Anthurium upalaense Croat & Baker

Brenesia 16 (Supl. l): 97. 1979.

TYPE: Costa Rica. Alajuela: along road between Canas (Guanacaste) and Upala, near Río Zapote, 1.8-2.7 km south of Río Canalete, ca. 100 m elev., disturbed margin of primary forest, 25 June 1976, Croat 36342 (MO-2381219, holotype; CR, F, isotypes; Live at MO).

Rosulate, epiphyte; stems less than 20 cm long, 1.5-2 cm diam.; leafscars hidden by persisting cataphylls and roots; roots moderately thick and long, 2-4 mm diam., brown, moderately dense and numerous; cataphylls coriaceous, 11-22 cm long, narrowly acuminate at apex, sometimes tinged lightly with red, turning brown on drying, remaining firm and intact, ultimately deciduous.

LEAVES erect-sprcading; petioles (6)10-39 cm long, 7-10 mm diam., subquadrangular, broadly and sharply sulcate, prominently 3-ribbed abaxially, the sides flat to somewhat concave; geniculum 1-1.5 cm long, sometimes reddish abaxially; blades narrowly oblong-elliptic to oblong-oblanceolate, moderately thick, gradually acuminale at apex (the acumen apiculate, down- lurned), broadly to narrowly acute at base, (30)40- 106 cm long, (4.5)8-23 cm wide, broadest at the middle or slightly above; both surfaces semi- glossy; the midrib flat at base above, narrowed and becoming triangular toward apex, prominently raised below, 3-ribbed near base; primary lateral veins more than 20 per side, departing midrib at ca. 60¡ angle, raised above and be- low, ± straight to collective vein, loop-connect- ing in upper half; the interprimary veins flat or slightly raised above, flat and slightly darker than surface below; lesser veins less conspicuous; col- lective vcin arising in the upper half, flat above and below, 3-5 mm from margin.

INFLORESCENCE erect to spreading or arching-pendent; peduncle (18)40-48 cm long, 4-9 mm diam., terete or l -ribbed below opening of spathe (faint- ly many ribbed), sometimes tinged purplish at base, longer than petioles; spathe moderately thick, yellow-green (B & K Yellow-green 6/7.5, 7/2.5, 7/5), sometimes heavily tinged with vi- olet-purple, oblong-lanceolate to lanceolate, (4.5) 8-20 cm long, 1-4 cm wide, broadest at base, gradually short-acuminate at apex, acute at base, inserted at 60¡-75¡ angle on peduncle; spa- dix sessile, green to purplish (B & K Yellow-green 6/5), sometimes red, 9-30 cm long, 6-15 mm diam. at base, 4-6 mm diam. at apex; the flowers square to rhombic, 2.3-3.4 mm long, 1.7-2.8 mm wide, the sides straight to weakly sigmoid; 7-15 flowers visible in the principal Spiral, 10- 20 flowers visible in the alternate Spiral; tepals matte, inconspicuously pale punctate, with drop-lets usually after 2 or more stamens emcrge, lat- eral tepals 0.8-1.8 mm wide, the inner margin straight to slightly convex; the pistil raised ca. 0.5 mm, green to purplish; Stigma elliptic, 0.4- 1.1 mm long, grecn, brushiike, exserted ca. 0.1 mm, with droplets for 5-15 days beginning 12- 22 days before the first stamens emerge, dry and black when stamens emerge; stamens emerging rapidly from the base or ncar the base, emerging one at a timc ca. l cm ahead ofone another on spadix, the apical slamens usually preceding the basal ones, cxserled on short greenish filaments, ca. 0.2 mm long, 0.5-1 mm wide, which retract to hold anthers in a light circle around pistil or separately at sides of pistil; anthers pale orange, 0.5-0.6 mm long, 0.7-1 mm wide; thecae ellipsoid, scarcely divaricatc, opening flat; pollen orange, fading to pale orange or lavender, white or tan.

INFRUCTESCENCE pendent; the spathe reflexed; the spadix to 55 cm long, 6 cm diam.; berries oblong, red, tapered to both ends, 9-15 mm long, acule at apex, emerging ± uniformly throughout spadix, well above tepals before fully mature; mesocarp mealy, sweet, white; seeds 2, oblong, to4 mm long, 3 mm wide, 2.5 mm thick, creamy white, obliquely truncate at apex with a short appendage, greenish at base. Figs. 211 and 212.

The species is known from the Atlantic slope of Nicaragua and Costa Rica at elevations from near sea level to usually less than 600 m in re- gions of tropical wet forest, premontane wet forest, and wetter parts of tropical moist forest life zones. A collection from Cartago Province, northeast of Río Grande de Orosi at 1,300 m is apparently also this species. Anthurium upalaense is a member of section Pachyneurium and is apparently most closely related to A. concolor K. Krause from Panama but differs in having the inflorescence pendent, the spadix long-tapcred and more or less green at anthesis, the spathe longerand recurled, the fresh anthers pale orange, the major lateral veins scarcely raised, offen forming a collective vein from near the middle or above, the reticulate veins clearly visible when fresh, weakly raised and prominulous when dried. Anthurium con- color, on the other hand, has the inflorescence stiffly erect, the spathe usually less than 10 cm long, usually not recurled at apex, the spadix not tapered, dark purple-violet, and usually less than 7 cm long at anthesis. Its flowers form minute globules of nectar on the tepals and the pollen is pale violet-purple, its major lateral veins are prominently raised and seldom form a collective vein except near the apex, and the reticulate veins are more or less obscure. Anthurium concolor is known only from the Atlantic slope in central Panama.

Map of Mesoamerican specimens with coordinates

Costa Rica Alajuela: 450 m, 10.45N 85.3W, 7-8 Nov 1975, Baker & Burger 9859 (CR, F, MO). Costa Rica Alajuela: 400 m,, 24 June 1976, Thomas B. Croat 36319 (MO). Costa Rica Alajuela: R’o Frio, 30-40 m, 11.2N 84.44W, 1 Aug 1949, Holm & Iltis 811 (B, G, K, P, U). Costa Rica Alajuela: 100 m, 11.2N 84.44W, 25 June 1976, Thomas B. Croat 36380 (MO). Costa Rica Alajuela: R’o Zapote, 100-200 m, 10.48N 85.2W, 12 Nov 1975, Burger & Baker 10001 (CR, F, MO). Costa Rica Alajuela: R’o Zapote, 100 m,, 25 June 1976, Thomas B. Croat 36342 (CR, F, K, MO). Costa Rica Alajuela:, , Taylor & Taylor 1170 (NY). Costa Rica Alajuela:, , Smith, Austin 1498 (NY). Costa Rica Alajuela:, , Molina et al 17264 (EAP, F). Costa Rica Alajuela: R’o Frio,, , Holm & Iltis 771 (G, NY). Costa Rica Alajuela: R’o Zapote,, , Burger & Baker 9964 (F). Costa Rica Alajuela: Lake Arenal,, , Brenes 18194 (CR). Costa Rica Alajuela: Lake Arenal,, , Brenes 18195 (CR). Costa Rica Alajuela: R.N.V.S. Ca–o Negro, 40 m,, 16 Apr 1995, Villalobos, R. 188 (INB). Costa Rica Cartago: 1300 m,, 27 May 1976, Utley & Utley 5062 (MO). Costa Rica Cartago:, , Pittier 12720 (G, US). Costa Rica Cartago: Lake Bonilla,, , Pacheco 92 (F). Costa Rica Costa Rica:, , N.E. Brown s.n. (K). Costa Rica Costa Rica:, , Bull 467/1873 (K). Costa Rica Guanacaste: R’o Aguacate, 500 m,, 16 Feb 1984, Khan et al. 1085 (BM). Costa Rica Guanacaste:, , Standley & Valerio 45328 (US). Costa Rica Heredia: Finca La Selva, 100 m,, 16 March 1983, Chacon 462 (MO). Costa Rica Heredia: Finca La Selva, 100 m,, 5 Dec 1982, McDowell 1058 (MO). Costa Rica Heredia: Finca La Selva, 100 m,, 7 Oct 1982, McDowell 389 (MO). Costa Rica Heredia: Puerto Viejo Region, 20 m,, 27 May 1976, Thomas B. Croat 35708 (MO). Costa Rica Heredia: Finca La Selva, 100 m,, 5 Jan 1978, Thomas B. Croat 44297 (MO). Costa Rica Heredia: Finca La Selva, 100 m,, 3 Jan 1978, Thomas B. Croat 44263 (MO). Costa Rica Heredia: Puerto Viejo Region,, , Burger & Matta 4343 (F, CR). Costa Rica Heredia: Finca La Selva,, , Folsom 9558 (TEX). Costa Rica Heredia: 10.26N 84.01W, 15 April 1986, Thomas B. Croat 61218 (B, MO, WU). Costa Rica Lim—n: 10-40 m, 10.40N 83.40W, 13-14 Sep 1986, Warren Douglas Stevens 24336 (MO). Costa Rica Lim—n: Cerro Coronel, 20-170 m, 10.41N 83.38W, 16-23 Jan 1986, Warren Douglas Stevens 23726 (MO). Costa Rica Lim—n: 20-170 m, 10.41N 83.38W, 15-20 Sep 1986, Warren Douglas Stevens 24501 (MO). Costa Rica Lim—n: Cerro Coronel, 10-100 m, 10.40N 83.40W, 24 Jan 1986, Warren Douglas Stevens 23923 (MO). Costa Rica Lim—n: 10-40 m, 10.40N 83.40W, 13-14 Sep 1986, Warren Douglas Stevens 24270 (MO). Costa Rica Lim—n: 300 m, 9.24.30N 82.58.15W, 8 July 1989, Abelardo Chac—n 131 (MO). Costa Rica Lim—n: 5 m, 09.38N 82.39W, 2 Nov 1984, Michael Grayum & Willie Burton 4324 (MO). Costa Rica Lim—n: 10-20 m, 10.39N 82.42W, 1 Aug 1984, Michael H. Grayum 3638 (MO). Costa Rica Lim—n: 25-40 m, 09.35N 82.39W, 19 Nov 1984, Michael Grayum 4501 (MO). Costa Rica Lim—n: 1000 m,, 6 May 1983, Gomez et al. 20543 (CR, MO, MSC, WIS). Costa Rica Lim—n: 40 m, 10.30N 83.47W, 22 Aug 1979, Davidson & Donahue 8727 (MO). Costa Rica Lim—n: 100 m,, 8 May 1977, Ocampo 1985 (MO). Costa Rica Lim—n: 40 m, 10.30N 83.47W, 21 Aug 1979, Davidson & Donahue 8608 (MO). Costa Rica Lim—n: 40 m, 10.30N 83.47W, 14 Aug 1979, Davidson & Donahue 8272 (MO). Costa Rica Lim—n: 0-10 m, 09.57N 83.2W, 27-30 Nov 1975, Baker & Burger 17 (CR, F, MO). Costa Rica Lim—n: R’o Catarata, 50 m,, 12 Aug 1977, Thomas B. Croat 43221 (F, MO, NY, RSA, SEL). Costa Rica Lim—n:, , Utley & Utley 660 (F). Costa Rica Lim—n: Valle de Talamanca,, , Tonduz 9507 (BR, CR). Costa Rica Lim—n: Cordillera de Talamanca,, , Pittier 9509 (BR). Costa Rica Lim—n:, , Tonduz 9228 (BR). Costa Rica Lim—n: R’o Reventaz—n,, , Standley & Valerio 48869 (US). Costa Rica Lim—n:, , Burger & Burger 8446 (CR, F). Costa Rica Lim—n:, , Baker & Burger 174 (F). Costa Rica Lim—n: 200 m, 10.01N 83.25W, 28 October 1988, Gerardo Herrera 2245 (MO). Costa Rica Lim—n: 10-80 m, 10.40.30N 8.39.30W, 17-18 Sep 1986, Gerrit Davidse & Gerardo Herrera 31352 (MO). Costa Rica Lim—n: 5 m, 10.41.30N 83.39.30W, 15-17 Sep 1986, Gerrit Davidse & Gerardo Herrera 31232 (MO). Costa Rica Lim—n: 10-120 m, 10.39.00N 83.40.40W, 15-16 Sep 1986, Gerrit Davidse & Gerardo Herrera 31271 (MO, W). Costa Rica Lim—n: 850 m, 09.14730"N 83.10530"W, 07 Apr 1989, Herrera & Madrigal 2548 (CR, MO). Costa Rica Lim—n: 4 m, 10.31N 83.30W, 24 December 1987, Rafael Robles 1496 (MO). Costa Rica Lim—n: Parque Tortuguero, 4 m, 10.31N 83.30W, 24 November 1987, Rafael Robles 1331 (F, L, MO). Costa Rica Lim—n: Parque Tortuguero, 4 m, 10.31N 83.30W, 24 November 1987, Rafael Robles 1332 (MO). Costa Rica Lim—n: 40 m, 10.25N 83.34W, 28 October 1987, Rafael Robles 1177 (MO). Costa Rica Puntarenas: Canton of Coto Brus, 1200 m, 08.49N 82.58W, 6-7 March 1984, Croat 57257 (CR, MO). Costa Rica San José:, , Tonduz 13319 (G, US). Nicaragua Chontales: 390 m, 12.17N 85.15W, 14 Sep 1982, Warren Douglas Stevens 21826 (MO). Nicaragua Chontales: 250 m, 12.18N 85.07W, 6 Abril 1984, A. Grijalva & D. Soza 3749 (MO). Nicaragua Chontales: 700 m,, 20 July 1975, Atwood & Neill AN35 (MO). Nicaragua Chontales: Cerro Oluma, 600-700 m, 12.18N 85.23W, Sept. 23, 1983, Micheal Nee 28359 (MO). Nicaragua Jinotega: R’o Coco, 200 m, 14.00N 85.20W, 14 March 1980, Stevens et al. 16823 (MO). Nicaragua Matagalpa: Quebrada Las Carpas, 580 m, 13.15N 85.34W, 11 March 1981, Stevens & Moreno 19715 (MO). Nicaragua Matagalpa: Cerro Musun, 500-800 m,, 20-21 Apr. 1977, Neill 1782 (MO). Nicaragua Matagalpa: R’o Bilampi, 200-500 m, 13.00N 85.14W, 15 May 1980, Araquistain, M. & Moreno, P. 2658 (L, MO). Nicaragua Matagalpa: Cerro Musœn, 200-500 m, 13.00-01N 85.14W, 16 May 1980, Araquistain, M. & Moreno, P. 2739 (BM, LE, MBM, MO). Nicaragua Matagalpa: Cerro Musun, 200-500 m, 13.00-01N 85.14W, 16 May 1980, Araquistain, M. & Moreno, P. 2744 (MO). Nicaragua Matagalpa: 600 m, 13.15N 85.35W, 13 septiembre 1982, Pedro P. Moreno 17218 (MO). Nicaragua Río San Juan: 40-60 m, 11.14N 84.26W, 30 Nov-2 Dec 1984, Warren Douglas Stevens 23437 (MO). Nicaragua Río San Juan:, , Neill 1552 (MO). Nicaragua Río San Juan: R’o San Juan del Norte, 5 m,, 2 Dec. 1982, Araquistain 3401 (MO). Nicaragua Río San Juan: R’o San Juan, 10 m,, 29 Nov. 1982, Araquistain 3283 (MO). Nicaragua Río San Juan: 0-2 m, 10.55N 83.43W, 5 Sept. 1982, Riviere 222 (MO). Nicaragua Río San Juan: R’o San Juan, 0 m, 10.55N 83.43W, 5 September 1982, Martínez & Riviere 1952 (MEXU). Nicaragua Río San Juan:, 29 Nov. 1982, Araquistain, M. 3302 (MO). Nicaragua Río San Juan: R’o Indio, 11.06N 83.54.30W, 6 Sept 1982, Martínez & Riviere 1957 (MEXU, MO). Nicaragua Río San Juan: 10.56N 83.46W, 1 Julio 1994, Ricardo Rueda, Blas Hern‡ndez y Edith Palma 1560 (MO). Nicaragua Río San Juan: 11.03N 83.50W, 5 Julio 1994, Ricardo Rueda, Blas Hern‡ndez y Edith Palma 1806 (CM, MO). Nicaragua Río San Juan: 10.42N 083.46W, 12 Febrero 1996, Ricardo Rueda, Alfredo Grijalva, Rolando Dolmus y Milton Castrillo 4118 (MO, PMA). Nicaragua Río San Juan: 11.02N 083.48W, 29 Julio 1996, Ricardo Rueda, Indiana Coronado y Norlan Tercero 4636 (MO). Nicaragua Río San Juan: 11.02N 083.48W, 29 Julio 1996, Ricardo Rueda, Indiana Coronado y Norlan Tercero 4637 (MO). Nicaragua Río San Juan: 11.02N 083.48W, 29 Julio 1996, Ricardo Rueda, Indiana Coronado y Norlan Tercero 4659 (MO, WU). Nicaragua Zelaya: Cerro Waylawas, 13.30.20N 84.45.25W, 28 Octubre 1982, A. Grijalva & F. Burgos 1685 (MO). Nicaragua Zelaya:, 19 Jul 1975, Atwood & Neill AN233 (MO). Nicaragua Zelaya: 140 m, 14.01N 84.34W, 20 February 1979, W.D. Stevens 12317 (MO). Nicaragua Zelaya: 200-400 m, 12.150-55N 85.10-5W, 13 Feb. 1979, Stevens 12143 (MO). Nicaragua Zelaya: 160-180 m,, 10 May 1978, Vincelli 328A (MO). Nicaragua Zelaya: 200-400 m, 12.50N 85.00W 12.55N 85.05W, 14 February 1979, W.D. Stevens 12206 (MO). Nicaragua Zelaya: 200-400 m, 12.50N 85.00W, 14 Feb. 1979, Stevens 12182 (MO). Nicaragua Zelaya: 100-200 m, 13.36N 85.05W, 18 March 1978, W.D. Stevens 7524 (MO). Nicaragua Zelaya: 10 m, 11.36N 83.51W, 06 Feb. 1982, Moreno, P. and Sandino, J. 14933 (MO). Nicaragua Zelaya: 10 m, 11.35N 83.42W, 25 octubre 1981, Pedro P. Moreno 12466 (MO). Nicaragua Zelaya: R’o Punta Gorda, 10 m, 11.34N 84.01W, 10 noviembre 1981, P.P.Moreno y J.C.Sandino 12755 (MO). Nicaragua Zelaya: 260-300 m, 13.38N 85.02W, 23 diciembre 1982, Pedro P. Moreno 19190 (MO). Nicaragua Zelaya: 10 m, 11.33N 83.48W, 4 febrero 1982, P.P. Moreno & J.C. Sandino 14655 (MO). Nicaragua Zelaya: 10 m, 11.33N 83.48W, 4 febrero 1982, P.P. Moreno & J.C. Sandino 14688 (MO). Nicaragua Zelaya: Siuna, 300-345 m, 13.46N 84.46W, 12 marzo 1984, F. Ortiz 1773 (MO).

Map of South American Specimens with coordinates Colombia Narino: La Planada, 1700 m, 1.06N 77.53W, 10 March 1990, Thomas B. Croat 43305 (AAU, CR, G, GH, K, LE, MO, NY, P, RSA, UCLA, W, WIS).