 X 1. 9-4.3 cm, elliptic, ehartaceous, dry¬ ing cinereous, opaque, the base cuneate, the apex acuminate, tip apiculate, the margin entire and thickened, venation brochidodromous with veins slightly prominulous on both sides. Inflorescence thyrsoid-paniculate, 1.1— 1.8 X 1. 1-1.8 cm, with ca. 15 flowers, axillary; bracts 0.4— 0.6 mm long, triangular, the margin entire; peduncle obsolete; in¬ florescence branches arising front an axillary tuft of six triangular sc ales; branches opposite or alter¬ nate, terete; pedicels 2.9 mm long, terete, minutely punctulate; bracteoles 2, opposite at base or middle of pedicel, triangular, 0.3 mm long. Flowers green, 4.6 mm wide at anthesis; sepals 5, 0.8-0.9 X 1.2- 1.3 mm, triangular or elliptic, unequal, ± carnose, the margin minutely erose; petals 5, 1.9— 2.1 X 2 mm, ± circular, ± carnose, spreading at anthesis, the margin entire to minutely erose; disc annular, short-tubular, ca. 0.3 mm high, 0.2 mm thick, ear- nose, free from reproductive parts, the margin en¬ tire; stamens 3, ca. 0.7 mm long, in open flower exserted and reflexed, the filaments flattened, the connective adaxially thickened, darkened when dry, not surpassing the thecae, the anthers ca. 0.22 X 0.48 mm, oblong, slightly emarginate at base, basifixed, dehiscence extrorse by almost transverse apical slits; pistil 0.6 X 0.7 mm, conical, ovary slightly 3-lobed, 3-locular, with 2 subapical ovules in each locule, style ca. 0.3 mm long after anthesis. 318 Novon of stamens. Drawn from the isotype Kollmann 5055 et al. (BHCB). central, terete; stigmas obscure. Mature drupes not seen, immature drupes 3.4— 4.8 X 2.8— 4.2 cm, ± pyriform, the epicarp crustaceous, smooth, drying brown-pruinose, minutely alveolate; seeds ea. 5, 2.3 X 1 cm, reniform. Phenology. Collected with dowers in Novem¬ ber; collected with immature fruits in April anil May. Distribution and habitat. Peritassa longifolia occurs at altitudes of ca. 700 m in the rain forests of Espfrito Santo, on the southeastern Atlantic coast of Brazil, and is known only within the limits of the Augusto Ruschi Biological Reserve, an area with 4000 ha of primary rain lorest. This area is one of the lew intact and relatively large forest fragments in Espfrito Santo. The Biological Reserve provides protection for this species, although the population status is unknown. This new species resembles Peritassa mexiae by the disc, stamens, and pistil, and the dried leaf color and texture, but may be distinguished by the blade apex (acuminate vs. acute), smaller inflores¬ cences (1.1— 1.8 X 1.1— 1.8 cm vs. (0.8— )2. 3— 5.0 X (0.8-)2.8— 7.1 cm), color of flowers (green vs. yel¬ low), shape of petals (± circular vs. elliptic), num¬ ber of ovules in each locule (2 vs. 1), and bigger fruits (immature 3.' 4—4.8 X 2.8— 4.2 cm vs. mature 2.4— 2.7 X 1.6— 2.2 X 1.1 cm). Peritassa longifolia is also similar to Peritassa sadleri described herein, with the almost identical flowers. However, the two species are distinguished by their habit (liana vs. small tree), by the bigger pedicels (2.9 mm vs. 1.2— Volume 14, Number 3 2004 Lombardi Celastraceae from Southeastern Brazil 319 1.7 mm), and by different fruits (immature 3.4— 4. 8 X 2.8— 4.2 cm, subpyriform vs. mature 4—6.8 X 2.6 X 2.1 cm, ellipsoid). Paratypes. BRAZIL. Espfrito Santo: Sarila Teresa. Nova Lombardia, Res. Biol. Augusto Rusclii, 8 May 2002. R. R. Vervloet 230 & E. Hausen (BHCB, MBML); trilha Roda D'Agua, 2 Apr. 2002, R. R. Vervloet 53, E. Hausen & W. Pizziolo (BHCB, MBML). Comments on the two Peritassa species described herein. The transversally dehiscent anther locule and inconspicuous connective group Peritassa lon- gifolia and P. sadleri with P. calypsoides (Cambes- sedes) A. C. Smith, P. rnexiae, and the former Ton- telea glabra, whose combination in Peritassa is here proposed; these five species stand apart from remaining Peritassa species by their almost trans¬ verse anther dehiscence slits and connective not exceeding thecae. However, an intermediary mor¬ phology is found in P hatschbachii Lombardi, with its oblique anther dehiscence slits and a connective which slightly or not at all surpasses the thecae. Salaeia nemorosa Lombardi, sp. nov. TYPE: Bra¬ zil. Espfrito Santo: Santa Teresa, Res. Biol. Santa Lucia, trilha do palmiteiro, 27 Jan. 2000, V. Demuner 621 & E. Hausen (holotype, MBML; isotype, BHCB). Figure 3. filter species austro-orientales Brasilienses inflorescen- tia forma singulari distincta. Salaciae mosenii disci forma similis, sed disci externa margine suleo praedito, floribus maioribus et fructibus minoribus pyriformibus distincta. frees, ca. 10—12 m, DBH 35 cm, glabrous, branches terete, lenticellate, sulcate, smooth, with seasonal growth marked by scale-like cataphylls. Leaves opposite; stipules 0.6 mm, persistent, tri¬ angular, margin denticulate; petioles 0.6— 0.7(— 0.8) cm, canaliculate; blades (8— )8.9— 10.6 X 3-4(— 4.9) cm. elliptic, chartaceous, drying brown, the base cuneate, the apex acuminate or acute, tip slightly emarginate, the margin entire and thickened, cream-colored when dry, venation broehidodromous with veins slightly prominulous on both sides, cream-colored when dry. Inflorescence thyrsoid-pa- niculate, 1.8— 2.8 X 1—2.5 cm, with ca. 18 flowers, axillary at leafless nodes; bracts 0.7-1. 3 mm long, triangular, the margin erose; peduncle 0.8— 1.3 cm, terete, all branches opposite, terete; pedicels (1.9— )2.9 mm long, terete, minutely punctuate; bracteoles 2, opposite at base of pedicel, triangular, 0.7 mm long. Flowers yellow, 4.7 mm w ide at anthesis; calyx syn- sepalous, sepals 5, lobes 0.5 X 1 mm, triangular or elliptic, coriaceous, the margin entire, involute; petals 5, 2.2 X 1—1.2 mm, elliptic, ± chartaceous, spreading at anthesis, the margin undulate; disc ca. 0.6 X 0.4 mm, conical, carnose, free from repro¬ ductive parts, the margin grooved, entire; stamens 3, ca. 1 mm long, at first appressed together around pistil, after exserted and reflexed, the filaments flat¬ tened, the connective adaxially thickened, the an¬ thers ca. 0.17 X 0.4 mm, oblong, slightly emargin¬ ate at apex, basifixed, dehiscence extrorse by oblique almost transverse apical slits; pistil 0.65 X 1 mm, conical, ovary 3-lobed, 3-locular, with 2 ax¬ illary ovules in each locule, style ca. 0.5 mm long after anthesis, central, conical, 3-angled, stigmas obscure. Immature drupes 5.4 X 3.5 cm, pyriform, sometimes ribbed at base, the epicarp crustaceous, drying brown-pruinose, minutely rugose; seeds I or 2. 2.8 X 2.3 X 1.5 cm, reniform. Phenology. Collected with flowers in January; collected with immature fruits in October. Distribution and habitat. Salaeia nemorosa oc¬ curs at altitudes of ca. 650—850 m in the rain for¬ ests of Espfrito Santo, on the southeastern Atlantic coast of Brazil. This taxon is known only from Santa Teresa municipality, which includes the Santa Lu¬ cia and Augusto Rusclii Biological Reserves, areas with 2500 and 4000 ha, respectively, of primary and secondary rain forests inside the city limits. The biological reserves provide adequate protection for this species, which have reproductive popula¬ tions in different places in Santa Teresa. This new' species is distinguished from all other southeastern Brazilian species by its pedunculate inflorescence; Salaeia elliptica (Martins ex Schul¬ tes) G. Don, Salaeia crassifolia (Martius ex Schul¬ tes) G. Don, and Salaeia grandifolia (Martius ex Schultes) G. Don have fasciculate inflorescences, while Salaeia mosenii A. C. Smith has chiefly ses¬ sile thyrsoid-paniculate inflorescences and Salaeia arborea (Schrank) Peyritseh has simple or com¬ pound dichasia. It resembles Salaeia mosenii in its conic disc, but it differs because the disc has a flattened outer margin instead of a groove and larg¬ er flowers (4.7 mm vs. 2.4— 4.3 mm) and different fruit shape (pyriform vs. spherical). Paratypes. BRAZIL. Espfrito Santo: Santa Teresa, Alto Sao Lourengo, Sftio da Cachoeira, 25 Oct. 2000, V. Demuner 1490, E. Hausen & W. Pizziolo (BHCB, MBML); Santa Teresa, Est. Biol, de Santa Lucia, 22 Sep. 1993, L. I). Thomaz 1722 (BHCB, MBML); Santa Teresa, Res. Biol. Augusto Ruschi, estr. Alto Goiapaba— Agu, 10 Jan. 2002, L Kollrnann 5294 & E. Hausen (BHCB, MBML), 10 Jan. 2002, L. Kollrnann 5299 & E. Hausen (BHCB, MBML); trilha da cachoeira, 29 Jan. 2002, /.. Kollrnann 5361 & E. Hausen (BHCB, MBML); Santa Teresa, Nova Lombar¬ dia, Res. Biol. Augusto Rusclii, estr. Goiapaba— Agu, 24 Oct. 2002, R. R. Verrloet 1272, E. Hausen & W. Pizziolo (MBML). 320 Novon Figure 3. Solaria nemorosa Lombardi. — A. Lateral view of flower. — lb branch with leaves and inflorescences. — C. Lateral and adaxial views of stamens. — D. Lateral view of middle section of open flower showing disc, two stamens, and pistil. Drawn from the isotype Demuner 621 & Hausen (BHCB). Iii addition to the aforementioned novelties, a new combination is proposed. IVritussa glabra (A. C. Smith) Lombardi, comb, nov. Basionym: Tontelea glabra A. C. Smith, Brittonia 3: 500. 1940. TYPE: Bolivia. La Paz: San Carlos, Mapiri region, 850 m, 5 Feb. 1927, O. Buchtien 1728 (holotype, NY; iso¬ type, F). Smith (1940) revised the Hippocrateoideae (as Hippocrateaceae) from the Neotropics, recognizing I 15 species, including Peritassa mexiae A. C. Smith and Peritassa calypsoides (Cambessedes) A. C. Smith. These species have an inconspicuous an¬ ther connective and oblique dehiscence clefts, al¬ most transverse, and are distinguished from other fleshy-fruited genera (tribe Salacieae, Mennega, 1997) because the anther locules, although dehis¬ cent by transversal clefts, are not confluent at the apex. The type of Peritassa calypsoides was not studied hv Smith, who confused this species with a previously undescribed Peritassa species, P. hatschbachii Lombardi (Lombardi & Temponi, Volume 14, Number 3 2004 Lombardi Celastraceae from Southeastern Brazil 321 1999). Peritassa mexiae was described based on a single specimen with detached “imperfect” flowers found between the inflorescence branches (Smith, 1940: 518). I examined the isotype specimens (K Mexia 5206, A, BM, F, GH, NY, VIC), and the flowers referred to by Smith appear to be older dried flowers that had fallen between the inflores¬ cence branches and remained on the sheet. At the same time. Smith described Tontelea glabra A. C. Smith (1940: 500), but did not perceive the simi¬ larity of this species with Peritassa mexiae. I went through the types of Tontelea glabra and a number of Amazonian collections from several herbarium collections (CTES, F, IAN. I PA. MG, NY. R, RB, S, U, UB, US) and did not encounter the anther character of “confluent horizontal apical clefts" (Smith. 1940: 501) used by the author for inclusion of this taxon in Tontelea. Tontelea in Smith’s (1940: 355) key is distin¬ guished from Peritassa by the dehiscence of anthers by confluent slits, while Peritassa dehiscence was by non-confluent slits. Almost all species in Ton¬ telea have oblong, relatively large anthers (0.2— 0.4 X 0.3— 0.6(0.8) mm) with clearly transverse dehis¬ cence slits and 3-lobed stigmas, alternate or op¬ posite with stamens, except for the aberrant species Tontelea cylindrocarpa, T. myrsinoides, T. nectan- drifolia, and T. glabra, which have relatively small¬ er anthers (0.1— 0.3 X 0.2— 0.3(0. 5) mm) and unlo- bed stigmas. The transfer of T. glabra to the genus Peritassa diminishes the morphological discrepan¬ cy within Tontelea ; the generic placement of the other three divergent species remains to be studied. Acknowledgments. The author thanks T. S. Fil- gueiras for the Latin diagnoses, Victoria Hollowell, Jon Ricketson, and Mark Simmons for critically re¬ viewing the manuscript, GNPq for the research fel¬ lowship grant (523026/96-0), and FAPEMIG for a financial research grant (CBS-2080/96) as well as the support of tin* author’s participation in the Bra¬ zilian Botanical Congress in 2003. I thank the cu¬ rators of the following herbaria for the loans of specimens: A, BM, CTES, F, GH, IAN, IPA, MBM, MBML, MG, MO, NY, R, RB, S, SPF, U, UB. US, VIC. Drawings are by the author. Literature Cited Hall6, N. 1990. Cdlastracdes (Hippoerateoiddes). Pp. 1— 247 in 15. Satabie & P. Morat, Flore du Cameroun 32. Ministdre de l’Enseignement Supdrieur de l’lnformatique et de la Recherche Scientifique, Yaoun¬ de. Hedin, J. P. T. 1999. Systematic Studies of the Neotropical Species of Salacia L. (Hippocrateaeeae) and Its Rela¬ tives. Ph.D. Thesis, Washington University, St. Ixmis. [Unpublished.] Lombardi, J. A. & L. G. Temponi. 1999. A new species of Peritassa Miers (Hippocrateaeeae) from southern Bra¬ zil, and notes on two confused species. Novon 9: 221— 226. Mennega, A. M. W. 1997. Wood anatomy ol the llippo- crateoideae (Celastraceae). IAWA .1. IB: 331—368. Miers, J. 1B72. On the Hippocrateaeeae of South America. Trans. Linn. Soc. London 2B: 319-432. Peyritisch, J. 1878. Hippocrateaeeae. Pp. 125-164 in C. F. P Martius & A. G. Eiehler, Flora Brasiliensis I 1(1). Frid. Fleischer, Lipsiae. Simmons, M. P., C. C. Clevinger, V. Savolainen, R. H. Archer, S. Mathews & J. F. Doyle. 2- 05-014-87 from COLCIENCI AS to the Universidad del Valle (Jorge E. Ramos-Perez. principal inves¬ tigator). Some of the collections were made during class field trips financed bv the Departamento de Biologfa of the Universidad del Valle. I .iterature Cited Calderon. E., (7 Galeano & N. Garcfa (editors). 2002. Libro rojo de plantas fanerdgamas de Colombia, Vol. I. Chrysobalanaceae, Dichapetalaceae y Lecythidaeeae. Instituto Alexander von Humboldt, Institutode Ciencias Naturales-Universidad Nat ional de Colombia, y Minis- terio del Medio Ambiente, Bogota. Greuter, W., J. McNeill, F. R. Barrie, II. M. Burdet, V. Demoulin, T. S. Filgueiras, D. II. Nicolson, P. C. Silva, J. E. Skog, P. Trehane, N. J. Turland & I). L. Hawk- sworth (editors). 2000. International Code of Botanical Nomenclature (Saint Louis Code). Regnum Veg. 138. Hailing. (7 1958. Monograph of the Cyelanthaceae. Acta Horti Berg. 18: 1—428. - . 1973. Cyelanthaceae. In: G. Harling & B. Sparre (editors). Flora of Ecuador I: 1-48. - & R. Eriksson. 1998. Cyelanthaceae. Pp. 471- 486 in J. A. Steyermark, P. E. Berry & B. K. Holst (editors), Flora of the Venezuelan Guayana, Vol. 4. Mis¬ souri Botanical Garden Press, St. Louis. Festuca dinirica and F. guaramacalana (Poaceae, Loliinae), Two New Species from the Venezuelan Andes Daniel Stancik Department of Botany, Faculty of Science, Charles University, Benatska 2, 128 01 Prague, Czech Republic, dan_stancik@yahoo.com Abstract. Two new species, Festuca dinirica Standfk and F. guaramacalana Stancfk (Poaceae, Loliinae), from the Venezuelan Andes are de¬ scribed. Festuca dinirica is found in the paramo zone and seems best placed in subgenus Festuca sect. Aulaxyper ; Festuca guaramacalana is found in humid mountain forests and belongs in Festuca subg. Subulatae sect. Subulatae. Both species are endemic to Venezuela, known from small areas of Dinira National Park and Guaramacal National Park, respectively. Resumen. Se presentan dos especies nuevas, Festuca dinirica Stancik and F. guaramacalana StanCfk (Poaceae, Loliinae), endemicas de los An¬ des de Venezuela. La especie F. dinirica pertenece al subg. Festuca sect. Aulaxyper y se conoce solo de la zona paramuna del Parque Nacional Dinira. La especie F. guaramacalana es un representante del subg. Subulatae sect. Subulatae , eudemica de la zona del bosque humedo altoandino del Parque Nacional Guaramacal. Key words: Andes, endemic. Festuca, Loliinae, Poaceae, Venezuela. The genus Festuca L. is a highly diversified cos¬ mopolitan grass genus, represented by over 140 species (Darbyshire et ah. 2003) in the Andes of South America and one extra-Andean enclave in southeastern Brazil and northeastern Argentina. Revision of Festuca in Venezuela has resulted in recognition of two new species. Festuca dinirica and F guaramacalana. Type collections of these new species come from recently organized field studies in little known regions of Dinira National Park (Stale Lara) and Guaramacal National Park (State Trujillo), and represent first collections of Festuca from these areas (for comparison, see Al¬ exeev. 1986; Briceno & Morillo, 1994; Dorr et ah, 2000; Vareschi, 1970). Both species appear to be endemic to the Venezuelan Andes. Festuca dinirica belongs in subgenus Festuca L. sect. Aulaxyper Dumort. This section is represented by only a few (probably 2 to 4) species in South America and remains insufficiently known. The geographically nearest species with clear affinity to section Aulaxyper is the markedly distinct species Festuca andicola Kunth from swampy habitats in Colombia and Ecuador. Festuca guaramacalana belongs in subgenus Subulatae (Tzvelev) E. A I exeev sect. Subulatae Tzvelev represented in Venezuela by two other spe¬ cies (F. ulochaeta Nees ex Steudel and F. coromo- tensis Briceno). An artificial key for distinguishing these species is provided. The survey of known South American members of Festuca sect. Subula¬ tae was published recently (Stancfk & Peterson, 2002) and included eight species. Festuca dinirica Stanfnk, sp. nov. TYPE: Vene¬ zuela. Lara: Munic. Humocaro Alto. NP Di¬ nira, 09°35'38.7"N, 70°07'12.2"W, 3170 m, 30 Nov. 2000, Standik 4287 (holotype, PRC; iso¬ types, CAR, COL). Figure If-j. Haec species a Festuca tolucensi innovationibus mixtis non intravaginalibus, ligula foliari 0.5 (non 1 .8—3.5) mm longa, foliis tenuioribus 0.3— 0.5 (non 0.5— 0.7) mm latis, glabris (non scabris), sclerenchymate abaxiali cum fasci- culis vascularibus non junctis differt. Tussocks, 50—60 cm tall; innovations mixed; culms erect, glabrous, with 1 to 3 nodes in basal half. Sheaths coriaceous-membranous, brownish gray, glabrous, fibrous; ligule 0.5 mm long, shortly ciliate; leaf blades linear, 20—30 cm long, 0.3— 0.5 mm wide, involute, pungent, glabrous, green. Pan¬ icle contracted, lanceolate, 6—10 cm long and 0.7- 1 cm wide; branches scabrous. Spikelets 8-10 mm long, lanceolate, with 3 to 4 perfect florets; rachilla scabrous or sparsely covered by bairs; glumes membranous, upper Vs scabrid; lower glume 3.5— 4.2 mm long, lanceolate, acute, I -nerved; upper glume 6-7 mm long, lanceolate, acute, 3-nerved; lemmas 6—7 mm long, 5-nerved, membranous, lan¬ ceolate, scabrous, awned; awn 1-3 mm long; callus glabrous; palea membranous, 2-carinate, scabrous, markedly 2-dentate, as long as lemma; lodicules lanceolate, 2-dentate; stamens 3, anthers 2—2.5 mm Novon 14: 341-344. 2004. 342 Novon Figure ]. a— e. Festuca guaramacalana Stanci'k (StanFik 4286, PRC). — a. Habit (bar: 9 cm). — b. Fragment of panicle (bar: 32 mm). — c. Spikelet (bar: 14 mm). — d. Glumes (bar: 7 mm). — e. Cross section of leaf (bar: 0.4 mm), f— j, Festuca dinirica Stani'fk (Stanfi'k 4287. PRC). — f. Habit (bar: 5 cm). — g. Panicle (bar: 2.5 cm). — h. Spikelet (bar: 10 mm). — i. Glumes (bar: 9 mm). — j. Cross section of leaf (bar: 0.3 mm). Volume 14, Number 3 2004 Stancik Species from Venezuelan Andes 343 long; ovary apex glabrous. Caryopsis lanceolate; hi- lum linear, %—% of total. Leaf blade section with 5 vascular bundles and 3 to 5 ribs above; scleren- chyma under abaxial and adaxial epidermis dis¬ continuous, vascular bundles free; adaxial epider¬ mis with microhairs about 0.09 mm long. Observations. Spikelet structure in the new species resembles Festuca tolucensis, another fre¬ quent species of the Venezuelan paramos. However, F. dinirica differs clearly in organization of the tus¬ socks (small, with mixed innovations in contrast to the big tussocks with intravaginal innovations of F. tolucensis), in structure of sheaths (brown, striate, and fibrous sheaths in contrast to the stramineous and integrated sheaths), length of the ligule (0.5 mm vs. 1 .8— 3.5 mm), and in the anatomic structure of the leaves (abaxial sclerenchyma isolated and all vascular bundles free in contrast to the continuous abaxial sclerenchyma and some vascular bundles united with sclerenchyma). The formation of the mixed innovations and brown, striate sheaths (partly disintegrated into fi¬ bers) make Festuca dinirica close to section Aulax- yper. In South America, this section is not very fre¬ quent and only poorly studied. The Colombian and Ecuadorian species Festuca andicola is (geograph¬ ically) the nearest species with clear affinity to sec¬ tion Aulaxyper. Festuca andicola differs from F. di¬ nirica by growing in swampy areas, forming only solitary culms with long rhizomes, and in having distinctive spikelets (lower glume 1.4— 1.8 mm long vs. 3.5— 4.2 mm; upper glume 2—2.5 mm long vs. 6—7 mm; lemma 5—5.5 mm long vs. 6—7 mm; awn 0.5— 0.7 mm long vs. 1—3 mm; anthers 0.8— 1.1 mm long vs. 2-2.5 mm). Distribution and habitat. Festuca dinirica is endemic to Venezuela and known only from the type locality — Dinira National Park in Lara. This species forms small tussocks in shrubby-grassy pa¬ ramos, associated with Blechnum L., Espeletia Mu- tis ex Bonpland, Hypericum L., Calamagrostis Adanson, Chusquea Kunth, Rhynchospora Vahl, etc., at an altitude of ca. 3200 m. No more infor¬ mation about the distribution of this species is available. It is probably steno-endemic and vulner¬ able to extinction. The specific epithet refers to the type locality, Dinira National Park. Paratypes. VENEZUELA. Lara: II umocaro Alto. NP Dinira, 09°35'38.7"N, 70°07' 1 2.2"W, grass paramo with shrubby patches, 3170 m, 30 Nov. 2000, Stancik 4289 (CAR, COL, PRC), Stan¬ cik 4288 (CAR, COL). Key to the Venezuelan Species Festuca sect. SUBULATAE la. Lemma with awn markedly flexuous, 7—15 mm long . 1 . F. ulochaeta lb. Lemma awnless or with straight awn 0.5—1 mm long . 2 2a. Spikelets 9—11 mm long, with 3 to 4 (lorets, awn 0—0.5 mm long . 2. F. coromotensis 2b. Spikelets 15—17 mm long, with 5 to 7 florets, awn 0.5-0. 1 mm long . . 3 . F. guaramacalana Festuca guaramacalana Stancik. sp. nov. TYPE: Venezuela. Trujillo: Bocono, PN Guaratnacal, 09°14'15. 1"N, 70°1 I '14.3"W. 2880 m, 29 Nov. 2000, Stancik 4286 (holotype, PRC; isotypes, CAR, COL). Figure la-e. Haec species a Festuca coromotensi ligula folii acuta non truncata, 1 — 1.5 (non 0.5— 0.7) mm longa, spiculis 15— 17 (non 9—11) mm longis, lemmate 9.5—10.5 (non 7.5— 8.5) mm longo distinguitur. Rhizomatous grass, forming small tussocks, 100- 130 cm tall; innovations extravaginal; culms erect, glabrous, with 3 or 4 nodes in distal half. Sheatli membranous-coriaceous, purplish brown, striate, fi¬ brous at base, margins lree; auricles absent; ligule 1 — 1.5 mm long, (coriaceous)membranous, acute; leaf blades linear, involute, 30—40 cm long and 0.6— 0.7 cm wide, (olive-)green, glabrous. Panicle branched, flexuous, pendent, elongate, 15—20 cm long and 2—5 cm wide; branches finely scabrous. Spikelets narrowly lanceolate, 15—17 mm long, with 5 to 7 perfect florets; rachilla pilose; glumes pur¬ plish, narrowly lanceolate, acute, coriaceous-mem¬ branous, sparsely scabrous; lower glume 3.7— 4.7 mm long, 1 -nerved; upper glume 5—7 mm long, 3- nerved; lemmas coriaceous-membranous, lanceo¬ late, 9.5—10.5 mm long, 5-nerved, purplish, apex entire, papillose, mucronate or shortly awned, the aivn 0.5—1 mm long; callus sparsely pilose; palea two-carinate, finely pilose, almost as long as lemma, apex shortly two-dentate; lodicules ovate, two-den¬ tate; stamens 3, anthers 2.5— 2.8 mm long; ovary apex glabrous. Caryopsis lanceolate; hilum linear, K-, of total length. Leaf blade cross section with ca. 1 I vascular bundles and 5 ribs above; sclerenchyma under both abaxial and adaxial epidermis discon¬ tinuous; unit with some vascular bundles forming girders, bulliform cells absent; adaxial epidermis with scattered microhairs, 0.09 mm long. Observations. This new species belongs to the section Subulatae, and among the South American members of this section Festuca guaramacalana has the largest spikelets and longest glumes (Stan- Cfk & Peterson, 2002). Similarly, as in the case of F. sodiroana Hackel ex E. B. Alexeev and versus 344 Novon typically long awns elsewhere in section Subulatae, the lemma is only shortly awned (0.5—1 mm). Mor¬ phologically, Festuca coromotensis B. Briceno ap¬ pears to he the most similar with truncate (vs. acute), shorter ligules (0.5— 0.7 mm long vs. 1-1.5 mm), shorter spikelets (9—11 mm long vs. 15—17 mm) with 3 to 4 (vs. 5 to 7) florets, and a shorter lemma (7.5— 8.5 mm long vs. 9.5—10.5 mm). Distribution and habitat. This species is en¬ demic to Venezuela. It is known only from the type collection from the National Park Guaramaeal (Tru¬ jillo), where it occurs at the margin of a brook with Neurolepis Meisner and Cortaderia Stapf in Andean mountain forest, at about 3000 m. This species probably occurs rarely, as no additional exemplars were found in the field or in herbarium collections. It is possibly a steno-endemic species vulnerable to extinction. The specific epithet refers to the type locality region, Guaramaeal National Park. Acknowledgments. I thank the staff of all the herbaria visited (AAU, B. C, CAR, COL, F, HFRZ, MER, MERC, MERF, MY. PORT, PRC, U. US. YEN, W). Thanks are due to Silvia Llamozas (YEN), B. Briceno (Universidad de los Andes, Me¬ rida), and Giuseppe Colonnello (CAR) for their spe¬ cial help and assistance. The investigation was fi¬ nancially supported by a grant from the Czeeh Grant Agency GACR (No. 206/03/0197). Literature Cited Alexeev, E. B. 1986. Festuca I.. (Poaceae) in Venezuela, Colombia A Ecuador. Novosti Sist. Vyssli. Hast. 23: 5— 23. [In Russian.] Briceno, B. & G. Morillo. 1994. El genero Festuca I.. (Po¬ aceae) en los paramos de Merida. Ernst ia 4(3-4): 73— 88. Darbyshire S. J.. B. J. Soreng, I). Stanffk & S. I). Koch. 2003. Festuca. In: R. J. Soreng, G. Davidse, P. M. Pe¬ terson, F. 0. Zuloaga, E. J. Judziewicz, T. S. Filgueiras & (). Morrone (editors), Catalogue of New World Grass¬ es (Poaceae): IV. Subfamily Pooideae. Contr. U.S. Natl. Herb. 48: 1-730. Dorr, L. J., B. Stergios, A. R. Smith & N. L. A. Cuello. 2000. Catalogue of the Vascular Plants of Guaramaeal National Park, Portuguesa and Trujillo States, Venezue¬ la. Contr. U.S. Natl. Herb. 40: 1-155. Stanffk, I). & P. M. Peterson. 2002. Two new species in Festuca from South America (Poaceae: Loliinae: Sect. Subulatae). Sida 20(1): 21-29. Vareschi, V. 1970. f lora de los pdramos de Venezuela, pp. 404—407. Universidad de Los Andes, Merida. Venezue- Sarocalamus , a New Sino-H imalayan Bamboo Genus (Poaeeae: Bambusoideae) Chris M. A. Stapleton l he Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 BAR, United Kingdom, c.stapleton@rbgkew.org.uk Grainne Nf Chonghaile and Trevor R. Hodkinson Department of Botany, University of Dublin, Trinity College, Dublin 2, Ireland. hodkinst@tcd.ie Abstract. Sarocalamus Stapleton, a new genus distributed from the eastern Himalayas to the Sich¬ uan Province of China, is described. Among all Asian bamboo genera it shares the greatest number of morphological characters with the North Ameri¬ can genus Arundinaria Michaux. However. DNA sequencing has suggested that other genera arc probably more closely related to Arundinaria, while Sarocalamus appears instead to be a possible sister group to PhyUostachys Siebold & Zuccarini. a ge¬ nus with some similarities in branching but very different flowers. Biogeograph ical implications arc discussed, and the new combinations Sarocalamus racemosus (Munro) Stapleton, S. faberi (Rendle) Stapleton, and .S'. spanostachyus (T. P. Yi) Stapleton are made. key words: Arundinaria , Bhutan, biogeography, China. India. Nepal, North A meriea, Sarocalamus. Characterization of the Asian temperate woody bamboos for the Flora oj Bhutan (Stapleton, 1994, 2000) and the forthcoming English-language ver¬ sion of the Flora of China has established that three species of Sino-Himalayan bamboo are mor¬ phologically most similar to the geographically dis¬ tant type species of Arundinaria Michaux, A. gi- gantea , from the southeastern United States (Texas to Maryland). One species , Arundinaria racemosa Munro, is from eastern Nepal, the Indian States of West Bengal and Sikkim, and Bhutan. The others, Arundinaria jaheri Rendle and Bashania spanos- tachya E P. \i, are from isolated mountains in southern Sichuan Province, and northeastern Yun¬ nan Province, Southwest China. Bashania fansi- panensis T. (,). Nguyen from northern Vietnam may also belong in this genus, but is not well known. Apart from Sarocalamus, all other Asian bam¬ boos can be separated clearly from the type species of Arundinaria on the basis of floral or vegetative morphological characters. This includes species in the genus Pleioblastus Nakai, which is still fre¬ quently synonymized within Arundinaria but differs in its fused prophyll margins (Stapleton, 1997). Al¬ though more than 500 species names have been pu blished in Arundinaria , only four, poorly known Asian species do not already have combinations in other genera and a different, modern placement (Ohrnberger, 1999). More than 30 Asian genera have been described by Japanese, Chinese, and European botanists to accommodate species origi¬ nally described in Arundinaria, and the number of genera currently recognized in this group ranges from 19 (Li, 1997) to 28 (Wang, 1997). In a mod¬ erate treatment Ohrnberger (1999) included 23 genera. Although the circumscription of Arundi¬ naria has remained controversial, with the number of Asian species included in the genus by different authors ranging from 16 (Chao & Chu, 1981) to none (Wang, 1997; Keng & Wang, 1996; Ohrnber¬ ger, 1999), it is clear that a large number of tem¬ perate Asian genera an; now recognized as being (list inct from that genus. Asian bamboos morphologically closest to the North American type species, Arundinaria gigan¬ tea, such as species of Sarocalamus, Pleioblastus, and Bashania P. C. Keng & T. P. Yi, arc often still included in Arundinaria (Clayton & Renvoize, 1986; Chao & Renvoize, 1989; Ei, 1997). However, molecular evidence from ITS, trn\ . intron, trn\,~ F intergenic spacer, and rpl 16 intron DNA sequence data (Zhang, 1996; Nf Chonghaile, 2002) has sug¬ gested that such species are not necessarily its closest relatives. Instead, Pseudosasa japonica (Steudel) Nakai has been suggested as the closest Asian relative of Arundinaria gigantea from both ndhV and rplAG sequence data (Zhang, 1996; Nf Chonghaile, 2002), a clade comprising A. gigantea and P. japonica receiving 86% BS (Nf Chonghaile, 2002), even though P. japonica has never been eon- Novon 14: 345-349. 2004. 346 Novon sidered a particularly close relative of A. gigantea on morphological grounds. Pseudosasa Nakai lias received almost universal recognition (McClure, 1973; Clayton Si Renvoize, 1986; Chao, 1989; De- moly, 1996; Keng & Wang, 1996). Although Sarocalamus shares the principal dis¬ creet macro-morphological character states with Arundinaria, it still differs in several, more quan¬ titative characters. These include shorter, more cy¬ lindrical. more separated and less imbricate florets of more consistent size, less than 10 X 2 mm; a more sinuous rhachilla with much longer inter¬ nodes; completely glabrous pedicels; and more del¬ icate, smoother, rugose rather than ribbed, less closely spaced leaf blades. The three species of Sarocalamus have been placed in four other Asian genera when not included in Arundinaria : Fargesia Franchet, Yushania P. C. Keng, Bashania, and Gel- idocalamus T. H. Wen. Sarocalamus differs quite profoundly from Fargesia and Yushania in having pachymorph rather than leptomorph rhizomes. It is closer to Bashania and Gelidocalamus. However, while those genera do have leptomorph rhizomes, they also have decurrent, strongly pulvinate synllo- reseence paraelades, and a much greater degree of compression of the basal internodes on their central branches, giving a higher number of branches at each culm node. Bashania also has thickened, prominent, long-ciliate prophyll keels, greater branchlet ramification and tougher leaves, while Gelidocalamus has much smaller florets, com¬ pressed rhachillas, swollen supra-nodal ridges, sol¬ itary, more lanceolate leaves, and its 7 to 12 sub¬ equal branches do not re-branch at all. Molecular evidence is generally lacking at the generic level in Asian woody bamboos, with low or absent support for many well-recognized taxonomic groupings. Combined analyses of data from differ¬ ent genes is starting to provide slightly better res¬ olution, however. What support has been shown at the generic level is generally congruent with mor¬ phological distinctions and current taxonomic treat¬ ments of Asian bamboos. In the most comprehen¬ sive study of woody bamboos (Nf Chonghaile, 2002), one species of Sarocalamus was included along with Arundinaria gigantea, species of Ba- shariia, Fargesia, Yushania, and 33 other genera. While the two species of Bashania, B. fargesii (E. C. Camus) P. C. Keng Si T. P. Yi and B. qing- chengshanensis P. (7 Keng Si T. P. Yi, formed a fairly well-supported elade (70% RS in parsimony analysis, 82% RS in N.l analysis), Sarocalamus fa- heri (included there as Bashania fangiana (E. (7 Camus) P. C. Keng & T. H. Wen) was not resolved within the Bashania elade and did not associate with any other genera in parsimony analysis. It re¬ mained a largely unresolved member of the well- supported monophyletic group of mainly Asian temperate bamboos first reported by Watanabe et al. (1994). It is now known to include 25 or more genera from Africa, Asia, and North America adapted to subtropical or temperate habitats in both hemispheres (Nf Chonghaile, 2002). hut is widely referred to as the “north temperate elade.” The generic name Sarocalamus means broom bamboo and is derived from saron (trapov: broom, sweepings) and calamus (Ka\ap.o£: reed, bamboo). It alludes to the very erect branching, reminiscent of a traditional broom or besom, and the frequent use of these bamboos for sweeping. Sarocalamus Stapleton, gen. nov. I \ PE: Saroca¬ lamus racemosus (Munro) Stapleton. Bashaniae et Arundinariae rhizomatibus tenuibus, staminibus 3, inflorescentia ebracteata, gemmis apertis similis, ab ilia ramis inflorescentiae erectis, intemodiis ramorum longioribus, foliis tenuibus, ab hac flosculis an- gustis, ab uterque pedicellis glabris, differt. Similar to Bashania and Arundinaria in its lep¬ tomorph rhizomes, 3 stamens, ebracteate inflores¬ cences and open buds. Differing from the former in its erect inflorescence branches, longer branch in¬ ternodes and thinner leaves, from the latter in its narrower florets, and from both in its glabrous ped¬ icels. Temperate self-supporting bamboo; rhizomes leptomorph; culms tillering (plurieaespitose), erect to drooping; internodes smooth, terete; nodes not swollen, unarmed; supra-nodal ridge well devel¬ oped; branch buds lanceolate, on promontory, en¬ closed within single, 2-keeled prophylls, always open at the front; branches initially I to 3, very erect, appressed, sulcate, basal internodes progres¬ sively increasing in length, basal internode often long, lateral branching often distant from culm; complement proliferating to become broom-like, at¬ tenuate; lateral branch axes always subtended by sheaths, no replication of lateral branches. Leaf blades linear-lanceolate, thin, less than 12 cm long, with prominent cross-veins (tessellate). Synfloresc- enee initially terminal or lateral to leafy branches, later in leafless complements; synflorescenee branches glabrous, without pulvini, remaining very erect and appressed, branching racemose or rarely partially paniculate, not fasciculate, subtended by very small remnants of sheaths or lings of hairs, rarely with very small pulvini. Spikelets 3 to 6 per synflorescenee, on glabrous promontory (pedicel); prophyll represented by lower glume; glumes 1 to 2, both small, lower glume usually very small, usu- Volume 14, Number 3 2004 Stapleton et al. New Sino-Himalayan Bamboo 347 ally distant from lower lemma, without subtended buds (flowering semelauctant); rhachilla often sin¬ uous, strongly flattened, more than half length of palea; palea keels curving, appressed to rhachilla; stamens 3; stigmas 3. Flowering gregarious. Distribution. East Nepal, India (West Bengal & Sikkim), Bhutan, Western China (Tibet, Sichuan and Yunnan Provinces), North Vietnam, 2800- 3900 m. 1. Sarocalamus racemosus (Illunro) Stapleton, comb. nov. Basionym: Arundinaria racemosa Munro, Trans. Linn. Soc. London 26: 17. 1868, emend. Gamble in Kew Bull. Misc. Inform. 1912: 138. 1912. Fargesia racemosa ( Munro) T. P. Yi, J. Bamboo Res. 2(1): 39. 1983. Yu- shania racemosa (Munro) R. B. Majumdar, in S. Karthikeyan et al., FI. Ind. Enumerat.— Monocot.: 283. 1989. TYPE: Nepal/Sikkim (India). Islumbo Pass |Chiya Banjyang], 13 Oct. 1857, Thomson s.n., Kew distribution number (K 6738) (lectotype, selected by Gam¬ ble (1912: 137). K). Arundinaria racemosa Munro was first described from collections of two species from the eastern Himalayas (Munro, 1868). It was emended by Gam¬ ble (1912), who separated the lower altitude col¬ lections from 6000 to 9000 ft. in the vicinity of Darjeeling into a new species, Arundinaria moling , now known as Yushania moling (Gamble) R. B. Ma- jumdar, leaving only the “high-level form,” found above 10.000 ft., in Arundinaria racemosa. The Munro syntypes were discussed in detail by Gam¬ ble (1912: 137) when describing his new species. He stated “it seems quite evident that Thomson’s Islumbo Pass flowering specimen is the real type of A. racemosa.'' This can be taken as an effective lectotypification under Article 7.1 1 of the St. I /mis Code (Greuter et al., 2000), as a type* element was indicated by direct citation including the term “type.” Chao and Renvoize (1989) later designated Munro’s first syntype, a Thomson collection made in 1857 from 6000 ft. on Birch Hill in Darjeeling, as lectotype of A. racemosa. They apparently did not realize that the species had already been lec- totypified by Gamble, even though they concur¬ rently lectotypified ,4. moling , in the protologue of w hich Gamble had made his lectotypification of A. racemosa. According to the label of Thomson’s Birch H ill collection, the lectotype designated by Chao and Renvoize was a “pale blue” bamboo. Two bamboos from Darjeeling, A. moling and tin* rarer / / i m alayacal amus hookerianus (Munro) Stapleton, both have pale blue culms, w hile culms of A. ra¬ cemosa are green or yellow. Gamble (1912: 137) had even stated that “Birch Hill is the end of the Darjeeling Ridge . . . and is a very unlikely locality for the high-level A. racemosa." However, as Gam¬ ble (1912: 137) had also pointed out, Thomson’s collection of some other, pale blue bamboo truly from Birch Hill would seem to have become lost, and the Birch Hill label is currently attached to a sheet bearing only a flowering specimen from Is¬ lumbo Pass, which bears the correct label as well. It would appear, therefore, that Chao and Renvoize 1 1989), by simply designating Munro’s first syntype as lectotype, may have inadvertently designated a long-lost collection of A. raiding or Himalayacala- mus hookerianus from Darjeeling as lectotype of A. racemosa, but fortunately there can be no doubt that the earlier lectotypification of Gamble (1912) takes priority. The Islumbo Pass material shows the distinctive branching and synflorescence of Saro¬ calamus and is accepted here as lectotype of A. racemosa Munro and type of the genus Sarocala¬ mus. It was collected by Thomson in 1857 in woods above the Islumbo Pass, now known in Nepali as Chiya Banjyang, on the Singalila Range, which separates East Nepal and Sikkim. The name Arun¬ dinaria racemosa is still often misapplied to Arun¬ dinaria moling Gamble in India (Tewari, 1993; Seethalakshmi & Muktesh Kumar, 1998), even though that species has been transferred by the Bo¬ tanical Survey of India as Yushania moling (Gam¬ ble) R. B. Majumdar. Habitat and distribution. Sarocalamus racemo¬ sus is found at an altitude of 2900-3800 m in the understory of Abies densa and mixed coniferous and rhododendron forests, in degraded forest, and in cleared areas of pastureland, in which il can be the principal species. It is often found on more freely drained sites than Yushania microphylla, which has hollow rhizomes and extends to a higher altitude. The type collection was made on a pass that con¬ stitutes the border between Nepal and Sikkim. Fur¬ ther collections have been made in Sikkim, where it was reported to be common in forests around Sen- clial (Gamble, 1896), and across western, central, and eastern Bhutan. It is to be expected that it will also be found in further localities in the eastern¬ most districts of Panchtar and Taplejung in Nepal. It has also been reported from Tibet (Yi, 1983) and is to be expected in the Indian State of Arunachal Pradesh. Additional specimens examined. INDIA. Sikkim: Sin¬ galila Range, Sukumbum, May 1889, l)r. dummies Col¬ lectors s.n. (K). BHUTAN. Puro: Chile La, 1 June 1971, Bedi 32 (K); Thimphu: Dorchula, 22 Dec. 1989 Stapleton 872 (TH1M). Tongsa: Yotong La, 15 Aug. 1989, Stapleton 348 Novon 832 (TIUM). Hiiiuthniig: Lame Gompa, 23 June 1990, lliirgi s.n. (K, THIM). Mongar: Thrumsingla, 11 Oct. 1990, Wood 7348 (E). Trashigung: Taahiyangtse, E side of Pang U, 7 June 2