A revision of Coccothrinax, Hemithrinax, Leucothrinax, Thrinax, and Zombia (Arecaceae) Author Henderson, Andrew text Phytotaxa 2023 2023-09-19 614 1 1 115 http://dx.doi.org/10.11646/phytotaxa.614.1.1 journal article 10.11646/phytotaxa.614.1.1 1179-3163 8389307 1. Coccothrinax Sargent (1899: 87) . Type :— Coccothrinax jucunda Sargent (= Coccothrinax argentata (Jacquin) Bailey ) Thrincoma Cook (1901: 539) . Type :— Thrincoma alta Cook (= Coccothrinax barbadensis (Loddiges ex Martius) Beccari ) Thringis Cook (1901: 544) . Lectotype (designated by Moore 1963 ):— Thringis latifrons Cook (= Coccothrinax barbadensis (Loddiges ex Martius) Beccari ) Haitiella Bailey (1947a: 7) . Type :— Haitiella ekmanii (Burret) Bailey (= Coccothrinax ekmanii Burret ) Coccothrinax subgen. Eucoccothrinax Beccari (1931: 336) . Type :—Not designated. Coccothrinax series Brevispadicae León (1939: 110) . Type :—Not designated. Coccothrinax series Brevispadicae subseries Cernuae León (1939: 110) . Type :—Not designated. Coccothrinax series Brevispadicae subseries Curvatae León (1939: 110) . Type :—Not designated. Coccothrinax series Longispadicae León (1939: 110) . Type :—Not designated. Coccothrinax series Longispadicae subseries Pauciramosae León (1939: 110) . Type :—Not designated. Coccothrinax series Longispadicae subseries Multiramosae León (1939: 110) . Type :—Not designated. Coccothrinax section Coccothrinax subsection Haitiella (Bailey) Muñiz in Muñiz & Borhidi (1982: 319) . Type :— Coccothrinax ekmanii Burret Coccothrinax section Longispadicae ( León ) Muñiz in Muñiz & Borhidi (1982: 316) . Type :— Coccothrinax rigida (Grisebach & Wendland) Beccari Coccothrinax section Longispadicae subsection Multiramosae León ex Muñiz in Muñiz & Borhidi (1982: 316) . Type :— Coccothrinax gundlachii León Coccothrinax section Longispadicae subsection Pauciramosae León ex Muñiz in Muñiz & Borhidi (1982: 317) . Type :—Not designated. Stems 4.5(0.03–20.0) m long and 9.2(2.9–50.0) cm diameter, solitary or clustered. Leaves persistent, giving a ‘skirt’ of dead leaves covering the stem, or more or less deciduous or only leaf bases persisting on stem; leaf sheaths not split at the base; leaf sheath fibers 1.0(0.1–7.5) mm diameter, thin (usually < 0.5 mm diameter), closely woven, forming persistent, triangular ligules at the apices, or thin (usually < 0.5 mm diameter), closely woven, not forming persistent ligules and soon disintegrating at the apices, or thin (usually < 0.5 mm diameter), flimsy, loosely woven, free and greatly elongate at the apices, or stout (usually> 1 mm diameter), woody, loosely woven, ± joined or briefly free at the apices, or stout (usually> 1 mm diameter), woody, loosely woven, the inner and outer layer combining at the apices to form erect , spine-like fibers, or stout (usually 0.3–0.5 mm diameter), loosely woven and forming a loose, hexagonal mesh, initially forming ligules at the apices, or thin (usually ca. 0.4 mm diameter), curled toward the apex, loosely woven and forming a loose, hexagonal mesh, truncate at the apex; petioles 10.7(2.0–26.4) mm diameter just below the apex; palmans 12.1(1.0–43.0) cm long and relatively short, with the adaxial veins prominent and terminating in a slight raised ridge and distinct pulvinus, or relatively long, without prominent adaxial veins; interfold filaments absent; leaf blades wedge-shaped, or not wedge-shaped; segments 35(12–71) per leaf, the middle ones 49.3(17.2–102.5) cm long and 2.9(0.8–6.5) cm wide; segments pendulous at the apices, giving a three-dimensional appearance to the leaf, or not pendulous at the apices, giving the leaf a flat appearance; middle leaf segments relatively long and narrow, tapering from base to apex, scarcely folded, flexible and not leathery, a shoulder or constriction absent or poorly developed, the apices thin, deeply splitting and breaking off, or relatively short and broad, abruptly narrowed (shoulder) toward the apex, otherwise parallel-sided, often strongly folded, stiff and leathery, the apices briefly splitting, or tapering from base to apex, often folded, stiff and leathery, with or without scarcely developed shoulders, the apices sharply pointed and briefly splitting, or relatively long and broad, abruptly narrowed (shoulder) about the middle, stiff and leathery, the apices elongate beyond the shoulder, briefly splitting; middle leaf segment apices blunt and rounded, or attenuate; leaf segments with a persistent, dense, whitish layer of wax adaxially, or not waxy or sometimes with a deciduous, thin layer of wax adaxially; leaf segments densely indumentose abaxially, with irregularly shaped, persistent, interlocking, fimbriate hairs, each one with a conspicuous, reddish-brown, pale brown, or greenish elliptic center, or densely indumentose abaxially, with irregularly shaped, persistent, interlocking, fimbriate hairs, each one with a rounded, raised, light green to greenish-brown or reddish-brown center, or densely indumentose abaxially, with irregularly shaped, semi-persistent, interlocking, fimbriate hairs without an obvious center, or with a thin layer of wax abaxially, or without indumentum, scales, or wax abaxially; leaf segments without or with poorly developed transverse veinlets, or with well-developed transverse veinlets. Inflorescences erect, at least initially, amongst or above the leaves, with few to numerous partial inflorescences, or erect amongst the leaves, with few partial inflorescences at apex of inflorescence, or curving, arching, or pendulous amongst the leaves, with few to numerous partial inflorescences; rachis bracts narrow, closely sheathing, sparsely tomentose, usually without hairs at the apex, or somewhat flattened, loosely sheathing, usually tomentose with a dense tuft of erect hairs at the apex; partial inflorescences 4(2–9); proximalmost rachillae straight or recurved, 8.0(2.5–22.0) cm long and 1.2(0.4–2.4) mm diameter in fruit; rachillae glabrous at or near anthesis, or uneven with lines of warty outgrowths, these often becoming more pronounced as fruits develop; flowers spirally arranged; stamens 8(5–13), elongate, spreading irregularly at anthesis, the anthers latrorse; fruit pedicels 1.7(0.1–6.2) mm long; fruits 7.7(3.0–16.4) mm long and 7.8(3.8–15.4) mm diameter, black, blackish, brown, rusty-brown, grayish-brown, blue-black, purple-black, dark purple, deep purple, purplish, purple, red-purple, burgundy, red-pink, rose-red, wine color, magenta, reddish, dark red, red-black, orange-yellow, yellowish, yellow-white, tan, cream, or white; fruit surfaces densely muricate, or smooth or sometimes with projecting fibers; seed surfaces deeply lobed, the lobes running from base of seeds almost to apices, or lobed, the lobes running from base of seeds approximately to equator, rarely smooth or scarcely lobed; seeds in longitudinal section not intruded by the hilum. Distribution and habitat: — Coccothrinax species are widely distributed throughout the Caribbean and adjacent mainland areas, from 26°40’N ( Bahamas ) to 10°03’S ( Trinidad ) and from 89°32’W ( Mexico ) to 59°34’W ( Barbados ) ( Fig. 7 ). The island with the highest number of species, 24, is Cuba followed by Hispaniola with 16. Species occur from sea level to 1,850 m elevation ( C. montana in the Dominican Republic). However, most specimens are from low elevations. Of the 246 specimens with elevation data, 128 (52%) occur at or below 200 m . Only 13% of specimens occur at or above 1,000 m — nearly all either C. montana or C. scoparia in Hispaniola. Species occur in a variety of habitats. In general they are found in areas of low rainfall and often in arid areas. On smaller islands and adjacent mainland areas they occur at low elevations near the sea, often in open forest or scrub forest on sandy or coralline soils. On both Hispaniola and Cuba , several local endemics are confined to serpentine or limestone soils, often on savannas or more open areas with low forest. FIGURE 7 . Distribution of Coccothrinax . Key to the species of Coccothrinax 1. USA ( Florida ), Bahamas , Turks and Caicos Islands , Haiti ( Tortuga island), Dominican Republic (Morro de Monte Cristi ), Jamaica , Cayman Islands , Mexico ( Quintana Roo , Yucatán ), Belize , Colombia (San Andrés, Providencia) and Honduras (Islas del Cisne). ......................................................................................................................................................................................... C. argentata - All other areas .................................................................................................................................................................................. 2. 2. Puerto Rico , Virgin Islands , Lesser Antilles, Trinidad and Tobago , and northeastern Venezuela ............................. C. barbadensis - Cuba and Hispaniola ........................................................................................................................................................................ 3. 3. Cuba ................................................................................................................................................................................................. 4. - Hispaniola ...................................................................................................................................................................................... 27. 4. Leaf segments pendulous at the apices, giving a three-dimensional appearance to the leaf; leaf segments densely indumentose abaxially, with irregularly shaped, persistent, interlocking, fimbriate hairs, each one with a conspicuous, reddish-brown, pale brown, or greenish elliptic center; widespread in coastal areas ..................................................................................... C. argentata - Leaf segments not as above ............................................................................................................................................................. 5. 5. Leaf sheath fibers thin, closely woven, not forming persistent ligules and soon disintegrating at the apices; leaf segments with poorly developed transverse veinlets; Guantánamo ........................................................................................................ C. hioramii - Leaf sheath fibers not as above; leaf segments without transverse veinlets .................................................................................... 6. 6. Leaf sheath fibers thin, curled toward the apex, loosely woven and forming a loose, hexagonal mesh, truncate at the apex; Santiago de Cuba ............................................................................................................................................................................ C. fragrans - Leaf sheath fibers not as above ....................................................................................................................................................... 7. 7. Fruit surfaces densely muricate; Guantánamo .................................................................................................................. C. munizii - Fruit surfaces not densely muricate ................................................................................................................................................. 8. 8. Leaf sheath fibers thin, flimsy, loosely woven, free and greatly elongate at the apices .................................................................. 9. - Leaf sheath fibers not as above ..................................................................................................................................................... 10. 9. Leaves persistent, giving a ‘skirt’ of dead leaves covering the stem; Matanzas ......................................................... C. borhidiana - Leaves more or less deciduous or only leaf bases persisting on stem; Cienfuegos, Pinar del Río, Sancti Spíritus ........... C. crinita 10. Leaf sheath fibers with the inner and outer layer combining at the apices to form erect , spine-like fibers .................................. 11. - Leaf sheath fibers ± joined or briefly free at the apices ................................................................................................................ 17. 11. Inflorescences erect, at least initially, amongst or above the leaves, with few to numerous partial inflorescences, or erect amongst or above the leaves, with few partial inflorescences at apex of inflorescence .............................................................................. 12. - Inflorescences curving, arching, or pendulous amongst the leaves, with few to numerous partial inflorescences ...................... 15. 12. Inflorescences erect, at least initially, amongst or above the leaves, with few to numerous partial inflorescences; Villa Clara, Sancti Spíritus, Ciego de Ávila, Camagüey ............................................................................................................................... C. clarensis - Inflorescences erect amongst or above the leaves, with few partial inflorescences at apex of inflorescence ............................... 13. 13. Leaves more or less deciduous or only leaf bases persisting on stem; Holguín, Santiago de Cuba ........................ C. pauciramosa - Leaves persistent, giving a ‘skirt’ of dead leaves covering the stem ............................................................................................. 14. 14. Stems solitary; Holguín ................................................................................................................................................... C. garciana - Stems clustered; Camagüey ...................................................................................................................................... C. pseudorigida 15. Middle leaf segments attenuate; Camagüey, Holguín, Las Tunas ................................................................................. C. salvatoris - Middle leaf segment apices blunt and rounded; Guantánamo ....................................................................................................... 16. 16. Leaf blades not wedge-shaped ..................................................................................................................................... C. yunquensis - Leaf blades wedge-shaped .......................................................................................................................................... C. microphylla 17. Middle leaf segment apices blunt and rounded; Guantánamo, Holguín ........................................................................ C. orientalis - Middle leaf segment apices attenuate ............................................................................................................................................ 18. 18. Inflorescences erect amongst or above the leaves ......................................................................................................................... 19. - Inflorescences curving, arching, or pendulous amongst the leaves ............................................................................................... 21. 19. Leaf segments with a persistent, dense, whitish layer of wax adaxially; Guantánamo ...................................................... C. torrida - Leaf segments without a persistent, dense, whitish layer of wax adaxially .................................................................................. 20. 20. Rachis bracts somewhat flattened, loosely sheathing, usually tomentose with a dense tuft of erect hairs at the apex; Granma, Santiago de Cuba .......................................................................................................................................................... C. gundlachii - Rachis bracts narrow, closely sheathing, sparsely tomentose, usually without hairs at the apex; Guantánamo ........ C. baracoensis 21. Leaf segments with a persistent, dense, whitish layer of wax adaxially; Sancti Spíritus .............................................. C. spirituana - Leaf segments without a persistent, dense, whitish layer of wax adaxially .................................................................................. 22. 22. Leaf segments without indumentum, scales, or wax abaxially; Holguín ............................................................................ C. rigida - Leaf segments densely indumentose abaxially, with irregularly shaped, persistent, interlocking, fimbriate hairs, each one with a rounded, raised, light green to greenish-brown or reddish-brown center ...................................................................................... 23. 23. Stems solitary; throughout Cuba ................................................................................................................................................... 24. - Stems clustered; Guantánamo ....................................................................................................................................................... 26. 24. Proximalmost rachillae recurved; Guantánamo ............................................................................................................. C. alexandri - Proximalmost rachillae straight ..................................................................................................................................................... 25. 25. Fruits 4.8(4.0–5.4) mm long and 4.8(4.0–5.7) mm diameter; seed surfaces lobed, the lobes running from base of seeds approximately to equator; Granma, Guantánamo, Santiago de Cuba ...................................................................................................... C. saxicola - Fruits 7.4(6.2–10.6) mm long and 7.7(6.2–11.2) mm diameter; seed surfaces deeply lobed, the lobes running from base of seeds almost to apices; western and central Cuba ................................................................................................................. C. miraguama 26. Leaf blades wedge-shaped; Santiago de Cuba .................................................................................................................. C. fagildei - Leaf blades not wedge-shaped; Guantánamo ..................................................................................................................... C. pumila 27. Leaf sheath fibers closely woven, forming persistent, triangular ligules at the apices; throughout Hispaniola ............. C. argentea - Leaf sheath fibers not forming persistent, triangular ligules at the apices .................................................................................... 28. 28. Leaf sheath fibers thin (usually < 0.5 mm diameter), closely woven, not forming persistent ligules and soon disintegrating at the apices ............................................................................................................................................................................................. 29. - Leaf sheath fibers stout (usually> 1 mm diameter), woody, loosely woven, ± joined or briefly free at the apices, or stout (usually> 1 mm diameter), woody, loosely woven, the inner and outer layer combining at the apices to form erect , spine-like fibers, or stout (usually 0.3–0.5 mm diameter), loosely woven and forming a loose, hexagonal mesh, initially forming ligules at the apices ... 30. 29. Stems 4.4–15.0 cm diameter, not swollen; at 10–26 m elevation in southeastern Dominican Republic ( La Altagracia , San Pedro de Macoris ) ..................................................................................................................................................................... C. barbadensis - Stems 11.0–50.0 cm diameter, sometimes swollen; throughout Hispaniola ....................................................................... C. spissa 30. Fruits 8.3–16.4 mm long; at higher elevations .............................................................................................................................. 31. - Fruits 3.9–7.9 mm long; at lower elevations ................................................................................................................................. 32. 31. Palmans 8.3(3.5–15.3) cm long, relatively short, with the adaxial veins prominent and terminating in a slight raised ridge and distinct pulvinus; Dominican Republic (Sierra Martín García and Sierra de Baoruco ) and Haiti (Massif de la Hotte and probably Massif de la Selle) ........................................................................................................................................................... C. scoparia - Palmans 20.1(12.0–26.50) cm long, relatively long, without prominent adaxial veins; Dominican Republic (Cordillera Central, Cordillera Septentrional, Sierra de Neiva) ...................................................................................................................... C. montana 32. Fruit surfaces densely muricate ..................................................................................................................................................... 33. - Fruit surfaces smooth or sometimes with projecting fibers.............................................................................................................35. 33. Leaf sheath fibers combining at the apices to form erect , spine-like fibers; southeastern Haiti ( Sud-Est ) and southwestern Dominican Republic ( Barahona , Pedernales ) ..................................................................................................................................... C. ekmanii - Leaf sheath fibers ± joined or briefly free at the apices ................................................................................................................ 34. 34. Leaf blades not wedge-shaped; Dominican Republic ( Azua ) ....................................................................................... C. boschiana - Leaf blades wedge-shaped; northeastern Dominican Republic ( Samaná ) .................................................................. C. samanensis 35. Inflorescences erect, at least initially, amongst or above the leaves; north-central Dominican Republic ( Espaillat ) .. C. bonettiana - Inflorescences curving, arching, or pendulous amongst the leaves ............................................................................................... 36. 36. Leaf blades not wedge-shaped ....................................................................................................................................................... 37. - Leaf blades wedge-shaped ............................................................................................................................................................. 39. 37. Leaf segments without indumentum abaxially; southern Haiti ( Department Sud-Est ) ................................................... C. concolor - Leaf segments densely indumentose abaxially .............................................................................................................................. 38. 38. Fruits 3.8–4.7 mm diameter; seed surfaces lobed, the lobes running from base of seeds approximately to equator; northeastern Dominican Republic ( El Seibo , Hato Mayor ) ..................................................................................................... C. montgomeryana - Fruits 7.1–7.2 mm diameter; seed surfaces deeply lobed, the lobes running from base of seeds almost to apices; northwestern Haiti ( Nord-Ouest ) ...................................................................................................................................................................... C. gracilis 39. Leaf segments densely indumentose abaxially; Haiti ( Artibonite ) ............................................................................ C. gonaivensis - Leaf segments without indumentum abaxially .............................................................................................................................. 40. 40. Leaf sheath fibers 0.3(0.2–0.4) mm diameter, forming a loose, hexagonal mesh, initially forming ligules at the apices; Dominican Republic ( Independencia ) ................................................................................................................................................ C. jimenezii - Leaf sheath fibers 1.3(1.1–1.5) mm diameter, ± joined or briefly free at the apices; Dominican Republic ( Azua ) ... C. landestoyii