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 1.23. Coccothrinax miraguama (Kunth) Beccari (1907: 295) . Corypha miraguama Kunth in Humboldt, Bonpland & Kunth (1816: 298) . Copernicia miraguama (Kunth) Kunth (1841: 244) . Thrinax miraguama (Kunth) Martius (1853: 320) . Lectotype (designated by Stauffer & Stauffer 2017 ):— CUBA . Inter urbem la Trinidad, Puerto Casilda et ostia fluminis Guaurabo, no date, A. Humboldt & A. Bonpland 1356 ( holotype P n.v ., P image!). Plate 17 Coccothrinax acuminata Sargent ex Beccari (1907: 313) . Lectotype (designated by Moya & Méndez 2018a ):— CUBA . No locality, no date, C. Wright 3966 ( lectotype K!, isolectotypes BRU n.v. , FI!, GH!, NY!, P n.v. , P image!, US !, V n.v. ). Coccothrinax miraguama var. novogeronensis Beccari (1931: 336) . Lectotype (designated by Moya & Méndez 2018a ):— CUBA . Isla de Pinos, 1904, A. Curtiss 423 ( lectotype FI!, isolectotypes A!, BH!, CM n.v. , G n.v. , GH!, HAC!, K!, LE n.v. , M n.v. , M image!, MO n.v. , MT n.v ., NY!, US !). Coccothrinax miraguama var. arenicola León (1939: 115) . Coccothrinax miraguama subsp. arenicola (León) Borhidi & Muñiz (1971a: 175) . Lectotype (designated by Moya & Méndez 2018a ):— CUBA . Pinar del Río , Sabanalamar, El Sábalo, 20 August 1934 , Fr. León 16146 ( lectotype HAC!, isolectotypes A!, US !). Coccothrinax miraguama var. havanensis León (1939: 116) . Coccothrinax miraguama subsp. havanensis (León) Borhidi & Muñiz (1971a: 175) . Lectotype (designated by Moya 2020 ):— CUBA . ( Habana ), Playa de Jibacoa, 13 May 1937 , Fr. León 16834 ( lectotype HAC!, isolectotypes A!, BH!, GH!, HAJB!, US !). Coccothrinax miraguama var. roseocarpa León (1939: 117) . Coccothrinax miraguama subsp. roseocarpa (León) Borhidi & Muñiz (1971a: 175) . Lectotype (designated by Moya 2020 ):— CUBA . ( Matanzas ), Loma de los Botinos, 27 August 1937 , Fr. León 16958 ( lectotype HAC!, isolectotypes A!, BH!, GH!, MICH n.v . , MICH image!, US !). Coccothrinax miraguama var. cupularis León (1939: 117) . Coccothrinax miraguama subsp. cupularis (León) Borhidi & Muñiz (1971a: 175) . Coccothrinax cupularis (León) Muñiz & Borhidi (1981 publ. 1982: 449). Type:— CUBA . Cienfuegos , Santa Clara, Monte de la Playa de Patado, Castillo de Jagua, 21 June 1932 , Fr. León 15595 ( holotype HAC!). Coccothrinax miraguama var. macroglossa León (1939: 118) . Coccothrinax macroglossa (León) Muñiz & Borhidi (1981 publ. 1982: 450). Coccothrinax miraguama subsp. macroglossa (León) Borhidi & Muñiz (1971a: 175) . Lectotype (designated by Moya 2020 ):— CUBA . Oriente, Cerro de Dumañuecos, Manatí, 23 July 1937 , Fr. León 16869 ( lectotype HAC!, isolectotype US !). Coccothrinax yuraguana León (1939: 119) . Lectotype (designated by Moya 2018):— CUBA . Pinar del Río , entre le Pan de Guajaibon et Las Pozas, April 1934 , G . Nateson 16103 ( lectotype HAC !, isolectotypes A !, AJBC n.v. , US n.v. ). Coccothrinax camagueyana Borhidi & Muñiz in Muñiz & Borhidi (1981 publ. 1982: 441). Lectotype (designated by Moya 2021 ):— CUBA . Camagüey , Tuabaquey, 28 October 1977 , O. Muñiz & A. Borhidi 106 ( lectotype HAC!, isolectotype BP n.v. ). Coccothrinax trinitensis Borhidi & Muñiz (1985: 228) . Type:— CUBA . Sierra de Escambray , falda norte del Pico Potrerillo, 8 July 1984 , O. Muñiz & M. Fernández 15061 ( holotype HAC n.v .) . Stems 4.5(3.0–7.0) m long and 6.5 cm diameter, solitary. Leaves more or less deciduous or only leaf bases persisting on stem; leaf sheath fibers 1.2(0.4–3.1) mm diameter, stout, woody, loosely woven, ± joined or briefly free at the apices; petioles 12.4(6.3–20.8) mm diameter just below the apex; palmans 11.6(2.4–28.5) cm long, relatively short, with the adaxial veins prominent and terminating in a slight raised ridge and distinct pulvinus; leaf blades not wedge-shaped; segments 34(22–49) per leaf, the middle ones 44.6(20.5–78.5) cm long and 3.8(2.0–6.5) cm wide; segments not pendulous at the apices, giving the leaf a flat appearance; middle leaf segments relatively short and broad, abruptly narrowed (shoulder) toward the apex, otherwise parallel-sided, often strongly folded, stiff and leathery, the apices briefly splitting; 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, persistent, interlocking, fimbriate hairs, each one with a rounded, raised, light green to greenish-brown or reddish-brown center, without transverse veinlets. Inflorescences curving, arching, or pendulous amongst the leaves, with few partial inflorescences; rachis bracts somewhat flattened, loosely sheathing, usually tomentose with a dense tuft of erect hairs at the apex; partial inflorescences 3(2–7); proximalmost rachillae straight, 8.9(4.7–15.0) cm long and 1.2(0.8–1.8) mm diameter in fruit; rachillae glabrous at or near anthesis; stamens 9(7–10); fruit pedicels 2.1(0.5–6.0) mm long; fruits 7.4(6.2–10.6) mm long and 7.7(6.2–11.2) mm diameter, black, purplish, reddish, orange-yellow, or yellow-white; 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:— Cuba ( Fig. 14 ) in pinelands, savannas, shrubby areas, on serpentine or limestone soils, and persisting in disturbed areas, at 694(200–900) m elevation. Taxonomic notes:— Seven preliminary species ( C. acuminata , C. camagueyana , C. cupularis , C. macroglossa , C. miraguama , C. trinitensis , C. yuraguana ) share a unique combination of qualitative character states and are recognized as a single phylogenetic species, the earliest name for which is C. miraguama . Coccothrinax miraguama is widespread and variable in Cuba and appears to exist in numerous disjunct populations in a variety of habitats, each slightly different from the others. Subspecific variation :—From the west, there is a population in western Pinar del Río , growing in savannas or on limestone rocks. The type of Coccothrinax miraguama var. arenicola is from this population (placed as a synonym of C. acuminata by Moya & Méndez 2018a ). It was said to differ by its leaves with shorter sheath apices and segments with pale reddish scales abaxially. This population has 3-layered sheath fibers and larger leaves. However, as pointed out by Nauman & Sanders (1991a) , the distinction between 2- and 3-layered fibers breaks down in C. miraguama and this variable is not used here to distinguish species. Slightly further east, there is a population in southeastern Pinar del Río growing in pinelands or savannas. This population has 2-layered fibers and smaller leaves. Further to the east, at or near the border between Pinar del Río and Artemesia provinces, there is a population represented by a single specimen ( Wright 3966 ), the type of Coccothrinax acuminata . This specimen has 3-layered sheath fibers, and exceptionally large leaves and inflorescences. Moya & Méndez (2018a) considered that the type locality of C. acuminata could be at Balestena in Artemesia. PLATE 17 . Coccothrinax miraguama , Cuba, with middle leaf segments short and broad, abruptly narrowed (shoulder) toward the apex, otherwise parallel-sided, strongly folded, stiff and leathery. On the Isla de la Juventud there is a population growing in savannas or pinelands. This population has 2-layered sheath fibers, smaller leaves, and shorter pedicels. The type of Coccothrinax miraguama var. novogeronensis is from this population (placed as a synonym of C. acuminata by Moya & Méndez 2018a ). It was said to differ by its sessile flowers. Specimens determined as preliminary species C. acuminata differ from other specimens of C. miraguama from western Cuba in only one variable (leaf segment length) with specimens of C. acuminata having a lower value ( t- test, P <0.05). On the north coast of Pinar del Río , and just reaching Artemesia, there is a population growing in xeromorphic woodland on serpentine soils. The type of Coccothrinax yuraguana is from this population. León (1939) considered that it differed from C. miraguama by its smaller leaves with fewer segments, shorter palmans, and shorter apices (see also Moya 2018). Leaf sheaths have 2- or 3-layered fibers. Two fertile specimens from exactly the same locality differ greatly in size; one ( Verdecia 1232 ) is reported as being 3.5 m tall and has much larger leaves than the other ( Verdecia 1231 ), which is reported as being 5 m tall. This perhaps an example of ontogenetic changes, with smaller plants having larger leaves. Coccothrinax yuraguana is somewhat tentatively included here. In some specimens, leaf segment apices could almost be scored as rounded and as such are somewhat similar to C. orientalis (and both C. orientalis and its synonym C. moaensis were originally described as varieties or subspecies of C. yuraguana ). However, the type locality of C. yuraguana is approximately 800 km from the nearest occurrence of C. orientalis . The problem is unresolved. Specimens of preliminary species C. yuraguana do not differ from specimens of preliminary species C. acuminata in any variable, but do differ from other specimens of C. miraguama from western Cuba in four variables (fiber width, number of segments, leaf segment length, rachillae length) with specimens of C. yuraguana having lower values ( ttest , P <0.05). Around Havanna there is a population growing on hill slopes on serpentine soils. The type of C. miraguama var. havanensis is from this population. León (1939) considered that it differed by its longer segment apices. Leaves of this variety are noticeably larger than those of the following, var. roseocarpa . According to Craft (2017) , var. havanensis occurs on alkaline soils and var. roseocarpa occurs on serpentine soils. To the east of Havanna , on the north coast of Matanzas , there is a population growing on serpentine hills. The type of C. miraguama var. roseocarpa is from this population. This was said to differ by its rose-purple fruits. Given the range in fruit color in Coccothrinax , and the range in color during ripening, this is not considered to have any taxonomic significance. There is a large population of C. miraguama in Cienfuegos, Villa Clara, and Sancti Spíritus provinces, growing in savannas or serpentine barrens, or at higher elevations on dry hills or rocky hillsides. Specimens have 2-layered or 3-layered leaf sheath fibers. The type of C. miraguama is from this population. León (1939) wrote that he travelled to the exact locality where Humboldt and Bonpland collected the type specimen near Trinidad . The specimen ( León 16932 ) that León collected has an exceptionally large leaf and inflorescence. Craft (2017) wrote that the species no longer occurs at the type locality. Two other specimens have been named from this population. A specimen from the coast near Cienfuegos was described as C. miraguama var. cupularis , and was said to differ in its “filamentos….. unidos comúnmente en la mitad de su longitud ±, formando una cúpula“ ( León 1939 ). However, the amount of connation of the filaments at the base is quite variable and does not appear to have any taxonomic significance. Two other specimens ( Britton 4589, Jack 7691 ) from at or near the type locality have filaments only briefly united at the base, and another specimen ( Britton 5476 ), determined as C. miraguama and from the Trinidad mountains has the filaments forming a short cupule. Nauman & Sanders (1991a) reported that the filaments of C. miraguama subsp. havanensis also form a cupule. A specimen from the Sierra de Escambray was described as Coccothrinax trinitensis . Moya (2021) wrote that the type specimen has not been found at HAC. However, based on the protologue, it does not appear not to differ from C. miraguama . There is a gap in the distribution of C. miraguama in the central part of Cuba , and it reappears in Camagüey , Las Tunas , and Holguín provinces. Two names have been applied to this population, C. camagueyana and C. macroglossa . Muñiz & Borhidi (1981) described C. camagueyana as having seeds with a basal embryo, which is not the case, and compared it with C. gundlachii , presumably because of its large size. Moya (2021) reported that only one specimen of C. camagueyana is known, the type , and it has not been collected again. Quantitatively, C. camagueyana is similar to preliminary species C. macroglossa , and the type locality is within the range of C. macroglossa . León (1939) distinguished C. macroglossa (as C. miraguama var. macroglossa ) by its larger hastulas and leaves. Risco Villalobos et al. (2017) determined three specimens ( León 16802, Shafer 946, 2859 ) from the vicinity of Nuevitas in Camagüey as C. macroglossa . León 16802 appears to be C. macroglossa, Shafer 2859 appears to be C. salvatoris , and Shafer 946 , although lacking leaf sheath fibers, is also included there. Risco Villalobos et al. (2017) determined three specimens ( León 18422, Shafer 500, 2922 ) from north of Camagüey as C. miraguama . They were here determined as preliminary species C. macroglossa . Craft (2017) wrote that C. macroglossa has an early deciduous waxy layer on the adaxial surface of the leaves. He also reported hybrids between C. macroglossa and C. pseudorigida in Camagüey . Specimens of preliminary species C. macroglossa differ significantly from those of C. miraguama in only three variables (palman length, leaf segment length, pedicel length), with C. macroglossa having higher values for all variables ( t- test, P <0.05). In particular, specimens of preliminary species C. macroglossa approach those of C. miraguama var. havanensis in size, differing only in the longer leaf segments of C. macroglossa .