A revision of Spondias L. (Anacardiaceae) in the Neotropics Author Mitchell, John D. Institute of Systematic Botany, The New York Botanical Garden, 2900 Southern Blvd., Bronx, NY 10458 - 5126 johndanmitchell@gmail.com Author Daly, Douglas C. Institute of Systematic Botany, The New York Botanical Garden, 2900 Southern Blvd., Bronx, NY 10458 - 5126 text PhytoKeys 2015 2015-08-05 55 1 92 http://dx.doi.org/10.3897/phytokeys.55.8489 journal article http://dx.doi.org/10.3897/phytokeys.55.8489 1314-2003-55-1 FF8CFFB4FFD52937FFDFFFB2FF93FC06 576322 Spondias dulcis Parkinson, J. voy. South Seas 39. 1773. Figs 2 , 15 , 16 , 20 Poupartia dulcis (Parkinson) Blume, Bijdr. fl. Ned. Ind. 1161. 1826-27. Evia dulcis (Parkinson) Blume, Mus. Bot. 1(15): 233. 1850. Type. based on Spondias dulcis Parkinson. Spondias cytherea Sonn., Voy. Indes orient. 3: 242, t. 123. 1782. Spondias dulcis Type. Mauritius (cultivated), Commerson s.n. (P!). Spondias dulcis Parkinson var. commersonii Engl. in A. DC & C. DC., Monogr. phan. 4: 247. 1883. Spondias dulcis Type. Several syntypes cited. Spondias dulcis Parkinson var. mucroserrata Engl. in A. DC. & C. DC., Monogr. phan. 4: 247. 1883. Spondias dulcis Type. Mexico, w/o date, Pavon 744 (G n.v.; GH-photo!, NY-photo!). Spondias dulcis Parkinson var. integra Engl. in A. DC. & C. DC., Monogr. phan. 4: 248. 1883. Spondias dulcis Type. Indonesia. Amboin, w/o date, Reinwardt s.n. (W!). Type. TAHITI. (without date), Capt. Cook [Banks & Solander] s.n. (lectotype, BM-793299 n.v., designated by A. C. Smith 1985 : 453). Description. Hermaphroditic, trees , reproductive height 8-25 m. Trunk 20-40 cm diam.; outer bark light gray or light brown, thin, smooth to moderately rough, lenticellate, shed in small thin plates. Plant entirely glabrous except for some capitate glandular hairs . Leaves sometimes partially deciduous, 4-12-jugate, 11-60 cm long; petiole 9-15 cm long; lateral petiolules 2-8 mm long, the terminal one 10-30 mm long; basal leaflets 4.3-7.5 x 1.3-3.5 cm, other laterals 5-15 x 1.7-5 cm, all laterals oblong or lanceolate to ovate, terminal leaflet 5-9 x 1.9-3.5 cm, (narrowly) elliptic with acute base; leaflet apex acuminate or occasionally acute, the acumen 4-13 mm long, apex tip acute and glandular-mucronate; lateral leaflets medially subsymmetrical, basal width subsymmetrical, base insertion symmetrical and cuneate or obtuse, decurrent; margin slightly revolute and usually serrulate or crenulate, when present teeth concave-convex, sinus spacing regular, sinus glandular; leaflets chartaceous, adaxial surface sometimes glossy. Inflorescences usually developing with new leaf flush, terminal and axillary, congested at branchlet apex, 9-32.5 cm long, 3-7 mm diam at base; secondary axes to 11.5 cm long; bracts 0.4-5 mm long, linear to lanceolate, bracteoles 0.3-0.9 mm, linear to ovate; pedicel 1-3 mm long, portion distal to articulation 1-2 mm, sometimes the upper bracts and bracteoles and pedicel with scattered capitate glandular hairs. Calyx 0.7-1.2 mm long, aestivation apert, divided nearly to base, the lobes 0.5-1 mm long, deltate; petals 2-3 x 0.5-1.1 (1.3) mm, oblong to ovate or deltate, apex acute to slightly acuminate, cream-colored or white or whitish green, glabrous, reflexed at anthesis; stamens spreading, antesepalous and antepetalous ones 1.7-2.1 and 1.3-1.5(1.9) mm long, respectively, the anthers 0.7-0.8 mm long, in dorsiventral view elliptic to ovate, in lateral view oblong; disk 0.3-0.5(0.7) mm tall, 0.2-0.4 mm thick, summit undulate and outer margin sulcate, yellow; pistil ca. 1.3 mm long, depressed-ovoid to subcylindrical overall, divided most of its length into very thickly subulate, apically connivent styles ca. 0.8 mm long, the stigmas obovate, slightly extrorse. Fruit 4-10 x 3-8 cm when dry, ellipsoid, obovoid or oblong, maturing yellow or orange, base of fruit basicrescent over distal portion of pedicel, the endocarp lacking a fibrous matrix but provided with spiny projections extending into the mesocarp. Leaflet venation : Fimbrial vein absent; secondary veins 12-20 pairs, usually darker than blade abaxially, usually straight and nearly perpendicular to midvein, spacing regular or sometimes decreasing toward base, angle increasing toward apex and base, insertion on midvein decurrent; intersecondaries ca. 1 per pair of secondaries and parallel to them, long and straight; intercostal tertiaries few, principally admedially branching parallel to secondaries but some irregular-reticulate, also some admedial tertiaries branching from intramarginal vein; quaternaries irregular-reticulate, FEVs highly branched, dendritic, tracheoid idioblasts absent; marginal secondary present; on abaxial side the midvein prominulous to prominent, secondaries flat; on adaxial side the midvein prominulous, secondary veins impressed to prominulous. Distribution. Broadly cultivated in lowland moist forest regions throughout the Neotropics. Ecology. Given this species' broad distribution, its known phenology is broken down by region. Central America: fruiting Aug-Sep; West Indies: flowering Mar-May, fruiting Nov-Jul; NW South America W of the Andes: flowering and fruiting Dec; Amazonia: flowering Aug-Oct, fruiting Aug-Mar; extra-Amazonian Brazil: flowering Oct-Apr, fruiting Nov; Venezuela: flowering Apr; Guianas: flowering May. There are reports in the literature that the fruits are dispersed by two species of large fruit-eating bats in the genus Artibeus ( Lobova et al. 2009 ). Common names. Brazil, Rio de Janeiro: caja manga ( Angeli 704 , NY); Dominican Republic: manzana de oro ( Zanoni & Mejia 16387 , NY); Ecuador, Napo: mauca (Yacu Indians, Irvine 653 , F); Guadeloupe: pomme cythere ( Pere Duss 3760 , NY, pro parte); Guyana: golden apple ( Omawale & Persaud 94 , NY); Jamaica: Jew plum ( Howard & Proctor 13531 , A) Nicaragua, Rio San Juan: jocote yuplon ( Sandino 3599 , NY); Panama, Panama : mangoteen ( Miller & Merello 230 , NY); Peru, Loreto: tapiriba ( Martin & Plowman 1781 , ECON), San Martin : taperiba ( Scolnik 1193 , NY), kapiniwa ( Berlin 870 , NY); Puerto Rico: ambarella, jobo de la India (Little 14914, NY); Venezuela, Delta Amacuro: jobo de los indios (Wurdack 315, NY). Economic botany. Spondias dulcis (often referred to as Spondias cytherea in the literature and on herbarium specimens) has been in cultivation for so long that its native range in Asia is difficult to determine. It was introduced to Jamaica from the South Pacific in the 18th Century ( Popenoe 1948 ), and it is planted in home gardens throughout the humid neotropics. In the American tropics, the only significant use of the species is for its juice and as a flavoring for ice creams and sorbets, although it is used to flavor yogurts in the Caribbean; it reportedly has a high Vitamin C content ( Lim 2012 ). Selected specimens examined. BELIZE. Toledo District, Temash River, ca. 11 km W of Caribbean Sea and ca. 3.5 km N of Belize/Guatemala border, ca. 15.949536°N , 89.033408°W , elev. 1 m, 8 Jun 1996, Atha & Romero 1372 (NY). BRAZIL. Acre : Mun. Tarauaca , Tarauaca town, 8.2°S , 70.8°W , 25 Sep 1994, Daly et al. 8361 (NY, HUFAC); Rio de Janeiro : Parque da Cidade de Gavea , 10 Aug 1986, C. Angeli 704 (GUA, NY). COLOMBIA . Amazonas : Leticia, 19 Sep. 1966, ForeroGonzalez 582 (NY). DOMINICAN REPUBLIC . Prov. Cristobal : at CESDA property, just outside of city of San Cristobal , 27 Jul 1981, Zanoni 15549 (NY). ECUADOR . Esmeraldas : Quininde , Bilsa Biological Station, Mache Mountains, 35 km W of Quininde , 5 km W of Santa Isabel, 2 lotes north of reserve, 400-600 m, 0°21'N , 79°44'W , 7 Dec. 1994, J. Clark 372 (NY); Napo : San Jose de Payamino, 40 km W of Coca, 0°30'S , 77°20'W , elev. 300-600 m, 20 Jan 1984, Irvine 653 (F). FRENCH GUIANA : Commune de Remire , Ile de Cayenne, 4°52S , 52°16'W , 25 Jul 1992 Wittingthon 44 (NY). GRENADA : St. George, Annandale Falls, 12°05'N , 61°43'W , 11 June 2001, Hawthorne et al. 459 (FHO, NY). GUADELOUPE : Grande-Terre, Grands-Fonds, Sainte-Anne, 12 Jul 1982, Barrier 3712 (NY). GUYANA : Diamond, east bank of Demerara River, 28 May 1970, Omawale & Persaud 94 (NY). JAMAICA . St. Anne Parish : grounds of Windsor Hotel (cult.), near St. Anne's Bay, 20-31 Dec 1953, Howard & Proctor 13531 (A). NICARAGUA . Rio San Juan, San Carlos, house S of cemetery (cult.), 16 Sep 1982, Sandino 3599 (NY). PANAMA . Panama : Barro Colorado Monument, Frijoles train stop, 9°10'28"N , 79°47'48"W , elev. 37 m, 25 Aug 2001, Miller & Merello 230 (NY). PERU . Amazonas : Huampami, Rio Cenepa, village, elev. 800 ft., 10 Feb. 1973, B. Berlin 870 (NY). PUERTO RICO : Mun. Isabela, Arenales Altos, along Hwy. 112, 3.4 miles NE of junction with Hwy. 444, 18°25'30"N , 67°02'W , 8 Nov 1993 , Nee 44157 (NY). TRINIDAD : Campus, University of the West Indies, 28 Jun 1916, Nevling 289 (A). VENEZUELA . Delta Amacuro : Rio Grande between Curiapo and Pta. Cangrejo, 10 Apr 1955, Wurdack 315 (NY). Conservation status. Considering that this taxon is native to Asia/Oceania and rather widely cultivated in tropical America, it can be considered of Least Concern, at least for the Neotropics. Discussion. According to Smith (1985) , the type specimen was made from a plant cultivated in Mauritius but grown from seed brought by Commerson from Tahiti in 1768. The earliest effective publication of Spondias dulcis is by Parkinson (J.voy. South Seas, 1773), a botanical artist who accompanied Banks and Solander in Captain Cooks's first expedition to the Southern oceans. A later publication of this name by G. Forster (Pl. esc. 33. 1786) is considered an isonym as it is based on the same type as Parkinson's name. Spondias cytherea Sonn. was once considered the earliest valid name of this species, on the basis that the names in Parkinson's publication were considered invalidly published ( Airy Shaw and Forman 1967 ). A more recent examination of Parkinson's work has shown Spondias dulcis to be a valid name and therefore to have priority over Spondias cytherea ( Fosberg 1960 ). Following this argument, Smith (1985) lectotypified Spondias dulcis with a collection considered the voucher of Parkinson's illustration of this species (and therefore a typotype). In 1869, Marchand subsumed Spondias acida , Spondias amara , and Spondias pinnata under Spondias dulcis as varieties. Here Spondias acida and Spondias pinnata are maintained as species, and Spondias amara is considered a synonym of S. pinnata. Spondias dulcis sensu Blanco is attributable to Spondias purpurea .