Spondias radlkoferi Donn. Sm., Bot. Gaz. (Crawfordsville) 6: 194. 1891.

Mitchell, John D. & Daly, Douglas C., 2015, A revision of Spondias L. (Anacardiaceae) in the Neotropics, PhytoKeys 55, pp. 1-92 : 40-45

Spondias radlkoferi Donn. Sm., Bot. Gaz. (Crawfordsville) 6: 194. 1891. Figs 2, 5, 15, 23

Spondias nigrescens Pittier, Contr. U.S. Natl. Herb. 18: 75, Fig. 82. 1914.

Spondias radlkoferi Type. Costa Rica. Nicoya, May 1900, A. Tonduz 13925 (holotype: US-861287!; isotypes: GH!, K! (2 sheets), MO! (fragment)).

Type.

GUATEMALA. Esquintla [Escuintla]: Esquintla, Apr 1890, J. Donnell Smith 2087 (lectotype: US-1381173!, here designated; isolectotypes: GH!, K!, US-1381174!).

Description.

Hermaphroditic trees, reproductive height 5-30 m. Trunk 2.5-60 (90) cm diam.; outer bark (brownish) gray, smooth or occasionally rough, with shallow longitudinal fissures, sometimes with colums of warts; inner bark (brownish) red with white striations. Trichomes of three types: (1) curved (rarely flexuous), erect or appressed whitish hairs 0.3-0.6 mm long; (2) erect, fine, sharp white bristles to 0.05 mm long; and (3) erect, thick, blunt hairs to 0.05 (0.1) mm long. Leaves sometimes facultatively deciduous, 4-14-jugate, 13-58 cm long; petiole 3.2-11.5 cm long, petiole and rachis glabrous or more often with dense curved hairs; lateral petiolules 2-13 mm long, the terminal one 10-20 mm long, petiolules with dense curved hairs, less often glabrous except for scattered to sparse shorter curved hairs to 1 mm long (also on rachis between leaflets); basal leaflets 1.5-6.5 × 1.1-3.1 cm, other laterals (2.5) 3-13.6 × 1.6-6 cm, all laterals medially asymmetric, acroscopic side semi-ovate to semi-lanceolate, basiscopic side (narrowly) semi-elliptic; terminal leaflet 3.6-8.5 × 1.8-3 cm, slightly (ob)ovate to oblanceolate or rarely elliptic; apex either (1) obtuse, rounded, or retuse or sometimes broadly short-acuminate (populations in Petén, Veracruz, parts of Oaxaca), or (2) abruptly and narrowly long-acuminate (remainder of range), the acumen 3-14 mm long, often mucronate; lateral lamina usually basally asymmetrical, the acroscopic side obtuse to slightly cordate, the basiscopic side acute to attenuate, basal insertion asymmetrical; leaflet margin sometimes revolute, entire to slightly crenate, occasionally with a few concave-convex teeth, sometimes ciliate with sparse curved hairs; leaflets chartaceous to membranaceous, both surfaces dull. Inflorescences subterminal, developing along with new flush of leaves, 16-33 (60) cm long, 2.8-10 mm diam at base, flowers congested toward ends of axes, secondary axes 2-22 cm long, axes (sub)glabrous (populations in Veracruz, Petén, parts of Oaxaca) or with scattered bristles and usually with dense to sparse curved hairs (elsewhere); bracts on primary and secondary axes ca. 2.5-6 mm long, lorate to subulate, bracts on higher-order branches and bracteoles 0.3-0.5 mm long, ovate or deltate, semi-clasping, the apex acuminate, all bracts ciliate with curved hairs; pedicel 0.7-2.5 (3.5) mm long, portion distal to articulation 0.4-1 (0.9) mm long, glabrous or pubescent as on inflorescence axes (but curved hairs only to 0.2 mm). Calyx 0.5-0.6 mm long overall, aestivation apert or slightly imbricate, divided nearly to base, the lobes 0.4-0.5 mm long, broadly rounded-ovate, glabrous or with sparse bristles, the margin ciliate with bristles or short blunt hairs; petals 1.8-2.2 × 0.8-0.9 mm, oblong-elliptic, apex acute to slightly acuminate, variously reported as white, greenish yellow, rose-white, or greenish cream, glabrous, reflexed at anthesis; stamens spreading, antesepalous and antepetalous ones 1.8-2.2 and 1.5-1.6 mm long, respectively, the anthers 0.5-0.6 mm long, in dorsiventral view oblong, in lateral view oblong(-elliptic); disk 0.4-0.6 mm tall, 0.4-0.7 mm thick, summit slightly undulate and outer margin sulcate, dark purple (Wendt et al. 316, NY), surface markedly papillate; pistil 0.4-0.8 mm long, depressed-globose overall, divided nearly to base into subulate, apically slightly divergent styles, often with a few hairs to 0.2 mm long, the stigmas extrorse and broadly vertically oblong. Fruits 2.2-4 × 1.5-2 cm (dry), oblong to slightly obovoid, the apex distinctly umbonate (dry), sometimes oblique at base, orange to green when ripe, surface dull, not lenticellate but sometimes warty. Seedlings (from Garwood 2009): cotyledons ligulate, entire; first two eophylls opposite and trifoliolate, then alternate, the leaflets ovate and sparsely but regularly toothed; petiole, petiolules, midvein, and margin with dense thin stiff hairs.

Leaflet venation: Intramarginal vein present or sometimes appearing to have a marginal secondary, occasionally hidden by revolute margin. Secondary veins in 5-12 pairs, often arcuate but straight near base, spacing decreasing toward apex, the angle almost uniform but decreasing toward apex; insertion on midvein abruptly decurrent or less often excurrent; some inter-secondaries present, 0-1 per pair of secondaries and usually perpendicular to the midvein, long and reticulating or basiflexed; epimedial tertiaries present, short, parallel to secondaries or perpendicular to midvein, reticulating; intercostal tertiaries alternate-percurrent and irregular-reticulate with some admedial branching; quaternaries irregular-reticulate and freely ramified, areolation at tertiary or quaternary ranks, FEVs 2-3-branched, dendritic, terminating in highly branched sclereids; on abaxial side the midvein and secondaries prominulous to prominent and discolorous, higher-order veins flat to prominulous, on adaxial surface the midvein narrowly prominulous and the secondaries and higher-order veins flattened to slightly impressed, on both sides the midvein and secondary veins often with dense to scattered curved hairs, rest of surfaces glabrous or with sparse to scattered hairs, glabrescent with age except abaxial surface usually with hairy-tuft domatia in the axils of secondary veins.

Distribution.

Spondias radlkoferi has been recorded from Mexico (Mexico State) S to NW Colombia and Venezuela (Zulia), with one record from Los Ríos in Ecuador. As noted above, different leaflet forms are associated with different parts of its range.

Ecology.

This species is rather versatile ecologically, growing in primary to secondary formations or even roadsides, in seasonally dry tropical forest, tall evergreen forest, and pluvial forest, on limestone, black clay, and reddish brown stony soils. It occurs on slopes and in valleys at elevations ranging from 10-1000 m.

Given this species’ relatively broad distribution, its known phenology is broken down by region. Mexico: flowering Mar-May, fruiting May-Dec; Central America: flowering Dec-Jul, fruiting May-Jan.

In central Panama, populations of Spondias radlkoferi often flower 4-6 weeks later than Spondias mombin populations ( Croat 1974a). The species is known to flower Apr-Jul (peaking in May-Jun) and to fruit Sep-Dec with a peak in Oct-Nov. Croat and others have suggested the fruit becomes an important source of food for mammalian species in times of food scarcity ( Croat 1974a, b) and/or forest fragmentation (spider monkeys; Chaves et al. 2012).

As with the other Spondias species whose dispersal has been documented, the fruits of Spondias radlkoferi are often dispersed by frugivorous bats ( Bonaccorso 1978, Bonaccorso and Humphrey 1984, Medellín and Gaona 1991); the endocarps of this species accounted for 50.5% of diaspores collected beneath the leaf tents of the tent-making bat Artibeus watsoni ( Melo et al. 2009).

Common names.

Belize: hog plum, ho-bo (Arvigo 899, NY), pook (Maya, Balick 1824, GH, NY), rum-p’ok (Kekchi Maya, Arvigo 627, NY); Honduras: jovo (Hagen & amp; Hagen 1096, NY), ciruela monte (Hagen & Hagen 1259, NY); Salvador: jocote (verde)(Rosales 394, NY).

Economic botany.

References to the economic botany of Spondias radlkoferi in the literature are scarce, because the species was treated as a synonym of Spondias mombin in the Flora of Panama ( Blackwell and Dodson 1967) and the Flora of Ecuador ( Barfod 1987), so some of the uses ascribed to Spondias mombin in these references should be applied to Spondias radlkoferi .

According to herbarium specimen data from Belize, the fruits of this species are edible; the bark is used to treat diarrhea, skin rashes, and fevers; a decoction is used as a mouthwash (Arvigo 899, NY); the leaves are boiled and drunk for bladder infections; a drink is prepared from the bark for internal bruises (Arvigo 825, NY); a tea from roots bark and buds is used to treat diarrhea; a tea from roots, bark, and buds is used to treat gonorrhea; boiled roots, bark and buds are used as an eye-wash; leaves and bark are boiled to make a tonic bath during pregnancy; and an infusion of leaves is used as a gargle for sore throats and to treat skin sores (Balick 1824, GH, NY).

Selected specimens examined.

BELIZE. Belize District: South of Yalbac Hills, Terra Nova Medicinal Plant Reserve, 17°21'N, 89°55'W, elev. 40 m, 19 July 1995, Walker et al. 1497 (NY); Toledo District: Bladen Watershed, Quebrada de Oro tributary, 16°35'N, 88°45'W, 16 Mar 1988, Brokaw 42 (NY). COLOMBIA: Antioquia: Mun. Dabeiba, km 4 Dabeiba Chigorodo road, 30 Jul 1987, Callejas et al. 4767 (MO, NY); Caldas: Mpio. Norcasia, Magdalena Medio, Hacienda Playa Alta, 260 m, 22 Jul 2001, Garzón & Lopera 103 (NY); Chocó: km 41-56 on QuibdóBolívar road, 5°47'N, 76°35'W, 11 Jun 1982, Gentry & Brand 36710 (NY); Córdoba: Mpio. Tierralta, installations of Urrá Dam, elev. 260 m, 9 Jun 2003, Fonnegra-G. et al. 7850 (NY). COSTA RICA. Alajuela: La Garita Dam, 25 Jul 1967, Lent 1148 (NY); Heredia: La Selva (OTS Field Station), on Río Puerto Viejo just E of confluence with Río Sarapiqui, elev. ca. 100 m, 8 Jun 1985, Jacobs 3293 (GH, BM, NY); Puntarenas: km 15 RincónPuerto Jiménez road, 8°33'N, 83°23'W, 4 Mar 1985, Croat & Grayum 59795 (MO, NY). ECUADOR. Los Rios: Vinces, Jauneche forest, km 70 Quevedo-Palenque, via Mocachi, 1°16'S, 79°42'W, elev. 70 m, (no date), Dodson et al. 8837 (GUAY, MO, SEL). EL SALVADOR. Dept. Ahuachapán, San Francisco Menéndez, El Corozo [Coroso], Mariposario, 13°49'N, 89°59'W, elev. 380 m, 24 Mar 2000, Rosales 394 (NY). GUATEMALA. Alta Verapaz: SE of Finca Yalpemech, near Alto Verapaz-Petén boundary, 23 Mar 1942, Steyermark 45214 (TEX); Petén: road to Melchior, 10 Jul 1965, Aguilar 37 (NY); Tikal National Park, Tikal, near airfield, 20 Aug 1959, Contreras 85 (GH, TEX). HONDURAS. Atlantida: Lancetilla Valley, near Tela, elev. 20-600 m, 6 Oct-20 Mar 1928, Standley 54022 (A); Yoro: Subirana, Oct 1937, C. von Hagen & W. von Hagen 1096 (NY); MEXICO. Campeche: Tuxpeña, 2 Nov 1931, Lundell 894 (GH, NY); Mpio. Calakmul, km 17 S of gate house for entry to Calakmul, 18°23'29"N, 89°54'W, 27 Jul 1998, Madrid et al. 1264 (MO); Chiapas: Mpio. Ocosingo, 0.2 km W of Nuevo Guerrero, 16°59'11"N, 91°17'14"W, elev. 210 m, 8 May 2002, Calónico Soto et al. 23370 (TEX); Quintana Roo: km 5 Las PanterasMargarita Maza road, on short cut to Mérida, 5 Aug 1982, E. Cabrera & H. Cabrera 3331; Vera Cruz: Mpo. San Andrés Tuxtla, Estación de Biología Tropical Los Tuxtlas, 18°3436'N, 95°0409'W, 15 Apr 1975, Calzada 1813 (NY). NICARA GUA. Boaco: Comarca San Isidro, ca. 17 km N of Camoapa, ca. 12°33'N, 85°30'W, 17 Jul 1984, Estrada et al. 9 (MO). Zelaya: Awas Tingni, 40 km S of Waspán, 14°23'N, 83°57'W, elev. 20 m, 20 Mar 1971, Little 25273 (MO). PANAMA. Canal Zone: Barro Colorado Island, 18 Apr 1968, Croat 4929 (NY), Colón: Distrito Portobelo, banks of Río Guanche, elev. 100 m, 9°31'N, 79°40'W, 18 Jan 1995, Galdames & Guerra 1928 (NY); San Blas: Comarca de San Blas, Playón Chico, aqueduct trail, 9°17'N, 78°15'W, 11 Sep 1994, Herrera & Arosemena 1834 (NY). VENEZUELA. Zulia: Sierra de Perija, vicinity of Kasmera ( Estación Biológica de la Universidad del Zulia), SW of Machiques, 25 Aug 1967, Steyermark & Fernández 99734 (NY).

Conservation status.

We consider this species to be of Least Concern because of its broad range and relatively large populations in Central America, S Mexico and Colombia, although the population(s) in the heavily deforested region of Zulia, Venezuela may be endangered.

Discussion.

Spondias radlkoferi most closely resembles Spondias mombin because of its usually densely fissured bark, leaves 3-14-jugate, the midvein of the leaflet usually prominent abaxially, and the petals glabrous abaxially. The former can be distinguished by the intramarginal secondary vein (sometimes (sub)marginal) (vs. always removed from the margin); the costal secondary veins usually distinctly arcuate with excurrent insertion on midvein, sometimes with hairy tuft domatia in the axils abaxially (vs. essentially straight to very slightly arcuate with decurrent insertion on midvein, without hairy tuft domatia); the pedicel 0.7-2.5 (3.5) mm long (vs. 2-4.5 mm); and the fruit maturing green (rarely orange), obovoid with abruptly short-acuminate apex (vs. maturing yellow or orange(-brown), oblong to ellipsoid to globose, apex rounded to truncate).

Moreover, the tertiary veins are alternate-percurrent and irregular-reticulate (vs. irregular-reticulate and/or admedially ramified); FEVs 1-2-branched, terminating in highly branched sclereids (vs. 3+-branched and not terminating in branched sclereids); on pedicel the portion distal to the articulation almost always shorter than basal portion (vs. distal portion longer); the sepals slightly imbricate at base (vs. calyx apert); the disk markedly papillate (vs. not), the pistil often with with a few trichomes to 0.2 mm long (vs. glabrous).