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 270518 10.11646/phytotaxa.614.1.1 143c8e81-3541-4298-b233-d3ce4e79d26c 1179-3163 8389307 1.5. Coccothrinax barbadensis (Loddiges ex Martius) Beccari (1907: 328) . Thrinax barbadensis Loddiges ex Martius (1853: 320) . Copernicia barbadensis (Loddiges ex Martius) de la Devansaye (1875: 35) . Neotype (designated here):— BARBADOS . Rim of Welshman Hall Gully, 19 June 1974 , R. Read 74-214 (neotpye US !). Plate 5 Coccothrinax alta (Cook) Beccari (1907: 331) . Thrincoma alta Cook (1901: 540) . Type:— PUERTO RICO . Road from Utuado to Arecibo , 14 June–22 July 1901 , L. Underwood & R. Griggs 848 ( holotype NY!). Coccothrinax laxa (Cook) Beccari (1907: 333) . Thringis laxa Cook (1901: 545) . Type:— PUERTO RICO . Punas Afuerto, Veja Baja, 8 December 1899 , O. Cook & G. Collins 1041 ( holotype US !). Coccothrinax latifrons (Cook) Beccari (1907: 326) . Thringis latifrons Cook (1901: 545) . Type:— PUERTO RICO . Propre Coamo in monte Calabaro, 30 December 1885 , P. Sintenis 3278 ( holotype NY!, isotypes G n.v. , G image!, GH!, LE n.v. , LE image!, M, n.v. , M image!, US !). Coccothrinax sanctae-thomae (“ sancti-thomae ”) Beccari (1907: 303) . Type :— US VIRGIN ISLANDS . Saint Thomas , 1905, Boergsen s.n. ( holotype C n.v. , C image!) . Coccothrinax eggersiana Beccari (1907: 321) . Type :— US VIRGIN ISLANDS . St. Jan ad Kings Hill , 900’, 26 December 1887 , H. Eggers 3117 ( holotype C n.v. , C image!). Coccothrinax eggersiana var. sanctae-crucis Beccari (1907: 323) . Type :— US VIRGIN ISLANDS . St. Croix , no date, O . Benzon s.n. ( holotype C n.v. , C image!) . Coccothrinax martinicaensis Beccari (1907: 324) . Lectotype (designated here):— MARTINIQUE . Vallée de St. Pierre , September 1873 , L. Hahn 1531 ( lectotype G n.v. , G image!, isolectotype FI!). Coccothrinax australis Bailey (1947b: 365) . Type :— TRINIDAD . St. Patrick , Quinam Bay , no date, L. Bailey 118 ( holotype BH !) . Coccothrinax discreta Bailey in Bailey & Moore (1949: 104) . Lectotype (designated by Reveal & Nixon 2013 ):— PUERTO RICO . Vieques , near Isabel II, May 1932 , L. Bailey 671 ( lectotype BH!, isolectotype MO n.v. , MO image!). Coccothrinax dussiana Bailey in Bailey & Moore (1949: 109) . Lectotype (designated here):— GUADELOUPE . Grands Fonds du Morne á l’Eau de Sainte Anne , 1894, Père Duss 3798 ( lectotype NY!). Coccothrinax sabana Bailey in Bailey & Moore (1949: 110) . Type :— SABA . Bottom , 7 February 1948 , L. Bailey 406 ( holotype BH !) . Coccothrinax boxii Bailey in Bailey & Moore (1949: 113) . Lectotype (designated by Reveal & Nixon 2013 ):— ANTIGUA AND BARBUDA . Barbuda , The Highlands, 15 May 1937 , H. Box 669 ( lectotype BH!). Stems 5.5(1.2–15.2) m long and 9.0(4.4–15.0) cm diameter, solitary. Leaves more or less deciduous or only leaf bases persisting on stem; leaf sheath fibers 0.4(0.1–0.6) mm diameter, closely woven, not forming persistent ligules and soon disintegrating at the apices; petioles 12.9(8.5–19.7) mm diameter just below the apex; palmans 26.7(13.3–43.0) cm long, relatively long, without prominent adaxial veins; leaf blades not wedge-shaped; segments 54(37–62) per leaf, the middle ones 74.7(50.5–102.5) cm long and 3.4(2.1–4.7) cm wide; segments 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; middle leaf segment apices attenuate; leaf segments not waxy or sometimes with a deciduous, thin layer of wax adaxially, densely indumentose abaxially, with irregularly shaped, semi-persistent, interlocking, fimbriate hairs without an obvious center, with well-developed transverse veinlets. Inflorescences curving, arching, or pendulous amongst the leaves, with few to numerous partial inflorescences; rachis bracts somewhat flattened, loosely sheathing, usually tomentose with a dense tuft of erect hairs at the apex; partial inflorescences 5(3–6); proximalmost rachillae straight, 10.8(5.0–17.0) cm long and 1.4(0.9–1.9) mm diameter in fruit; rachillae glabrous at or near anthesis; stamens 10(8–12); fruit pedicels 2.4(0.6–4.5) mm long; fruits 8.1(6.3–10.1) mm long and 8.2(6.5–10.8) mm diameter, purple, dark purple, red-purple, reddish-purple, purple-black, red-pink, red-black, grayish-brown, blackish, or black; fruit surfaces smooth or sometimes with projecting fibers; seed surfaces deeply lobed, the lobes running from base of seeds almost to apices. Distribution and habitat:— Southeastern Dominican Republic ( La Altagracia , San Pedro de Macoris ), Puerto Rico , Virgin Islands , Lesser Antilles, Trinidad and Tobago , and northeastern Venezuela ( Fig. 9 ) in deciduous or semi-deciduous forest, xeromorphic forest, dry forest, or secondary forest, on limestone soils, at 120(8–325) m elevation. No specimens from St. Vincent or Granada have been seen, nor are these two islands listed in Read (1979) . However, Read noted that many populations had been eliminated after the arrival of humans on the islands. Coccothrinax barbadensis also occurs on Margarita Island, Venezuela ( Braun 1984 ), although no specimens from there have been seen. Taxonomic notes:— Two preliminary species ( Coccothrinax alta , C. barbadensis ) share a unique combination of qualitative character states and are recognized as a single phylogenetic species, the earliest name for which is C. barbadensis . Type specimens of C. australis , C. boxii , C. discreta , C. dussiana , C. eggersiana , C. eggersiana var. sanctae-crucis, C. latifrons , C. laxa , C. martinicaensis , C. sabana , and C. sanctae-thomae share the same combination of character states and are treated as synonyms of C. barbadensis . Bailey (1949) gave C. dussiana as new name for C. barbadensis but according to POWO (2023) it is a heterotypic synonym of C. barbadensis . PLATE 5 . Coccothrinax barbadensis , cultivated, Miami, with inflorescences pendulous amongst the leaves. The protologue of C. barbadensis consists of a single, short sentence “258. Th. barbadensis Lodd. 257 n. 6. Insula Barbadensis”, and there is no description, nor illustration, nor type. According to Bailey (1949) the name was based on a young, cultivated palm of uncertain genus. Given that Coccothrinax does occur on Barbados and Thrinax does not, it is assumed that the protologue referred to a Coccothrinax , and a neotype from Barbados is designated. Coccothrinax barbadensis is a widespread and variable species occurring throughout the eastern Caribbean. Subspecific variation :—From the west, the status of specimens from southeastern Hispaniola is still unclear. Mejía et al. (1998) identified these as C. barbadensis (illustrated in Fernández & Gottschalk 2017 ). However, these specimens appear to have short pedicels, unlike C. barbadensis and more like C. argentea . There may be some hybridization here between C. barbadensis and C. argentea . In the remainder of the range, two preliminary species were recognized, C. alta from Puerto Rico and the Virgin Islands , and C. barbadensis from the Lesser Antilles, Trinidad and Tobago , and northeastern Venezuela . Specimens of C. alta differ significantly from those of C. barbadensis in four variables (petiole length, number of segments, proximalmost rachillae length, pedicel length) with specimens of C. barbadensis having higher values for all variables ( t- test, P <0.05). There are other differences. Specimens of C. barbadensis appear to have more conspicuous transverse veinlets. They have longer pedicels and usually the bracteole of the pedicel is situated on the pedicel itself, whereas specimens of C. alta have shorter pedicels and usually the bracteole is situated at the base of the pedicel. However, within these two areas ( Puerto Rico and the Virgin Islands ; Lesser Antilles, Trinidad and Tobago , and northeastern Venezuela ) there is evidence for numerous, local populations rather than two large populations. For example, even within Puerto Rico there appear to be two populations, a northern and a southern one (with the specimens from extreme eastern Puerto Rico included in the southern population). Specimens from the northern population occur on “moist/wet northern limestone”, while specimens from the southern and eastern population occur on “dry-volcanic limestone” ( Helmer et al. 2002 ). Specimens from the southern population appear to be more similar to specimens from the Lesser Antilles, often having conspicuous transverse veinlets and longer pedicels with the bracteole of the pedicel situated on the pedicel itself. Specimens from St. Croix appear to be similar (although there is some doubt whether some of these specimens are from cultivated plants). For the smaller islands of the Lesser Antilles, both Beccari (1907) and Bailey in Bailey & Moore (1949) pointed out inter-island differences. Bailey in Bailey & Moore (1949) wrote “There is strong indication that differences of specific value inherent to various islands and that the species are endemic”. Beccari (1907) noted on that on the island of St. Thomas specimens appeared to have indistinct transverse veinlets, and this is confirmed by examination of the specimens known from there. In summary, rather than comprising various subspecies, C. barbadensis appears to comprise two populations in Puerto Rico and numerous, disjunct, island populations in the Lesser Antilles. Subspecies are not recognized for the same reasons discussed under C. argentata ; too few specimens from most of the smaller islands to test for differences in quantitative variables, dozens of populations and thus a potentially unwieldly number of subspecies, and evidence of dispersal and hybridization.