A Conservation Framework for the Critically Endangered Endemic Species of the Caribbean Palm Coccothrinax

A conservation framework for the Critically Endangered endemic species of the Caribbean palm Coccothrinax

B RETT J ESTROW,BRÍGIDO P EGUERO,FRANCISCO J IMÉNEZ,RAÚL V ERDECIA L ISBET G ONZÁLEZ-OLIVA,CELIO E. MOYA,WILLIAM C INEA,M.PATRICK G RIFFITH A LAN W. MEEROW,MIKE M AUNDER and J AVIER F RANCISCO-ORTEGA

Abstract With  threatened species ( categorized as plant exploration initiatives, taxonomic revisions, outreach, Critically Endangered and  as Endangered, sensu IUCN), and fundraising. The ultimate aim of this review is to provide Coccothrinax (c.  species) is the flagship palm genus for baseline information that will develop conservation synergy conservation in the Caribbean Island Biodiversity Hotspot. among relevant parties working on Coccothrinax conserva- Coccothrinax has its centre of taxonomic diversity in these tion in Cuba, Haiti and the Dominican Republic. Such colla- islands, with c.  endemic species. We present a conservation borations could also benefit through partnerships with framework for the  Critically Endangered species, found botanists working in other countries. in Cuba, Haiti or the Dominican Republic. Only two species Keywords Antilles, Arecaceae, IUCN, plant biodiversity, (C. jimenezii, C. montana) occur in more than one country red lists, taxonomy, tropical islands (Haiti and the Dominican Republic). Immediate threats include oil drilling and nickel mining, intrusion of saline water into soil, urban and agricultural development, low population recruitment, uncontrolled fires, interspecific hy- Introduction bridization, and unsustainable ethnobotanical practices. Coccothrinax bermudezii, C. borhidiana, C. crinita ssp. crini- alms are an iconic feature of the Caribbean landscape ta, C. leonis and C. spissa are not conserved in protected areas. Pand are associated with strong folk and ethnobotani- Coccothrinax bermudezii, C. jimenezii, C. leonis and C. nipen- cal traditions (Liogier, ; Leiva Sánchez, ). The sis are not part of ex situ collections. Based on results from a Caribbean Islands hold  endemic species of palms, with conservation project targeting C. jimenezii, we recommend Cuba () and Hispaniola () hosting the majority (Moya international cooperation between the three range states to & Leiva Sánchez, ; Roncal et al., ; Freid et al., ). implement integrative conservation management plans, The islands have two endemic palm genera: Hemithrinax (four species) from Cuba and the monotypic Zombia from Hispaniola. The most widespread genera with endemic species can be divided into two major groups. The first group BRETT JESTROW Fairchild Tropical Botanic Garden, Miami, USA (c.  endemic species in  genera) has few species in the BRÍGIDO PEGUERO and FRANCISCO JIMÉNEZ Jardín Botánico Nacional, Santo Domingo, Dominican Republic Caribbean Islands but many taxa from the Neotropical mainland. The second group comprises genera with their RAÚL VERDECIA Jardín Botánico Cupaynicú, Guisa, Granma, Cuba centre of diversity in these islands (c.  species in six gen- LISBET GONZÁLEZ-OLIVA Herbario Nacional de Cuba, Instituto de Ecología y Sistemática, La Habana, Cuba era: Coccothrinax, Copernicia, Gaussia, Pseudophoenix, Roystonea and Thrinax) and with few species occurring CELIO E. MOYA Sociedad Cubana de Botánica, Sección de Palmas, Boynton Beach, Florida, USA on the mainland (Zona et al., ; Roncal et al., ).

WILLIAM CINEA Jardin Botanique des Cayes, Cayes, Haiti Among them, Coccothrinax is the genus with the highest number of species in the Caribbean Islands. M. PATRICK GRIFFITH Montgomery Botanical Center, Coral Gables, Miami, USA − Coccothrinax is regarded as being taxonomically difficult ALAN W. MEEROW United States Department of Agriculture Agricultural  Research Service−Subtropical Horticultural Research Station, National and in need of further systematic revision (Zona, ). Two Germplasm Repository, Miami, USA prior taxonomic revisions focused on species from Cuba

MIKE MAUNDER The Eden Project, Bodelva, Cornwall, UK, and National Tropical (León, ) and the southern Greater Antilles (Bailey, Botanical Garden, Hawaii, USA ). Prior to these two works, Burret () provided a JAVIER FRANCISCO-ORTEGA* (Corresponding author) Department of Biological taxonomic account of the taxa on Cuba and Hispaniola. Sciences, International Center for Tropical Botany, Kimberly Green Latin ’  American and Caribbean Center, Cuban Research Institute, Florida However, Burret s( ) publication was based solely on International University, Miami, USA. E-mail [email protected] material collected by the Swedish botanist and plant collect- –  *Also at: Fairchild Tropical Botanic Garden, Miami, USA or Leonard Ekman ( ). In the s the Hungarian – Received  December . Revision requested  January . botanist A. Borhidi ( ) and the Caribbean botanist Accepted  April . First published online  September . O. Muñiz (–) described new species for Cuba,

resulting in a palm catalogue for the island (Muñiz & Islands, covering () relationships between taxonomy and Borhidi, ). Additional attempts were made to clarify conservation, () integrative species conservation manage- the taxonomy of Coccothrinax (Nauman & Sanders, a, ment, () in situ and ex situ conservation, () plant ex- b), based on phylogenetic analyses of morphological traits ploration, () the role of DNA data in understanding mostly observed from cultivated plants. phylogenetic relationships and patterns of genetic diversity, The current taxonomic uncertainties within this genus and () outreach and environmental education. arise, in part, from the work of Henderson et al. (), Based on our own conservation initiatives we assert that whose influential field guide for New World palms recog- international collaboration within the region provides the nized only  species. Nevertheless, most palm taxonomists best possible approach to deal with these six issues. Similar and botanists working on the Caribbean Islands (e.g. Moya conservation challenges are faced by other threatened ende- & Leiva Sánchez, ; Acevedo-Rodríguez & Strong, ; mics of the Caribbean Islands (reviewed by Maunder et al., Greuter & Rankin, ) do not follow that narrow taxo- , , and Carey et al., ), and we propose that con- nomic view and still recognize many species that were servation initiatives centred on Coccothrinax could be ap- synonymized by Henderson et al. (). For instance, plied effectively to other taxa. A main goal of this review is C. jamaicensis and C. proctorii are currently regarded as dis- to provide a framework for land managers, conservationists, tinct species endemic to Jamaica and the Cayman Islands, palm biologists and environmental educators that will offer a respectively (Proctor, ; Duno de Stefano & Moya, regional perspective for what we consider the flagship palm ). Likewise, botanists from Cuba and Hispaniola still ac- genus for conservation in the Caribbean Islands. This is par- cept Coccothrinax species that Henderson et al. () syno- ticularly relevant as this Biodiversity Hotspot comprises sev- nomized (e.g. González-Oliva et al., , ; Peguero eral countries with a range of historical and sociological et al., b; Verdecia, ). backgrounds (Maunder et al., , ). The current available taxonomic framework for Data collection pertinent to ex situ conservation of Coccothrinax is not the result of a single monographic en- these threatened species in botanic gardens, as well as deavour, and therefore as a working taxonomy for our con- literature reviews, were conducted during January – tribution we follow the classification system currently March . Field observations were conducted during accepted by most palm biologists and plant taxonomists –. working in the region (Fig. ). Therefore, we consider that   the genus has species, with of these endemic to the Ecology, biology, and conservation biology of the  Caribbean Islands (Fig. ). These island endemics are re- Critically Endangered species of Coccothrinax stricted to the Bahamas and the Greater Antilles (except Puerto Rico). The taxonomic placement of C. alta, thought Red List assessments of Coccothrinax species to be endemic to Puerto Rico and the Virgin Islands (Acevedo-Rodríguez & Strong, ), needs further study. Zona et al. () assessed the conservation status of species In this review, C. alta is considered a synonym of C. barba- of Coccothrinax of the Caribbean Islands using the IUCN densis following the recent treatment of Acevedo-Rodríguez Red List categories and criteria (IUCN, , ). Other & Strong (). Red List assessments have targeted Coccothrinax and other Two Caribbean species (C. argentata and C. barbadensis) endemic plant species of the Cayman Islands (Burton, are found both on the islands and on the mainland, and only ), Cuba (Rankin Rodríguez & Areces Berazaín, ; one species (C. readii) is restricted to the mainland (Fig. ). Berazaín Iturralde et al., ; González-Oliva et al., , The vast majority of Caribbean Island endemic species are ; González-Torres et al., ) and Hispaniola (Peguero single-island endemics, with the exception of C. fragrans &Jiménez,; Peguero et al., b). Of the  species of (found on Hispaniola and Cuba). Reports of the presence Coccothrinax endemic to the Caribbean Islands,  are cate- of C. jamaicensis on the small islands of Providencia, gorized as Endangered and  as Critically Endangered, with Colombia (Galeano-Garces, ), and Swan, Honduras  of the latter restricted to Cuba (Fig. ). (Nelson & Proctor, ), need to be validated by additional The Cuban endemic C. camagueyana was originally taxonomic studies. For intraspecific taxa, C. alexandri, described by Muñiz & Borhidi (), and is reported C. clarensis, C. crinita and C. salvatoris have one subspecies from the south of Sierra de Cubitas, Camagüey province each, and C. miraguama has three subspecies. (Méndez Santos et al., ). It was categorized as Given its high number of species, many with restricted Critically Endangered by Zona et al. (); however, its ranges in the Caribbean Islands, Coccothrinax provides a taxonomic status is unclear, and more recently it was cate- good case to study regional species conservation issues in gorized as Data Deficient (González-Oliva et al., ). Field the Caribbean. We provide a review of conservation chal- exploration led by CM in  failed to find this species in lenges and perspectives for the  Critically Endangered spe- the location from which it was originally reported (Moya cies of Coccothrinax that are restricted to the Caribbean et al., a).

FIG. 1 Geographical distribution and Red List status of Coccothrinax species: (a) Caribbean Island endemics; (b) species found both on the Neotropical mainland and on the islands. Red List categorizations (LC, Least Concern; VU, Vulnerable; EN, Endangered; CR, Critically Endangered; DD, Data Deficient) follow the IUCN (, ) guidelines and come from Zona et al. (), Burton (), González-Oliva et al. (, ) and Peguero et al. (b). Taxonomy and geographical distribution follow Moya & Leiva Sánchez (), Hoyos Fernández & Braun (), Acevedo-Rodríguez & Strong (), Proctor (), Duno de Stefano & Moya (), Freid et al. (), Peguero et al. (a,b), Verdecia (), Greuter & Rankin (), Moya et al. (b) and Jestrow (unpubl. data).

All of the Critically Endangered species have been as- Among the inland species, the Hispaniolan endemic signed to this category based on the criteria Bab or Bab C. montana occurs at the highest elevations, to , m; (IUCN, , ; Table ); i.e. they have an extent of occur- it was described from the Cordillera Central in the   rence ,  km or an area of occupancy ,  km . They Dominican Republic and the Massif de la Selle in Haiti are restricted to a single locality or to a fragmented site. (Burret, ). However, its distribution is poorly under- In addition, they show continuing decline in their extent stood, with populations reported from three disjunct sites of occurrence, area of occupancy, habitat quality, number in the Dominican Republic (Cordillera Septentrional, of sites, or number of mature individuals (IUCN, , Cordillera Central, and Sierra de Bahoruco) and two sites ). Eight of these species are shown in Plates  and . in southern Haiti (Massif de la Hotte and Massif de la Selle) (Burret, ; Judd, ; B. Peguero, unpubl. data). We have no recent data on population size and distribution Ecology, distribution and demography ranges for C. montana except from the Massif de la Hotte, where there are two populations of fewer than seven indivi- Of the  Critically Endangered species of Coccothrinax, duals each (B. Jestrow, unpubl. data), and from Bahoruco, most occur on limestone soils, with six Cuban taxa (five spe- where a single population with very few individuals is cies and one subspecies) occurring on serpentine soils known to exist. Fieldwork is needed to determine the taxo- (Table ). Only five species occur in coastal environments nomic and distribution status of this species both in Haiti (Table ). One of the two populations of C. jimenezii occurs and the Dominican Republic. Reports of the Hispaniola en- on the fringes of Lago Enriquillo in the Dominican demic C. spissa in northern Haiti (Bailey, ) need to be Republic, a hypersaline lake  m below sea level, which is confirmed (Henderson et al., ). currently expanding (Romero Luna & Poteau, ), and We have population estimates for only five of the  where the wild populations are threatened by saline intru- Critically Endangered species of Coccothrinax (Table ). sions (Peguero et al., a). Coccothrinax cupularis, C. leonis, C. nipensis and

Medio Ambiente y Recursos Naturales, ) and will be a major concern for in situ conservation of threatened Coccothrinax species. Tourism is an important economic activity in the Greater Antilles (Gayle & Goodrich, ), and is a growing industry particularly in Cuba (Hingtgen et al., ), where the sector is likely to undergo significant development following the easing of U.S. embargo restric- tions and the establishment of formal diplomatic relationships in July  (Hershberg & LeoGrande, ). This may result in an intensification of human pressure on Cuban ecosystems, with a potential negative impact on endemic palms (González-Torres et al., ). Five of the Critically Endangered taxa are not recorded FIG. 2 Distribution of the  Critically Endangered species of from protected areas (Table ), and those found within pro- Coccothrinax. (a) Cuban taxa; the three sites for C. cupularis tected areas of the Dominican Republic are also of major represent the distribution range of the highly fragmented conservation concern. From our field observations it ap- population of this species. (b) Hispaniolan taxa. pears that most of these protected areas (with the exception of Parque Nacional Nalga de Maco and Reserva Científica C. pauciramosa each have a single but dispersed population Quita Espuela) have poor conservation enforcement comprising scattered individuals; fieldwork is required to (Powell & Inchaustegui, ; Pasachnik et al., ). confirm their distribution range, population sizes and Eleven of the Critically Endangered taxa are cultivated in demographic status. botanic gardens (Table ). These collections are located Population recruitment has been recorded for six taxa in Cuba (five botanic gardens), the Dominican Republic (Table ). Based on preliminary field observations, both C. (one), Spain (one), and USA (two) and they are based yuraguana and C. pauciramosa also appear to have recruit- on wild collected germplasm. Four of the Critically ment in their single locations. Coccothrinax jimenezii has Endangered species of Coccothrinax are not held in ex situ low levels of recruitment, with seedlings found only in the conservation collections (Table ). Dominican Republic population (Peguero et al., a). There are no population size estimates for eight species. Conservation biology research

Conservation challenges Only four taxa have been the focus of conservation research that included population size studies, ethnobotanical sur- Our review identified six major threats (Table ). Two of veys, ecological studies or seed germination protocols these threats (population decline; urban, agricultural and (Table ). Coccothrinax jimenezii is the only Critically forest development) affect  of the Critically Endangered Endangered species that has been the subject of conserva- species. Threats from invasive alien plant species, unsustain- tion genetics studies (Jestrow et al., a). This work was able ethnobotanical exploitation, and uncontrolled fires are based on microsatellite markers and included the two relevant to seven taxa. Poaching for the horticulture trade known populations of the species, with the majority of indi- has been reported only for C. victorinii. Four species are at viduals being sampled. Levels of genetic differentiation be- risk because of nickel mining or oil drilling. One species tween the two populations of C. jimenezii are unusually high (C. crinita) has been detected to hybridize with at least compared to those reported among populations of other another species of the genus, and one is facing the threat palm species. This suggests that species delimitation within of saline intrusion in soils. Based on our experience in Coccothrinax needs to be re-evaluated, and a clear taxonom- botanic gardens, interspecific hybrids are produced easily ic framework for conservation initiatives is needed. Despite in Coccothrinax. These threats are likely to intensify in the their small size, both populations of C. jimenezii have a near future; for example, nickel mining and oil drilling are relatively large number of unique alleles and there is no major development priorities for the Cuban economy evidence of genetic bottlenecks. Only the population in (Wacaster et al., ). Furthermore, invasive alien plant Haiti exhibits evidence of inbreeding, with a moderately species are regarded as a major challenge for plant bio- high positive inbreeding coefficient value (Fis = . vs diversity management in the Caribbean (Maunder et al., Fis = . for the Dominican Republic population). , ; reviewed for Cuba by Oviedo Prieto & Recent habitat fragmentation coupled with long generation González-Oliva, , and González-Torres et al., ; re- times could explain the unusually high levels of genetic di- viewed for the Dominican Republic by Ministerio de versity in these Caribbean species (Jestrow et al., a).

Oryx, 2018, 52(3), 452–463 © 2017 Fauna & Flora International doi:10.1017/S0030605317000588 Downloaded from https://www.cambridge.org/core. IP address: 170.106.40.219, on 27 Sep 2021 at 03:19:29, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0030605317000588 https://www.cambridge.org/core/terms Downloaded from 5 .Jsrwe al. et Jestrow B. 456 https://www.cambridge.org/core . https://doi.org/10.1017/S0030605317000588

   . IPaddress: TABLE 1 Red List assessment criteria (sensu IUCN, , ), ecology, number of populations, demographics, and the in situ and ex situ conservation status of the Critically Endangered species of Coccothrinax.

IUCN criteria for 170.106.40.219 Critically Endangered No. of populations/ Ex situ conservation1 Taxon status Ecology No. of individuals In situ conservation (No. of individuals) C. baracoensis B1ab(i,ii,iii,v)+2ab(i, Scrub on serpentine, 10–200 m (Cuba)2 1/Unknown2 Cuchillas de Toa UNESCO Man & the JBT3 ii,iii,v)2 Biosphere Reserve , on C. bermudezii B1ab(ii,iii,v)+2ab(ii, Scrub & pine forest on serpentine, 500–700 m 14/, 10,0002 Unprotected Unprotected 27 Sep2021 at03:19:29 iii,v)2 (Cuba)2 C. borhidiana A4ace;B1ab(i,ii,iii,iv, Coastal scrub on limestone, 15–20 m (Cuba)6 14,5 /3386,7 Unprotected FTBG (4), JBM (7), Oryx v)+2ab(i,ii,iii,iv,v)6 JBN-C (3), JBT(15), MBC 08 23,452 52(3), 2018, , (25), PSCT (15) C. boschiana A4acd;Blab(i,ii,iii,iv, Coastal scrub on limestone, 5–200 m 15/Unknown9 Parque Nacional Sierra Martín García FTBG (5)3, JBN-DR (6), v)8 (Dominican Republic)9 MBC3, PSCT (11) , subjectto theCambridgeCore termsofuse,available at C. crinita ssp. B1ab(iii)+2ab(iii)2 Scrub on serpentine & human-modified sa- 15/c. 5002,10 Unprotected JBC (9), JBI (11), JBN-C 10 – –

463 crinita vannah, 20 40 m (Cuba) (70), JBT (20) C. crinita ssp. A2acde;B1ab(i,ii,iii,v) Semi-deciduous forest & human-modified 14/1762 Área Protegida de Recursos Manejados JBI (8), JBT (5) © 2 11 07Fua&FoaItrainldoi:10.1017/S0030605317000588 International Flora & Fauna 2017 brevicrinis +2ab(i,ii,iii,v);C1 savannah on limestone, 300 m (Cuba) Yaguanabo C. cupularis B2ab(ii,iii,v)2 Coastal, sub-coastal scrub & evergreen forest 14,5 /Unknown2 Área Protegida de Recursos Manejados JBN-C (11), JBI (7), JBT on limestone, 5–15 m (Cuba)2 Península de Zapata & Paisaje Natural (4) Protegido Guajimico C. jimenezii A2abcd+4abcd;B1ab Scrub on limestone fringe of hypersaline lake, 15/6112 Dominican Republic: Parque Nacional Lago Unprotected (i,ii,iii,iv,v)+2ab(i,ii,iii, & coastal scrub on limestone, 45 m (below sea Enriquillo e Isla Cabritos iv,v);C2a(i)8 level, Dominican Republic), 70–80 m (Haiti)12 C. leonis B2ab(ii,iii,v)2 Scrub & human-modified savannah on lime- 14/Unknown2 Unprotected Unprotected stone, 100–600 m (Cuba)2 https://www.cambridge.org/core/terms Downloaded from Oryx 08 23,452 52(3), 2018, , https://www.cambridge.org/core

Table 1 (Cont.) –

463 Taxon IUCN criteria for 1

© Critically Endangered No. of populations/ Ex situ conservation . https://doi.org/10.1017/S0030605317000588 07Fua&FoaItrainldoi:10.1017/S0030605317000588 International Flora & Fauna 2017 status Ecology No. of individuals In situ conservation (No. of individuals) C. montana B2ab(i,ii,iii,iv,v);C1 Montane evergreen forest, cloud forest, & pine 54/Unknown Haiti: Parc National Pic Macaya, Parc PSCT (2) 8 – . IPaddress: +2a(i) forest on limestone, 500 1700 m (Haiti & National Morne la Visite; Dominican Dominican Republic)13,14 Republic: Reserva Científica Quita Espuela, Parque Nacional Nalga de Maco C. nipensis B1ab(ii,iii,v)+2ab(ii, Scrub & pine forest on serpentine, 400–600 m 14/Unknown2 Parque Nacional La Mensura–Pilotos Unprotected 170.106.40.219 iii,v)2 (Cuba)2 C. pauciramosa B2ab(ii,iii,v)2 Scrub & pine forest on serpentine, 400–900 m 14/Unknown2 Parque Nacional La Mensura–Pilotos FTBG (4), JBT (5), PSCT (Cuba)2 (2) C. spissa A2ace+4ace;B1ab(i,ii, Scrub on limestone, 20–600 m (Dominican 34,5 /4,500 Unprotected JBN-DR (9), MBC (28), , on iii,iv,v)+2ab(i,ii,iii,iv, Republic)15,16 PSCT (9) 27 Sep2021 at03:19:29 v)8 C. victorinii A2c;B1ab(ii,iii,v)+2ab Coastal scrub on limestone, 0–3 m (Cuba)2 2/c. 2502 Parque Nacional Desembarco del Granma JBC (4) (ii,iii,v)2 C. yuraguana B1ab(ii)+2ab(ii)6 Scrub & pine forest on serpentine, 30–300 m 14/unknown6,17 Área Protegida de Recursos Manejados Mil JBN-C (7), JBT3 (Cuba)17 Cumbres  , subjectto theCambridgeCore termsofuse,available at Botanic gardens with ex situ collections based on wild collected individuals are as follows: FTBG, Fairchild Tropical Botanic Garden, USA; JBC, Jardín Botánico de Cupainicú, Cuba; JBI, Jardín Botánico de Cienfuegos; JBN-C, Jardín Botánico Nacional, Cuba; JBN-DR, Jardín Botánico Nacional, Dominican Republic; JBM, Jardín Botánico de Matanzas, Cuba; JBT, Jardín Botánico de las Tunas, Cuba; MBC, Montgomery Botanical Center, USA; PSCT, Palmetum de Santa Cruz de Tenerife, Spain  González-Oliva et al. ()  Ex situ collection has seedlings that have not yet been transplanted to the field  Highly fragmented population(s)  Population(s) exhibit(s) recruitment for framework Conservation  González-Oliva et al. ()  Enríquez Rodríguez et al. ()  Peguero et al. (b)  Mejía & García ()  Leiva Sánchez et al. ()  Suárez Oropesa ()  Peguero et al. (b)  Burret ()  Judd ()  Peguero et al. ()  Veloz et al. () Coccothrinax  Urquiola Cruz et al. () 457 458 B. Jestrow et al.

PLATE 1 Critically Endangered species of Coccothrinax: (a) C. jimenezii, (b) C. borhidiana (in ex situ collection of Montgomery Botanical Center), (c) C. spissa, (d) C. victorinii (in ex situ collection of Jardín Botánico de Cupainicú). Photograph credits: PLATE 2 Critically Endangered species of Coccothrinax (all in (a) Francisco Jiménez, (b) Patrick Griffith, (c) Scott Zona, and habitat): (a) C. yuraguana, (b) C. montana, (c) C. boschiana, (d) (d) Raúl Verdecia. C. jimenezii (heavily harvested to make brooms). Photograph credits: (a) Lisbet González-Oliva, (b–d) Brett Jestrow. Future genetic studies may find that at least some of the re- maining Critically Endangered species of Coccothrinax for which there are currently no population genetic data avail- Cuba, the Dominican Republic and Haiti. Conservation ef- able still harbour high genetic diversity. forts for C. jimenezii have been structured within the con- text of broad international cooperation led by the Jardín Botánico Nacional (Dominican Republic) and Jardin Conservation and research agenda Botanique des Cayes (Haiti) in association with botanists from South Florida (Fairchild Tropical Botanic Garden, The conservation management of threatened plants faces Montgomery Botanical Center, U.S. Department of particular challenges in the Caribbean Islands (Brown Agriculture, and Florida International University). There et al., ; Maunder et al., , ; Carey et al., ), have been extensive plant exploration expeditions in both most notably the diversity of political, cultural and socio- Haiti and the Dominican Republic, including demographic economic systems. We propose the following conservation studies and conservation assessments (Peguero et al., a). and research framework for Coccothrinax. Outreach outputs included the production of printed educa- tional material presented during the VI Simposio Flora de La  Establish a regional conservation approach Española in June at Santo Domingo, Dominican Republic. Furthermore, we have established joint initiatives Regional cooperation is required to utilize and coordinate for professional progression of young Caribbean botanists, the existing skills and resources of the various Caribbean na- including two workshops on plant systematics in Haiti, tions more effectively. With endemic Critically Endangered training of one graduate student from the Dominican species in three countries, a comprehensive regional ap- Republic at Florida International University (Rodríguez, proach to the conservation of Coccothrinax will require col- ), and developing grant-supported projects for palm laboration between managers and plant biologists from conservation through the Prince Bernhard Nature Fund,

Oryx, 2018, 52(3), 452–463 © 2017 Fauna & Flora International doi:10.1017/S0030605317000588 Downloaded from https://www.cambridge.org/core. IP address: 170.106.40.219, on 27 Sep 2021 at 03:19:29, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0030605317000588 https://www.cambridge.org/core/terms Downloaded from Oryx 08 23,452 52(3), 2018, , https://www.cambridge.org/core TABLE 2 Main conservation challenges and conservation biology research conducted for the  Critically Endangered species of Coccothrinax (Table ).

Nickel min- Urban, agricultural ing or oil Invasive Population & forest Uncontrolled Unsustainable Conservation biology – 463 Taxon drilling alien plants decline developments fires ethnobotany Other challenges research

https://doi.org/10.1017/S0030605317000588 C. baracoensis Nickel Yes Yes Yes Yes No 07Fua&FoaItrainldoi:10.1017/S0030605317000588 International Flora & Fauna 2017 C. bermudezii No1 No1 Yes1 Yes1 No1 No1 C. borhidiana Oil2,3 No2 Yes2 Yes2 Yes2 Yes2 Ecology3 4,5 4,5 4,5 4,5 4,5 4,5 4,5 . IPaddress: C. boschiana No No No No No No C. crinita ssp. No1 Yes1 No1 Yes1,6 No1,6 Yes6 Ethnobotany6, ecology7,8 crinita 1 1 1,9 1 1 1 10 9

170.106.40.219 C. crinita ssp. No Yes Yes Yes Yes Yes Hybridization Ecology brevicrinis C. cupularis No1 Yes1 Yes1 Yes1 Yes1 Yes1 C. jimenezii No4 No4,11 Yes4,11 No No4,11 Yes (Plate 2)11 Saline intrusion in soils11 Conservation genetics12, 11

, on ecology 1 1 1 1 1 1

27 Sep2021 at03:19:29 C. leonis No Yes Yes Yes Yes No C. montana No4 No4 Unknown Yes4 Unknown Unknown C. nipensis Nickel1 No1 Yes1 Yes1 Yes1 No1 C. pauciramosa Nickel1 Yes1 Yes1 Yes1 Yes1 No1 C. spissa13,14 No4 Unknown Yes13,14 Yes13,14 No13,14 Yes13,14 Ecology13,14 , ethno- botany13,14 , seed germination13,14 , subjectto theCambridgeCore termsofuse,available at C. victorinii No1 Yes1 Yes1 Yes1 No1 Yes1 Adult plants are transplanted to gardens in urban & resort developments C. yuraguana No2,15 No2,15 Yes2,15 Yes2,15 No2,15 Yes2,15   González-Oliva et al. ( ) for framework Conservation  González-Oliva et al. ()  Enríquez Rodríguez et al. ()  Peguero et al. (b)  Mejía & García ()  Martínez Betancourt & Miranda (–)  Leiva Sánchez et al. ()  Martínez Betancourt ()  Suárez Oropesa et al. ()  Suárez Oropesa ()  Peguero et al. (a)   Jestrow et al. ( b) Coccothrinax  Peguero et al. ()  Veloz et al. ()  Urquiola Cruz et al. () 459 460 B. Jestrow et al.

the International Palm Society, and the Mohamed bin Zayed the Dominican Republic have maintained a tradition of bo- Species Conservation Fund. tanical fieldwork and research, Haiti has not had sustained Botanists from the Dominican Republic have developed support for field botany. Both the Dominican Republic and successful protocols for seed germination of C. spissa Haiti have a single botanic garden, and are key to developing (Peguero et al., ; Veloz et al., ), which may also be a framework for future conservation initiatives. Through applicable for the Critically Endangered species from our inter-institutional collaborations (mostly supported by Cuba. Across the distribution range of Coccothrinax there the Mohamed bin Zayed Species Conservation Fund), are field botanists and palm taxonomists who are familiar members of our team have been exploring areas of Haiti sel- with its ecology and morphology. Their preliminary surveys dom visited since Ekman was there in the s (Jestrow suggest that hybridization detected in botanic gardens et al., b). Fieldwork led by RV resulted in the rediscov- is also common in areas where species overlap (Suárez ery of C. rigida on limestone cliffs near Sagua de Tánamo Oropesa, ). Caribbean island conservation biologists (Holguín province) in  (Verdecia et al., ). This spe- working with this genus have also gained experience regard- cies had not been seen since it was described by Grisebach ing unsustainable ethnobotanical practices, and are familiar () based on material collected by the American plant ex- with the challenges faced by land managers and administra- plorer Charles Wright (–). Recent plant exploration tors regarding implementation of effective protection in the activities led to the discovery of a new species, Coccothrinax protected areas of Cuba, the Dominican Republic and Haiti. spirituana, in Cuba by CM and RV (Moya et al., b). This Preliminary molecular results suggest that the microsatellite new species is currently known in a single locality in the loci used for C. jimenezii (Jestrow et al., a) are also ap- province of Sancti Spiritus in an area of serpentine soils. plicable to other species of this genus. The experiences Studies focusing on its demography and distribution range already gained by individual teams provide a framework are in progress. for development of a biodiversity strategy for There are four Coccothrinax species that have not been Coccothrinax in the region. located since they were originally discovered. The first, C. acunana, was described by León (), and plants of this species were reported to occur only in the highest Establish a scientific framework for conservation mountains of Cuba, near Pico Turquino, in the province of Santiago de Cuba. The species is categorized as A phylogenetic analysis and taxonomic review would clarify Vulnerable on the Red List for the Cuban flora (González- the conservation status of Coccothrinax taxa. For instance, Torres et al., ); however, its conservation status needs to C. jimenezii was described as a new species in  (Mejía be reassessed. The second species, C. microphylla, is another & García, ) but molecular studies revealed uncertainties Cuban endemic, described by Muñiz & Borhidi (). It ap- in the taxonomy of the two populations of this species pears to be restricted to limestone cliffs of Abra de Mariana (Jestrow et al., a), highlighting the need for a compre- in the lowlands of Guantanamo province. The third species, hensive taxonomic revision for Coccothrinax. The morpho- C. concolor, is known from only one herbarium collection, logical phylogenetic study of the genus undertaken by by Ekman in . The species was described by Burret Nauman & Sanders (a,b) identified three major groups, () and reported from low-elevation areas with volcanic each one including Critically Endangered species, with high soils near Jacmel, southern Haiti (Bailey, ). The fourth levels of homoplasy for the morphological characters. species, C. camagueyana, not only has an uncertain taxo- Molecular phylogenetic reconstructions for Coccothrinax nomic status but has not been found since it was described (Roncal et al., ) did not resolve major clades. (Moya et al., b). However, the conservation genetics study of C. jimenezii (Jestrow et al., a) demonstrated the utility of microsatellite DNA markers for insights into both genetic diversity and taxonomy. There are still many gaps in knowledge of Ensure wild populations are protected in situ, with the biology of the species, and there have been no studies supporting ex situ collections focused on breeding systems, plant–animal interactions, The Critically Endangered species of Coccothrinax are fa- competition from invasive alien plant species, or population cing a variety of threats. Five of the species are not present viability analyses. in protected areas, and four are not included in ex situ germplasm collections. An integrative approach to conservation  Assess and monitor wild populations is the most immediate priority (Esposito, ; Petriello & Wallen, ). We are aware that in many cases official pro- There is a need to expand botanical field work in the range tected areas are not sufficiently resourced to ensure viability nations, particularly in Hispaniola, where there are still of wild populations; however, both Cuba and the under-collected areas, mostly in Haiti. Although Cuba and Dominican Republic have protected areas with effective

conservation management. Ex situ conservation is necessary de Santa Cruz de Tenerife, and Scott Zona for sharing in some cases; for instance, the Dominican Republic insights concerning taxonomic challenges associated with population of C. jimenezii is facing an immediate threat Coccothrinax and allied genera. This is contribution num- from the increasing water level of the hypersaline lake ber  from the Tropical Biology Program of Florida Lago Enriquillo, and the small population in Haiti is International University. overharvested. Establishing an ex situ resource should be part of an integral approach for conservation planning that involves Author contributions all stakeholders (Pritchard et al., ; McGowan et al., ). Ideally this resource would be held primarily in the BJ, BP, FJ and WC led field work in Hispaniola and reviewed range state, and should ensure adequate genetic representa- conservation challenges for Haiti and the Dominican tion (Oldfield, ). There are botanic gardens located out- Republic. CEM, RV and LGO led field work in Cuba and re- side the Caribbean Islands that have ex situ conservation of viewed conservation challenges for Cuba. MPG made sig- palms as a mission priority (Maunder et al., ). Several of nificant contributions pertinent to ex situ conservation. these (e.g. Fairchild Tropical Botanic Garden, Montgomery AWM provided insights regarding conservation genetics. Botanical Center, and Palmetum of Tenerife) are already BJ, JFO and MM developed the proposed conservation collaborating with botanists from Hispaniola and Cuba to and research agenda and wrote the first draft of the article. develop representative living collections that hold duplicates BJ and JFO led grant proposals to undertake this project, of living germplasm collections and can mitigate issues re- and designed and developed the structure of this study. garding limited space and resources for ex situ conservation.

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