Spondias globosa J. D. Mitch. & Daly, 2015

Mitchell, John D. & Daly, Douglas C., 2015, A revision of Spondias L. (Anacardiaceae) in the Neotropics, PhytoKeys 55, pp. 1-92 : 23-28

Spondias globosa J. D. Mitch. & Daly sp. nov. Figs 1, 2, 10, 13, 15, 16

Diagnosis.

Canopy or emergent tree to 40 m tall, inner bark red with narrow white striations; similar to Spondias mombin because of the similar indumentum, the inflorescences highly branched, disk short and thick, and fruits of similar size; Spondias globosa differs by the outer bark lacking spinose projections (vs. corky, tubercular, or spinose projections), intersecondary veins parallel to secondaries and strong, often reaching intramarginal vein (vs. intersecondaries reticulating and weak, not usually reaching intramarginal vein), intercostal tertiaries arising at or near intramarginal vein (vs. intercostal tertiaries primarily irregular-reticulate), fruits globose to perdepressed-ovoid, rarely very slightly oblong of obovoid (vs. oblong or less often ellipsoid or slightly oblong-ovoid).

Type.

BRAZIL. Acre: Mun. Santa Rosa, Alto Rio Purus, left bank, Seringal Mamuriá, ca. 9°05'05"S, 69°59'07"W, 25 Mar 1999, D. C. Daly, H. Kuchmeister, D. Gomes da Silva, L. Lima & E. Consuelo 10039 (holotype: HUFAC!; isotypes: AAU!, MO!, NY!).

Description.

Hermaphroditic trees, reproductive height 8-40 m. Trunk 10-105 cm diam; outer bark (light) gray to brown, usually thin, usually with many long, broad, shallow, wavy fissures, sometimes rough but lacking spinose projections, also with small white lenticels, shed in flat, narrow, regular plates; inner bark red (less often orange) with narrow white (less often beige) striations, or red-and-white striate, thick. Trichomes of two types: straight or slightly curved erect white hairs to 0.15 mm long; and short, usually straight, erect, whitish bristles to 0.05 mm long. Leaves (1) 3-6 (7)-jugate,13-40 cm long; petiole 3.8-8 cm long, petiole and rachis glabrous or with sparse bristles, flanks of petiolules with dense longer hairs and sparse bristles; lateral petiolules 3-10 mm long, the terminal ones 12-32 mm long, petiolules with slightly curved white hairs; the basal leaflets 3.2-8.5 × 1.8-4.3 cm, (broadly) elliptic to (broadly) ovate, sometimes broadly obovate or almost rotund, other laterals (3.2) 6.2-11.5 (14.5) × 2-4.9 (5.3) cm, sometimes slightly obliquely ovate or lanceolate but more often strongly asymmetrical and the acroscopic side semi-ovate to semi-lanceolate and the basiscopic side semi-(oblong-)elliptic, sometimes broadly so; terminal leaflet 3.7-9.5 × 1.8-5 cm, (broadly) elliptic, obovate, or oblanceolate; leaflet apex usually abruptly and narrowly long-acuminate or sometimes broadly short-acuminate, the acumen (3) 6-18 cm long, often the apex tip mucronate; lateral lamina medially and basally slightly to strongly asymmetrical, the acroscopic side truncate to rounded or rarely obtuse, the basiscopic side cuneate to attenuate; basal insertion often asymmetrical, both sides abruptly decurrent; margin entire, sometimes slightly revolute, sparsely ciliate with longer hairs; leaflets chartaceous to subcoriaceous, sometimes glossy adaxially. Inflorescences (sub)terminal, produced with leaf flush, 15-26 cm long, 3-5 mm diam near base, broadly branched, secondary axes 1.8-14.5 cm long, these axes with dense to sparse bristles, higher-order axes with sparse bristles, also with sparse longer hairs; bracts on primary axes ca. 4 mm long, lanceolate, those on secondary axes 0.6-1.6 mm long, lanceolate to deltate, with dense bristles on both surfaces, bracteoles 0.25-0.5 mm long; pedicel 1.2-3 mm long overall, portion distal to articulation 0.5-1.3 mm long, pedicel and both sides of calyx with sparse bristles (denser toward base). Calyx 0.8-1 mm long overall, aestivation apert, the lobes 0.4-0.7 mm long, deltate to narrowly ovate, calyx with pubescence as on pedicel, the margin often ciliate; petals 2.4-3.3 × 1.2-1.5 mm, lanceolate, apex slightly acuminate, whitish to yellowish or cream, abaxial surface glabrous, reflexed at anthesis; stamens spreading, the antesepalous and antepetalous ones 2-2.3 and 1.4-1.6 mm long, respectively, the anthers 0.8-1.1 mm long, in dorsiventral view oblong, in lateral view oblong to el liptic; disk 0.3-0.5 mm tall, 0.2-0.3 mm thick, summit markedly undulate and outer margin deeply sulcate; pistil (1) 1.5-1.8 mm long, depressed-ovoid overall, divided ca. 2/3 its length into broadly subulate, apically slightly divergent styles 0.8-1.2 mm long, extrorse, stigmas vertically elliptic. Fruits 1.6-3 × 2.2-3 cm diam (to 4 cm diam fresh), usually (depressed-) globose, rarely very slightly oblong or obovoid (then the apex obtuse), maturing yellow, surface smooth, dull. Seedlings (Pennington et al. 17244, NY): cotyledons 2.1-2.4 cm long, with several parallel veins; first eophylls opposite, trifoliolate, petiolules with sparse curved hairs, the leaflets ovate, margin glabrous, sparsely toothed, the teeth concave-convex.

Leaflet venation: Fimbrial vein absent; secondary veins in 9-15 pairs, straight to slightly arcuate, the spacing decreasing toward apex and base, the angle decreasing toward the apex and increasing toward base, insertion on midvein decurrent; intersecondaries occasional, parallel to secondaries and almost reaching the intramarginal vein; intercostal tertiaries few, most of them arising from near the intramarginal vein and forming strong composite admedials parallel to secondaries, with some irregular reticulation; quaternaries irregular-reticulate and freely admedially ramified; areolation usually at quaternary rank, FEVs 5+- branched, dendritic, tracheoid idioblasts absent; marginal ultimate venation usually looped (sometimes incompletely); on abaxial side all veins narrowly prominent or sometimes the secondaries and tertaries prominulous, occasionally discolorous; on adaxial side all veins narrowly prominulous to almost flat or occasionally all but the midvein slightly impressed; on both surfaces the midvein with scattered longer hairs and bristles near the base and (sub)glabrous distally, sometimes glabrescent.

Distribution.

Spondias globosa is a western Amazon element, apparently disjunct to Zulia and Barinas in western Venezuela.

Ecology.

This is very much a lowland taxon, ranging only between 100-350(500) m elevation. It is most often found in formations such as floodplain forests or tahuampa forest on poorly drained, periodically or seasonally inundated soils, although it has been reported from a range of soils including not just black alluvial soils but also oxisols, lateritic soils, and red and yellow clay soils. Apart from flooded formations it is found in primary forests in well-drained soils, including on undulating or hilly terrain. Occasionally it grows in secondary forest, bamboo-dominated forest, or rarely pasture or shrubby disturbed vegetation.

In SW Amazonia, this species is known to flower in Sep-Nov and fruit Oct-May, but in NW Amazonia the collection data indicate that it can be found flowering and fruiting all year.

The yellow-footed tortoise, Geochelone (Chelonoidis) denticulata , has been observed dispersing the fruits of Spondias globosa (as Spondias venulosa ) in Amazonian Peru and Colombia ( Rodríguez-Bayona and Rylander 1985 and Stevenson et al. 2007, respectively). These fruits comprise an important food source in times of scarcity for primates such as the woolly monkey ( Stevenson 2005).

Common names.

Brazil, Acre: cajá (Cid Ferreira & Nelson 3066, NY), taperibá (Daly et al. 10039, NY), taperebá (Silveira et al. 1622, NY); Venezuela: jobo (Steyer mark 102015, NY); Ecuador: aurumuyo (Quichua, Zuleta 191, NY) azua muyo (Quichua, Moya & Reyes 146, NY); mientuhue (mientuhuem for fruit)(Huaorani, Miller et al. 703, NY); mamantunim (Shuar, Jua (RBAE) 69, NY); mïyëtowëmo (Wao, Ríos 576, NY); mientohuemo (Huaorani, Aulestia & Gonti 1769, NY); mientuhueno (Aulestia et al. 3020, NY); ovu muyo (Huaorani, Aulestia et al. 402, NY); mijentuemo (Huaorani, Dik & Andi 906, NY); Peru, Loreto: ubos (Martin & Lau-Cam 1252, ECON), huvos (Torres 88, GH), hubus (Schunke 250, A), hubos (Torres 350, ECON), ubos colorado (Chota 5, NY).

Economic botany.

Fruit edible (Schunke 250, A); bark cooked with water taken for diarrhea (Plowman et al. 7257, ECON); fruit pulp used to make a fruit juice (Daly et al. 10039, NY); for chronic diarrhea, make tea from 1 kg of finely chopped bark and drink twice daily, or use liquid concoction as vaginal douche to treat flor blanca ('yeast infections?'), or apply to infected wounds (Chota 5, NY); fruits are edible, much appreciated and frequently sold in Iquitos market (Peters & Hammond 164, NY); branches and trunk used as firewood (Jua (RBAE) 69, NY); eaten by a number of animals (Miller et al. 703, NY), eaten by game animals (Lizarralde ML307, NY). In Acre, Brazil, Kainer and Duryea (1994) observed preparation of a type of tucupi sauce that combines hot peppers with the juice of Spondias globosa fruits.

Etymology.

The specific epithet refers to the usually globose fruits characteristic of this taxon.

Selected specimens examined.

BOLIVIA. Beni: Prov. Ballivián, Estación Biológica Beni, 56 km E of Río Maniqui on road to Trinidad, then 18 km NNE to Estancia 07, then 6 hrs to Río Maniquicito, 250 m, 14°44'S, 66°20'W, 6 Nov 1985, Solomon 14593 (NY); Pando: Manuripi, vicinity of La Conquista, elev. 160 m, 19L FH95, 30 Jan 1983, Fernández-Casas & Susanna 8566 (MO, NY); Santa Cruz: Prov. Ichilo, E side of Río Tapacani at junction with Río Surutu, 0.5 km upstream and S from bridge over Río Yapacani at Villa Tapacani, 17°24'S, 63°50'W, 30 Oct 1990, M. Nee 39607 (MO, NY, TEX). BRAZIL. Acre: Mun. Sena Madureira, basin of Rio Purus, Rio Iaco, right bank, Nova Olinda, between Igarapé Santo Antônio and Ig. Boa Esperança, 10°07'S, 69°13'W, 22 Oct 1993, Daly et al. 7836 (HUFAC, NY, TEX); Amazonas [erroneously sited in Acre state on label]: Mun. Boca do Acre trail from W bank of Rio Iaco to Rio Purus, 3 km above confluence, 5 Oct 1968, Prance et al. 7873 (GH, MG, NY, R). COLOMBIA. Amazonas: Aduche, Asentamiento Muinane, south bank of río Caquetá, 0°41'30"S, 72°06'00"W, 11 May 1999, Arévalo & Reyes 57 (NY); Meta: Parque Nacional Natural Tinigua, Serranía Chamusa, Centro de Investigaciones Primatológicas La Macarena, 7 Mar 1990, Stevenson 109 (COL). ECUADOR. Morona - Santiago: Centro Shuar-Yukutais, 3°30'S, 78°10"W, 18 Apr 1989, Bennett & Gómez A. 3711 (NY); Napo: Orellana, Parque nacional Yasuní, km 46-52 Maxus road under construction, elev. 250 m, 00°47'S, 76°30'W, 1-11 Sep 1993, Aulestia et al. 402 (NY); Pastaza: “Moretecocha” oil well of ARCO, río Landayacu, 75 km E of Puyo, elev. 580 m, 1°34'S, 77°25'W, 4 Dec 1990, Gudiño 1158 (NY); Sucumbios: Cuyabeno, Parroquia Tarapoa, Siona community of Sototsiaya, 50 min. downstream from Poza Honda on Río Aguarico, 00°14'27"S, 76°26'15"W, elev. 230 m, 25 Feb 2005, Miranda & Moya 446 (MO). PERU. Amazonas: Río Santiago, behind community of Caterpiza, elev. 200 m, 4 Sep 1979, Huashikat 392 (NY); Huánuco: Prov. Puerto Inca, Dtto. Yuyapichis, Unidad Modelo de Manejo y Producción Forestal Dantas, 9°40'S, 75°02'W, 16-30 Nov 1989, Kröll 694 (NY); Loreto: Río Nanay, Puerto Almendras, ca. 20 km WSW of Iquitos, ca. 3°46'S, 73°20'W, 15 Mar 1989, Chota 5 (NY); Madre de Dios: Prov. Tambopata, Zona Reservada Tambopata-Candamo, along trails of Explorer’s Inn, 12°49'S 69°18'W, 22 Apr 1991, Phillips & Chávez 636 (NY); Ucayali: Prov. Coronel Portillo, Carretera Marginal, 22 km S of km 86 on PucallpaTingo Maria Highway, 75°00'W, 8°41'S, 11 Feb. 1981, Gentry et al. 31215 (NY). VENEZUELA. Barinas: Reserva Forestal Caparo, 16-18 km SE of Campamento Cachicamo, E of El Cantón, elev. 100 m, 9 Apr 1968, Steyermark et al. 102015 (NY); Zulia: along Quebrada Perayra, tributary of Río Tokuku (Tocucu), SW of Misión de Los Angeles de Tokuku, SW of Machiques, 29 Aug 1967, Steyermark 99828 (NY).

Conservation status.

This taxon is widespread in Amazonia and can be considered of Least Concern except in Zulia, Venezuela (Maracaibo watershed), where very little lowland forest remains and where it has been collected only once.

Discussion.

Although Spondias globosa is geographically sympatric with Spondias mombin in many localities,the two appear to have undergone niche partitioning: in interviews with forest residents in the middle Ucayali and upper Purus rivers, they readily recognized the two as distinct taxa long before botanists came to the same conclusion, pointing out not only differences in the bark and fruits but also that Spondias globosa tends to keep to the floodplains (vs. terra firme) and flowers and fruits later in any given locality. In the middle Ucayali the prevailing common name for Spondias globosa is "uvos colorado," referring to its mostly red (versus usually pale pink) inner bark.

Morphologically, the two can be distinguished by Spondias globosa lacking corky tubercular or spinose projections, the inner bark usually (pale) red-and-white striate (vs. inner bark pale red to pink to orange, sometimes striate with beige), the leaves 3-5 (-7)-jugate (vs. 3-7 (-12)-jugate), the leaflets with composite admedial tertiaries arising at or near the intramarginal vein (vs. tertiary veins primarily irregular-reticulate, some admedial branching), fimbrial vein absent and the marginal ultimate venation incompletely looped (vs. fimbrial vein present), the flower pedicel 1.2-3 mm long (vs. 2-4.5 mm long), the fruit usually (depressed-)globose, rarely very slightly oblong or obovoid (vs. oblong or less often ellipsoid or slightly oblong-ovoid), and occurring in W Amazonia plus Zulia and Barinas in Venezuela (vs. central Mexico S to SE Brazil and widely cultivated in the moist tropics).

Table 4 summarizes the morphological characters that separate the two species rather consistently. The existence of possible hybrids between Spondias mombin and Spondias globosa is discussed in the section on Hybridization and Intermediates in the Introduction; some examples are Grández & Jaramillo 2042 (MO, NY), Spichiger & Encarnación 1095 (MO), and Vásquez et al. 4873 (MO, NY).