Flora Mesoamericana, Volme 2 (1), , page 1 of 7

First published on the Flora Mesoamericana Website, 19 Dec. 2011.

70. OPILIACEAE Descripción de la familia por P. Hiepko.

Trees or shrubs (rarely vine-like), root-parasites. Leaves without stipules, petiolate, distichous, simple; blades often coriaceous, in dried state mostly finely tubercled by cystoliths located in the mesophyll, usually extremely variable in form and size, entire, mostly glabrous, venation pinnate. Inflorescences axillary, racemose (Mesoamérica); bracts very small, often scale-like, caducous. Flowers pedicellate, actinomorphic, 4-5-merous, unisexual and dioecious (Mesoamérica). Perianth with valvate, free tepals. Stamens as many as and opposite to the tepals (in female flowers wanting), anthers introrse, 2-celled, longitudinally dehiscent. Disc intrastaminal, lobed (lobes alternating with the stamens), or annular. Pistil 1; ovary superior, 1- celled, stigma sessile, entire or shallowly lobed; ovule 1, pendulous from the apex of a central placenta, anatropous and unitegmic, sometimes integument not differentiated (). Fruit drupaceous, pericarp rather thin, mesocarp fleshy-juicy, endocarp woody or crustaceous. Seed large, conform to the drupe, embryo terete, embedded in rich, oily endosperm, nearly as long as the seed, with 3-4 linear cotyledons, radicle often very small. 10 gen., 33 spp. Trópicos del Viejo y del Nuevo (1 gen., 10 spp.) Mundo. Bibliografía: Hiepko, P. Fl. Neotrop. (in prep.). Sleumer, H. en Engler, A. & Prantl, K. Nat. Pflanzenfam. ed. 2, 16b: 33-41 (1935). Standley, P.C. J. Washington Acad. Sci. 10: 505-508 (1920); Trees & shrubs Mexico, Contr. U.S. Natl. Herb. 23(2): 235-236 (1922). Nevling, L.I., Jr. Fl. Panama, Ann. Missouri Bot. Gard. 47: 291-292 (1960). Williams, L.O. Ciencia (Mexico) 24: 227-228 (1966).

1. Agonandra Miers ex Benth. Por P. Hiepko.

Dioecious trees or shrubs, sometimes vinelike. Leaves usually glabrous, base of lamina decurrent on petiole. Inflorescence a raceme with 1-3(-4) flowers per bract, bracts slightly peltate,

1 Flora Mesoamericana, Volme 2 (1), Opiliaceae, page 2 of 7 caducous, bracteoles (if present) minute, rachis glabrous, papillate or puberulous. Flowers pedicellate, 4-5-merous, unisexual. Tepals free, glabrous, papillate or puberulous. Stamens with filiform filament, exceeding the perianth. Disc lobed or annular to urceolate. Pistil in male flowers rudimentary. Drupe ellipsoid, sometimes more or less globular, to 2.5 cm long, pedicel cylindrical or obconical. 10 spp. Mexico, Central and South America.

1. Racemes at least partly with 3(-4) flowers per bract. 2. Trees, to 50 m tall, in wet forest; racemes glabrous but minutely papillate. 2. A. goldbergiana 2. Small trees, 3-12(-20) m tall, in savanna or dry deciduous forest; racemes glabrous or puberulous. 3. Racemes and flowers densely puberulous; 3 flowers per bract . 1. A. brasiliensis subsp. brasiliensis 3. Racemes and flowers glabrous; 3-1 flower per bract. 3. A. racemosa 1. Racemes with 1 flower per bract. 4. Young branches and racemes glabrous or minutely papillate 4. A. macrocarpa 4. Young branches, petioles, and racemes more or less puberulous (rarely glabrous). 5. Young branches and petioles puberulous to glabrous, midrib glabrous; fruits 8-10 × 5-7 mm; growing from 0 to 500 m. 5a. A. obtusifolia subsp. obtusifolia 5. Young branches, petioles, and lower part of midrib beneath pubescent; fruits 15-18 × 12 mm; growing above 1000 m. 6. Shrubs to 3.5 m tall; leaves usually smaller than 3 × 1 cm. 5b. A. obtusifolia subsp. conzattii 6. Trees, c. 10 m tall; leaves 5-8 × 3-6 cm. 6. A. ovatifolia

1. Miers ex Benth. in Benth. & Hook.f., Gen. Pl. 1: 349 (1862). Lectotipo (designado por Hiepko, 1993): Brazil, Gardner 1503 (K!). Ilustr.: Nevling, Fl. Panama, Ann. Missouri Bot. Gard. 47: 291, f. 83 (1960).

1a. Agonandra brasiliensis subsp. brasiliensis.

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Small trees, (1-)3-12(-30) m tall, with thick corky, longitudinally furrowed, pale yellow- grey bark; young branches minutely papillate to puberulous. Leaves chartaceous to coriaceous, broadly ovate to narrowly elliptic, 4-9(-11) × 2-5(-6) cm, apex acute to shortly acuminate, base rounded to attenuate, shining above, dull below, midrib flat above, prominent beneath, lateral veins 6-8 per side, prominulous above and (less) beneath; petioles(3-)5-20(-30) mm long. Racemes axillary, 1-2(-6, in male specimens only) per axil, often at defoliated nodes, puberulous, male racemes 3-6(-7) cm, female racemes (1.5-)2-4 cm long; bracts broadly ovate to angular-ovate, c. 2 mm long, upper half with ciliate margin. Flowers 3 (rarely 4) per bract (in female inflorescences often only in the lower half in groups of 3) with 2 very small bracteoles at the base of the pedicels of the lateral flowers. Pedicels and flowers puberulous, pedicels 1-2.5 mm long. Male flowers: Tepals oblong, acute, 2-3 mm long. Stamens 2.5-3.5 mm long, anthers oval, 0.5 mm. Disc lobes fleshy, 0.5-0.7 mm long, apex irregularly toothed. Rudimentary pistil cylindric, c. 1 mm long. Female flowers: Tepals oblong,0.5 mm long. Disc annular, thinly fleshy, up to 0.2 mm high. Pistil conical, 0.5 mm long; stigma shallowly 3-lobed. Drupe 2.3 × 1.7 cm, pedicel up to 14 mm long. In savannas and deciduous forests, sometimes in riparian forests. P (Williams 368, NY). 0-300 m. Mesoamerica, Colombia, Venezuela, Guyana, Bolivia, Brazil, Paraguay.)

2. Agonandra goldbergiana Hiepko, Willdenowia 27: 225 (1997). Izabalaea excelsa Lundell, non Agonandra excelsa Griseb. Holotipo: Guatemala, Contreras 6791 (LL!). Trees, to 50 m tall, 1 m diam.; young branches minutely papillate. Leaves chartaceous to coriaceous, elliptic to narrowly elliptic, 7-10(-12) × 2-4.5 cm, apex obtuse to acute, base attenuate, midrib flat above, prominent beneath, lateral veins 5-7 per side, very slender and inconspicuous, hardly prominulous on both sides; petiole 2-5 mm long, minutely papillate at base adaxially. Racemes axillary, 1-3 per axil, mostly on defoliated nodes, rachis densely papillate, male ones 1-2.5 cm, female ones 2-2.5 cm long; bracts depressed-orbicular, c. 3 mm wide, upper half ciliolate; flowers usually 3 per bract, without bracteoles at the base of the pedicels of the lateral flowers. Pedicels papillate like the rachis, flowers glabrous, pedicels 2-2.5 mm long. Male flowers: Tepals oblong, acute, 2-2.5 mm long Stamens 4 mm long, anthers oval, 0.5 mm long. Disc lobes thinly fleshy, 1-1.5 mm long, apex extrorsely irregularly toothed. Rudimentary pistill cylindric, c. 1 mm long. Female flowers: Tepals 1 mm long. Disc annular,

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thinly fleshy, o.3 mm high. Pistil conical, 1 mm long, stigma cushion-shaped. Drupe (not yet ripe) 2.5 × 1.7 cm, pedicel 7-11 mm long. Flowering Mar; fruiting Apr. In tall riparian forests. G (Contreras 19065, LL). (Endémica.) This species was originally decribed by Lundell as a new monotypic genus of .

3. Agonandra racemosa (DC.) Standl., J. Washington Acad. Sci. 10: 507 (1920). Schaefferia racemosa DC., Prodr. 2: 41 (1825). Holotipo: Mexico, "Schweinizia racemosa" (unpublished illustration, Sessé & Mociño, at Hunt Institute for Botanical Documentation, copies at G, photo IDC 713, vol.1, tab. V). N.v.: Limoncillo, Ch; ciprés silvestre, pino silvestre, ES. Loranthus anomalus M.E.Jones. Small trees or shrubs (rarely vine-like), 2.5-10(-15) m tall, branches drooping, brittle, glabrous; trunk to 25(-100?) cm diam., bark brown-black, slightly corky and fissured. Leaves chartaceous to coriaceous, narrowly ovate to narrowly elliptic, rarely broadly ovate or elliptic, 4- 8(-10.5) × (1-)2-3(-4.5) cm, apex acute to shortly acuminate, base cuneate to rounded, midrib flat above, slightly prominent beneath, lateral veins 4-6 per side, curved ascending, mostly slightly prominulous on both sides, sometimes hardly visible; petioles 5-10(-12) mm long. Racemes terminal or axillary, 1 per axil, 1.5-4(-5.5) cm long (terminal inflorescences often with racemes instead of flowers in the basal part thus forming a double raceme), rachis glabrous, in dry state longitudinally ridged, 3, 2 or 1 flowers per bract; bracts broadly cordate, 1 × 1 mm, acute, glabrous, bracteoles linear, very small, at the base of the pedicels of the lateral flowers. Pedicels and flowers glabrous, pedicels 1.5-3 mm long. Male flowers: Tepals oblong, acute, c. 2 mm long. Stamens 2.5-3 mm long, anthers oval, c. 0.5 mm long. Disc lobes thinly fleshy, 1 mm long, sometimes more or less fused at the base (up to ¼ of their length), apex irregularly toothed. Rudimentary pistil thinly cylindric with capitate stigma, not exceeding the disc lobes. Female flowers: Tepals c. 1 mm long. Disc annular, thinly fleshy, 0.5 mm high, often with some higher lobes, which are sometimes as long as the pistil. Pistil ovoidal, c. 1 mm long, stigma cushion- shaped, shallowly 3- to 4-lobed. Drupe ellipsoid (sometimes more or less globular), yellow to whitish with purple, (7-)9-11 × 6-8 mm, pedicel 5-10(-12) mm long. Flowering Oct-June; fruiting Dec-June. In seasonally dry deciduous forests and in gallery forests. Ch (Breedlove

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42232, MEXU); G (Aguilar 1032, F); H (Rodriguez 2569, F); ES (Sandoval 138, LAGU). 600- 3500 m. (Mexico [western Mexico: from Sonora and Chihuahua to Chiapas]; Mesoamerica.)

4. Agonandra macrocarpa L.O. Williams, Ciencia (Mexico) 24: 227 (1966). Holotipo: Honduras, Standley 25878 (F!). Ilustr.: Burger, Fl. Costaric., Fieldiana, Bot., n.s. 13: 15, t. 3, 28 (1983). N.v.: Pakalché, Y; palo de melón, N. Agonandra loranthoides L.O. Williams; A. obtusifolia sensu Standley (1930). Small trees or shrubs, 3-10(-15) m tall, bark gray-brown with deep fissures; young branches obscurely papillate-puberulent. Leaves coriaceous, often hard and wrinkled when dry, elliptic to narrowly ovate to rhombic, (2-)3-5.5(-7) × 1-3 cm, apex obtuse, base attenuate, midrib flat above, prominulous beneath, lateral veins 3-4 per side, steep ascending, rather obscure on both sides; petiole glabrous, 4-7(-10) mm long. Racemes axillary, often on defoliated nodes, 1-4 per axil, male ones 1-2.5 cm, female ones 0.5-1 cm long, rachis minutely papillate or glabrous, 1 flower per bract; bracts crescent shaped, 1.5 mm wide, ciliolate, no bracteoles. Pedicels minutely papillate like the rachis, flowers glabrous, pedicels 1.5-2 mm long. Male flowers: Tepals oblong, acute, 2-2.5 mm long. Stamens 3-4 mm long, anthers oval, 0.5 mm long. Disc lobes thinly fleshy, c. 1 mm long, apex irregularly toothed. Rudimentary pistil cylindric with capitate stigma, c. 1 mm long. Female flowers: Tepals more or less triangular, c. 1 mm long. Disc annular, fleshy, margin undulating, c. 0.3 mm high. Pistil ovoidal, c. 1 mm long, stigma cushion-shaped. Drupe greenish, 1.8-2.1 (fresh up to 3 cm)x 1.4-1.9 cm, pedicels 5-6 mm long. Flowering Dec-Apr; fruiting Feb-May. In seasonally very dry deciduous or semideciduous forests. Y (Gaumer 23609, F); C (Shepherd 58, MICH); QR (Miranda 8159, MEXU); H (Molina R. 2724, BM); N (Laguna 232, HNMN); CR (Gomez-Laurito 10036, CR). 50-900 m. (Mesoamerica.) "The fruit of this tree is much sought for by the deer and many thousands of them are killed under these trees" (Gaumer 23609).

5. Agonandra obtusifolia Standl., J. Washington Acad. Sci. 10: 507 (1920). Holotipo: Mexico, Palmer 421 (US!). Ilustr.: Acevedo Rosas & Martínez y Pérez, Fl. Veracruz 84: 5, t. 1 (1995). Shrubs or small trees, 1-5(-8) m tall, often compact with many lateral branches and stiff, thorne-like branchlets, bark dark gray, fissured; young branches pubescent or finely puberulous.

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Leaves coriaceous, often hard and wrinkled when dry, elliptic to narrowly oblong, ovate or obovate, (2-)3-4(-6) × (0.7-)1-2 cm, apex obtuse or slightly mucronulate, base attenuate, midrib prominulous on both sides, lateral veins 2-3(-4) per side, steep ascending, prominulous above or rather obscure on both sides; petiole pubescent (sometimes also lower part of midrib beneath pubescent) or puberulous to more or less glabrous, 2-4 mm long. Racemes axillary, sometimes on defoliated nodes, 1-2(-3) per axil, male ones 0.5-1.5 cm, female ones 0.4-0.7 cm long, rachis finely puberulous to minutely papillate, 1 flower per bract; bracts broadly ovate, 1-1.5 mm wide, ciliolate, no bracteoles. Pedicels minutely papillate to finely puberulous like the rachis or glabrous, flowers glabrous, pedicels 1-1.5 mm long. Male flowers: Tepals oblong, acute, 2 mm long Stamens 2.5-3(-4) mm long, anthers oval, 0.5-1 mm long. Disc lobes thinly fleshy, c. 1 mm long, apex irregularly toothed. Rudimentary pistil cylindric with capitate stigma, c. 1 mm long. Female flowers: Tepals narrowly triangular, c. 1 mm long. Disc annular, fleshy, margin slightly undulating, c. 0.3 mm high. Pistil ovoidal, c. 1 mm long, stigma cushion-shaped. Drupe dark yellow, 8-18 × 5-12 mm, pedicel 3-4.5(-6) mm long, cylindrical or slightly obconical. On sand dunes and in seasonally dry deciduous forests or thorny thickets, often in hedgerows. Ch. 0-1900 m. (Mexico [Tamaulipas and Veracruz to Puebla, Oaxaca], Mesoamerica.)

5a. Agonandra obtusifolia subsp. obtusifolia. Shrubs or small trees, 1-5(-8) m tall. Leaves (2-)3-4(-6) × 1-2 cm; petiole puberulous to more or less glabrous. Drupe 8-10 × 5-7 mm, pedicel 3-4.5 mm long, cylindrical. Ch (Breedlove et al. 54190, CAS). 0-500 m. (Mexico, [Tamaulipas and Veracruz to Oaxaca], Mesoamerica.)

5b. Agonandra obtusifolia subsp. conzattii (Standl.) Hiepko, Willdenowia 27: 226 (1997). A conzattii Standl., J. Washington Acad. Sci. 10: 508 (1920). Holotipo: Mexico, Oaxaca, Conzatti 3558 (US no. 1012311!). Shrubs, 1.5-3.5 m tall. Leaves 2-3(-3.5) × 0.7-1(-1.5) cm; petiole and lower third of midrib beneath pubescent. Drupe 15-18 × 10-12 mm, pedicel 3-6 mm long, obconical, base c. 1 mm, apex 2 mm thick. In seasonally dry deciduous forests or thorny thickets. Ch (Miranda 6034, MEXU). 1000-1900 m. (Mexico [Puebla, Oaxaca], Mesoamerica.)

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6. Agonandra ovatifolia Miranda, Anales Inst. Biol. Univ. Nac. México 24(1): 71 (1953). Lectotipo (designado aquí): Mexico, Miranda 6906 (F!). Ilustr.: Miranda, Anales Inst. Biol. Univ. Nac. México 24: 72, t. .2 (1953). N.v.: Aceituna, Ch. Trees, c. 10 m tall. Leaves coriaceous, hard and wrinkled when dry, broadly ovate to elliptic, 5-8 × 3-6 cm, apex obtuse, base rounded to attenuate, midrib flat above, prominent beneath, hairy in the lower third beneath, lateral veins 3-5(-6) per side, rather obscure on both sides; petiole hairy below, glabrous above, c. 8 mm long. Racemes axillary, usually on defoliated nodes, 1-3 per axil, male ones not known, female one 5-7 mm long, rachis papillate, 1 flower per bract, bracts broadly ovate, 1.5 mm wide, ciliolate. Pedicels minutely papillate like the rachis, 1.5-2 mm long. Male flowers: not known. Female flowers: Tepals c. 1 mm long. Disc annular, c. 0.3 mm high, pistil ovoidal, c. 1 mm long. Drupe not known. Ch (Miranda 6906, F). 1700 m. (Endémica.) Known only from the type. The female flowers and inflorescences resemble those of Agonandra obtusifolia subsp. conzattii and also the indument of the petioles and the lower part of the midrib beneath is typical for that taxon too, but A. ovatifolia is a much bigger tree and the shape and size of the leaves is also extremely different. According to Miranda (1953) the very hard, yellowish wood is used as timber.

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