Spondias macrocarpa Engl. in Mart., Fl. bras. 12(2): 375, pl. 78. 1876.

Mitchell, John D. & Daly, Douglas C., 2015, A revision of Spondias L. (Anacardiaceae) in the Neotropics, PhytoKeys 55, pp. 1-92 : 29

Spondias macrocarpa Engl. in Mart., Fl. bras. 12(2): 375, pl. 78. 1876. Figs 2, 15, 16, 19

Spondias dulcis Parkinson var. macrocarpa (Engl.) Engl. in A. DC. & C. DC., Monogr. phan. 4: 247. 1883. Type. Based on Spondias macrocarpa Engl.

Type.

BRAZIL. Rio de Janeiro: Canta Gallo [Cantagalo], 1859, Peckolt 224 (Lectotype: BR-572018, here designated).

Description.

Hermaphroditic trees, reproductive height 12-22 m. Trunk 24-60 cm diam; outer bark brown, rough, thin, scaly, shed in usually large irregular plates; inner bark red with tan striations. Trichomes of three types: long, white, flexuous to nearly straight, sometimes uncinate, 0.7-1 mm long on vegetative parts; short, erect hairs usually less than 0.1 mm long, primarily on basal portions of inflorescence; and orange capitate glandular hairs to 0.05 mm long (these rare). Leaves 7-12jugate, 12-30 cm long; petiole 3-5 cm long, petiole and rachis with dense long white hairs; lateral petiolules 1-3 mm long, the terminal one 3-20 mm long, petiolules with hairs as on rachis; basal leaflets 1.7-4.8 × 0.7-1.8 cm, ovate or less often lanceolate, other laterals 5-7 × 1-2 cm, (falcate)lanceolate to elliptic, terminal leaflet 2-6 × 0.7-1 cm, ovate to lanceolate; leaflet apex long-acuminate, the acumen 4-13(20) mm long; lateral lamina medially and basally asymmetrical, acroscopic side obtuse to cordate, basiscopic side attenuate to cuneate, basal insertion on petiolule (sub)symmetrical and excurrent or slightly decurrent; margin flat to slightly revolute (often slightly so at base) and sparsely serrulate, the teeth usually concave-convex, sinus appearing glandular; leaflets membranaceous to chartaceous, both surfaces dull. Inflorescences (sub)terminal, developing with leaf flush, 5-15 cm long, ca. 4 mm diam at base, secondary axes to 2 cm long, axes with sparse to dense short erect hairs toward base, distal portions glabrescent, bracts on inflorescence axes to 0.5-1.2 mm long, bracteoles 0.3-0.5 mm long, all bracts ovate to subulate; pedicel 2.5-3.5 mm long overall, portion distal to articulation 0.7-2.7 mm. Calyx 0.5-0.6 mm long overall, aestivation apert, divided nearly to base, the lobes 0.3-0.5 mm long, (depressed-)deltate, margin occasionally papillate; petals 2.5-2.6 × 1.3-1.6 mm, essentially elliptic, acute or usually slightly acuminate, white or cream, glabrous, reflexed at anthesis; stamens spreading, the antesepalous and antepetalous ones 2.9-3.0 and 2.5-2.6 mm long, respectively, the anthers 0.7-0.9 mm long, oblong in both dorsiventral and lateral views; disk (0.1) 0.3-0.6 mm tall, 0.1 mm thick, summit shallowly undulate and outer margin nearly entire; pistil 1.5 mm overall, depressed-ovoid to subcylindrical overall, divided halfway to 2/3 its length into 5 broadly subulate, apically connivent styles 0.8-1 mm long, stigmas extrorse, obovate. Fruits 3.5-4.2 × 2.3-2.5 cm (dry), maturing yellowishgreen to (orange-)yellow, oblong to slightly (ob)ovoid, surface sparsely whitishlenticellate, mesocarp whitish, sweetsour.

Leaflet venation: Fimbrial vein absent; secondary veins 10-15 pairs, straight to slightly arcuate, insertion on midvein excurrent or abruptly decurrent, spacing irregular, angle usually irregular, slightly acute to nearly perpendicular; intersecondaries and/or epimedial tertiaries sometimes present, parallel to secondaries, nearly reaching intramarginal vein, branching admedially; intercostal tertiaries few per secondary vein, strongly admedially branched, sometimes also sparsely branched toward margin; quaternary veins irregular-reticulate and freely ramified; areolation at tertiary and quaternary ranks; FEVs 3+-branched, dendritic, terminating in tracheoid idioblasts; marginal ultimate venation incompletely looped; on abaxial side all veins narrowly prominent, on adaxial side the midvein narrowly prominulous (almost keeled) but sunk in a groove, rest of veins flat to impressed and obscure; both sides sparsely to densely pubescent.

Distribution.

Spondias macrocarpa is native to moist upland forests of the Mata Atlântica Complex, in southern Bahia, Rio de Janeiro, Espírito Santo, and extreme southeastern Minas Gerais.

Ecology.

This species appears to be rare where it does occur. It has been recorded in mussununga forest (dense forest with discontinuous canopy 8-15 m high, on level terrain, in sandy soils (spodosols) that are often poorly drained)(Stefano et al. 200, NY), and tabuleiro forest (dense forest with continuous canopy 20-25 m high, on level terrain, in sandy clay soils (oxisols))(Stefano et al. 225, NY). The species is known to flower Jun-Feb and to fruit Mar-Apr (Jun).

Common names.

Brazil. Espírito Santo: cajá mirim (Farias 475, NY); Rio de Janeiro: acajá (Peckolt s.n. (BR-571916, BR), cajá (Peckolt 224, BR).

Selected specimens examined.

BRAZIL. Bahia: São Paulinho, on road to Catolesinho, 9 Nov 1942, Fróes 12671/37 (A, NY); by Itatinga road, [15°15'S, 40°15'W] 7 Oct 1945, Fróes 20079 (IAN, NY); Mun. Santa Cruz Cabrália, Estação Ecológica do Pau Brasil-ESPAB, ca. 16 km W of Porto Seguro, BR-367 (Porto Seguro-Eunápolis) highway, 18 Dec 1987, F. Souza Santos 820 (CEPEC); Mun. Juçari, Fazenda Sto. Antônio Alciato de Carvalho, ca. 6 km N of Juçari, 0.5 km from "Fazenda de Cacau," 22 Jun 1991, Thomas et al. 6823 (CEPEC, NY); Espírito Santo: Linhares, Reserva Natural Vale, next to native plant nursery, 26 Nov 1991, Farias 475 (CVRD, NY); Mun. Linhares, Reserva Natural Vale, 1700 m on Farinha Seca Road, 27 Apr 1992, Folli 1614 (CVRD, NY); Mun. Linhares, Reserva Florestal da Sooretama, 10 Feb 1993, Hatschbach & Silva 60062 (TEX); Mun. Pinheiros, Pinheiros, km 12 of Pinheiro-Montanha road, 50 m from asphalt, 23 Nov 1991, V. Souza 267 (CVRD, NY); Reserva Florestal da CVRD, Linhares, 11 Nov 1977, J. Spada 013/77 (CVRD, NY); Mun. Linhares, Reserva Vale (BR-101 Norte, km 122), Estrada Flamengo, 19°7'14"N, 39°54'59"W, 1 Mar 2011, Stefano et al. 200 (CVRD, NY, RB); Minas Gerais: Alegria, near Caraça, 12 Oct 1882 (1883 on BR sheet), Glaziou 13678 (BR [on-line image seen], K, P, R); Serra da Carajá, 12 Aug 1882, Glaziou 13679 (P); Coronel Pacheco, Estação Experimental do Café, 6 Sep 1940, Heringer 60 (RB); Rio de Janeiro: Canta Gallo [Cantagalo], 1859, Peckolt 224 (BR 571985) (BR), Jan 1860, Peckolt s.n. (BR 571916)(BR), 1860, Peckolt s.n. (BR 571982)(BR).

Conservation status.

We classify this species as “Vulnerable,” although it is relatively widespread in Atlantic Forest Complex of Brazil, because of the relatively small number of known specimens represented in a well-collected region. It should be noted that many of the collections are rather old and may be from localities that are no longer forested.

Discussion.

The species referred to as Spondias macrocarpa in Árvores Brasileiras ( Lorenzi 1998), Brazilian Trees ( Lorenzi 2002), and Brazilian Fruits and Cultivated Exotics ( Lorenzi et al. 2006) is in fact Spondias expeditionaria ; these books were published before discovery of the latter. Indeed, Spondias macrocarpa most closely resembles Spondias expeditionaria ; a comparison of the two appears under the latter species.