A Synopsis of Rhynchospora (Cyperaceae) in Mesoamerica

Wm. Wayt Thomas

Se describe e ilustra Eleocharis reznicekii (Cyperaceae, subg. Eleocharis, serie Eleocharis), especie con características morfológicas intermedias entre las de Eleocharis densa y las de dos especies de la subserie Eleocharis (E. macrostachya y E. palustris). Se distingue por tener tallos conspicuamente comprimidos (3-5 veces más anchos que gruesos en fresco) y estilopodio sésil sobre una base anular fina. De E. densa se distingue además por tener espiguillas menos densas, estilos en su mayoría bífidos y aquenio casi liso a 30x, más redondeado hacia el ápice. De E. macrostachya y E. palustris lo hace por su hábito más robusto, espiguillas oblongas a linear-oblongas, gluma proximal envolviendo únicamente 1/3 a poco más de 1/2 de la base de la espiga, y por tener la gluma proximal y 1 a 2(3) glumas subproximales sin flor. Hasta ahora, E. reznicekii se conoce solamente del estado de Durango, en el Altiplano Mexicano y piedemonte de la Sierra Madre Occidental.

Systematic studies of scirpoid species in the Andes showed the necessity to exclude one species each from Scirpus and Carex. They are combined in a new genus Zameioscirpus and a third new species is described. The autonomous generic position of Zameioscirpus within the Scirpeae is supported by a phylogenetic analysis based on rbcL and trnL-F sequencing data and by conspicuous morphological similarities.

Cyperaceae is a cosmopolitan family with over 5,000 species, which are often found in wet habitats. In this study we found 31 hydrophylic species of Cyperaceae, belonging to 10 genera (Ascolepis, Calyptrocarya, Cyperus, Eleocharis, Fimbristylis, Fuirena, Lipocarpha, Pycreus, Rhynchospora and Scleria), that occur in the Araguaia River Basin, located mainly within the Brazilian Cerrado (savanna). Short descriptions, geographic distribution, illustrations, identification keys and comments are presented.

A synopsis of Rhynchospora (Cyperaceae) in Mesoamerica WM. WAYT THOMAS Thomas, Wm. Wayt (The New York Botanical Garden, Bronx, NY 104585126). A synopsis of Rhynchospora (Cyperaceae). Brittonia 44: 14-44. 1992.This treatment provides a key, brief synonymy, habitat descriptions, and distributions for the 81 species of Rhynchospora known from Mesoamerica. Included are descriptions and illustrations of three new species and one new subspecies (R. davidsei, R. rosae, and R. fascicularis subsp, sierrensis described by the author and R. talamancensis described jointly with J. G6mez-Laurito). In addition, the achenes of 17 species of Rhynchospora are illustrated as an aid in identification. Key words: classification, Cyperaceae, Mesoamerica, Rhynchospora. In 1931, Paul Standley published a treatment o f the Cyperaceae of Central America which included 28 species of Rhynchospora. In the ensuing 60 years, botanical exploration and a greater understanding of the genus have almost tripled the known number o f species in the region. This synopsis is intended as a companion to the Spanish treatment to appear in Flora Mesoamericana (Thomas, in press) and the region covered is the same: Central America and the tropical states of Mexico (Chiapas, Tabasco, Campeche, Yucatfin, and Quintana Roo). The format used here is similar to that o f Flora Mesoamericana (Davidse et al., 1982), but descriptions are lacking and 21 taxa are illustrated. For each species, a selected synonymy is presented. All cited specimens have been seen unless otherwise noted. While our understanding of Rhynchospora in Mesoamerica is much greater than it was in Standley's time, it is by no means complete. Throughout I have noted potential new species and taxa requiring further study. Much of the increase in known numbers o f species is from the coastal plain of the Gu lf o f Mexico, particularly in Belize and Nicaragua, where numerous species characteristic of the West Indies have been discovered. In addition, careful collecting, especially in Costa Rica, has revealed many Andean taxa previously known only from South America. R H Y N C H O S P O R A Vahl Rhynchospora Vahl, E n u m . pl. 2: 229. 1806. TYPE SPECIES"R. alba (L.) Vahl, from Schoenus albus L. Dichromena Michaux, F1. bor.-amer, l: 37. 1803. TYPE SPECIES: Dichromena leucocephala Michaux. Psitocarya Torrey, Ann. L y c e u m Nat. Hist. New York 3: 359. 1836. TYPE SPEOES: Psilocarya scirpoides Torrey. Vegetatively variable perennials or annuals. Spikelets ovoid or lanceoloid to terete or occasionally compressed, with several to many spirally arranged scales, the lower (1)2-several scales sterile. Flowers borne in axils of scales, the lower 1-several perfect, the upper 1-several functionally staminate. Stamens 1-3(12). Hypogynous bristles 0 6(20). Achenes usually lenticular to globose, topped with a persistent style base. Style undivided to deeply parted, the persistent base usually triangular to deltate, occasionally strap-like or subulate. Approximately 250 species. Primarily neotropical, but with species in temperate North and South America; also paleotropical and with a few species in temperate Europe and Asia. Brittonia, 44(1), 1992, pp. 14-44. 9 1992, by the New York Botanical Garden, Bronx, NY 10458-5126 ISSUED: 20 March 1992 1992] THOMAS: RHYNCHOSPORA Key to Mesoamerican 15 species of Rhynchospora Spikelets all sessile and arranged in l or more true capitula. 2 Capitula 2 or more per culm. 3 Capitula in a terminal, branching, inflorescence; spikelet scales usually brown to castaneous; hypogynous bristles exceeding achene. 4 Style base stout, as wide as or wider than achene, sulcate; largest leaf blades 15-21 m m wide .......................................................................................................................................................................................... 73. R. gigantea Link. 4 Style base slender, much narrower than width of achene (Fig. 8B), not sulcate; largest leaf blades 1-16 m m wide. 5 Leaves 7-16 m m wide; spikelets 8-11 m m long ........................................79. R. hassleri C. B. Clarke. 5 Leaves 1--4 m m wide, spikelets 3.2-7 m m long. 6 Achenes 2.5-3 m m long; capitula 1-2(3) per culm ............................................78. R. tracyi Britton. 6 Achenes 1.5-2 m m long; capitula (1)4-18 per culm .................................................................................. ..................................................................................................................... 77. R. holoschoenoides (L. C. Rich.) Herter. 3 Capitula axillary along culm; spikelet scales pale stramineous or pale brown; hypogynous bristles usually lacking or shorter than achene body, occasionally with 4 bristles exceeding achene. 7 Spikelet scales cartilaginous, shiny; mature spikelets 1.6-2.3 m m wide; achenes 2.8-3.2 m m long ................................................................................................................................ 61. R. torresiana Britton & Standley. 7 Spikelet scales papery, dull; mature spikelets 0.7-1.5 m m wide; achenes 1.5-2.2 m m long. ..................................................................................................................................................................................................... 25. R. exaltata Kunth. 2 Capitulum 1 per culm or reduced to a single spikelet. 8 Bracts subtending capitulum white basally. 9 Bracts subtending capitulum white adaxially and abaxially, glabrous or rarely ciliolate basally. t0 Plants cespitose; keels oflower spikelet scales and occasionally the margins ofthe bracts ciliolate. 11 Hypogynous bristles present; plant base slightly bulbous ...................................................................... ..................................................................................................................................... 76. R. oaxacana Kral & W. Thomas. 11 Hypogynous bristles absent; plant base not bulbous ................................................................................... ....................................................................................................................... 2. R. floridensis (Britton ex Small) Pfeiffer. 10 Plants with slender rhizomes; spikelet scales and bracts glabrous ........................................................ ................................................................................................................................................................................ 1. R. colorata (L.) Pfeiffer. 9 Bracts subtending capitulum white only adaxially, usually ciliate basally and often hirsute abaxially ...................................................................................................................................................... 3. R. nervosa (Vahl) Brckeler. 12 Plants rhizomatous; culms usually longer than the leaves; basal leaves 1-2.6 m m wide, the blades seldom hirsute abaxially .............................................................3a. R. nervosa subsp, nervosa. 12 Plants cespitose; culms usually shorter than the leaves; basal leaves 1.5--4.5 m m wide, the blades often hirsute abaxially .............................................3b. R. nervosa subsp, ciliata Koyama. 8 Bracts subtending the capitulum green, occasionally brown or stramineous or, rarely, reduced and scale-like. 13 Hypogynous bristles present; style base usually longer than broad. 14 Bracts subtending the capitulum greatly reduced, less than half as long as the spikelets. 15 Achene body strongly papillose and furrowed on 1 side; mature spikelets usually 5-20 per culm .......................................................................................... 69. R. mexicana (Liebmann) Steudel. 15 Achene body smooth, lenticular, not furrowed; mature spikelets usually 1-4 per culm .................................................................................................................................................................. 68. R. curvula Griseb. 14 Bracts subtending the capitulum mostly as long as o r longer than the spikelets. 16 Bracts subtending the capitulum ascending, enclosing the capitulum, stramineous to brown. 17 Bracts subtending the capitulum 3-7, cartilaginous, glabrous or glabrescent on margins; hypogynous bristles plumose ........................................................................................................ ........................................................................................................ 72. R. globosa (Kunth) Roemer & Schultes. 17 Bracts subtending the capitulum ca 12-25, chartaceous, densely ciliate; hypogynous bristles antrorsely scabrous ..................... 71. R. armerioides J. Presl. & C. Presl. 16 Bracts subtending capitulum deflexed to horizontal, usually green. 18 Largest inflorescence bracts 5-11 m m wide; capitulum 20-40 m m long; achenes lenticular without winged margins and with a narrowly triangular style base. ...................................................................................................................................................... 62. R. cephalotes (L.) Vahl. 18 Largest inflorescence bracts 0.5-2 m m wide; capitulum 7-17 m m long; achenes piano-convex with 2 incurved marginal wings or lenticular with a subulate style base. 19 Plants rhizomatous; style base subulate; achenes lenticular (rarely, R. holo- 16 BRITTONIA [VOL. 4 4 may key out here, but that species has achenes 1.5-2 mm, not 2.5-3 m m long) ..................................................................................................... 78. R . tracyi Britton. 19 Plants cespitose; style base narrowly triangular; achenes piano-convex with 2 incurved marginal wings (Fig. 1E) ............................... 70. R . b a r b a t a (Vahl) Kunth. 13 Hypogynous bristles absent; style base as broad as or broader than long. 20 Plants stoloniferous; spikelet scales white or white with a green nerve; spikelets 1--6 per culm; culms 0.6-13 crn tall. 21 Inflorescence of only 1 spikelet; spikelet scales entirely white, ascending and covering the achenes; culms 1--4.5 cm tall .................................................................... 8. R. a l b e s c e n s (Miq.) Kiik. 21 Inflorescence of 3-6 spikelets; spikelet scales white with a green nerve, spreading and often exposing the achenes; culms 0.6-13 cm tall ....................................................................... ....................................................................................................................................... 9. R . berteroi (Sprengel) C. B. Clarke. 20 Plants cespitose or rhizomatous; spikelet scales brown at maturity and spikelets 116(70) per culm or, rarely, spikelet scales white and then spikelets 3-9 per culm; culms 4-100 cm tall. 22 Spikelet scales white at maturity, slightly spreading; achenes strongly transversely rugose; culms 4-25 cm tall .............................................. 7. R . p u b e r a subsp, p a r v u l a W. Thomas. 22 Spikelet scales brown at maturity, appressed or rarely spreading; achenes transversely rugulose; culms 8-100 em tall. 23 Spikelet scales spreading; spikelets 7-70 per capitulum and 5-6.5 m m long ..... ...................................................................................................................................... 6. R . p o l y s t a c h y s (Turrill) Pfeiffer. 23 Spikelet scales appressed; spikelets 1-6 per capitulum and 7-12 m m long or 612 per capitulum and 11-24 m m long. 24 Spikelets 6-16, 11-24 m m long; largest eauline leaf blades 6-14 m m wide; bracts exceeding inflorescence 5-9 .......................... 5. R . w a t s o n i i (Britton) Davidse. 24 Spikelets 1-6, 7-12 ram long; largest cauline leaf blades 1.3-5 m m wide; bracts exceeding inflorescence 2-5 ........................................................................................................ ..................................................................................... 4. R . radicans (Schlechtend. & Cham.) Pfeiffer. 25 Spikelets single, rarely 2-3; culms often arching; bracts reflexed; style base 0.3-0.6 m m high ................................................... 4a. R . r a d i c a n s subsp, radicans. 25 Spikelets 3-6, central 1 largest; culms erect to slightly arching; bracts spreading to slightly reflexed; style base 0.15-0.35 m m high ................... ..... 4b. R . radieans subsp, m i c r o c e p h a l a (Bertero ex Sprengel) W. Thomas. 1 Spikelets, at least some, pedicellate and arranged in fascicles or panicles. 26 Hypogynous bristles present (occasionally easily deciduous or rarely vestigial). 27 Style base subulate, 2.3-10 m m long; achenes obovate or narrowly so, the surface punctate or rugulose. 28 Achenes biconcave, the upper part preach surface projecting in the shape of a rough inverted " U " (Fig. 8A) ........................................................................................................................................................ 75. R . s e u t e l l a t a Griseb. 28 Achenes lenticular to flattened, without an inverted " U . " 29 Athene surface rugulose, usually stramineous; mature spikelets 5.8-8.5 m m long; style base 4--4.8 m m long ........................................................................................................................................ 81. R . triflora Vahl. 29 Achene surface punctate, often reddish-brown or greenish; mature spikelets 8-12 mrn long; style base 3.5-9.2 m m long. 30 Style base 3.5-4.5 m m long; achenes usually greenish; mature spikelets 8-9 x 33.5 mm; spikelet scales usually olive with paler margins ............................................................... .................................................................................................................................................................. 79. R. hassleri C. B. Clarke. 30 Style base 6.2-9.2 m m long; achenes usually dark reddish-brown; mature spikelets 9.5-12 x 1.1-1.8 m m (not including exserted style base); spikelet scales reddishbrown ............................................................................................... 80. R . trispicata (Nees) Schrader ex Steudel. 27 Style base triangular, at least narrowly so, or even depressed, seldom more than 3 m m long; achenes usually widely obovate, the surface reticulate or sometimes rugulose. 31 Style base stout, sulcate, its base as wide as or wider than the achene; achene body often coarsely wrinkled, obovate to elliptic. 32 Spikelets sessile in many small compact capitula; achenes parallel-sided, punctate, with lighter colored margins, 2.1-2.8 m m long (Fig. 8B); leaves septate-nodulose between veins; spikelets 5-6 m m long .......................................................................................................... 73. R . g i g a n t e a Link. 32 Spikelets pedicellate, in fascicles; achenes with sides tapering to a point at base, rugulose to papillose or punctate with margins same color, 1.9-3.5 m m long; leaves smooth, not septate-nodulose; spikelets 4.5-8.5 m m long. 33 Achenes bi-concave or flattened laterally, with the upper part of each surface projecting in the shape of a rough inverted " U " ; style base with incurved sides, slightly umbonate (Fig. 8A) ........................................................................................... 75. R . s c u t e l l a t a Griseb. schoenoides 1992] THOMAS: RHYNCHOSPORA 17 33 A c h e n e s lenticular, irregularly coarsely wrinkled; style base with straight or convex sides (Fig. 8C) ............................................................................................................... 74. R. corymbosa (L.) Britton. 31 Style base slender or depressed, not sulcate, its base usually narrower than the achene; achene b o d y well-formed, not wrinkled, usually broadly obovate. 34 Spikelet scales pale, whitish or stramineous; widest leaves 2-10 m m wide and all cauline or 12-35 m m wide and either basal or cauline and basal. 35 Widest leaves (12)14-35 m m wide; leaves basal or basal and cauline and if some leaves cauline, then the spikelets 2-3 m m long. 36 Spikelets 6.3-7.7 m m long; culms 90-200 cm tall .............................................................................. ................................................................................................................ 63. R. comata (Link) R o e m e r & Schultes. 36 Spikelets 2-3.7 m m long, or if 5.6-9 m m long, then culms 20-75 cm tall and leaves with petiole-like bases. 37 Achenes 1.7-3.2 m m tong; spikelets 5.6-8(9) m m long; leaves gradually tapering to petiole-like bases ............................................................... 60. R. argentea Standley. 37 Achenes 1-1.8 m m long; spikelets 2-4 m m long; leaves not tapering to petiole-like bases. 38 Plants relatively slender; leaves mostly basal; culms below the lowest panicle 1.5-2 m m wide .........................................................59. R. pirrensis W. Thomas. 38 Plants robust: leaves cauline and basal; culms below the lowest panicle 4-7 m m wide. 39 Achenes 1.2-1.5 m m wide, often shiny and s m o o t h puncticulate. ........................................................................................................... 57. R. andresii Grmez-Laurito. 39 Achenes 0.9-1.1 m m wide, usually lustrous and puncticulate to longitudinally pitted .......................................................56. R. locuples C. B. Clarke. 35 Widest leaves 2-10 m m wide; leaves cauline; spikelets 3.7-7.5 m m long. 40 Achenes stramineous to boney white, lightly pitted, shorter than the style base (Fig. 5A); spikelets (5)5.5-7 m m long, often in fascicles o f 2-4 .................................. .................................................................................................................. 65. R. tuerckheimii C. B. Clarke ex Kiik. 40 Achenes ferrugineous to castaneous, _+ coarsely pitted, as long as to twice as long as the style base (Fig. 5B); spikelet 3.7-5.5(7.5) m m long, single or rarely in fascicles .................................................................................................................................. 64. R. polyphylla Vahl. 34 Spikelet scales pale brown or fuscous to castaneous or nearly black; leaves filiform to broad, 0.2-30 m m wide and mostly basal. 41 Hypogynous bristles 16-20, antrorsely barbed basaUy, retrorsely barbed distally (Fig. 2D) ................................................................................... 27. R. macra (C. B. Clarke ex Britton) Small. 41 Hypogynous bristles 1-6, antrorsely barbed throughout, or scabrous or plumose, never retrorsely barbed. 42 Hypogynous bristles plumose in part, at least with a distinct basal tuft o f hairs. 43 Style base bicornute; bristles plumose to ca height o f style base (Fig. 2A). .......................................................................................................................................... 24. R. diodon (Nees) Griseb. 43 Style base triangular to deltate; bristles plumose at least at very base (occasionally R. filifolia may key out here because o f the presence o f a few hairs at the base o f the bristles; it, however, has achenes 0.6-0.8 m m wide and a style base with scabrous margins). 44 Inflorescence o f 1-2 fine branchlets, each bearing 1-3 widely separated spikelets; achenes 1.4-1.7 m m wide ........................................................................................ ............................................................................................. 28. R. oligantha A. Gray var. oligantha. 44 Inflorescence o f 1-3 _+ aggregated, cylindrical to turbinate small fascicles; achenes 1-1.3 m m wide ...................................................29. R. plumosa Elliott. 42 Hypogynous bristles barbed or scabrous, never plumose. 45 Spikelets 2-3 m m long, stramineous to pale brown; widest leaf blades 1622 m m wide. 46 Achenes 1.3-1.5 m m wide, often shiny and s m o o t h puncticulate ............ ..................................................................................................................... 57. R. andresii Grmez-Laurito. 46 Achenes 0.9-1.1 m m wide, usually lustrous and puncticulate to longitudinally pitted ........................................................................... 56. R. locuples C. B. Clarke. 45 Spikelets (1.7)3-16 m m long, castaneous to pale brown; widest leaf blades (0.4)1.2-26 m m wide (if spikelets 3 m m long or less, then leaf blades less than 12 m m wide). 47 Inflorescence open and paniculate, corymbose or pyramidal, with many spikelets, or if not clearly so, then leaves more than 5 m m wide. 48 Panicles slender, narrowly ovoid, the branches ascending, 4-6 x 0.6-2.3 cm; leaves 7-11 m m wide ................................................................................. ................................................................................. 48. R. macrochaeta Steudel ex BSckeler. 18 BRITTONIA [VOL. 4 4 48 Panicles pyramidal to corymbose, the branches often divergent to reflexed, 2.5-20 x 3-18 cm; leaves 1.2-30 m m wide. 49 Panicles, at least terminal one, corymbose; lowest primary branches o f each panicle ascending, as long as or longer than the axis o f the panicle; bracts subtending the panicle branches often leaf-like. 50 Achenes 0.8-1.2 m m long; spikelets 3-8 m m long, often attenuate and slightly curved; leaves 4-12 m m wide ........ .................................................................................................................. 58. R. dives Standley. 50 Achenes 1.3-2.5 m m long, or if 1 . l - l . 3 m m long, then the leaves less than 3.6 m m wide; spikelets 4.5-16 m m long; leaves 2-23 m m wide. 51 Inflorescence ascending and leaves 2-4.5 m m wide; achenes l . l - l . 8 m m long. 52 Style base more than ~/2as long as the achene, 0.92.2 m m long; bristles 1.25-2x as long as the achene and style base (Fig. I D) ............................................. .............................46. R. marisculus Nees ex Lindley & Nees. 52 Style base seldom more than 1/2as long as the achene, 0.5--0.8 m m long; bristles longer than the achene to slightly exceeding achene and style base (Fig. IC) ................................................................ 47. R. rugosa (Vahl) Gale. 51 Inflorescence lax and leaves 3.5-10 m m wide or ascending and leaves 10-23 m m wide; achenes (1.5)1.82.5 m m long. 53 Largest leaf blades 3.5-8(10) m m wide; spikelets 6 11 m m long; peduncles o f lower corymbs slender, lax (Fig. 5E) .............................................55. R. aristata B6ckeler. 53 Largest leaf blades (7) 10-23 m m wide; spikelets 10 16 m m long; peduncles o f lower corymbs not slender, more or less erect ............................................................ ........................................................................... 54. R. hieronymi B/Sckeler. 49 Panicles pyramidal; lowest primary branches o f each panicle usually divergent or slightly reflexed, usually shorter than the axis o f the panicle; bracts subtending panicle branches often reduced. 54 Largest leaf blades 1.2-6(8) m m wide; spikelets 2.5-5(6) m m long; achenes 0.6-1( 1.1 ) m m wide, pale stramineous to brown or pinkish; culms 15-90 cm tall. 55 Style base (0.3)0.5-1(1.3) m m long; achenes 0.6-1.3(1.4) m m long, longitudinally ribbed or pitted, or nearly smooth ....................................................................... 51. R. vulcani B6ckeler. 55 Style base 0.3--0.5 m m long; achene 1.4-1.7 m m long, transversely rugulose (Fig. 4) ...... 43. R. rosae W. Thomas. 54 Largest leaf blades (4.5)7-26 m m wide; spikelets 4-12 m m long; achenes 0.9-2.2 m m wide, usually boney white to stramineous; culms (30)80--250 cm tall and, if less than 80 cm tall, then the achenes over 1.4 m m wide. 56 Spikelets pale brown, beige, or pale greenish; achenes brown to orange brown ............................................................................... ............................................63. R. comata (Link) R o e m e r & Schultes. 56 Spikelets castaneous; achenes boney white to stramineous. 57 Achenes 1.1-1.6(1.8) x 0.9-1.3(1.6) m m , boney white to pale stramineous, often dull, longitudinally pitted or ribbed (Fig. 5C); spikelets 3-6.5(8) m m long ....................................................50. R. ruiziana B6ckeler. 57 Achenes 1.6-2.3(2.6) x 1.4-2.1 m m , stramineous to golden brown (rarely boney white), shiny, lightly longitudinally pitted to smooth (Fig. 5D); spikelets (5)6-12 m m long ......................................................................... ....................................49. R. schiedeana (Schlechlend.) Kunth. 47 Inflorescence congested, fasciculate or cymose, often with few spikelets. 58 Achene surface smooth to cancellate. 1992] THOMAS: RHYNCHOSPORA 19 59 Style base triangular to narrowly so, 0.9-2.1 m m long. 60 M a r g i n s o f style base serrulate; spikelets in congested, usually turbinate fascicles ................................................32. R. harperi Small. 60 M a r g i n s ofstyle base s m o o t h a n d glabrous; spikelets in loose to congested, usually cylindrical to occasionally turbinate fascicles. 61 Inflorescence very open a n d lax, with remote spikelets; a c h e n e s 1-1.2 x 0.75-0.8 m m ....................................................... .........................................................................52. R. bo6viensis C. B. Clarke. 61 Inflorescence o f 1--4 congested panicles or corymbs; achenes 1.2-1.5 x 0.7-0.9 m m or 1.5-1.7 x 1.3-1.6 mm. 62 A c b e n e s 0.7-0.9 m m wide; c u l m s 10-35 c m tall (Fig. 7) ................................................................................................................ 967. R. talamancensis G 6 m e z - L a u r i t o & W. T h o m a s . 62 A c b e n e s 1.3-1.6 m m wide (Fig. 2(2); c u l m s 4 0 - 7 0 c m tall ........................................................34. R. gracilenta A. Gray. 59 Style base deltate to broadly so or rarely triangular, 0.3-0.7 m m long. 63 H y p o g y n o u s bristles vestigial, usually 1-2 less t h a n 0.3 m m long. 64 Leaves flliform, 0.4-0.5 m m wide; style base 0.3-0.5 m m long; achene castaneous with a pale center ........... .........................................................................30. R. brachychaeta C. Wright. 64 Leaves plicate, 1-1.6 m m wide; style base 0.9-1 m m long; achene uniformly s t r a m i n e o u s to pale grayishb r o w n .................................. 53. R. dissitiflora Steudel ex B~Sckeler. 63 H y p o g y n o u s bristles evident, 5-6. 65 Style base raised on narrowed neck o f a c h e n e , n o t confluent with achene s u m m i t ..................................................................... .........................................................42. R. globularis var. recognita Gale. 65 Style base confluent with achene s u m m i t . 66 Leaves canaliculate a n d stiff, ca 2-15 c m long .......... .................................................36. R. oreoboloidea G 6 m e z - L a u r i t o . 66 Leaves filiform a n d flexuous or involute or plicate, 10-30 c m long. 67 A c h e n e surface faintly rugulose, u n i f o r m l y stram i n e o u s to golden b r o w n (Fig. 3) ........................... . 3 5 b . R. fascicularis subsp, sierrensis W. T h o m a s . 67 A c h e n e surface glassy s m o o t h to faintly cancellate, golden b r o w n to dark b r o w n at least along margins, the center pale. 68 A c h e n e s 1.1-1.4 m m wide; largest blades 1-2.5(4) m m wide (Fig. 2B) ................................. .............................................35a. R. fascicularis (Michaux) Vahl subsp, fascicularis. 68 A c h e n e s 0.6-0.8 m m wide; largest blades 0.4-1 m m wide ................ 33. R. filifolia A. Gray. 58 A c h e n e surface rugose to rugulose. 69 Leaves filiform, 0.2-0.5 m m wide. 70 A c h e n e s 1-1.2 m m long; spikelets 1.7-2 m m long ..................... .............................. 38. R. lindeniana var. bahamensis (Britton) Gale. 70 A c h e n e s 1.3-1.7 m m long; spikelets 2.8-3.7 m m long. 71 Inflorescence open; achenes rugulose; leaves filiform, 0.2-0.5 m m wide .............. 37. R. rariflora (Michaux) Elliott. 71 Inflorescence congested; achene faintly rugulose (Fig. 3); leaves involute to plicate, 0.4-0.9(1.1) m m wide. ........................ 35b. R. fascicularis subsp, sierrensis W. T h o m a s . 69 Leaves involute to plicate, 1-4.5 m m wide. 72 Spikelets very dark castaneous to black .................................................. .......................................................................................39. R. kunthii Nees ex K u n t h . 72 Spikelets brown, ferrugineous, or castaneous. 20 BRITTONIA [VOL. 4 4 73 Style base clearly not confluent with the achene summit, either raised on the narrowed neck o f the achene or at least slightly expanded along basal edge. 74 Achenes 0.9-1.2 m m wide, pale stramineous to stramineous; style base 0.4-0.7 m m wide, discontinuous on s u m m i t o f achene (Fig. 4) ................. ............................................................................... 43. R. rosae W. Thomas. 74 Achenes 1.2-1.6 m m wide, ferrugineous to golden brown or castaneous; style base 0.7-0.9 m m wide, on narrowed raised neck o f achene ................................... ..............................................42. R. globularis var. recognita Gale. 73 Style base confluent with the achene summit. 75 Achene surface clearly rugose; spikelets 2-2.5 m m long. 76 Culms 65-130 c m tall; achenes flattened-lenticular; spikelets 0.8-1.1 m m wide ...................... ............................44. R. microcarpa Baldwin ex A. Gray. 76 Culms 20--60 cm tall; achenes lenticular; spikelets 1-1.4 m m wide .......................40. R. brittonii Gale. 75 Achene surface rugulose or, ifrugose, then the spikelets 3.3-3.5 m m long, otherwise spikelets 3-6.5 m m long. 77 Achenes strongly rugose, with 6-8 deeply etched transverse ridges; style base transversely deltare, 0.2-0.3 x 0.6-0.7 m m .......................................... ....................................................................... 45. R. jubata Liebmann. 77 Achenes rugulose to papillose, with 9-22 often obscured ridges; style base usually deltate to triangular, 0.5-2.2 x 0.6-1 m m . 78 Achenes with ridged s u m m i t at edge o f style base, dark castaneous, faintly rugulose, strongly convex; bristles t/2 as long as the achene to occasionally equalling or slightly surpassing it ............................................ 41. R. brownii subsp, americana Guaglianone. 78 Achenes with s u m m i t confluent with style base, golden brown to stramineous, rugulose to faintly so, slightly convex; bristies usually exceeding achene to 2 x as long as the achene and style base. 79 Style base more than 1/2 as long as the achene, 0.9-2.2 m m long; bristles 1.25-2 x as long as the achene and style base (Fig. 1D) ........... 46. R. marisculus Nees ex Lindley & Nees. 79 Style base seldom more than 1/2 as long as the achene, 0.5-0.8 m m long; bristles exceeding achene to slightly exceeding achene and style base (Fig. 1C) ...............................47. R. rugosa (Vahl) Gale. 26 Hypogynous bristles absent. 80 Achenes smooth to finely cancellate, papillose or wrinkled; style base usually less than 0.5 m m high. 81 Spikelets 9-12.5 m m long ................................................................................................................................ 17. R. filiformis Vahl. 81 Spikelets 2.2--4 m m long. 82 Largest blades 4-7 m m wide; style base reduced to an apiculus 0-0.1 m m high ............... .................................................................................................................................................................................. 21. R. brevirostris Griseb. 82 Largest blades 0.4-2 m m wide; style base 0.05-0.6 m m high. 83 Inflorescence o f 1-3 congested, o b o v o i d to turbinate fascicles. 84 Spikelets 2.2-2.5 m m long; fascicles solitary; style base 0.2-0.4 m m high ........... 31. R. chapmanii M. A. Curtis. ............................................................................................................................................ 84 Spikelets 2.5-3.2 m m long; fascicles 1-3; style base 0.3-0.5 m m high ...................... ........................................................................................................................................... 30. R. brachychaeta C. Wright. 1992] THOMAS: RHYNCHOSPORA 21 83 Inflorescence of 1-5 panicles or corymbs. 85 Culms and abaxial leaf surfaces hirsute or the blades at least ciliate above the leaf sheath; style base 0.1-0.2 m m high (Fig. 1A)........... 26. R. hirsuta (Vahl) Vahl. 85 Culms and leaves glabrous; style base 0.05-0.1 m m high ......................................................... ........................................................................................................... 19. R. divergens Chapman ex M. A. Curtis. 80 Achenes rugose to rugulose; style base various. 86 Style base tridentate ............................................................. 18. R. tenerrima Nees ex Sprengel subsp, tenerrima. 86 Style base deltate to triangular, or strap-like. 87 Achenes 1.1-1.9 m m wide. 88 Style base depressed, semilunate, no more than 0.2-0.5 m m high in the center. 89 Spikelets broadly ovoid to subglobose, the tips blunt; inflorescence branches spreading or divergent; achene base usually truncate; style base decurrent on the achene .................................................................................................................. 10. R. trichodes C. B. Clarke. 89 Spikelets usually ovoid, the tips acuminate or acute; inflorescence branches spreading obliquely, the spikelets often erect; achene base not truncate; style base decurrent or not. 90 Fertile spikelet scales 3.5-4.5 m m long; annuals; style base ends strongly upturned ....................................................................................................... 14. R. eximia (Nees) B6ckeler. 90 Fertile spikelet scales 6-9.5 m m long; perennials; style base ends slightly decurrent .......................................................................... 15. R. waspamensis Kral & W. Thomas. 88 Style base triangular, 0.8-1.6 m m high. 91 Achenes 1.7-2.4 m m long; style base 0.9-1.6 m m long ............................................................... .......................................................................................................................................... 13. R. robusta (Kunth) BiSckeler. 91 Achenes 1.1-1.4 m m long; style base 0.8-1.1 m m long ............................................................... ........................................................................................................................................ 12. R. velutina (Kunth) B6ckeler. 87 Achenes 0.5-1 m m wide. 92 Spikelets + globose, ca 1.5-2.5 m m long (Fig. 1B)............ 23. R. contracta (Nees) Raynal. 92 Spikelets lanceoloid to ovoid, 1.9-6 m m long. 93 Achenes with a distinct cuneate basal stipe ca 0.35 m m long .............................................. ..................................................................................................................................... 22. R. carrillensis G6mez-Laurito. 93 Achenes without a basal stipe. 94 Largest blades 3-10 m m wide; achenes subcircular in outline .................................. ........................................................................................................................................ 11. R. nitens (Vahl) A. Gray. 94 Largest blades 0.3-1.4 m m wide; achenes broadly obovate to ovate. 95 Leaves cauline; style base 0.6-1 m m long (Fig. 6) ........................................................ ................................................................................................................................ 66. R. davidsei W. Thomas. 95 Leaves basal or from lower half of stem; style base 0.05-0.4 m m long. 96 Style base 0.05-0.1 m m high; achenes 0.5-0.6 m m wide ......................... ............................................................................................................................... 20. R. pusilla Chapman. 96 Style base 0.2-0.4 m m high; achenes 0.7-1.1 m m wide ............................ ................................................................................................................................................ 16. R. tenuis Link. 1. RHYNCHOSPORA COLORATA (L.) P f e i f f e r Schoenus coloratus L., Sp. P1. ed. 1, h 43. 1753. R. colorata (L.) Pfeiffer, Repert. Spec. Nov. Regni Veg. 38: 89. 1935. Dichromena colorata (L.) Hitchc., Annual Rep. Missouri Bot. Gard. 4: 141. 1893. TYPE: Sloane, Voy. Jamaica 1:119, pl. 78, fig. 1. 1707. Schoenus stellatus Lam., Encycl. 1: 741. 1784. R. stellata (Lam.) Griseb., Abh. Krnigl. Ges. Wiss. Grttingen 8:271.1857. TYPE: UNITED STATES. Florida (HOLOTYPE:P--n.v.). Scirpus cephalotes Walter, F1. carol. 71. 1788. Dichromena cephalotes (Walter) Britton, Bull. Torrey Bot. Club 15: 100. 1888. TYPE: U N I T E D STATES. (BM--n.v.). Dichromena leucocephala Michaux, Flora bor.-amer. 1: 37. 1803. Dichroma leucocephala (Michaux) Persoon, Syn. pl. l: 57.1805. R. leucocephala (Michaux) Brckeler, Vidensk. Meddel. Dansk Naturhist. Foren. Kjoebenhavn ser. 3, 1: 144. 1869. TYPE: UNITED STATES. Carolina and Georgia, Bosc s . n . (HOLOTYPE:P; ISOTYPE" G). R. drummondiana Steudel, Syn. pl. giumac. 2: 135. 1855. TYPE: U N I T E D STATES. Louisiana. New Orleans, D r u m m o n d 394 (HOLOTYPE: BM). R. stellata forma intercedens Kiik., Bot. Jahrb. Syst. 75: 300. 1951. TYPE: WEST INDIES. Haiti. Portau-Prince, Mariani, 1 Mar 1926, E k m a n N H 5 6 5 7 (HOLOTYPE" B; ISOTYPES"GH, US). Pastures, roadsides or other slightly disturbed areas usually on sandy circumneutral soil, usually near sea level. Tabasco; Chiapas; Yucatfin; Belize; Guatemala; Costa Rica. 0-500 m. (Also the Atlantic coastal plain of the United States from North Carolina to Florida and Texas; West Indies; scattered along the Gulf Coast from Mexico to Venezuela.) 22 BRITTONIA 2. RHYNCHOSPORAFLORIDENSIS(Britton ex Small) Pfeiffer Dichromenafloridensis Britton ex Small, F1. s.e.U.S. !90. 1903. Dichromenafloridensis C. B. Clarke, Kew Bull. Misc. Inform., Add. Ser. 8: 32. 1908. Rhynchosporafloridensis (Britton ex Small) Pfeiffer, Repert Spec. Nov. Regni Veg. 49: 82. 1940. TYPE: UNITED STATES. Florida, Small & Nash 181 (HOLOTYPE: NY). Dichromena inauguensis Britton in Britton & Millsp., Bahama Flora 53. 1920. TYPE: WEST INDIES. Bahamas. Little Inagua, 21 Oct 1954, Nash & Taylor 1254 (HOLOTYPE: NY; ISOTYPE: US). Open pine woodlands, usually over limestone. Campeche; Quintana Roo; Belize. 0 200 m. (Also in extreme S Florida and the Bahamas.) 3. RHYNCHOSPORANERVOSA (Vahl) B~ckeler Dichromena nervosa Vahl, Enum. pl. 2: 241. 1806. R. nervosa (Vahl) B~ckeler, Vidensk. Meddel. Dansk Naturhist. Foren. Kjoebenhavn, ser. 3, 1: 143. 1869. TYPE: "'America meridionalis," von Rohr s . n . (HOLOTYPE"N~r microfiche ex C). Tabasco to Panama. (West Indies; Mexico to S Brazil and N Argentina.) 3a. RHYNCHOSPORA NERVOSA NERVOSA subsp. Dichromena persooniana Nees in C. Martius, Fl. bras. 2(1): 112. 1842. R. persooniana (Nees) Griseb., F1. Brit. W. I. 577. 1864. TYPE: WEST INDIES. Puerto Rico. Essequeho, M e y e r s . n . (HOLOTYPE" GOET--n.v.) Savannas or other natural, open grasslands. Tabasco; Chiapas; Quintana Roo; Guatemala; Honduras; E1 Salvador; Nicaragua; Costa Rica; Panama. 0-2100 m. (Throughout range of species but more common inland and in drier areas.) 3b. RHYNCHOSPORA NERVOSA subsp. CILIATA Koyama Dichromena ciliata Vahl, Enum. pl. 2: 240. 1806. R. ciliata (Vahl)Kiik., Bot. Jahrb. Syst. 56(Beibl. 125): 16. 1921. R. nervosa subsp, ciliata Koyama, Madrofio 20: 254. 1970. SYNTYPES: "America meridionali," Richard s.n.; Puerto Rico, West s.n. ( N Y microfiche of both ex C). Dichromena ciliata vat. cinnamomea Kiik., Repert. Spec. Nov. Regni Veg. 23: 200. 1926. R. nervosa var. cinnamomea (K~k.) Kfik., Bot. Jahrb. 75: 297. 1951. TYPE: VENEZUELA. Nueva Esparta. Isla [VOL. 44 de Margarita, San Juan Mts., 500 m, 2 Jul 1903, J. R. Johnston 201 (LECTOTYPEdesignated here, see explanation below: NY). Weedy areas, pastures, roadsides, and lawns. Tabasco; Yucatan; Belize; Guatemala; Honduras; E1 Salvador; Nicaragua; Costa Rica; Panama. 0-1400 m. (Throughout range of species but most common in costal regions and moist disturbed areas.) In an earlier treatment of this species (Thomas, 1984), I incorrectly chose as lectotype a specimen (Miller & Johnston 189) not cited in the original description; that error is rectified here. 4. RHYNCHOSPORARADICANS (Schlectend. & Cham.) Pfeiffer Dichromena radicans Schlechtend. & Cham., Linnaea 6: 28. 1831. R. radicans (Schlechtend. & Cham.) Pfeiffer, Repert. Spec. Nov. Regni Veg. 38: 93. 1935. TYPE: MEXICO. Serro Colorado, Schiede & Deppe s . n . (HOLOTYPE"BM). Moist, disturbed, usually partly shaded areas, forest edges, trail sides, and clearings. 0-1600 m or more. Tabasco; Chiapas; Belize to Panama. (West Indies; S Mexico to N South America and in the Andes to Bolivia.) 4a. RHYNCHOSPORA RADICANS subsp. RADICANS Chiapas; Belize; Guatemala; Honduras; Costa Rica; Panama. 0-500 m. (S Mexico to Honduras, scattered elsewhere in Central America and N South America.) 4 b . RHYNCHOSPORA RADICANS subsp. MICROCEPHALA (Bertero ex Sprengel) W. Thomas Dichromena microcephala Bertero ex Sprengel, Syst. Veg. 1: 202. 1824, non R. microcephala Britton ex Small, 1903. R. radicans subsp, microcephala (Bertero ex Sprengel) W. Thomas, Mem. New York Bot. Gard. 37: 60. 1984. TYPE: WEST INDIES. Jamaica, Bertero s.n. (ISOTYPE: MO). Tabasco; Chiapas; Quintana Roo; Belize; Guatemala; Honduras; E1 Salvador; Nicaragua; Costa Rica; Panama. 0-1600 m. (West Indies; S Mexico to the Guianas and in the Andes to Bolivia.) 1992] 5. RHYNCHOSPORA Davidse THOMAS: RHYNCHOSPORA WATSONII (Britton) Dichromena watsonii Britton, Bull. Torrey Bot. Club 15:101.1888. R. watsonii (Britton) Davidse, Ann. Missouri Bot. Gard. 61: 529. 1974. TYPE:GUATEMALA. S. Watson 153 (HOLOTYPE:NY). Moist forests, forest edges, clearings, and road cuts. Belize; Guatemala; Nicaragua; Costa Rica; P a n a m a . 0 - 1 6 0 0 m. (S Mexico to N South America.) 6. RI-IYNCI-IOSPORAPOLYSTACHYS (Turrill) Pfeiffer Dichromena polystachys Turrill, M6m. Soc. Sci. Nat. Neuchfltel 5: 348. 1914. R. polystachys (Tun-ill) Pfeiffer, Repert. Spec. Nov. Regni Veg. 38: 93. 1935. R. nervosa var. polystachys (Tun-ill) Kiik., Bot. Jahrb. Syst. 75: 298. 1951. TYPE:COLOMBIA. "pr6s de Angiopolis," E. M a y o r 618 (HOLOTYPE:K). Forest edges, trails, stream banks, and clearings. Honduras. 0 - 2 1 0 0 m. (Also in A n d e a n South A m e r i c a from N C o l o m b i a to S Peru.) This collection is the first recorded from Central America a n d represents a significant range extension from the next nearest k n o w n locality in the C h o c 6 o f Colombia. 7. RHYNCHOSPORAPUBERA(Vahl) Bfckeler subsp, p a r v u l a W. T h o m a s R. pubera (Vahl) B6ckeler subsp, parvula W. Thom- as, Mere. New York Bot. Gard. 37: 50. 1984. TYPE: COLOMBIA. Meta, F. Pennell 1501 (HOLOTYPE: us). Savannas or pastures in moist, acidic, sandy or peaty soils. Costa Rica, P a n a m a . 0 - 1 0 0 m. (Very rare in Central A m e r i c a and widely scattered in South A m e r i c a north o f the A m a z o n river.) The c o m m o n e r subspecies p u b e r a is larger with longer spikelets and larger achenes. It is found t h r o u g h o u t northern South America east o f the Andes. 8. RHYNCHOSPORAALBESCENS(Miq.) Kiik. Isolepis albescens Miq., Linnaea 19: 226. 1847. R. albescens (Miq.) Kiik., Bot. Jahrb. Syst. 75: 304. 1951. TYPE:SURINAM. Para Distr., Koropina Creek, Focke 1074 (HOLOTYPE:U). R. monostachya B6ckeler, Allg. Bot. Z. Syst. 2: 78. 23 1896. TYPE:BRAZIL. Pani. Santar6m near Paria, R. Spruce s.n. (ISOTYPES:B, NY). Open or partially shaded bare sand or m u d near edges o f black water streams and other acidic wet areas that m a y be inundated at some times o f year. Costa Rica. 0 - 4 0 0 m. (Otherwise k n o w n f r o m A m a z o n i a n Col o m b i a and Venezuela to Pargt and Amaph, Brazil.) In Central A m e r i c a k n o w n only from two collections from Buenos Aires, Puntarenas, Costa Rica. These two collections are disjunct approximately 1200 km f r o m the next nearest k n o w n locality in Meta, Colombia. 9. RHYNCHOSPORABERTEROI (Sprengel) C. B. Clarke Hypo~lytrum berterii Sprengel, Neue Entd. 1: 241. 1820. R. berteroi (Sprengel) C. B. Clarke in Urban, Symb. antill. 2:119. 1900. TYPE:WEST INDIES. Guadeloupe, Bertero s.n. (ISOTYPE:MO). Schoenus pusittus Sw., Prodr. 20. 1788. R. pusilla (Sw.) Griseb., Abh. K6nigl. Ges. Wiss. G6ttingen 7: 271. 1857, non Chapman, 1847. TWE: WEST INDIES. Jamaica, Swartz s.n. (HOLOTVPE:BM; ISOTYPE:M). Moist clearings in forests, usually on circumneutral soil. Belize. ( K n o w n also f r o m the Greater Antilles and Guadeloupe.) In Mesoamerica, k n o w n from only one collection, L u n d e l l 6 8 7 from H o n e y C a m p , Belize. 10. RHYNCrtosr~RA TRICHODESC. B. Clarke R. trichodes C. B. Clarke in Urban, Symb. antill. 2: 116. 1900. Schoenus hispidulus Vahl, Enum. pl. 2: 219. 1806. R. hispidula (Vahl) B6ckeler, Lin- naea 37: 604. 1873, non Griseb., 1866. TYPE: UNITED STATES. "Hab. in Carolina? ex herb. Lamarck," (HOLOTYPE;C?--n.v.). Marshes, savannas, and other open inundated areas. Belize; Costa Rica; P a n a m a . 0 - 3 0 0 m. (West Indies; N South America.) 1 1. RHYNCHOSPORANITENS (Vahl) A. G r a y Scirpus nitens Vahl, Enum. pl. 2:272.1806. R. nitens (Vahl) A. Gray, Manual, ed. 5: 568. 1867. Psilocarya nitens(Vahl) Wood, Amer. bot. ft. 364. 1870. TYPE:UNITED STATES. "Habitat in Carolina," Bosc s.n. (HOLOTYPE:C?--n.v.). LOW, wet, sandy or peaty open areas. Belize; Nicaragua. 0 - 1 0 0 m. (Coastal plain of 24 BRITTONIA the U n i t e d States from Massachusetts to Florida and Texas; West Indies.) 12. RHYNCHOSPORA VELUTINA ( K u n t h ) B6ckeler [VOL. 44 Sandy savannas or peaty clearings. Tabasco; Chiapas; Belize; Guatemala; H o n duras; Nicaragua; Costa Rica; Panama. 0 300 m. (S Florida; West Indies; Mexico to S Brazil; Africa.) Dichromena velutina Kunth, Enum. pl. 2: 282. 1837. Psilocarya velutina (Kunth) Nees in C. Martius, F1. bras. 2(1): 115.1842. R. velutina (Kunth) Brck- 15. RHYNCHOSPORA WASPAMENSIS Kral & W. T h o m a s eler, Vidensk. Meddel. Dansk Naturhist. Foren. Kjoebenhavn 149. 1869. TYPE: BRAZIL. "Brasilia meridionalis," Sellow s.n. (HOLO~E: P?.-- R. waspamensis Kral & W. Thomas, Brittonia 40: n.v.). Psilocarya rufa Nees in C. Martius, Fl. bras. 2(1): 117.1842. R. rufa (Nees) Brckeler, Vidensk. Med- del. Dansk Naturhist. Foren. Kjoebenhavn 149. 1869. SY~ZrYVES:BRAZIL. Goi~s. "Ad fluvium S. Marci," Pohl 781 (n.v.); GUYANA. Schomburgk 667 (ISOTYPE:K). Marshes, wet savannas, and roadside ditches. Tabasco; Honduras; Nicaragua. 0 700 m. (Mexico; West Indies; tropical South America.) R h y n c h o s p o r a v e l u t i n a and R . r o b u s t a are both variable and are very similar to one another; I have previously treated them, in the Flora o f Nicaragua, as s y n o n y m o u s (Kral & T h o m a s , in press). While they have different tendencies in a n u m b e r o f characters (achene width, style base size and shape, spikelet shape and size, a n d even elevation), they differ consistently only in achene length. In M e s o a m e r i c a at least, the style base characters used by K o y a m a (1972) do not separate the two species successfully. 13. RHYNCHOSPORA ROBUSTA ( K u n t h ) BSckeler Dichromena robusta Kunth, Enum. pl. 2: 283. 1837. Psilocarya robusta (Kunth) Nees in C. Martius, Fl. bras. 2(1): 116.1842. R. robusta (Kunth) Brckeler, Linnaea 37: 616. 1873. TYPE: BRAZIL. "Brazil meridionalis," Sellow s.n. 0SOTYPE:K). Marshes, bogs, and wet savannas or m e a d o w s . C h i a p a s ; Belize; G u a t e m a l a ; H o n d u r a s ; Panama. 4 0 0 - 2 2 0 0 m. (Also the West Indies and tropical South America.) See c o m m e n t s under R . v e l u t i n a . 14. RHYNCHOSPORAEXIMIA (Nees) Brckeler Spermodon eximius Nees in Seeman, Voy. Herald Bot. 222. 1857. R. eximia (Nees) B6ckeler, Linnaea 37: 601. 1873. TYPE:PANAMA. Seemann 140 (HOLOTYPE:BM). 34. 1988. TYPE:NICARAGUA. Zelaya. Waspam, G. Davidse & R. Pohl 2318 (HOLOTYPE:HNMN; ISOTYPE: MO). Wet p o n d margins. Nicaragua. 0 - 1 0 0 m. ( K n o w n only from Zelaya, Nicaragua.) 16. RHYNCHOSPORATENUIS Link R. tenuis Link, Jahrb. Gew~ichsk. 1(3): 76. 1820. TYPE: BRAZIL. Humboldt, in herb Willd. 1175, )Col. 1 (HOLOTYPE:B, destroyed, microfiche NY ex B). Open m e a d o w s and along streams, especially in open pine-oak woodlands. Chiapas; Belize; Guatemala; Honduras; Nicaragua. 8 0 0 - 1 8 0 0 m. (Also the West Indies and South America to S Brazil and N Argentina.) 17. RHYNCHOSPORAFILIFORMISVahl R. filiformis Vahl, Enum. pl. 2: 232. 1806. Dichrornena filiformis (Vahl)Kunth, Enum. pl. 2: 281. 1837. Spermodon filiformis (Vahl) Nees, in C. Martius, F1. bras. 2(1): 118. 1842. TYPE:WEST INDIES. Puerto Rico, A. Ledru s.n. (C--n.y.). R. podosperma C. Wright in Sauvalle, Anales Acad. Ci. Mrd. Habana 8: 87. 1871. TWE: WEST INDIES. Cuba. Between Pinar del Rio and La Coloma, C. Wright 3791 (ISOTYPE:GH--n.v., NY). R. longispicata Brckeler, Linnaea 37: 600. 1873. SYrcrypEs:BRAZIL. Campos Vitoria, Sellow s.n. (ISOSVa~q~CPE:K); FRENCH GUIANA. Sagot 389 (?P--n.v.), Leprieur 45 (?P--n.v.). Moist savannas and seasonal pools and marshes. Tabasco; Belize; Guatemala; H o n duras; Nicaragua; Costa Rica. 0 - 5 0 0 m. (West Indies; tropical South America.) 18. RHYNCHOSPORA TENERRIMA Nees ex Sprengel subsp. TENERRIMA R. tenerrima Nees ex Sprengel, Syst. veg. 4(2): Curae posteriores 26. 1827. TYPE:WEST INDIES [but cited as "Nov. Holl.']. Kohaut s.n., distributed as F. Sieber agrost, no. 116 (AWH--n.v.). Schoenus setaceus Bergius, Act. Helv. Phys.-Math. 1992] THOMAS: RHYNCHOSPORA 7: 130. 1772. R. setacea (Bergius) Brckeler, Vidensk. Meddel. Dansk Naturhist. Foren. Kjoebenhavn 159. 1869, non Vahl, 1806. TYPE: SURINAM. (n.y.). Marshes, wet savannas, seepage slopes, a n d edges o f s e a s o n a l p o o l s . T a b a s c o ; C h i a p a s ; Belize; G u a t e m a l a ; H o n d u r a s ; N i c a ragua; Costa Rica; Panama. 0-1000 m. (West Indies; South America south to Bol i v i a a n d Brazil.) Subspeciesfallax (Uitten) Koyama, which is f o u n d in V e n e z u e l a a n d S u r i n a m as w e l l as i n A f r i c a , differs i n h a v i n g t h e t w o l a t e r a l h o r n s o f t h e style b a s e m u c h d i m i n i s h e d , giving the achene summit a more truncate appearance. 19. RHYNCHOSPORA DIVERGENS C h a p m a n ex M. A. C u r t i s R. divergens Chapman ex M. A. Curtis, Amer. J. Sci. Arts 57: 409. 1849. TYPE: UNITED STATES. South Carolina. Santee, Ravenel s.n. (nOLOTYPE: NY). S a v a n n a s o r o c c a s i o n a l l y f o r e s t edges. C h i a p a s ; Belize. 0 - 2 0 0 m . ( A l s o t h e U n i t e d S t a t e s f r o m S o u t h C a r o l i n a to F l o r i d a a n d Alabama; The Bahamas.) 20. RHYNCHOSPORA PUSILLA C h a p m a n R. pusilla Chapman, F1. South. U.S. 528. 1865. R. pusilla Chapman ex Curtis, Amer. J. Sci. Arts 57: 409. 1849 [incidental mention], non (Sw.) Griseb. 1857. TYPE: UNITED STATES. Florida. Apalachicola, Chapman 4473 (HOLOTYPE:NY). R. intermixta C. Wright in Sauvalle, Anales Acad. Ci. Mrd. Habana 8: 88. 1871, reprinted in Sauvalle, F1. cubana 184.1873. TYPE:WEST INDIES. Cuba. "En pinares hfimedos cerca de Pinar del Rio, rodeadas de Rhync. lamprospermas," Wright 3794 (probable ISOTYPES:NY--2 sheets). Savannas and basco; Chiapas; ragua. 0-100 m. f r o m M e x i c o to o t h e r m o i s t o p e n areas. T a Belize; G u a t e m a l a ; N i c a (West Indies; the Gulf coast Nicaragua.) 21. RHYNCHOSPORA BREVIROSTRIS G r i s e b . R. brevirostris Griseb., Cat. pl. Cub. 246.1866. TYPE: WEST INDIES. "Cuba occ.," C. Wright 3410 [misprinted in protologue as 3413 which is R. cernua Griseb.] (GH--n.v.). R. clarkei Rose, Contr. United States Nat. Herb. 10: 464. 1908 [nom. nov.]. R. pringlei C. B. Clarke, Kew Bull. Misc. Inform., Add. Ser. 8: 89. 1908, ~aon Greenman, 1904. TYPE: MEXICO. Jalisco, 25 Pringle 2319 (LECrOTVPEdesignated by Koyama, 1972; K--n.v.). Savannas, marshes, and other open moist areas. C h i a p a s ; Belize; H o n d u r a s ; N i c a r a gua; C o s t a R i c a ; P a n a m a . 0 - 2 0 0 0 m. ( A l s o t h e W e s t I n d i e s , T r i n i d a d , a n d Brazil.) 22. RHYNCHOSPORA CARRILLENSIS G 6 m e z Laurito R. carrillensis G6mez-Laurito, Brenesia 22: 353. 1984. TYPE: COSTA RICA. San Josr. Parque Nacional Braulio Carrillo, 1100 m., C. Todzia et al. 1994 (ISOTVPE:NY). L o w e r m o n t a n e r a i n forests. C o s t a R i c a ; Panama. 1100-1500 m. Known from two collections from Costa Rica, the type and H e p p e r 133, a n d o n e f r o m P a n a m a , M o r i & Kallunki 6201. This species has the superficial appeara n c e o f R . h i r s u t a b u t t h e larger, s m o o t h e r achenes are quite distinct. 23. RHYNCHOSPORA CONTRACTA (Nees) J. R a y n a l . (Fig. 1B) Haloschoenus contractus Nees in C. Martius, F1. bras. 2(1): 123. 1842. R. contracta (Nees) J. Raynal, Adansonia 17(3): 277.1978. TYPE:BRAZIL. Piaui, Martius s.n. (M--n.v.). R. micrantha Vahl, Enum pl. 2: 231. 1806 [nora. illeg.]. Scleria minutiflora L. C. Rich. ex Spreng., Syst. veg. 3: 831. 1826 [nom. inval.]. R. minutiflora (L. C. Rich. ex Spreng.) Adams, Phytologia 21: 70. 1971. TYPE: "Carolina, Ins. Antill.," (HOLOTYPE:B?-- n.v.). M o i s t o p e n d i s t u r b e d a r e a s s u c h as r o a d sides, d i t c h e s , p a s t u r e s , a n d f o r e s t m a r g i n s . C h i a p a s ; Yucatfin; Belize; G u a t e m a l a ; H o n d u r a s ; E1 S a l v a d o r ; N i c a r a g u a ; C o s t a R i c a ; Panama. 0-1000 m. (Tropical America; moist tropical Africa.) 24. RHYNCHOSPORA DIODON ( N e e s ) G r i s e b . (Fig. 2A) Ptilochaeta diodon Nees in C. Martius, FI. bras. 2(1): 148, t. 8, fig. 2. 1842. R. diodon (Nees) Griseb., Cat. pl. Cub. 245. 1866. TYPE: BRAZIL. "In praeruptis viarum in campis prov. Piauhiensis, et ad flumen Itapicur6 prov. Maragnaniensis, Majo: M." Martius s.n. (HOLOTYPE:B, destroyed?). [Although two localities are listed, it is not clear whether Martius collected two specimens; in addition K has Gardener 2376 from Piaui noted as a type for this name.] 26 BRITTONIA J [VOL. 44 7 84 / / t / P i ! ii E FIO, 1. Rhynchospora achenes; scale bar = 1 mm. A. R. hirsuta (L. D. G6mez 19002). B. R. contracta (Kral 27736). C. R. rugosa (Davidse & Pohl 1224). D. R. marisculus (Breedlove 21522). E. R. barbata (Duke 8700). R. lunata Griseb., Cat. pl. Cub. 244. 1866. TYPE: WEST INDIES. "Cuba or.," 0SOTVPE:NY). Open savannas. Honduras. (West Indies; C o l o m b i a to Brazil.) 25. RHYNCHOSPORA EXALTATA K u n t h R. exaltata Kunth, Enum. pl, 2: 291. 1837. Dichromena exaltata (Kunth) Macbride, Publ. Field Co- lumbian Mus., Bot. Ser. 8:113. 1930. TYPE:BRAZIL. "Brasilia meridionalis," Sellow s, n. 0SOTYPE: K). R. triceps Steudel, Nomencl., ed. 2, 2: 456. 1841. TYPE: n . y . R. arundinacea B/Sckeler, Vidensk. Meddel. Dansk Naturhist. Foren. Kjoebenhavn 1879-80; 27. 1879-80. TYPE: BRAZIL. Rio de Janeiro. Rio de Janeiro, Glaziou s.n. (n.y.). Echinoschoenus beyrichii Nees, in C. Martius, FI. bras. 2(1): 132. 1842. R. beyrichii (Nees) Steudel, Syn. pl. glumac. 2: 144. 1855. TYPE: BRAZIL. Rio de Janeiro. "In sylvis urnbrosis humidis prope Norum Friburgum," Beyrich s.n. (HOLOTVPE:B, destroyed?). R. exaltata var. ovalis Kiik., Bot. Jahrb. Syst. 74(3): 441. 1949. TYPE: BOLIVIA. Regi6n de Mapiri, Copacabana, Buchtien 339 0SOa~CPE:NY). Mitrospora cephalophora Nees, in C. Martius, F1. bras. 2(1): 133.1842. R. cephalophora (Nees) Steudel, Syn. pl. glumac. 2: 146. 1855. SvawrvPES: BRAZIL. "Brasilia meridionali," Sellow s.n. (B, destroyed?); Minas Gerais: Itamb6, Poh13569 (B, destroyed?), Fanado, Pohl 5064 (B, destroyed?). P i n e s a v a n n a s a n d s l o p e s , o f t e n in s a n d y o r g r a n i t i c soil. Belize. 4 0 0 - 1 0 0 0 m . ( C u b a ; Brazil to Bolivia and Paraguay.) A w i d e s p r e a d a n d v a r i a b l e s p e c i e s in South America that deserves further study. As of yet I have been unable to discern patterns of variation that would permit the m a i n t e n a n c e as s e p a r a t e e n t i t i e s a n y o f t h e synonyms listed above. 26. RHYNCHOSPORA HIRSUTA (Vahl) V a h l . (Fig. 1A) Schoenus hirsutus Vahl, Eclog. amer. 1: 6. 1796. R. hirsuta (Vahl) Vahl, Enum. pl. 2: 231. 1806. TYPE: 1992] THOMAS: RHYNCHOSPORA 27 / 1 FIG. 2. Rhynchospora achenes; scale bar = 1 mm. A. R. diodon ( Williams & l~7lliams 44473). B. R. fascicularis subsp, fascicularis (Dwyer & Pippin 10039). C. R. gracilenta (Svenson 4757). D. R. macra (Svenson 4507). South America, yon Rohr s. n . (HOLOTYPE"C-- n.y., NY microfiche ex C). R. venezolana Kiik., Repert. Spec. Nov. Regni Veg. 53: 74. 1944. TYPE: (B, destroyed?). Moist savannas, open marshes, or pond m a r g i n s . T a b a s c o ; C h i a p a s ; Belize; G u a t e mala; Honduras; Nicaragua; Costa Rica; Panama. 0-500 m. (Also the West Indies and N South America.) 27. RHYNCHOSPORA MACRA (C. B. C l a r k e ex B r i t t o n ) S m a l l . (Fig. 2 D ) R. alba var. macra C. B. Clarke ex Britton, Trans. New York Acad. Sci. I1: 88. 1892. R. macra (Clarke) Small, Man. s. e. ft. 180. 1933. SYNTYPES: UNITED STATES. Florida, Chapman s.n. (NY); Texas, Drummond 281 (NY). Wet savannas. Nicaragua. 0-100 m. (Also t h e SE U n i t e d S t a t e s a n d P u e r t o R i c o . ) 28. RHYNCHOSPORA OLIGANTHA A. G r a y v a r . OLIGANTHA R. oligantha A. Gray var. oligantha, Ann. Lyceum Nat. Hist. New York 3: 212. 1835. SYrcrYPES: UNITED STATES. North Carolina. Fayetteville, Schweinitz s.n. (NY), near Wilmington, Curtis s. n. (NY). D a m p a r e a s in s a v a n n a s . Belize; N i c a ragua. 0 - 5 0 0 m . ( A t l a n t i c c o a s t a l p l a i n o f t h e U n i t e d S t a t e s f r o m N e w J e r s e y to G e o r gia a n d T e x a s . ) V a r . b r e v i s e t a is f o u n d in F l o r i d a a n d t h e West Indies. 29. RHYNCHOSPORA PLUMOSA E l l i o t t R. plumosa Elliott, Sketch bot. S. Carolina l: 58. 1816. TYPE: UNITED STATES. South Carolina. "Grows in dry pine barrens. Common around Beaufort. Near St. Mary's," BaMwin s.n. (ISOTYPE: NY). R. semiplumosa A. Gray, Ann. Lyceum Nat. Hist. New York 3:213.1835. TYPE: UNITED STATES. Louisiana. New Orleans, Ingalls s.n. 0SOTYPE:NY). R. penniseta Griseb., Cat. pl. Cub. 244. 1866. TYPe: WEST INDIES. "'Cuba occ.,'" C. Wright 3398 0SOTYPE: NY). Savannas and other open, moist sandy areas. Belize; H o n d u r a s ; N i c a r a g u a . 0-1 100 m. (Atlantic coastal plain of the United S t a t e s f r o m N o r t h C a r o l i n a to F l o r i d a a n d Texas; Cuba.) A c o m p l e x a n d v a r i a b l e species. F u r t h e r study may show that the Mesoamerican material may warrant separation into two taxa 28 BRITTONIA [VOL. 44 on the basis of whether or not the style base is confluent with the achene summit, whether the bristles are plumose only at the base or nearer the tip of the bristle, and the degree of elongation of the inflorescence. Moist savannas, pinelands, and pond margins. Tabasco; Belize; Nicaragua. 0-200 m. (Atlantic coastal plain of the United States from New Jersey to Florida and Texas; Cuba.) 30. 34. RHYNCHOSPORAGRACILENTAA. Gray. (Fig. 2C) R H Y N C H O S P O R A BRACHYCHAETA C . Wright R. brachychaeta C. Wright in Sauvalle, Anales Acad. Ci. M6d. Habana 8: 85. 1871. TYPE: WEST INDIES. Cuba, C. Wright 3782 (ISOTYPE:NY). R. blauneri Britton, Bull. Torrey Bot. Club 50: 56. 1923. TYPE: WEST INDIES. Puerto Rico. Sierra de Luquillo, Blauner 247 (HOLOTYPE:NY). Savannas or other moist open areas. Belize; Nicaragua. 0-100 m. (Greater Antilles.) Similar to R. chapmanii but differing in its acute spikelet tips, longer tubercle, smaller umbo, usually the presence of a rudimentary bristle, and often more elongate fascicles. 31. RI-IYNCHOSPORACHAPMANIIM. A. Curtis R. chapmanii M. A. Curtis, Amer. J. Sci. Arts, ser. 2, 7: 409. 1849. TYPE: UNITED STATES. North Carolina. Wilmington, Curtis s.n. 0SOTYPE:NY). Pine savannas and other moist open areas. Belize; Nicaragua. 0-500 m. (Atlantic coastal plain of the United States from North Carolina to Louisiana and Florida.) See comments under R. brachychaeta. R. gracilenta A. Gray, Ann. Lyceum Nat. Hist, New York 3:215.1835. TYPE:UNITED STATES. "'Pine barrens of New Jersey," Sep 1834, A. Gray s.n. (HOLOTYPE:NY). Moist savannas and other open moist areas. Chiapas; Belize; Nicaragua. (Atlantic coastal plain of the United States from New Jersey to Florida and Texas; Cuba.) 35. R H Y N C H O S P O R A FASCICULARIS (Mi- chaux) Vahl. Schoenusfascicularis Michaux, Fl. bor.-amer. 1: 37. 1803. R. fascicularis (Michaux) Vahl, Enum. pl. 2: 234. 1806. TYPE: UNITED STATES. "Hab. in Carolina," Bosc s.n. (HOLOTYPE:P, NY--frag.). Savannas, marshes, moist pinelands, and other moist open areas. Tabasco; Belize; Guatemala; Nicaragua. Low elevations. (Also the Atlantic coastal plain of the United States from Virginia to Florida and Texas; West Indies; N South America.) The following description of the typical subspecies is included to better distinguish it from the new subspecies described below. 3 5 a . RHYNCHOSPORA FASCICULARISsubsp. 32. RHYNCHOSPORAHARPERI Small R. harperiSmall, Man. s.e. ft. 182, 1503. 1933. TYPE: UNITED STATES. Georgia. Pulaski Co.: wet pine barrens about 3 mi E of Hawkinsville, Harper 1377 (NOLOTVPE:NY). Moist savannas and marshes. Belize. (Also the United States in Georgia, Florida and Alabama.) Davidse & Brant 32882 (MO, NY) represents the first record for this species outside of the United States. 33. RHVNCHOSPORAinLIFOLIAA. Gray R. fdifolia A. Gray in Torrey, Ann. Lyceum Nat. Hist. New York 3: 366. 1835. Svl,rrYPES:UNITED STATES. North Carolina, Curtis s.n. (NY); middle Florida, Chapman s.n. (NY). FASCICULARIS.(Fig. 2B) Stems 25-110 cm. Largest blades 150300 m m x 1-2.5(4) ram. Inflorescence of 1-2(4) dense, fasciculate clusters of spikelets; terminal cluster 0.8-2 x 1-2.5 cm, simply fasciculate or corymbosely compound. Spikelets 3.7-4.7 x 1.1-1.5 mm, ellipsoid to ovoid, castaneous, the scale tips attenuate to aristate. Hypogynous bristles usually 5, longest ones 1/2as long as to rarely equalling achene. Achene 1.2-1.6 x 1.1-1.4 mm, smooth and lustrous to very faintly cancellate, deep castaneous with paler, golden brown to stramineous disc. Style base 0.4-0.7 x 0.6-1 ram, deltate to broadly so, gray with pale ridge along interface with athene summit. 1992] THOMAS: RHYNCHOSPORA Savannas, marshes, moist pine forests, moist open areas. Tabasco; Belize; Nicaragua. 0-200 m. (Also the United States from Virginia to Texas; West Indies; N South America.) 36. 29 R H Y N C H O S P O R A OREOBOLOIDEA 6 6 = mez-Laurito R. oreoboloidea G6mez-Laurito, Phytologia 50(7): 459. 1982. TYPE: COSTA RICA. Cartago, A. S. Wheeler 5967a (HOLOTYPE: CR). R. paramora Mora, Mutisia 29: 4. 1966, non Stey- 35b. R h y n c h o s p o r a f a s c i c u l a r i s subsp. sierrensis W. Thomas, subsp, nov. (Fig. 3) ermark, 1951. TYPE:COLOMBIA. Dept. de Cundinamarca. P~iramode Palacio, alt. 3200 m, L. E. Mora 938 (n.y.). TYPE: G U A T E M A L A . Dept. Huehuetenango. Sierra de los Cuchumatanes, top o f Cerro Chemalito, 3.5 mi W o f Santa Eulalia, 3100-3150 m, 2 Aug 1942, S t e y e r m a r k 4 9 9 1 7 (HOLOTYPE: MO; ISOTYPE: N ' Y ) . Paramos. Costa Rica; Panama. 3 0 0 0 3400 m. (Also known from the paramos o f Colombia.) Planta glabra perennis; subsp.fascicularem simulans sed setae achaenii breviores et achaenium minus pallidum rugulosum. Culmi 12-28 cm alti; folia principalia filiformia, 100-150 x 0.4-0.9(1.1)mm; spiculae ellipsoideae, 2.8-3.2 x 1-1.2 mm, castaneae vel valde ferrugineae; setae 5-6; achaenium lenticulare obovaturn, 1.5-1.7 x 0.9-1.1 mm, transverse rugulosum; styli basis triangularis vel deltata, 0.6-0.7 x 0.5-0.7 mm. Glabrous cespitose perennial; stems 1228 cm tall. Largest blades 100-150 m m x 0.4--0.9(1.1) m m , filiform. Inflorescence o f 2-4 congested cymes o f ca 1-7 spikelets, the lowest cyme often o f one spikelet and hidden among the leaves; terminal cyme 3-8 x 3-8 m m , o f 2-7 spikelets. Spikelets 2 . 8 3.2 x 1-1.2 m m , ellipsoid, castaneous to deep ferrugineous. H y p o g y n o u s bristles 5 6, 1/2-3/4as long as (or rarely equalling) the achene. Achene 1.5-1.7 x 0.9-1.1 m m , obovate, golden brown to stramineous, faintly transversely rugulose, includes narrow basal stipe ca 0.2 m m long. Style base 0.6-0.7 x 0.5-0,7 m m , triangular to deltate with concave sides, continuous with s u m m i t o f achene. Mountain tops. Guatemala. 3 1 0 0 - 3 1 5 0 m. (Known only by the type.) The achenes are similar to those o f R. f a s c i c u l a r i s s u b s p . f a s c i c u l a r i s but are smaller, paler, and with evident rugosity and epidermal cells (i.e., the surface not smooth); the bristles are shorter, also. It is ecologically very different from subsp, fascicularis, occurring at elevations o f over 3000 m rather than at coastal elevations. T h e overall similarity o f the achenes, however, militate against its m a i n t e n a n c e as a separate species.. 37. RHYNCHOSPORA RARIFLORA (Michaux) Elliott Schoenus rariflorus Michaux, F1. bor.-amer. 1: 35. 1803. R. rariflora (Michaux) Elliott, Sketch hot. S. Carolina 1: 58.1816. TYPE: UNITED STATES. "Hab. in Georgia, herb. Michx." (HOLOTYPE:P). S a v a n n a s a n d p o n d m a r g i n s . Belize; Honduras; Nicaragua. 0-100 m. (Coastal plain o f the United States from N e w Jersey to Florida and Texas and scattered inland; West Indies.) RHYNCHOSPORA LINDENIANA Griseb. var. BAHAMENSlS(Britton) Gale 38. R. bahamensis Britton, Torreya 13: 217. 1913. R. lindeniana Griseb. var. bahamensis (Britton) Gale, Rhodora 41: 223. 1944. TYPE: WEST INDIES. Bahamas. New Providence, N. L. Britton & Brace 588 (HOLO~WPE:NY). Seasonally inundated deciduous forest. Quintana Roo; Guatemala. 0-120 m. (Also Cuba, Hispaniola, and Puerto Rico.) The typical variety is known only from Cuba and has longer spikelets and achenes. 39. RHYNCHOSPORAKLrNTHIINees ex Kunth R. kunthii Nees ex Kunth, Enum. pl. 2: 296. 1837. R. kunthii Nees, Linnaea 9: 297. 1834 [nomen]. TYPE: M E X I C O . " A d Lipaguira: H u m b o l t ' " (probable HOLOTYPE: P). R. schaffneri B6ckeler, Linnaea 37: 575-576. 1873. TYPE: MEXICO. San Luis Potosi, Schaffner 548 (HOLOTYPE:B destroyed?; ISOTVPE: K--n.y., this sheet cited as holotype by G6mez-Laurito, 1989). Sphagnum bogs, wet meadows, and openings in moist forests. Chiapas; Guatemala; Costa Rica; Panama. 2 3 0 0 - 3 3 5 0 m. (Central Mexico to Colombia and probably Peru.) Further investigation m a y reveal that R. I A. r I IImm. I ~Cm,] ]lmm. C. lmm. J. FIG. 3. Rhynchosporafascicularis subsp, sierrensis. A. Plant habit. B, Spikelet. C. Spikelet scales. D. Achene. (Drawn from holotype.) 1992] THOMAS: RHYNCHOSPORA 31 kunthii comprises two entities separable on the bases o f spikelet size, style base length, and bristle length. G 6 m e z - L a u r i t o (1989) has suggested that this m a y be the case and considers R. schaffneri the proper n a m e for Costa Rican material. I believe that further study throughout the range o f the species, including the Andes, as well as additional study o f the types is needed. Until these studies are possible, I consider both names to belong to a single species. sissippi River, southern shores o f the lower Great Lakes, and N California; West Indies.) Rhynchospora globularis var. globularis is a smaller plant with smaller achenes and spikelets and is found along the eastern coastal plain from Delaware to Florida and Texas; R. globularis var. pinetorum (Small) Gale has flatter, s m o o t h e r achenes and is found from Louisiana to Florida as well as in Cuba and Jamaica. 40. RHYNCHOSPORABRITTONII Gale 43. R h y n c h o s p o r a rosae W. Thomas, sp. nov. (Fig. 4) R. brittonii Gale, Rhodora 46: 241. 1944. TYPE: WEST INDIES. Cuba. Pinar del Rio, E. G. Britton 10023 (HOLOTYPE: NY). Savannas, sandy streamsides, and lake shores. Tabasco. 0 - 1 0 0 m. (Also W Cuba and the Isle o f Pines.) Two collections from Tabasco, Barlow s. n. (WISC) and Barlow 30/55 (BM, WISC), represent the first continental records for this species. BROWNII R o e m e r & Schultes subsp. AMERICANA Guaglianone 4 1 . RHYNCHOSPORA R. brownii subsp, americana Guaglianone, Darwiniana 22: 272. 1979. TYPE: MEXICO. Chiapas. Teopisca, 19 Aug 1966, D. Breedlove 15063 (HOLOTYPE." NY). Meadows, pinelands, and other moist open areas. Chiapas; Guatemala; H o n d u ras; Nicaragua; Panama. 0 - 2 0 0 0 m. (Mexico S along Andes to Chile; also central Argentina, Bolivia, Paraguay, and SE Brazil.) Very similar to R. rugosa (Vahl) Gale. Subspecies brownii is found in tropical and warm temperate Asia, Africa, and Australia. 42. RHYNCHOSPORA GLOBULARIS (Chapman) Small var. RECOGNITA Gale R. globularis (Chapman) Small var. recognita Gale, Rhodora 46: 245. 1944. TYPE: UNITED STATES. Virginia. Sussex Co.: 4 mi. NW of Waverley, Fernald & Long 6070 (HOLOTYPE:GH--n.v.). Savannas and open pine woodlands. Chiapas; Belize; G u a t e m a l a ; H o n d u r a s ; Nicaragua. 0 - 1 6 0 0 m. (Also U n i t e d States, along the Atlantic coastal plain from New Jersey to Florida and Texas, the lower Mis- TYPE: MEXICO. Chiapas. Mun. Jitotol; 3 mi S o f Jitotol, gentle grassy slope with Quercus and Pinus, elev. 5300 it, 29 May 1965, Breedlove 10115 (HOLOTYPE: NY; ISOTYPE: MICH). Planta gracilis caespitosa perennis; Rhynchosporam globularem simulans sed achaenium angustius pallidius et styli basis minor. Culmi 15-55 cm alti; folia principalia plieata, 70-200 • 1.2-2.8 ram; spiculae ellipsoideae vel ovoideae, 3-4 • 1-1.5 mm, castaneae vel ferrugineae; setae 6; achaenium lenticulare obovaturn, 1.4-1.7 • 0.9-1.2 ram, transverse rugulosum stramineum vel pallide stramineum; styli basis deltata vel late deltata, ab aehaenii apiee distineta. Slender, caespitose, glabrous perennial; stems 15-55 cm long. Leaves mostly basal or from near base, plicate; largest blades 7 20 cm x 1.2-2.8 m m . Inflorescence a series o f 2-4, _+ congested to open cymes with ascending branches, the branches terminating in 1-several spikelets; terminal cyme ca 1.2-4 x 0.6-4 cm. Spikelets 3-4 x 1-1.5 ram, ellipsoid to ovoid, castaneous to ferrugineous. Hypogynous bristles usually 6, 20-67% the length o f the achene, antrorsely scabrous. Achenes 1.4-1.7 x 0.9-1.2 mm, lenticular, obovate, stramineous to pale stramineous, transversely rugulose. Style base 0.3-0.5 x 0.4-0.7 mm, deltate to broadly so, clearly distinct on s u m m i t o f achene, not confluent. PARATYPES: MEXICO. Chiapas. Mun. Ocosingo, 16 km NE of Oxchuc along rd to Ocosingo, 1700 m, Breedlove 27774 (NY); mun. La Independencia, 6-10 km NNE of La Soledad along logging rd from Las Margaritas to Campo Alegre, elev. 1600 m, Breedlove 51262 (NY); 10 km S of Pueblo Nuevo Solistahuae~in, elev. 5000 ft, E. Lathrop 5976 (US); Rt. 195, 74 km from jet. with Rt. 190, near 74 km marker, Ozment, Altig 1 l l m m . 3Cm.[ A. t ,' iI C. l lmm. FIG. 4. Rhynchospora rosae. A. Plant habit. B. Spikelet. C. Spikelet scales. D. Achene. (Drawn from holotype.) 1992] THOMAS: RHYNCHOSPORA & Altig 490 (WISC); mun. Pueblo Nuevo Solistahuac~in, at Clinica Yerba Buena, 2 km NW of Pueblo Nuevo Solistahuacfin, elev. 5400 ft, Raven & Breedlove 19916 (NY); Lagunas de Montebello near Guatemala border, 30 km E ofLa Trinitaria, elev. ca 1200 m, Roe, Roe & Mori 971 (BM, WlSC--2 sheets). Open oak and pine forests on grassy slopes. 1200-1700 m. K n o w n only f r o m Chiapas. Differs from R. globularis var. recognita in its narrower, lighter achenes, the s u m m i t o f the achene without a distinctly narrowed neck, and the smaller style base. N a m e d in h o n o r o f Encarnaci6n Rosa Guaglianone, a meticulous student o f the Cyperaceae. 44. RHYNCHOSPORA MICROCARPA Baldwin ex A. Gray R. microcarpa Baldwin ex Gray, Ann. Lyceum Nat. Hist. New York 3: 202. 1835. TYPE: U N I T E D STATES. North Carolina. Wilmington, Curtis s. n. (HOLOTYPE: MY). 33 R. borinquensis Britton, Bull. Torrey Bot. Club 42: 387. 1915. TYPE: WEST INDIES. Puerto Rico. Naguabo: Rio Icaco and adjacent hills, 465-720 m elev., Shafer 3515 (HOLOTVeE:NY). Boggy areas in open pine woodlands. Chiapas; Belize; Guatemala; Honduras; E1 Salvador; Nicaragua; Costa Rica. 0 - 1 7 0 0 m. (West Indies; South America S to Argentina.) 47. RHYNCHOSPORA RUGOSA (Vahl) Gale. (Fig. 1C) Schoenus rugosus Vahl, Eclog. amer. 2: 5. 1798. R. glauca Vahl, Enum. pl. 2: 233. 1806. R. rugosa (Vahl) Gale, R h o d o r a 46: 275. 1944. TYPE: "America meridionale," Boca Chica no. 27, yon Rohr s . n . (HOLOTYPE:C--n.v., microfiche NY ex C). R. pungens Liebmann, Kongel. Danske Vidensk. Selsk. Naturvidensk. Math. Aih. ser. 2, 5(2): 253. 1851. TYPE: MEXICO. Mirador, July 1841, Liebmann s.n. (ISOTYPE: K). Savannas. Belize. 0 - 1 0 0 m. (Coastal plain o f SE United States; Cuba, Puerto Rico, Bahamas.) O p e n pine forests, m o i s t s a v a n n a s , marshes, and bogs. Tabasco; Chiapas; Belize; G u a t e m a l a ; H o n d u r a s ; N i c a r a g u a ; Costa Rica; Panama. 0 - 2 1 0 0 m. (West In4 5 . RHYNCHOSPORA JUBATA L i e b m a n n dies; Mexico S along Andes to Bolivia, SE Brazil.) R. jubata Liebmann, Kongel. Danske Vidensk. Selsk. Rhynchospora rugosa is the core o f a Naturvidensk. Math. Aih., ser. 2, 5(2): 254. 1851. widespread and variable species complex TYPE: MEXICO. Veracruz. Huatusco and Coscomatepec, Jan 1843, Liebmann s.n. (ISOLECTOTYPE (including R. marisculus, R. brownii, and R. designated by Guaglianone, 1979: K). jubata) in need o f further study. The opinLake shores. Chiapas. 1400-1600 m. In ion o f Guaglianone (1979) has been foladdition to the type, this species is known lowed except that R. pungens, which is inonly from one collection, A. Sharp 45463 t e r m e d i a t e b e t w e e n R. rugosa and R. from Chiapas, o f which I have examined brownii, is considered as part o f the former. Burger & Baker 9544 and Antonio 1548 from the M O sheet. While very similar to R. rugosa and R. Costa Rica and Panama, respectively, are marisculus, R. jubata can be distinguished examples o f a high elevation element which by its more strongly rugose achenes and its is intermediate between typical R. rugosa and R. marisculus. It has the more compact very short style base. inflorescence and smaller spikelets o f R . ru46. RHYNCHOSPORA MARISCULUS Nees ex gosa but has achenes, style bases, and bristles more like those o f R. marisculus. Lindley & Nees. (Fig. 1D) R. marisculus Nees ex Lindley & Nees in C. Martius, F1. bras. 2(1): 142.1842. R. marisculus Nees, Linnaea 9: 297. 1834 [nomen]. TYPE: BRAZIL. Rio de Janeiro. S e b a s t i a n 6 p o l i s , Martius 3193 (LECTOTYPE designated by Koyama, 1972: M - n.y.). R. tenuiseta Wright, in Sauvalle, Anales Acad. Ci. M6d. Habana 8: 83. 1871. TYPE: WEST INDIES. Cuba, C. Wright 3780 (ISOTYPE: NY). 48. RHYNCHOSPORAMACROCHAETASteudel ex Boeckeler R. macrochaeta Steudel ex Boeckeler, Linnaea 37: 632. 1873. TYPE: PERU. Near Agapata, 14/".Lechler 1819 (ISOTYPE: K). Rocky grassy slopes and paramos. Costa 34 BRITTONIA b [VOL. 44 1 FIG. 5. Rhynchospora achenes; scale bar = 1 mm. A. R. tuerckheimii (Skutch 3162). B. R. polyphylla (Denton 1262). C. R. ruiziana (Thomas 2186). D. R. schiedeana (Davidse et al. 23959). E. R. aristata (Hunt et al. 2184). Rica. 3200-3500 m. (Also along Andes from Colombia to Bolivia.) See commentary under R. schiedeana. 49. RHYNCHOSPORASCHIEDEANA(Schlechtend.) Kunth. (Fig. 5D) Chaetospora schiedeana Schlechtend., Linnaea 6: 28. 1831. R. schiedeana (Schlechtend.) Kunth, Enum. pl. 2: 300. 1837. TYPE: MEXICO. Schiede s.n. (rtOLOTYPE: B, destroyed, photo NY ex B). Clearings and steep slopes in wet forests. Chiapas; Guatemala; E1 Salvador; Costa Rica; Panama. 2000-3400 m. (Along Andes to Bolivia.) The complex of large, paniculate species including R. schiedeana presents the most difficult taxonomic problems in Rhynchospora. With study, related species such as R. vulcani, R. aristata, and R. hieronymii become, relatively speaking, clear. The remaining Mesoamerican material of this group appears to separate into three entities, although with some intergradation among the three. These I have called R. schiedeana, R. ruiziana, and R. macrochaeta, with the realization that further study of ranges of variation and type material of these and other names is necessary. Compared to R. schiedeana, R. macrochaeta is shorter with more compact inflorescences and is restricted to higher elevations, while R. ruiziana has shorter spikelets and achenes that are smaller, paler, and duller. 50. RHYNCHOSPORA RUIZIANA B/Sckeler. (Fig. 5C) R. ruiziana B6ckeler, Linnaea 37: 641. 1873. R. macrochaeta var. ruiziana (B6ckeler) Kiik. Bot. Jahrb. Syst. 74:393.1949. TYPE: PERU. Ruiz s.n. (HOLOTYPE: B, destroyed, photo NY ex B). Stream banks, boggy areas, and moist open fields. Costa Rica; Panama. 1700-3200 m. (Also S Mexico and Andean South America to Bolivia.) See comments under R. schiedeana. North of Costa Rica, this species is known from only one collection from Oaxaca, Mexico; further collecting will probably reveal its presence in the higher elevations of Chiapas and Guatemala. 1992] THOMAS: RHYNCHOSPORA 5 1. RHYNCHOSPORA VULCANI B r c k e l e r R. vulcani Brckeler, Linnaea 37: 638. 1873. TYPE: COSTA RICA. Volcfin Barba, C. Hoffmann 66 (HOLOTYPE: B, destroyed, photo NY ex B). R. lechleri Steudel ex Brckeler, Linnaea 37: 633. 1873. Steudel in Lechler, Berb. Amer. Austr. 56. 1857 [nomen]. TYPE: PERU. Tantanara, Lechler 35 1400-1450 m. (Rare; along the Andes from Venezuela and Colombia to Bolivia). H a m i l t o n & Krager 4006 is assigned to this species with trepidation and may, in fact, represent a new taxon. See comments under R. boliviensis. 2525 0SOTYPE:K). Various wet, open to partially shaded habitats at higher elevations, including paramos, open cloud forests, and lake shores. 1200-3800 m. Chiapas; Guatemala; Costa Rica; Panama. (Along Andes from S Mexico to Peru.) 52. RHYNCHOSPORA BOLIVIENSIS C. B. Clarke R. boliviensis C. B. Clarke, Kew Bull. Misc. Inform., Add. Ser. 8: 37. 1908. SY~TYPES: BOLIVIA. Unduavi, Rusby 82 (NY), Rushy 83 (NY). Cloud forests. Honduras. 1500-3000 m. (Rare; also Oaxaca, Mexico, along the Andes from Colombia to Bolivia). The species o f Section R a c e m o s a e C. B. Clarke in the Mesoamerican flora include R. boliviensis and R. dissitiflora. This group is quite variable and apparently quite rare. Thus, our knowledge o f the ranges of variation is poor. While some authors have treated R. boliviensis as a synonym o f R. dissitiflora (i.e., Koyama, 1972), there are enough differences in the limited material available that I have chosen to maintain them as separate. Study of more material, especially the types, is necessary to resolve these problems. KiJkenthal (1949) included R. lechleri Steudel ex Brckeler in Section Racemosae; it appears, however, that R. lechleri is the same species as R. vulcani which is correctly placed in Section Paniculatae B/Sckeler. 53. RHYNCHOSPORA DISSlTIFLORA Steudel ex Brckeler R. dissitiflora Steudel ex Brckeler, Linnaea 37: 598. 1873. TYPE: PERU. Tantanara, Lechler 2525a 54. RHYNCHOSPORAHIERONYMI Brckeler R. hieronymiBrckeler, Beitr. Cyp. 1: 30. 1888. TYPE: ARGENTINA. Tucumhn. Hieronymus & Lorentz 821 (HOLOTYPE: B--n.v., photo F ex B). Rain forest clearings and stream banks. Guatemala; Costa Rica; Panama. 14002500 m. (Also N Argentina, Uruguay, and S Brazil.) This species differs from the similar R. longiflora Presl of the West Indies in its divergent, non-fasciculate, and longer spikelets, wider leaves, and more robust habit. Rhynchospora hieronymi was studied in detail by Guaglianone (1981). 55. R H Y N C H O S P O R A ARISTATA Brckeler. (Fig. 5E) R. aristata Brckeler, Flora 40: 36. 1857. SV'NTYPES: MEXICO. Veracruz,Sartorius s.n. in Herb. C. H. Schultz(B?--n.v.);Jalapa, Schiede 865 (B?--n.v.); Mirador, Liebmann 594 0SOSYNTYPE:K), Liebmann 865 (n.y.). Moist forests in shade or in clearings. Chiapas; Guatemala; Honduras; Nicaragua; Costa Rica. 1200-2700 m. (Also central and S Mexico; possibly Greater Antilles and tropical montane South America, see discussion.) Although specimens from t h r o u g h o u t tropical America have been identified as R. aristata, I believe that the center of distribution of this species is central and southern Mexico and a d j a c e n t Central America. There are some South American and West Indian specimens which resemble this species but which ultimately may prove to be closer to R. hieronymi or R. schiedeana. 0SOTYPE:G). R. culmenicola Steyermark,FieldianaBot. 28(1):41. 1951. SYNTYPES:VENEZUELA. Monagas, Cerro Negro, Steyermark 62110 (K, NY); Sucre, Steyermark 62635 (NY). 56. RHYNCHOSPORALOCUPLESC. B. Clarke Boggy spots in cloud forests. Panama. Premontane tropical forest to cloud for- R. locuples C. B. Clarke, Bot. Jahrb. Syst. 34(Beibl. 78): 5. 1904. TYPE: COSTA RICA. Tonduz 11919 (ISOTYPE: US). 36 BRITTONIA est. Costa Rica; Panama. 650-2800 m. (Also the Andes from Colombia to Ecuador.) See comments under R. andresii. 57. RHYNCHOSPORA [VOL. 44 Although R. argentea may occur there, the specimens I have seen from Colombia identified as R. argentea are best referred to R. kuntzei C. B. Clarke. ANDRESII G6mez- Laurito R. andresii G6mez-Laurito, Phytologia 51(7): 477. 61. RHY-NCHOSPORATORRESIANABritton & Standley 1982. TYPE: COSTA RICA. San Jos& Parque Nacional Braulio Carillo, 900 m, G6mez-Laurito 8426 R. torresiana Britton & Standley, J. Washington Acad. 15: 473. 1925. TYPE: COSTA RICA. P. Standley 33864 (HOLOTYPE:US herb. no. 1152725). Open areas on forested slopes. Costa Rica; Panama. 600-900 m. (Known only from Costa Rica and Panama.) This species is very closely related to R. locuples from which it differs in its shorter achenes, obtuse and emarginate spikelet scale tips, and its shinier, paler, and smoother achenes. Moist forests. Costa Rica. 1200-1700 m. (Known only from Costa Rica.) 5 8. RHYNCHOSPORA DIVES Standley Forest edges and clearings and other partially shaded areas. Tabasco; Chiapas; Yucatfin; Belize; Guatemala; Honduras; Nicaragua; Costa Rica; Panama. 0-600 m. (Common throughout tropical South America.) See discussion under R. comata. (HOLOTYPE:CR). R. dives Standley, Field Mus. Nat. Hist., Bot. Ser. 17:315. 1938. TYPE: HONDURAS. Comayagua, T. Yuncker, R. Dawson & H. Youse 5900 (HOLOTYPE:F). R. orizabensis C. B. Clarke ex Kiik., Bot. Jahrb. Syst. 74(3): 406. 1949. SYNTYPES:MEXICO. Orizaba, Bourgeau 2977 (F, P), Botteri s.n. (NY), Miiller 1958 (K, NY); G U A T E M A L A . Coban, von Tuerckkeim 1256 (n.v.). Montane forests or adjacent clearings, often with Pinus, Liquidarnbar, or Quercus. Chiapas; Guatemala; Honduras; Costa Rica. 650-2300 m. (Veracruz, Mexico, to Costa Rica.) 59. RHYNCHOSPORA PIRRENSIS W . Thomas R. pirrensis W. Thomas, Brittonia 38: 314. 1986. TYPE: PANAMA. Dari6n. Serrania de Pirre, 27 Jul 1976, T. Croat 37889 (HOLOTYPE:MO). Lower m o n t a n e rainforests. Panama. 1300-1600 m. (Only known from Cerro Pirre, Panama.) 60. RHYNCHOSPORA ARGENTEAStandley R. argentea Standley, Contr. U.S. Nat. Herb. 18(3): 87. 1916. TYPE: PANAMA. Puerto Obaldia, H. Pittier 4307 (HOLOTYPE:US). Wet tropical forests, cloud forests, and associated road cuts. Panama. 100-1000 m. (Apparently endemic to Panama, see discussion.) 62. RHYNCHOSPORACEPHALOTES(L.) Vahl Scirpus cephalotes L., Sp. pl. ed. 2: 76. 1762. R. cephalotes (L.) Vahl, Enum. pl. 2:237.1806. TYPE: WEST INDIES. Herb. Linn. sheet no. 71.56 (LINN--n.v.). 63. RHYNCHOSPORA COMATA (Link) Roemer & Schultes Schoenus comatus Link, Enum. Hort. bot. berol. 1: 41. 1821. R. comata (Link) Roemer & Schultes, Mant. 2: 50. 1824. TYPE: BRAZIL. "Semina et specimena misit Sellow," (HOLOTYPE: B destroyed?). In thickets and forest clearings. Costa Rica; Panama. 50-800 m. (Widely scattered in tropical South America, rare in West Indies). Rhynchospora comata is a widely scattered and variable species comprising several elements which further scrutiny may show to deserve separate taxonomic status. Of these, the specimens from Mesoamerica, first brought to my attention by J. G6mezLaurito, plus several from the Choc6 and Santa Marta region of Colombia represent a distinct and robust element within this complex. Rhynchospora comata is closely related to R. cephalotes and R. cephalotoides Griseb. Koyama (1972) discusses the distinctions among these three. 1992] THOMAS: RHYNCHOSPORA 37 evenly distributed along stem, the distal ones equalling to surpassing the inflorescence; largest blades 7 0 - 1 3 0 m m x 0.7-1.5 m m , R. polyphylla Vahl, Enum. pl. 2: 230. 1806. TYPE: plicate. Inflorescence o f 2-4 open cymes; WEST INDIES. Monserrat, Ryan s.n. (HOLOTYPE cymes ca 1-2 x 0.5-1 cm, o f 2 - 7 spikelets C--n.y., microfiche NY ex C). each. Spikelets 4.1-4.9 x 0.8-1 mm, lanOpen areas in forests, disturbed areas, ceoloid, usually pedicellate, golden brown. r o a d s i d e s . C h i a p a s ; Belize; G u a t e m a l a ; Hypogynous bristles absent. Achene 1.2-1.3 Honduras; Nicaragua; Costa Rica; Panama. x 0.7-0.8 m m , deeply lenticular, obovate, 0 - 1 5 0 0 m. (Also West Indies; tropical South finely rugulose. Style base 0.6-1 x 0.3-0.4 America, especially along the Andes.) m m , subulate to narrowly triangular, sulSee c o m m e n t s u n d e r R. tuerckheimii. cate or at least obscurely so on both sides, _ raised and discontinuous with s u m m i t o f 65. RHYNCHOSPORA TUERCKHEIMII C. B. achene. Clarke ex Kiik. (Fig. 5A) On bare rocks in slight spray o f waterfall. Panama. 500 m. ( K n o w n only by the type.) R. tuerckheimii C. B. Clarke ex Kiik., Bot. Jahrb. Although its relationships are not clear, Syst. 74: 402--403. 1949. TYPE: GUATEMALA. Tuerckheim 1258 (HOLOTYPE: K). this species is probably close to R. dissitiflora with which it shares cauline leaves and Open areas or edges o f cloud forests or a similar achene. It is n a m e d in h o n o r o f rain forests. Guatemala; Honduras; Costa Gerrit Davidse, an energetic and perceptive Rica; Panama. 4 0 0 - 2 2 0 0 m. (Also S Mexcollector o f Cyperaceae. ico; possibly N South America.) Some specimens f r o m Chiapas and Gua67. Rhynchospora talamancensis G 6 m e z temala combine features o f this species and Laurito & W. T h o m a s , sp. nov. (Fig. 7) R. polyphylla (L. Williams et al. 40518 and TYPE: C O S T A R I C A / P A N A M A . Provs. Breedlove & Raven 13463). T h e y have the Puntarenas/Bocas del Toro" Cordillera de short spikelets and style bases o f R. polyphylla and the achene size, color, and tex- Talamanca, Cerro Echandi, on the inter82049'W, 22 Aug ture o f R . tuerckheimii. While I suspect that national border, ca 9~ 1983, G. Davidse, L. D. G6mez, A. Grijalva, they represent a distinct species, further I. Chac6n, R. Chac6n & G. Herrera Ch. study is required to detect whether or not 23895 (HOLOTYPE" CR; ISOTYPES: BM, MO, they are hybrids. 64. RHYNCUOS~RA POLYPHYLLAVahl. (Fig. 5b) NY). 66. Rhynchospora davidsei W. Thomas, sp. nov. (Fig. 6) TYPE: P A N A M A . Coc16. Distr. de la Pintada, Chorro Tile, on the Rio Tile, 11.5 airline k m N W o f E1 Cop6, Atlantic slope, 8~ 80~ 29 Sept 1983, G. Davidse 25064 (HOLOTYPE" N-Xxr; ISOTYPE" MO). Planta caespitosa gracilis perennis; culmi 10-15 cm alti, folia caulina plicam, laminae principales 70-130 • 0.7-1.5 mm; spiculae lanceoloideae, 4.1-4.9 • 0.81 mm, aureo-brunneae, plerumque pedicellatae, setae nullae; achaenium lenticulare obovatum, 1.2-1.3 x 0.7-0.8 mm, subtiliter rugulosum, styli basis subulata vel anguste triangularis, 0.6-1 x 0.3-0.4 mm, utrinque sulcata vel obscure sulcata, ab achaenii apice distincta. Slender, arching, caespitose perennial; stems 10-15 cm long. Leaves cauline, + Planta caespitosa perennis, rhizomatibus congestis; culmi recti, 10-35 cm alti, folia culmo dimidio breviora vel raro aequilonga, 1.3-1.8 m m lata, spiculae lanceolatae, 5-5.5 x 0.7-1 mm, castaneae; setae 3 antrorse hispidulae, nucem aequantes vel parum excedentes; achaenium lenticulare ellipticum, 1.2-1.5 x 0.7-0.9 mm, castaneum vel stramineum nitidum longitudinaliter cancellatum; styli basis triangularis, 1.32 x 0.5-0.6 mm, basaliter crassa. Cespitose perennial with knotted rhizomes; culms erect, 10-35 cm tall, compressed terete with a shallow groove on one face. Leaves basal or cauline near base, h a l f as long to rarely equalling culm in length, 1.3-1.8 m m wide. Inflorescence paniculate o f 2-4 partial panicles, ___ congested with ascending spikelets, 1.5-4 x 0.7-1.5 cm, o f ca 15-60 spikelets. Spikelets lanceoloid, 5 5.5 x 0.7-1 m m , castaneous. H y p o g y n o u s 38 BRITTONIA [VOL. 44 ] lrnrn. C. A. IfilmI . D. cmI FIG. 6. holotype.) Rhynchosporadavidsei.A. Plant habit. B. Spikelet. C. Spikelet scales. D. Achene. (Drawn from bristles 3, antrorsely scabrous, equalling or slightly exceeding achene. Achene elliptic, lenticular, 1.2-1.5 x 0.7-0.9 m m , castaneous to stramineous, shiny, short-stipitate, the surface longitudinally cancellate. Style base triangular, 1.3-2 m m long, 0.5-0.6 m m wide at base, the basal edge ridged a n d pron o u n c e d on slightly constricted s u m m i t o f achene. PARATYPES: COSTA RICA. Lim6n. Cordillera de Talamanca, Atlantic slope, Kfimuk massif, pfiramo NE of main Kfimuk peak, 3000-3300 m, 9~ ....... 3cm[ C. A, lrnrn. D. FIo. 7. Rhynchospora talamancensis. A. Plant habit. B. Spikelet. C. Spikelet scales. D. Achene. (Drawn from isotype, NY). 40 BRITTONIA 83~ 17-18 Sep 1984, Davidse & Herrera 29284 (MO, NY). COSTA RICA/PANAMA. Puntarenas/ Boeas del Toro. Same locality as type, 22 Aug 1983, Davidse et al. 23865 (MO). COLOMBIA. Cundinamarca. Near Bogota, pfiramo de Choachi, 3200-3400 m, 27 Sep 1917, Pennel12230 (NY). Restricted to paramos. Costa Rica; Panama. 3050-3160 m. (Known only from the Cordillera de Talamanca of Costa Rica and Panama and from Cundinamarca, Colombia.) This species is related to R. vulcani and its allies. It is described here in collaboration with Jorge G6mez-Laurito of the Museo Nacional, San Josr, Costa Rica. [VOL. 44 Savannas, open seepage slopes, and other grasslands. Tabasco; Chiapas; Campeche; Belize; Guatemala; Honduras; Nicaragua; Costa Rica; Panama. 0-1000 m. (Common in savannas throughout tropical South America; Hispaniola, Trinidad.) 71. RHYNCHOSPORAARMERIOIDES J. Presl & C. Presl R. armerioides J. Presl & C. Presl, Reliq. haenk, l: 197, t. 31, fig. 2. 1828. TYPE: PANAMA. Haenke s.n. (ISOTYPE"K). Savannas. Costa Rica; Panama. 0-500 m. (Scattered throughout tropical South America.) 68. RHYNCHOSPORACURVULAGriseb. R. curvula Griseb., Fl. Brit. W. I., 574. 1864, non (Nees) B6ckeler, 1873. TYPE: TRINIDAD. Lockhart 299 (ISOTYPE:NY). Savannas and open pinelands. Belize. 0 700 m. (Also the Guayanan region from A m a z o n i a n Colombia, Venezuela, and Trinidad to French Guiana and Parfi, Brazil.) 69. RHYNCHOSPORAMEXICANA(Liebmann) Steudel Haplostylis mexicana Liebmann, Kongel. Danske Vidensk. Selsk. Naturvidensk. Math. Afh., set. 2, 5(2): 249. 1851. R. mexicana (Liebmann) Steudel, Syn. pl. glumac. 2: 144. 1855. TYPE: MEXICO. Oaxaca. Between La Galera and Pochutla, Liebmann s.n. (C--n.v.). R. buchii Urban, Symb. antill. 7: 169. 1912. TYPE: WEST INDIES. Haiti. Near St. Michel, Buch 1047 (HOLOTYPE: B, destroyed?). Savannas and open pinelands. Belize; Guatemala; Honduras; Costa Rica; Panama. 0-1800 m. (Also West Indies; S Mexico; Venezuela.) 70. RHYNCHOSPORABARBATA(Vahl) Kunth. (Fig. 1E) Schoenus barbatus Vahl, Eclog. amer. 2: 4. 1798. R. barbata (Vahl) Kunth, Enum. pl. 2: 290. 1837. TYPE: WEST INDIES. yon Rohr s.n. (HOLOTYPE: C--n.y., NY microfiche ex C). Chaetospora pterocarpa HBK, Nov. gen. sp. 1: 230. 1816. R. pterocarpa (HBK) Roemer & Schultes, Syst. veg. 2: 89. 1817. TYPE: VENEZUELA. "Crescit in humidis prope Atures,juxta ripam Orinocensem," Humbolt & Bonpland s.n. (HOLOTYPE: P--n.v.). 72. RHYNCHOSPORAGLOBOSA(Kunth) Roemer & Schultes Chaetospora globosa Kunth in H.B.K., Nov. gen. sp. 1: 230. 1816. R. globosa (Kunth) Roemer & Schultes, Syst. veg. 2: 89. 1817, non Schoenusglobosus Kunth, 1816. TYPE: VENEZUELA. Orinoco river near San Fernando de Atabapo, Humbolt & Bonpland s.n. (HOLOTVPE:P). Savannas, marshes, and open woodlands. Belize; Guatemala; Honduras; E1 Salvador; Nicaragua; Costa Rica; Panama. 0-1000 m. ( C o m m o n , West Indies, tropical South America.) 73. RHYNCHOSPORAGIGANTEA Link. (Fig. 8B) R. gigantea Link, Jahrb. Gewiichsk. 1(3): 76. 1820. TYPE: BRAZIL. Hoffmannsegg s.n. (HOLOTYPE:B, destroyed, Willd. herb. no. 1129, microfiche NY ex B). Wet savannas, marshes, and open streamsides. Tabasco; Nicaragua. 0-100 m. (West Indies; the Guianas to southern Brazil.) 74. RHYNCHOSPORA CORYMBOSA (L.) B r i t - ton. (Fig. 8C) ScirpuscorymbosusL, Cent. pl. 2: 76. 1756. R. corymbosa (L.) Britton, Trans. New York Acad. Sci. 11: 84. 1892. TYPE: "Habitat in India" (n.y.). R. aurea Vahl, Enum. pl. 2: 229. 1809. TYPE: "Habitat in Surinamo, Jamaica, India orientali, China," (n.v.). Marshes, along streams, and other open wet areas. Tabasco; Chiapas; Belize; Gua- 1992] THOMAS: RHYNCHOSPORA 41 1 \ ,/! \ / / s~ \ 2, ,,x FIG. 8. Rhynchospora achenes; scale bar = 1 mm. A. R. scutetlata (Gentle 1220). B. R. gigantea (Svenson 4454). C. R. corymbosa (Hamilton & Stockwel13140). temala; Honduras; Nicaragua; Costa Rica; Panama. 0 - 1 6 0 0 m. (Pantropical.) 75. RHYNCHOSPORA SCUTELLATA Griseb. (Fig. 8A) R. scutellata Griseb., Cat~ pl. Cub. 246. 1866. TYPE: WEST INDIES. "Cuba occ.," C. Wright 3406 ( i s o ~ : NY). R. pringlei Greenman, Proc. Amer. Acad. Arts 39: 69. 1902. Tw,~: MEXICO. Michoacan. Swamps, Zamora, Pringle 8642 0SOTYVENY). R. indianolensis Small, F1. s.e.U.S. 193.1903. TYPE: UNITED STATES. Texas. " D a m p prairie near Indianola," Ravenel 160 (HOLOTYPE: NY). Marshes, moist savannas, and wet pastures. Tabasco; Yucatfin; Belize. Low elevations. (Southern U n i t e d States; West Indies; s c a t t e r e d in S o u t h A m e r i c a to Argentina.) 76. RHVNCnOSPORA OAXACANA Kral & W. Thomas 42 BRITTONIA R. oaxacana Kral & W. Thomas, Brittonia 38: 210. 1986. TYPE:MEXICO. Oaxaca. Mun. Ixtaltepec, 10.7 km NE of Lfizaro Cfirdenas on rd to Sta. Maria Chimalapa, IV. Thomas et al. 3584 (HOLOTYPE:MEXU; ISOTYPES:NY, VDB). Open or shrubby savannas. Chiapas. 2 0 0 850 m. (Endemic to the I s t h m u s o f Tehuantepec in Chiapas, Oaxaca, Tabasco, and Veracruz, Mexico.) Although very similar in overall appearance to species o f section D i c h r o m e n a , particularly R. floridensis, R . o a x a c a n a does not belong to the same subgenus as section D i c h r o m e n a and is m u c h m o r e closely related to the Brazilian R. c o n s a n g u i n e a (Kunth) B6ckeler o f section Pluriflorae. 77. RHYNCHOSPORAHOLOSCHOENOIDES(L. C. Rich.) Herter Schoenus holoschoenoides L. C. Rich., Actes Soc. Hist. Nat. Paris 1: 106. 1792. R. holoschoenoides (L. C. Rich.)Herter, Rev. Sudamer. Bot. 9: 157. 1953. TYPE: FRENCH GUIANA. D. LeBlonde s . n . (HOLOTYPE:P). Schoenus cyperoides Sw., Prod. veg. Ind. occ. 19. 1788, non Retz., 1786. R. cyperoides (Sw.) C. Martius, Denkschr. K6nigl. Acad. Wiss. Miinchen 6: 149. 1816/17. TYPE:n.y. Open moist areas, savannas, marshes, open woodlands, and disturbed areas. Tabasco; Chiapas; Quintana Roo; Belize; Guatemala; Honduras; Nicaragua; Costa Rica; P a n a m a . 0 - 1 5 0 0 m. ( T h r o u g h o u t tropical America; Africa and Madagascar.) 78. RHYNCHOSPORATRACYI Britton R. tracyi Britton, Trans. New York Acad. Sci. 11: 84. 1892. Ceratoschoenus capitatus Chapman, F1. South. U.S. 529. 1860, non R. capitatus Roemer & Schultes, 1817. TYPE:UNITED STATES. Florida, Hitchcock 418 (HOLOTYPE:NY). Marshes, shallow ponds, and wet savannas. Belize. 0 - 1 0 0 m. (Also coastal plain o f SE U n i t e d States; West Indies.) 79. RHYNCHOSPORAHASSLERIC. B. Clarke R. hassleri C. B. Clarke, Bull. Herb. Boiss., srr. 2, 3: 1023. 1903. TYPE: PARAGUAY. Funnen Opa [Flumen Apa], Hassler 8255 0SOTYPE:NY). R. viridi-lutea C. B. Clarke, Kew Bull. Misc. Inform., Add. Ser. 8: 36, 118. 1908. TYPE:GUYANA. Parker s.n. (LECTOTYPEdesignated by Koyama, 1972; K--n.y.). [VOL. 44 Open marshes and edges o f ponds, unc o m m o n . Belize; H o n d u r a s ; N i c a r a g u a ; Costa Rica. 0 - 1 0 0 m. (Sporadic in the West Indies, the Guianas, A m a z o n i a n Brazil, Bolivia, and Paraguay.) 80. RHYNCHOSPORA TRISPICATA Schrader ex Steudel (Nees) Ephippiorhynchium trispicatum Nees in C. Martius, F1. bras. 2(1): 136. 1842. R. trispicata (Nees) Schrader ex Steudel, Syn. pl. glumac. 2: 145. 1855. TYPE: BRAZIL. "In campestribus udis provinc. Piahy," Martius 3232 (HOLOTYPE:M--n.v.). M a r s h e s , wet s a v a n n a s , a n d s h a l l o w ponds. Tabasco; Chiapas; Belize; H o n d u ras; Nicaragua; Costa Rica. 0 - 3 0 0 m. (West Indies; tropical South America.) 81. RHYNCHOSPORATRIFLORA Vahl R. triflora Vahl, Enum. pl. 2: 232. 1806. TYPE:"Hab- itat in India orientali" (HOLOTYPE:C--n.v., microfiche NY ex C). Wet savannas and marshes. Belize; Guatemala; Nicaragua; P a n a m a . 0 - 1 0 0 m. (Also West Indies; tropical South America; tropical Africa and Asia.) Vegetatively almost identical to R. trispicata, but rarer and with rugulose rather than punctate achene surfaces. Acknowledgments I would like to thank those who carefully reviewed and edited earlier versions o f this manuscript, most particularly C. D. Adams, A. O. Chater, G. Davidse and J. Kallunki. In addition, I would like to t h a n k those experts w h o gave advice and assistance: E. R. Guaglianone, J. G 6 m e z - L a u r i t o , T. K o y a ma, and R. Kral. Jorge G 6 m e z - L a u r i t o brought to m y attention R . t a l a m a n c e n s i s which we jointly describe here. The artists deserve special mention: Bobbi Angell for so carefully rendering the illustrations o f the new taxa (Figs. 3, 4, 6, 7) and Pat K a y for bringing to life the achenes o f 17 species o f R h y n c h o s p o r a (Figs. 1, 2, 5, 8). Literature C i t e d Davidse, G., M. Sousa & A. O. Chater. 1982. Flora Mesoamericana, guia para autores. Missouri Botanical Garden, St. Louis, MO. 1992] THOMAS: RHYNCHOSPORA Grmez-Laurito, J. 1989. La sistemfitica de Rhynchospora Vahl subg. Rhynchospora (Cyperaceae) en Costa Rica. Brenesia 32: 33-72. Guaglianone, E . R . 1979. Sobre Rhynchospora rugosa (Vahl) Gale (Cyperaceae) y algunas especies afines. Darwiniana 22:255-311. .. 1981. Contribucirn al estudio del grncro Rhynchospora Vahl (Cyperaceae) III. Darwiniana 23: 489-506. Koyama, T. 1972. The botany of the Guayana Highland--Part IX. Cyperaceae--Rhynchosporeae and Cladieae. Mem. New York Bot. Gard. 23: 23-89. Kral, R. & W. Thomas. In press. Rhynchospora (Cyperaceae), In: D. Stevens, editor. Flora of Nicaragua. Missouri Botanical Garden, St. Louis. 43 Kiikenthal, G. 1949. Vorarbeiten zu einer Monographie der Rhynchosporoideae. Rhynchospora. Bot. Jahrb. Syst. 74: 375-509. Standley, P . C . 1931. The Cyperaceae of Central America. Publ. Field Columbian Mus. Bot. Ser. 8(4): 237-292. Thomas, W. W. 1984. The systematics of Rhynchospora section Dichromena. Mem. New York Bot. Gard. 37: 1-116. - - . In press. Rhynchospora (Cyperaceae). In: G. Davidse, M. Sousa & A. O. Chater, editors. Flora Mesoamericana. Missouri Botanical Garden, St. Louis. Appendix ALPHABETICAL LIST OF SPECIES 8. 57. 60. 71. 55. 70. 9. 52. 30. 21. 40. 41. 22. 62. 31. 1. 63. 23. 74. 68. 66. 24. 53. 58. 19. 25. 14. 35. 33. 17. 2. 73. 72. 42. 34, 32. 79. 54. albescens (Miq.) Kiik. andresii G6mez-Laurito argentea Standley armerioides J. Presl & C. Presl aristata Brckeler barbara (Vahl) Kunth berteroi (Sprengel) C. B. Clarke boliviensis C. B, Clarke brachychaeta C. Wright brevirostris Griseb. brittonii Gale brownii Roemer & Schultes subsp, americana Guaglianone carrillensis Grmez-Laurito cephalotes (L.) Vahl chapmanii M. A. Curtis colorata (L.) Pfeiffer comata (Link) Roemer & Schultes contracta (Nees) Raynal corymbosa (L.) Britton curvula Griseb. davidsei W. Thomas diodon (Nees) Griseb. dissitiflora Steudel ex B/Sckeler dives Standley divergens Chapman ex M. A. Curtis exaltata Kunth eximia (Nees) Brckeler fascicularis (Micbaux) Vahl 35a. subsp, fascicularis 35b. subsp, sierrensis W. Thomas filifolia A. Gray filiformis Vahl floridensis (Britton ex Small) Pfeiffer gigantea Link globosa (Kunth) Roemer & Schultes globularis (Chapman) Small var. recognita Gale gracilenta A. Gray harperi Small hassleri C. B. Clarke hieronymi B/Sckeler 26. 77. 45. 39. 38. 56. 27. 48. 46. 69. 44. 3. 1 l. 76. 28. 36. 59. 29. 64. 6. 7. 20. 4. 37. 13. 43. 47. 50. 49. 75. 67. 18. 16. hirsuta (Vahl) Vahl holoschoenoides (L. C. Rich.) Herter jubata Liebmann kunthii Nees ex Kunth lindeniana Griseb. var. bahamensis (Britton) Gale locuples C. B. Clarke macra (C. B. Clarke ex Britton) Small macrochaeta Steudel ex Brckeler marisculus Nees ex Lindley & Nees mexicana (Liebmann) Steudel microcarpa Baldwin ex A. Gray nervosa (Vahl) Brckeler 3b. subsp, ciliata Koyama 3a. subsp, nervosa nitens (Vahl) A. Gray oaxacana Kral & W. Thomas oligantha A. Gray 28a. var. oligantha oreoboloidea Grmez-Laurito pirrensis W. Thomas plumosa Elliott polyphylla Vahl polystachys (Turrill) Pfeiffer pubera (Vahl) Brckeler subsp, parvula W. Thomas pusilla Chapman radicans (Schlechtend. & Cham.) Pfeiffer 4b. subsp, microcephala (Bertero ex Sprengel) W. Thomas 4a. subsp, radicans rariflora (Michaux) Elliott robusta (Kunth) Brckeler rosae W. Thomas rugosa (Vahl) Gale ruiziana Brckeler schiedeana (Schlechtend.) Kunth scutellata Griseb. talamancensis Grmez-Laurito & W. Thomas tenerrima Nees ex Sprengel subsp, tenerrima tenuis Link 44 61. 78. 10. 81. 80. BRITTONIA 65. 12. 51. 15. 5. torresiana Brinon & Standley tracyi Britton trichodes C. B. Clarke triflora Vahl trispicata (Nees) Schrader ex Steudel [VOL. 44 tuerckheimii C. B. Clarke ex Kiik. velutina (Kunth) B6ckeler vulcani B6ckeler waspamensis Kral & W. Thomas watsonii (Britton) Davidse BOOK REVIEW The Evolutionary Process. A Critical Study of Evolutionary Theory. Second Edition. By Verne Grant, Columbia University Press, 562 West 113th Street, New York, NY 10025. ISBN 0-231-07324-0. 1991. 487 pp. $52 (cloth). Verne Grant is at least as well known for his lucid, beautifully written textbooks as for his research on evolutionary theory and the taxonomy of the Polemoniaceae. Here he presents another book that can be enthusiastically recommended to both students and professional colleagues. I read it with interest and pleasure. He has updated the first (1985 ) edition, and addressed himself to many ongoing controversies. Botanists will welcome the frequent use of plants as examples. Naturalistically inclined biologists like myself will be glad that one need not be a mathematician to understand what he has to say. Is there anything I would have liked to see done differently? Of course. More orthodox than I, Grant is a bit too defensive Books Received Bryozoan Evolution. By Frank M. McKinney and Jeremy B. C. Jackson. The University of Chicago Press, 5801 S Ellis Ave., Chicago, IL 60637. ISBN 0-226-560473. 1991. 238 pp. $15.95 (paper). [Reprint of 1989 edition.] Landscape Gardening and Rural Architecture. By A. J. Downing. Dover Publications, Inc., 31 East 2nd St., Mineola, NY 11501. ISBN 0-486-26737-7. 1991. 576 pp. $12.95 (paper) plus $2.50 postage and handling. [Facsimile reprint of the 7th edition (1865) of an influential horticultural treatise first published in 1841. A about traditional synthetic evolutionary theory for my taste, a bit too reluctant to accept new parts into the overall scheme. I would have liked to see more about regulatory genes in relation to macro-evolutionary changes and punctuated equilibria. I would have liked to see something about the work of Dover, Flavell, and their colleagues on gene conversion and duplicative transposition, even if only to discount their evolutionary importance. I would have liked to see some attention paid to the relative looseness of morphological integration in higher plants as compared to higher animals, the muted operation of competitive exclusion in plants, and the evolutionary consequences of these factors. The obvious reply to this wish-list is that it is Grant's book, not mine. Not everything can be addressed in a book of less than 500 pages, and the choice of what to include must be up to the author. The book is a good job, well done.-- ARTHUR CRONQUIST, The New York Botanical Garden. new (1990) introduction is provided by the architectural historian George B. Tatum.] Organic Gardening in Cold Climates. By Sandra Perrin. Mountain Press Publishing Co., 2016 Strand Ave., P.O. Box 2399, Missoula, MT 59806. ISBN 0-87842-2757. 1991. 143 pp. $7.95 (paper). The Subsidy from Nature. Palm Forests, Peasantry, and Development on an Amazon Frontier. By Anthony B. Anderson, Peter H. May, and Michael J. Balick. Columbia University Press, 562 West 113th St., New York, NY 10025. ISBN 0-23107222-8. 1991. 233 pp. $35 (cloth).

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