Academic editor: Dimitrios Koureas
In the course of the present revision, 353 herbarium collections, comprising approximately 540
Twenty-eight validly published names (corresponding to twenty-one typified taxa at the species level and below) are available in
During the 20th century, most taxonomic treatment of
All species of
In Cuban species such as
Leaf texture is usually coriaceous and occasionally membranaceous (individuals of
Leaf indument usually comprises at least few scattered trichomes at midribs, but leaves and individuals being completely glabrous are occasionally found in
In
Female and male flowers differ only with respect to formation of gynoecium and androecium, but not to perianth. Male flowers have very well developed stamens with (in outline) reniform functional anthers as demonstrated by the presence of pollen (visible with the stereo microscope at 60x magnification). Filaments are flattened, which is another unusual character within
Female flowers have smaller, shrivelled anthers lacking pollen (verified in SEM in the course of the present study) on short filaments. The ovary is globose and exhibits the coenocarpous-syncarpous architecture usually developed in
Sex distribution is dioecious, which is a very unusual trait within
Flowers are sometimes fragrant (as noted on herbarium specimens) indicating zoophily like in many other
Most
Delimitation of
Species of
We adopted a rather broad morphological species concept for
Different from many other taxa of woody borages, species of
Vegetative traits help to distinguish between species that occur sympatrically, but the degree of homoplasy is high when considered over the entire geographic range. Individual specimens of, for example,
In this revision, we provide comprehensive nomenclatural data about all names published for species of
Shrubs or trees, rarely lianas, thorny, short shoot galls present or absent; bark striate, with superficial grooves, longitudinally fissured, grey light to dark brown, with lenticels; indument absent at maturity or puberulent, trichomes simple, sometimes glandular, rarely branched (
Shrubs up to 4.0 m tall, rarely trees 4.0–7.0 m tall, galls absent; twig indument absent or sericeous, glabrescent at maturity, trichomes simple; bark grey whitish to grey dark or brown, sometimes peeling; thorns 0.5–0.8 cm long, slender, acute, simple, numerous, axillary, alternate, glabrous through variously sericeous. Leaves fasciculate; petiole 0.05–0.2 cm long, slender, hirsute; blade 0.5–1.5 cm long, 0.2–0.3 cm wide, elliptic to obovate, widely obovate or circular (particularly distal immature leaves), coriaceous, primary vein prominent, with scattered trichomes, secondary veins 3–6, tertiary veins arcuate or reticulate; base cuneate, rarely rounded; apex rounded or obcordate, occasionally retuse, emarginated or cleft; adaxial surface hirsute to pubescent, with simple trichomes and cystolith-like structures in subsidiary cells, abaxial surface bright, variously hirsute to glabrous. Inflorescence axillary, flowers in clusters of 2 or rarely 3 or solitary, pedicels up to 0.02-0.03 cm long. Calyx 0.40–0.50 cm long, 0.45–0.50 cm wide, coriaceous, glabrous or with few scattered trichomes outside, sometimes hirsute, glabrous inside with a tuft of trichomes at the distal part of lobes, lobes 0.35–0.40 cm long, 0.40–0.43 cm wide, ovate to broadly ovate, apex acute, margin strigose. Corolla 0.40–0.50 cm long, white, yellowish or pale yellow, rarely green (as in
Coastal habitats and scrublands across Caribbean islands between Haiti in the West and Guadeloupe in the East, with high abundance in the Dominican Republic, Puerto Rico and the Virgin Islands (symbol "⨀" in Fig.
Flowering throughout the year (Jan–Dec); fruiting Oct–Mar, May–Aug, but probably contiguously.
Traditionally,
Morphologically,
Representative specimens examined. — HAITI.
“bois d’ébène”, “ébénier noir”, “galle-galle”, “gratte-galle” in Haiti, “corazón de paloma” (Span., pigeon heart), “ébano” and “trejo” in Dominican Republic, “juso” in Puerto Rico and “greenheart ebony” in US Virgin Islands (
The trunk is employed to make posts for fences (
Shrubs or small trees up to 1.5–4.5 m tall, galls absent; twig indument puberulent, trichomes simple; bark greyish white to brown, longitudinally fissured, scaly, wood very brittle; thorns 0.3–0.8 cm long, slender, distally somewhat acute, simple, scattered, alternate, axillary or rarely terminal, sericeous. Leaves fasciculate, rarely alternate (
Across multiple islands of The Bahamas archipelago (Crooked Island, Great Exuma, Inagua, Long Island, Mayaguana, San Salvador) and putatively also in western Cuba (Pinar del Río, Havana) in scrub lands, rocky coppice hills and thicket edges, at sea level or slightly above (symbol "▲" in Fig.
Flowering Feb–Mar, Oct–Dec; fruiting Dec–Jan, Oct.
The 3 sterile collections from Cuba are tentatively placed under
unknown.
Shrubs or small trees 1.5–10.0 m tall, branches spreading, galls absent; indument glabrescent, slightly pubescent when immature, trichomes simple; bark greyish white, longitudinally fissured; thorns 0.5–0.7 cm long, slender, acute, simple, scattered, axillary, alternate, glabrescent. Leaves fasciculate; petiole 0.4–1.5 cm long, glabrescent; blade 1.9–3.4(–6.0) cm long, 1.2–3.0 cm wide, obovate, sometimes circular, coriaceous, primary veins prominent, lateral veins in pairs of 4–5, arching, tertiary veins absent; base cuneate; apex rounded or slightly emarginate; adaxial surface with cystolith-like structures in epidermal cells, mostly glabrous, but sometimes also with scattered, simple, bent trichomes, abaxial surface with scattered trichomes on the midrib, immature leaves sometimes barbellate. Inflorescence axillary and terminal, branching sympodial, branches slender, glabrescent, pedicel 0.20–0.40 cm long. Calyx 0.35–0.45 cm long, coriaceous, hirsute outside, glabrous inside, sometimes with a few clustered trichomes on distal part, lobes 0.30–0.35 cm long, 0.32–0.35 cm wide, obovate, apex rounded to slightly acute, margin strigose. Corolla 0.35–0.40 cm long, yellow through light orange, occasionally greenish (
Restricted to islands of the Lesser Antilles (Guadeloupe, Marie Galante, Montserrat, Martinique) and eastern Puerto Rico (symbol "●" in Fig.
Flowering Apr–May, Jul, Sep–Oct, Dec; fruiting Dec–Jan, May–Jun.
"bois vert" in Martinique (noted on
Shrubs 1.0–3.0 m tall or small trees up to 6.0 m tall, prickly, branches arching, short shoot galls occasionally present, 0.8–1.0 cm long; indument glabrescent, immature branches finely pubescent, trichomes simple; bark greyish white through grey brown, with longitudinal crevices; thorns 0.3–0.5 cm long, slender, acuminate, dichotomously branched, numerous, axillary, glabrescent. Leaves fasciculate, dark green; petiole nearly sessile or up to 0.2 cm long, glabrescent; blade 0.4–0.9 cm long, 0.3–0.5 cm wide, elliptic, sometimes obovate, coriaceous, primary veins pinnate, secondary veins 5–9, tertiary veins reticulate; base cuneate, rarely round; apex rounded, occasionally emarginated; adaxial surface with distinct cystoliths in epidermal cells, glabrous or sometimes covered with grey, simple, long trichomes (visible by naked eye) emerging from a cystolith-like structure, abaxial surface mostly glabrous, sometimes with scattered trichomes distally and on midrib. Inflorescence axillary or terminal, flowers usually in clusters of 2 or solitary, pedicel 0.02–0.2 cm long. Calyx 0.35–0.40 cm long, 0.30–0.40 cm wide, coriaceous, glabrous or with few scattered trichomes outside, sometimes hirsute, glabrous inside, strigose at tips, lobes 0.30–0.35 cm long, 0.40–0.45 cm wide, divided at the base, ovate to widely ovate, apex slightly acute. Corolla 0.35–0.40 cm long, pale yellow, membranaceous, glabrous on both surfaces, tube 0.20–0.25 cm long, funnel-shaped, lobes 0.30–0.35 cm long, ovate, slightly ciliate at tips. Male flower unknown; anthers of female flower 0.05–0.07 cm long, oblong, filaments 0.07–0.10 cm long, adnate to corolla tube for further 0.05–0.08 cm, pollen absent. Ovary of female flower globose, 0.25–0.32 cm long, stylodia 2, 0.25–0.35 cm long, filiform, glabrous, ovules present, stigmas cotyliform. Fruit 0.20–0.40 cm tall, 0.30–0.50 cm wide, globose; style accrescent, occasionally persistent, pyrene 0.30–0.40 cm tall, 0.23–0.33 cm wide, 0.13–0.20 cm deep, adaxial surface cutate.
Cuba and Jamaica (symbol "+" in Fig.
Flowering Mar, Jun–Jul; fruiting Mar, Jul–Sep, Dec–Jan.
With respect to leaf size and shape,
“bronce”, “carbonero”, “espuela de caballero” (Span. knight’s spur), sargento (
Shrubs up to 2.5 m tall or small trees up to 8.0 m tall, galls absent; indument variously sericeous through glabrescent, trichomes simple; bark grey light through grey dark or brown, superficial grooves present; thorns 0.7–1.0 cm long, slender, acute, simple or rarely branched, scattered, alternate, axillary or rarely terminal, glabrous. Leaves fasciculate, rarely opposite or alternate; petiole 0.3–0.8 cm long, slender, hirsute or sometimes glabrescent; blade 1.5–3.5(–6.6) cm long, 0.5–3.0(–3.5) cm wide, obovate to widely obovate, occasionally orbicular, membranaceous, primary veins prominent, hirsute, secondary veins 4–11, tertiary veins reticulate; base cuneate or rounded; apex rounded, obcordate, sometimes retuse, rarely cleft; adaxial surface brightly glabrescent, with very rare cystoliths-like structures in epidermal cells, ciliate at tips, trichomes emerging from a swelled cystolith cell giving the impression of undulate leaf margin, abaxial surface brightly rugose, with scattered trichomes. Inflorescence axillary or terminal, branches slender, hirsute to glabrescent, pedicel 0.30–0.35 cm long. Calyx 0.35–0.40 cm long, 0.30-0.45 cm wide, coriaceous, hirsute outside, glabrous inside with scattered trichomes at distal blade, lobes 0.30–0.35 cm long, 0.28–0.40 cm wide, divided from the base, obovate, apex rounded. Corolla 0.40–0.55 cm long, membranaceous, glabrous on both surfaces, tube 0.25–0.30 cm long, funnel-shaped, lobes 0.25–0.35 cm long, obovate, slightly cilliate distally. Male flower unknown; anthers of female flower 0.07–0.08 cm long, oblong, filaments 0.03–0.04 cm long, adnate to the corolla tube for further 0.03–0.04 cm, pollen absent. Ovary of female flower globose, 0.25–0.30 cm long, style divided right from the base, branches 0.18–0.25 cm long, stigma cotyliform. Fruit 0.50–0.60 cm tall, 0.50–0.60 cm wide, globose; pyrene 0.45–0.50 cm tall, 0.25–0.30 cm wide, 0.10–0.13 cm deep, abaxial surface cutate.
Jamaica, and presumably (with very few collections:
Flowering Mar, Jul, Sep, Nov; fruiting Mar, Jul.
Across Caribbean islands,
Olof P. Swartz’ names were not properly typified before the present revision (
"green ebony" and "bois vert" in Jamaica.
Lianas or shrubs up to 4.5 m tall or trees up to 10.0 m tall, branches arching, galls absent; indument almost glabrescent at maturity, hirsute when young, trichomes simple; bark whitish grey, grey brown or brown dark, with superficial crevices; thorns 0.6–1.5 cm long, robust, acute and slightly curved at tips, simple, scattered (also on older branches), alternate, axillary, hirsute. Leaves fasciculate; petiole 0.3–2.3 cm long, glabrescent; blade (2.0–)5.0–10.5(–13.0) cm long, 1.5–4.5(–6.0) cm wide, obovate, rarely elliptic, coriaceous, primary and secondary veins prominent, secondary veins 6–9, tertiary veins arching; base cuneate; apex acute, retuse, obcordate, sometimes round or cuspidate; adaxial surface bright, with cystolit-like structures in epidermal cells, glabrous, sometimes with scattered, bent trichomes, abaxial surface shiny, papillate, glabrous, sometime barbellate or rarely with trichomes apically. Inflorescence axillary and terminal, branching sympodial, branches slender, hirsute to glabrous, pedicel 0.20–0.50 cm long. Calyx 0.40–0.45 cm long, 0.45–0.49 cm wide, coriaceous, hirsute outside, glabrous inside, lobes 5 or occasionally 4, 0.35–0.40 cm long, 0.35–0.43 cm wide, very widely ovate, apex obtuse, margin strigose. Corolla 0.35–0.60 cm long, yellow, white, dull white or sometimes greenish (
Abundant in Mexico, but also present in adjacent and other countries in Central America (Belize, Costa Rica, Guatemala, Nicaragua), as well as western Cuba (symbol "⬡" in Fig.
Flowering Jan, Apr–Aug; fruiting Feb–Mar, May–Dec.
The species has a lianescent habit, which is unusual for
Still in the 60ies of the past century,
“palo dulce” in Mexico and “carey de costa” in Cuba.
Shrubs, short shoot galls occasionally present, 0.7–1.0 cm long; indument glabrous, trichomes simple; bark greyish white, slightly fissured; thorns 0.7–1.1 cm long, relatively robust, acute, simple or dichotomously branched, axillary or terminal, glabrous. Leaves fasciculate; petiole 0.4–1.0 cm long, robust, glabrous; blade 1.1–3.9 cm long, 0.7–1.5 cm wide, oblong-ovate, sometimes elliptic, coriaceous, primary veins very prominent, secondary veins 10–16, tertiary veins absent; base cuneate; apex retuse or obcordate; adaxial surface bright, with numerous cystoliths and simple, bent trichomes emerging from an inflated cystolith basal cell, abaxial surface bright, striate, very rarely with scattered trichomes, midrib and tips with a few trichomes. Inflorescence axillary or terminal, secondarily branched, branches slender, with diffusely scattered trichoms, pedicel 0.50–0.60 cm long. Calyx 0.25–0.30 cm long, 0.28–0.30 cm wide, coriaceous, with scattered trichomes outside, glabrous inside, but with a few scattered trichomes at blade tips, lobes 0.20–0.40 cm long, 0.30–0.35 cm wide, broadly obovate to triangulate, apex obtuse. Flowers at anthesis unknown. Mature fruit unknown; pyrene 0.35-0.40 cm tall, 0.28-0.30 cm wide, 0.20-0.22 cm deep, abaxially completely smooth.
Endemic to eastern Cuba (symbol "⌾" in Fig.
Fruiting Jun.
The species is very similar to Jamaican
The species is endemic to Southern Cuba and is known only from five herbarium collections. According to the Red List of Cuban Plants (
none.
Trees up to 10.0 m tall or smaller shrubs 2.0–5.0 m tall, crown arching, galls absent; indument glabrescent, young twigs variously pilose through glabrescent; bark grey light to grey brown, longitudinally fissured, wood light; thorns 0.80–1.0 cm long, robust, somewhat acute, sometimes slightly curved at tips, simple, scattered, alternate, glabrous. Leaves fasciculate, occasionally stipulate (2 deciduous stipules, as stated by
Northern South America (symbol "□" in Fig.
Flowering Jan, Mar–May, Jul, Sep, Nov; fruiting Feb–Dec.
“macarao”, “pega paloma” and “tachuelo” in Venezuela and “cruceto macho” and “tatacun o revienta puerco” in Colombia.
In Venezuela, the species is occasionally cultivated and forms hedgerows together with
Shrubs or small trees 3.0–5.0 m tall, galls absent; indument tomentose, especially on young organs, trichomes simple or stellate; bark grey whitish to light brown, longitudinally fissured; 0.7–1.3 cm long, slender, acute, successively branched at 2 or 3 levels, numerous, alternate or terminally of twigs, variously tomentose through glabrescent. Leaves fasciculate; petiole 0.4–0.6 cm long, robust, covered with a dense layer of stellate trichomes; blade 0.6–3.1 cm long, 0.4–2.1 cm wide, elliptic, widely elliptic or sometimes widely ovate, coriaceous, primary veins prominent, secondary veins 5–8, tertiary veins arcuate; base rounded or cuneate; apex rounded, retuse, obcordate, rarely mucronulate; adaxial surface bright, with cystoliths, rugose to almost glabrous or with a few scattered trichomes on the midrib, cilliate at tips, or with long (visible by the naked eye), simple, curved trichomes emerging from an inflated cystolith cell, abaxial surface densely tomentose, occasionally almost glabrous. Inflorescence axillary, flowers in clusters of 2 to 8, pedicels 0.10-0.15 cm long. Calyx 0.20–0.25 cm long, coriaceous, densely stellate outside, ciliate at tips, sometimes with simple trichomes scattered inside towards the distal part, lobes 0.18–0.22 cm long, 0.13–0.15 cm wide, shape triangular, apex acute, occasionally tridentate. Flowers at anthesis unknown. Fruit 0.40–0.50 cm tall, 0.40–0.50 cm wide; style 0.18–0.20 cm long, globose, divided in the proximal part, branches 2, 0.08–0.10 cm long, smaller than fruit, persistent (also the stigmas); pyrene 0.35–0.40 cm tall, 0.18–0.20 cm wide, 0.12–0.13 cm deep, abaxial surface with 5-6 longitudianal ridges.
Flowering Mar–Aug, Nov–Dec; fruiting: Apr–Aug.
Within his
Some individuals of
“espino de costa” in Holguín and “carey de costa” in Santiago de Cuba.
1 | Abaxial leaf surface with stellate (i.e., multi-branched) trichomes. |
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– | Abaxial leaf surface with simple trichomes or glabrous. |
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2 | Thorns (at least some) branched; plants from Cuba. |
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– | Thorns simple, plants from elsewhere (if from Cuba, then mature leaf length <3.0 cm except |
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3 | Mature leaf length < 3.0 cm; flowers sub-sessile; pyrenes abaxially ornamented. |
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– | Mature leaf length > 3.0 cm; flowers distinctly pedicellate; pyrenes abaxially smooth. |
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4 | Flowers sessile, 1–2 at an individual node. |
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– | Flowers distinctly pedicellate. |
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5 | Mature leaf length > 8.0 cm. |
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– | Mature leaf length < 6.0 cm. |
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6 | Lianas from Central America and westernmost Cuba; style divided in proximal half. |
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– | Trees and shrubs from elsewhere; 2 stylodia. |
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7 | Leaves (occasionally > 6.0 cm) predominantly membranaceous, mainly obovate. |
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– | Leaves predominantly coriaceous, very widely obovate to orbicular. |
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8 | Plants from The Bahamas and western Cuba, with dense cystolith-like structures adaxially. |
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– | Plants from the Leeward Islands and Eastern Puerto Rico, adaxial surface mostly glabrous. |
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We thank the curators of the herbaria B, F, G, GH, K, M, MO, P and US for providing us with material and/or information, Hermann Manitz (Jena) for literature and access to herbarium JE, Stan Huzarewicz (Storrs) for providing a *.pdf file of M.W. Lefor\'s thesis, Mia Ehn (Stockholm), Laura Guglielmone (Turin) and Mats Hjertson (Uppsala) for sending images of (putative) type material, Lucian Ionuț Roşu (Iaşi) for assistance with the distribution map and Denis Filer (Oxford) for his support on the BRAHMS online system. Hartmut H. Hilger (Berlin) and another anonymous reviewer provided helpful comments to this revision that are greatfully acknowledged here.
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♂ | – | 1 | 3 | 4 | 4 | 3 | 1 | 1 | – | 1 | 1 | – |
♀ | 2 | 2 | 6 | 3 | 2 | 7 | 3 | 2 | – | 1 | 3 | 3 |
fruit | 5 | 7 | 7 | 4 | 9 | 12 | 12 | 13 | 9 | 7 | 9 | 6 |