Corresponding author: Geoffrey A. Levin (
Academic editor: Lyubomir Penev
A synopsis of the New World species of
The genus
Although no formal groups below the sectional level have been recognized among New World members of section
The study was based on the examination of specimens from the following herbaria: A, ARIZ, BM, CAS, CM, CR, DAV, F, G, G-DC, GH, GOET, HAJB, ILLS, K, LL, MEXU, MICH, MO, NY, P, SD, TEX, U, UC, UCR, and US. All cited specimens were seen by the author unless otherwise indicated. Identifications were made by comparison with the original descriptions and, when available, with the type material or photographs of types. Details of the flowers and fruits were examined under a stereoscopic microscope. All descriptions and data on flowering times, habitats, and distribution are based on the herbarium material examined. Countries in the Selected Specimens sections are listed alphabetically.
1 | Fruiting pedicels (8–)10–15 mm long; Costa Rica | 2. |
– | Fruiting pedicels 4–10 mm long; Mexico and West Indies | |
2 | Staminate pedicels 7–14 mm long; drupes 12–15 mm long; Mexico | 3. |
– | Staminate pedicels 3–6 mm long; drupes 9–10(–13) mm long; West Indies | [1. |
3 | Stigmas sessile; Hispaniola, Puerto Rico, U.S. Virgin Islands | 1a. |
– | Stigmas borne on style 1 mm long; Cuba, Jamaica, Lesser Antilles | 1b. |
[Haiti.] Île de Tortue, s.d.,
Haiti, Dominican Republic, Puerto Rico, and U.S. Virgin Islands.
Forests from sea level to 1000 m, primarily on calcareous soils, but in Puerto Rico also on serpentine-derived soils.
Flowering primarily January to April, rarely as early as November. Fruiting February to August.
Least concern.
Cuba, occ., 1863 (fr),
Cuba, Jamaica, and the Lesser Antilles (Antigua, Guadeloupe, Martinique, Barbados).
Forests on limestone and schist, from sea level to 1100 m.
Flowering October to March. Fruiting December to July.
Least concern.
The plants I include in
The only character that seems consistently to differentiate
Differs from the other New World
Costa Rica. Puntarenas: Cove at NE base of peninsula, Punta Quepos (3 km S of Puerto Quepos),
Trees 6–20 m, to 35 cm dbh; bark with longitudinal fissures; branches brown when young, becoming gray, glabrous or sparsely minutely puberulent with spreading hairs. Leaves: stipules 0.5 × 0.5 mm, deltate, puberulent; petiole 3–10 × 0.3–1 mm, glabrous; blade elliptic to lanceolate, straight or somewhat curved, 4–12 × 1.5–4.5 cm, base asymmetrical, acute, margins entire or minutely crenulate-serrulate, often undulate, apex attenuate, surfaces glabrous, 2° veins 6–9/side. Inflorescences axillary fascicles; staminate 25–40-flowered, bracts 0.5 mm, puberulent, pedicels 5–8 × 0.2 mm, glabrous; pistillate (known only in fruit) 2–6-flowered, bracts 0.25 × 0.25 mm, deltate, puberulent, pedicels (8–)10–15 × 0.4–0.8 mm, glabrous. Staminate flowers: sepals 5–6, narrowly triangular to narrowly lanceolate, 1 × 0.3 mm, spreading and slightly incurved at apex, apex acute, margins ciliate, abaxial surface glabrous except puberulent at apex, adaxial surface densely to sparsely puberulent; stamens 5(–6), irregularly alternate and opposite sepals, filaments 1.5–2 mm × 0.1 mm, glabrous, anthers 0.4–0.5 × 0.4–0.5 mm, glabrous, latrorse; disc lobed between stamens, densely puberulent. Pistillate flowers unknown, but remnant sepals (below fruits) ovate-elliptic, 1.5 × 0.6 mm, apex acute and slightly incurved, abaxial surface glabrous, adaxial surface densely puberulent; disc annular, densely puberulent; ovary unknown; style becoming subapical during fruit development, 0.5 mm; stigma subreniform, 0.5 × 1 mm, glabrous. Drupes (immature) green, 1-carpellate, ovoid-globose, 12–18 × 7–10 × 6–8 mm, apex strongly asymmetrical, sparsely to densely puberulent with very short hairs (0.1 mm). Seed 1.
The specific epithet refers to the strongly asymmetrical drupes, which are unique among Central American
Known only from Costa Rica, where it is found from the north central part of the country to the central west coast. It may also be expected in extreme southern Nicaragua.
Forests at elevations from sea level to 750 m.
Flowering January (possibly longer, but only a single flowering specimen known). Fruiting March to June (possibly longer as only immature fruits are known).
Probably of Least Concern. The range of
All the pistillate specimens studied have immature fruits, so their full size and color at maturity are unknown. The label on
Mexico. Sinaloa: Capadero, Sierra Tacuichamona, rocky canyon under basaltic rim, 3500 ft., 13 Feb 1940 (fr),
Western Mexico, in the Sierra Madre Occidental from near 27° N in Chihuahua and Sonora to about 19° N in Colima.
Tropical deciduous forests at about 100–1100m.
Flowering December–February. Fruiting December–June.
Least Concern.
When
Trees 8–25 m, often with multiple trunks from near base, to 20–100 cm dbh; bark scaled and with longitudinal fissures; branches brown when young, becoming gray, minutely puberulent with spreading hairs, becoming glabrous. Leaves: stipules 0.5–0.6 × 0.7–1 mm, deltate, puberulent; petiole 6–12 × 0.7–1 mm, puberulent with spreading hairs or glabrous; blade elliptic to lanceolate, straight or somewhat curved, 4–15 × 1.5–4 cm, base asymmetrical, acute to narrowly obtuse, margins subentire to crenulate-serrulate, often undulate, apex attenuate, surfaces glabrous or very sparsely pubescent with appressed hairs especially near base, 2° veins 6–9/side. Inflorescences axillary fascicles; staminate 20–40-flowered, bracts 0.5 × 0.5 mm, deltate, puberulent, pedicels 7–14 × 0.2 mm, glabrous; pistillate 1–6-flowered, bracts 0.5 x0.5 mm, deltate, puberulent, pedicels 3–10 × 0.4–0.5 mm, puberulent when young, becoming glabrous. Staminate flowers: sepals 5(–6), linear to narrowly triangular, 1.2 × 0.4 mm, spreading and slightly incurved at apex, apex bluntly acute, margins ciliate, abaxial surface glabrous except puberulent at apex, adaxial surface puberulent; stamens 5(–6), mostly opposite sepals, filaments 1.6–2.2 mm × 0.1 mm, glabrous, anthers 0.8–1 × 0.5–0.6 mm, glabrous, latrorse; disc lobed between stamens, densely puberulent. Pistillate flowers: sepals 5, narrowly triangular to linear, 1–1.2 × 0.3–0.4 mm, spreading, entire, apex bluntly acute, abaxial surface glabrous to sparsely puberulent but densely puberulent at apex, adaxial surface densely puberulent; disc annular, densely puberulent; ovary densely puberulent; style absent; stigma apical at anthesis, becoming subapical during fruit development, subreniform, 0.8 × 1.2 mm, glabrous. Drupes (immature) green, 1-carpellate, obovoid, 12–15 × 7–9 × 6–8 mm, apex strongly asymmetrical, densely puberulent with very short hairs (0.1 mm). Seed 1.
The mature fruits are described as white (
I thank Jochen Heinrichs for providing high-resolution scans of specimens at GOET, and the staff of the other herbaria listed for providing access to the specimens in their care. Lynn Gillespie and three anonymous reviewers provided helpful comments on the manuscript. Figure 1 was prepared by Yevonn Wilson-Ramsey.
Abbott, W.L. 2571, 2932 (alba var. alba)
Acevedo-Rodríguez, P. 4710, 4781, 7724 (alba var. alba)
Alvarez, A. 28778, 45576 (alba var. latifolia)
Areces, A. 25812 (alba var. latifolia)
Bello, E. 35, 58 (asymmetricarpa)
Bisse, J. 16899, 22625, 26478, 34469, 46192, 47203, 47259, 53290 (alba var. latifolia)
Britton, E.G. 5154 (alba var. alba)
Britton, N.L. 2457, 2748, 4128, 4208 (alba var. alba); 5459 (alba var. latifolia); 8104, 9676 (alba var. alba); 15305 (alba var. latifolia)
Bye, R.A. 3401, 3557, 3558, 3958, 6066, 9707, 12847 (gentryi)
Duke, J.A. 7350 (alba var. alba)
Duss, A. 50, 80 (alba var. latifolia); 3628 (alba var. latifolia and Drypetes glauca Vahl); 4142 (alba var. latifolia)
Eggers, H.F.A. 2612 (alba var. alba)
Ekman, E.L. 4146 (alba var. latifolia); 4179 (alba var. alba); 4543, 4568, 4838, 5006 (alba var. latifolia); 5528 (alba var. alba); 6694, 8544, 9764, 9913 (alba var. latifolia); 12160 (alba var. alba); 12462, 18213 (alba var. latifolia); H.3512, H.3870, H.4178, H.4179, H.5134, H.5528, H.6061, H.9614, H.12160 (alba var. alba)
Felger, R.S. 94-56, 97-118 (gentryi)
Gentry, A.H. 50716 (alba var. alba)
Gentry, H.S. 3618, 5597 (gentryi)
Grayum, M.H. 6614 (asymmetricarpa)
Haenke, T. 1605 (gentryi)
Hammel, B.E. 17046 (asymmetricarpa)
Harmon, W.E. (asymmetricarpa)
Harris, W. 7098 (alba var. latifolia)
Herrera, G. 1693 (asymmetricarpa)
Holdridge, L.R. 149 (alba var. alba)
Jack, J.G. 7809 (alba var. latifolia)
Jiménez Almonte, J. de J. 1981, 4360, 5058 (alba var. alba)
León, Bro. 6081, 11800 (alba var. latifolia)
Leonard, E.C. 11589, 14257, 14268, 15686 (alba var. alba)
Lesueur, D.H. 1419 (gentryi)
Levin, G.A. 1973, 1975, 2001, 2014, 2015 (gentryi)
Liogier, A.H. 1017, 1261, 1275, 2997, 5719 (alba var. latifolia); 10466, 10714, 10779, 14325, 15661, 20858, 21293, 21600, 22628, 24490, 24537, 25846, 26463, 26835 (alba var. alba)
Little, E.L., Jr. 13297, 13366 (alba var. alba)
Luna, A. 15 (alba var. latifolia)
McVaugh, R. 1647, 25404 (gentryi)
Pérez J., L.A. 413 (gentryi)
Proctor, G.R. 9956 (alba var. latifolia)
Ricksecker, A.E. 462 (alba var. alba)
Sagra, R. de la 607 (alba var. latifolia)
Sanders, A.C. 1096 (gentryi)
Schiffino, J. 15 (alba var. alba)
Shafer, J.A. 3693, 4403 (alba var. latifolia)
Sintenis, P.E.E. 1502 (alba var. alba)
Stehlé, H. 416 (alba var. latifolia)
Valeur, E.J. 837 (alba var. alba)
Van Devender, T.R. 93-1381 (gentryi)
Wilson, P. 9287 (alba var. latifolia)
Wright, C. 498, 501 (alba var. alba and Drypetes lateriflora (Sw.) Krug & Urb.); 593 (alba var. latifolia and D. lateriflora); 1112 (alba var. latifolia); 1927 (alba var. latifolia and D. lateriflora); 1928, 1929, 1929, 593b (alba var. latifolia)
Zanoni, T.A. 36401 (alba var. alba)