Spondias L., Sp. pl. 371. 1753.

Mitchell, John D. & Daly, Douglas C., 2015, A revision of Spondias L. (Anacardiaceae) in the Neotropics, PhytoKeys 55, pp. 1-92 : 13-14

Spondias L., Sp. pl. 371. 1753.

Cytheraea Wight & Arn., Prodr. Fl. Ind. Orient. 1: 173. 1834, nom. prov. = Spondias dulcis Parkinson.

Wirtgenia Jung. ex Hassk., Flora 27: 624-625. 1844, (non Sch. Bip.) pro parte quoad Wirtgenia decandra = Spondias pinnata (Koenig ex L.f.) Kurz.

Evia Comm. ex Blume, Mus. Bot. 1(15): 233. 1850. Type: Evia dulcis (Parkinson) Comm. ex Blume = Spondias dulcis Parkinson.

Warmingia Engl. in Mart., Fl. Bras. 12(2): 281. 1874 (non Rchb. f.). Type: Warmingia pauciflora = Spondias purpurea L.

Type.

Spondias mombin L.

Description.

Small to large trees (rarely shrubby and broadly branching), usually hermaphroditic (except Spondias purpurea ). Simple, thick plank buttresses to 100 cm high sometimes present. Outer bark brown or usually gray, densely to broadly fissured, sometimes thick, usually rough, often with raised lenticels, rarely (some Spondias mombin and Spondias purpurea ) with large, corky, tooth-like projections. Inner bark usually broadly striate (white and rose, red, orange, or brown). Resin viscous and usually clear or less often cloudy ( Spondias globosa ). Trichomes of various simple hairs and capitate glandular hairs. Leaves alternate, aggregated toward branch tips, sometimes deciduous (sometimes facultatively so), usually imparipinnate, petiolulate; leaflets (sub)opposite, the apex often apiculate; margin entire/cren(ul)ate/serr(ul)ate (Figs 2 - 4), the sinuses often glandular; intramarginal vein present (see Figs 1 - 10 for leaflet venation); intercostal tertiaries usually irregular-reticulate and often branching admedially, areoles often poorly developed; freely ending veinlets dendritic, often highly branched, leaflets often glandular-punctate. Inflorescences produced either before leaf flush (e.g., Spondias purpurea ) or during leaf flush, terminal or axillary, paniculate (reduced to a pseudoracemose botryoid in Spondias purpurea , Barfod 1988); inflorescence bracts subtending secondary axes early-caducous; pedicel articulated. Flowers usually hermaphroditic but strongly protandrous (dioecious in Spondias purpurea , Fig. 14), pentamerous, calyx green (often red in Spondias purpurea ), shorter than the disk, lobes usually apert (quincuncial at the base and valvate at the apex where they meet in Spondias purpurea , Bachelier and Endress 2009); corolla valvate, petals yellowishwhite (usually red in Spondias purpurea ); patent or strongly reflexed at anthesis (spreading in Spondias purpurea ), often terminating in a swollen and strongly inflexed apiculum, the margin papillate; stamens (8)10, in two unequal series, outer series opposite the carpels, filaments linear to subulate, glabrous, anthers dorsifixed, yellow; disk intrastaminal, annular, summit and outer margin variously shaped (Fig. 15), usually yellow or less often pink (some Spondias purpurea ) or purple (some Spondias radlkoferi ); pistillode ( Spondias purpurea only) reduced to 3-5 subulate styles; at anthesis the ovary usually rudimentary, developing after the anthers dehisce (Fig. 11), pistil (3-) 5locular, surmounted by (3-)5 short, free styles ( Spondias purpurea ) or usually the styles (broadly) subulate and separate at least half the length of the pistil, connivent at least basally, stigmas usually extrorse (introrse in Spondias purpurea and Spondias radlkoferi ), discoid to linear; each locule with one apical ovule, the ovules crassinucellar and bitegmic, styles separating as fruit develops. Fruit (Fig. 16) a (1) 3-5-seeded, green, yellow, orange, or red drupe, oblong, (depressed-)globose, obovoid, or ellipsoid, remnants of styles widely separated and often visible in developing fruits; exocarp usually smooth (densely lenticellate in Spondias testudinis ); mesocarp fleshy, edible, sweet and sour, endocarp bony and enveloped by a fibrous matrix (with spiny projections and growing over apex of pedicel in Spondias dulcis ), at maturity the fruit often breaking away at articulation of pedicel. Pollen (based on Roubik and Moreno P. 1991): 41-56 × 22-44 µ, prolate or subprolate, pores 6-15 µ diam. Seedlings: germination hypogeal, tap root and hypocotyl emerge from blunt end of fruit; hypocotyl emerges, curved at first, then carries cotyledons aloft; phanerocotylar; cotyledons opposite, linear, green, sessile, somewhat fleshy; first eophylls opposite and imparipinnate ( Spondias testudinis , Fig. 17H) or trifoliolate (pers. obs., Vogel 1980, Garwood 2009). Chromosomes : n = 16, 2n = 32 ( Simmonds 1954, Bawa 1973, Mehra 1976, Guerra 1986, Pedrosa et al. 1999, Almeida et al. 2007).

Of the ca. 18 species in Spondias , ten are native to the New World, distributed from Mexico to southern Brazil, one is native to Madagascar, and seven are native to Asia and the South Pacific, from Malesia (sensu Flora Malesiana) to tropical China, Sri Lanka, Indochina, Thailand, India (except extreme north), Myanmar (Burma), Solomon Islands east to Polynesia. Spondias dulcis is cultivated in tropical America and the Antilles; Spondias mombin and Spondias purpurea are both introduced throughout Tropical West Africa and Asia (and in the West Indies, where they are not found in primary vegetation and may not be native); Spondias mombin is often adventive in Tropical West Africa.

Some of the species native to the Neotropics have restricted distributions. Spondias testudinis is restricted to southwestern Amazonia, and Spondias admirabilis and Spondias expeditionaria both are known from very few localities in Brazil’s Atlantic Coastal Forest, while the other two Atlantic Forest species, Spondias macrocarpa and especially Spondias venulosa , are somewhat more broadly distributed in that region; both Spondias expeditionaria and Spondias macrocarpa are rare as well. The natural distribution of Spondias tuberosa is the arid caatinga vegetation of Northeastern Brazil, and Spondias globosa is a Western Amazon species. Spondias radlkoferi ranges from Mexico through Central America to Colombia and NW Venezuela; there is an unconfirmed report from Los Ríos in W Ecuador (Dodson 8837, MO). Spondias purpurea is native to N Mexico through Central America and may be native to SW Ecuador. Spondias mombin is native to moist forests through much of northern South America, although it is uncertain whether the populations in Brazil’s Atlantic Coastal Forest are native.