Molecular Systematics of Threatened Seed Plant Species Endemic in the Caribbean Islands Nora Oleas, Brett Jestrow, Michael Calonje, Brígido Peguero, Francisco Jiménez, Rosa Rodríguez-Peña, Ramona Oviedo, Eugenio Santiago-Valentín, et al. The Botanical Review ISSN 0006-8101 Volume 79 Number 4 Bot. Rev. (2013) 79:528-541 DOI 10.1007/s12229-013-9130-y 1 23 Your article is protected by copyright and all rights are held exclusively by The New York Botanical Garden. This e-offprint is for personal use only and shall not be selfarchived in electronic repositories. If you wish to self-archive your article, please use the accepted manuscript version for posting on your own website. You may further deposit the accepted manuscript version in any repository, provided it is only made publicly available 12 months after official publication or later and provided acknowledgement is given to the original source of publication and a link is inserted to the published article on Springer's website. The link must be accompanied by the following text: "The final publication is available at link.springer.com”. 1 23 Author's personal copy Bot. Rev. (2013) 79:528–541 DOI 10.1007/s12229-013-9130-y Molecular Systematics of Threatened Seed Plant Species Endemic in the Caribbean Islands Nora Oleas1,2,3 & Brett Jestrow2 & Michael Calonje1,4 & Brígido Peguero5 & Francisco Jiménez5 & Rosa Rodríguez-Peña1,2,5 & Ramona Oviedo6 & Eugenio Santiago-Valentín7 & Alan W. Meerow8 & Melissa Abdo2 & Michael Maunder1 & M. Patrick Griffith4 & Javier Francisco-Ortega1,2,9 1 Department of Biological Sciences, Florida International University, Miami, FL 33199, USA Kushlan Tropical Science Institute, Fairchild Tropical Botanic Garden, Coral Gables, Miami, FL 33156, USA 3 Current Address: Centro de Investigación en Biodiversidad y Cambio Climático, Universidad Tecnológica Indoamérica, Quito, Ecuador 4 Montgomery Botanical Center, Coral Gables, Miami, FL 33156, USA 5 Departamento de Botánica, Jardín Botánico Nacional, Santo Domingo, República Dominicana 6 Herbario, Instituto de Ecología y Sistemática, 10800 La Habana, Cuba 7 Jardín Botánico de Puerto Rico, Universidad de Puerto Rico, San Juan, PR 00936, USA 8 United States Department of Agriculture, Agriculture Research Service, National Germplasm Repository, Miami, FL 33158, USA 9 Author for Correspondence; e-mail: ortegaj@fiu.edu 2 Published online: 27 August 2013 # The New York Botanical Garden 2013 Abstract A review of available Caribbean Island red-lists species (CR and EN categories based on the IUCN guidelines from 2001, and E category established according to the IUCN guidelines from 1980) is presented. A database of over 1,300 endemic species that are either Critically Endangered or Endangered sensu IUCN was created. There are molecular systematic studies available for 112 of them. Six of these species (in six genera) are the only members of early divergent lineages that are sister to groups composed of a large number of clades. Seven of the species (in seven genera) belong to clades that have a small number of taxa but are sister to species/genus-rich clades. Ten of the species (in six genera) are sister to taxa restricted to South America or nested in clades endemic to this region. Fifty-seven of the species (in 35 genera) are sister to Caribbean Island endemic species. Erigeron belliastroides, an Endangered (EN) Cuban endemic, is sister to the Galapagos genus Darwiniothamnus. The phylogenetic placement of four of the threatened species resulted in changes in their taxonomic placement; they belong to polyphyletic or paraphyletic genera. Keywords Biodiversity hotspots . Tropical islands . Phylogenies . Spermatophyta . IUCN . Red lists Electronic supplementary material The online version of this article (doi:10.1007/s12229-013-9130-y) contains supplementary material, which is available to authorized users. Author's personal copy Caribbean Island Threatened Seed Plants 529 Introduction Caribbean Island Endemic Red Lists Revealing the phylogenetic and taxonomic placement of threatened species helps to establish conservation priorities and to increase public awareness for conservation (Purvis et al., 2005). The Caribbean Islands are a major focus for conservation because they are one of the 34 biodiversity hotspots (Smith et al., 2004). These islands have been severely impacted by human activities, and in the last 500 years there has been a dramatic reduction of the original vegetation cover primarily because of habitat loss (Maunder et al., 2008, 2011). The Caribbean Islands harbor a native flora of approximately 12,000 species of vascular plants, and over 8,000 of them are endemic to the region (Santiago-Valentín & Olmstead, 2004). There is no comprehensive red list for this endemic flora based on the IUCN (2001) categories of threat. Available information on the conservation status of these Caribbean endemics is primarily limited to the few taxa found in the Red List of Threatened Species of the World Conservation Union (IUCN, 2011) and those listed in 14 publications focusing on threatened plant species of Antigua, Barbuda, British Virgin Islands, Cayman Islands, Cuba, Dominican Republic, French Lesser Antilles, Jamaica, and Puerto Rico (Borhidi & Muñiz, 1983; Kelly, 1988; Government of Jamaica, 2000; Rankin-Rodríguez & Areces-Berazaín, 2003; Gargominy, 2003; Pollard & Clubbe, 2003; Berazaín-Iturralde et al., 2005; González-Torres et al., 2007; Burton, 2008; Pratt & Lindsay, 2008; Urquiola Cruz et al., 2010; Centro Nacional de Biodiversidad, 2011; Peguero & Jiménez, 2011; U.S. Fish & Wildlife Service, 2011). The most recent red lists are from Cayman Islands, Cuba and Dominican Republic (Rankin-Rodríguez & Areces-Berazaín, 2003; Berazaín-Iturralde et al., 2005; GonzálezTorres et al., 2007; Burton, 2008; Centro Nacional de Biodiversidad, 2011; Peguero & Jiménez, 2011; U.S. Fish & Wildlife Service, 2011), and these are the only ones that follow the latest conservation categories and criteria developed by the World Conservation Union (IUCN, 2001). The remaining studies do not distinguish between the “Critically Endangered” and “Endangered” categories and they place those taxa belonging to these two groups within the broad “Endangered” rank [as defined by IUCN (1980)]. In order to differentiate between the “Endangered” category sensu IUCN (2001) or sensu IUCN (1980), in our paper the code “E” will be used when the “Endangered” conservation status was developed based on the criteria proposed by IUCN in 1980. We use the acronym “EN” when this category refers to the most recent guidelines established by IUCN in 2001. It is noteworthy that the “Critically Endangered” category is not part of the threat ranks delineated by IUCN in 1980. In a previous review Torres-Santana et al. (2010) reported that approximately 1,553 of the single-island-endemic vascular plant species belong to the Vulnerable (VU), Endangered (EN), or Critically Endangered (CR) IUCN categories. Unique Molecular Phylogenetic Patterns Revealed in Our Studies—Review Aims Because of the geographical location of our institutes and their research history, studies pertinent to plant biodiversity (including genetic diversity and molecular phylogenetics) of the Caribbean Islands have been a main focus of our teams (e.g., Author's personal copy 530 N. Oleas et al. Bogler & Francisco-Ortega, 2004; Santiago-Valentín & Olmstead, 2004; FranciscoOrtega et al., 2007; Meerow et al., 2007, 2012; Santiago-Valentín & FranciscoOrtega, 2008; Maunder et al., 2008, 2011; Andrus et al., 2009; Namoff et al., 2010a). We found that previous molecular systematic research conducted in our laboratories demonstrated that at least six threatened species have: (1) unusual biogeographical connections or (2) are part of early divergent lineages that are sister to a large assemblage of genera or species. One of the most striking examples is provided by Feddea cubensis. This is a Critically Endangered species endemic to the unique nickel-rich serpentine soil environments of eastern Cuba. This taxon is sister to the Heliantheae, one of the largest clades of the family [ca. 482 genera and 5,700 species, Cariaga et al. (2008)]. Based on both molecular and morphological data Cariaga et al. (2008) accommodated this genus in its own unigeneric tribe, the Feddeeae. A similar pattern is followed by Microcycas, a Critically Endangered unispecific genus, confined to a few “mogotes” of western Cuba, which is sister to Zamia (Bogler & Francisco-Ortega, 2004; Zgurski et al., 2008). The latter is the largest genus of the Zamiaceae, and the second largest in the order Cycadales (Osborne et al., 2012). Zamia is found from Bolivia northward through tropical America to southeastern United States and the Caribbean Islands. The third example is provided by Erigeron bellidiastroides (Asteraceae), an Endangered (EN) herbaceous species confined to quartz sand areas of western Cuba which is sister to Darwiniothamnus (Asteraceae), a genus endemic to the Galápagos Islands that has a woody habit (Andrus et al., 2009). The fourth example concerns the recently described genus endemic to Hispaniola: Garciadelia (Euphorbiaceae). This genus has four species, all of them are threatened and it is sister to a clade composed of two Caribbean Island genera, Lasiocroton and Leucocroton (Euphorbiaceae). These two genera have over 30 species, the majority of which are in Leucocroton and are endemic to serpentine areas of Cuba suggesting that the species growing on limestone areas to have the ancestral ecological conditions (Jestrow et al., 2008, 2010, 2011). Studies within this group showed the value of molecular data to address taxonomic questions. The fifth study from our teams is for a Critically Endangered palm species endemic to Haiti. Attalea crassispatha is the only member of the genus to occur in the Caribbean Islands, and a molecular phylogeny based on seven unlinked nuclear loci strongly supports this species as sister to a clade that includes the Scheelea and Orbignya groups (Meerow et al., 2009). It is estimated that these two groups have over 45 species. These molecular phylogenetic studies have identified a sixth case in which an assemblage of four endemic genera [Coeloneurum (unispecific), Espadaea (unispecific), Goetzea (two species), and Henoonia (unispecific)] form a monophyletic group within the Solanaceae that is recognized as an important part of the subfamily Goetzeoideae (Olmstead et al., 2008). One of the species of this generic alliance, G. elegans, is Endangered (EN) (Caraballo-Ortiz & Santiago-Valentín, 2011), and it is sister to the Hispaniolan endemic G. ekmanii O. E. Schulz (Olmstead et al., 2008). In this study we provide a review of molecular phylogenies that have included Caribbean Island endemics that are most threatened [E, EN and CR categories of the IUCN (1980, 2001) (Fig. 1)]. Our study aimed to: (1) evaluate to what extent these high conservation priority endemics have been the subject of molecular systematic studies, (2) to know if the phylogenetic patterns detected in our laboratories are also reported in other molecular systematic studies that included threatened species, and (3) to determine Author's personal copy Caribbean Island Threatened Seed Plants 531 Fig. 1 Images of threatened Caribbean Islands endemics that have been the subject of molecular phylogenetic studies: a Croton trigonocarpus (photo B. Van Ee); b Pereskia quisqueyana (photo R. Briones); c Phyllomelia coronata (photo J. Molina); d Rhytidophyllum grandiflorum (photo F. Jiménez); e Portlandia albiflora (by courtesy of Fairchild Tropical Botanic Garden, photo J. Valdés) the impact that molecular studies are having in our current understanding of the systematic relationships of these taxa. We are aware that under the system established by IUCN (2001) the term “Threatened” applies to taxa belonging to three categories: Vulnerable (VU), Endangered (EN), and Critically Endangered (CR). However, in our review the concept of “Threatened” refers only to the Critically Endangered (CR) and Endangered (EN and E) categories as defined by IUCN (1980, 2011). Author's personal copy 532 N. Oleas et al. Methods Using the available red lists for the Caribbean Islands (see above) we compiled a list of threatened endemic species. This initial list did not include the work of Borhidi and Muñiz (1983) as we found many inconsistencies between the conservation categories of this study and those provided by most recent studies (i.e., Rankin-Rodríguez & ArecesBerazaín, 2003; Berazaín-Iturralde et al., 2005; González-Torres et al., 2007; Urquiola Cruz et al., 2010; Centro Nacional de Biodiversidad, 2011). Based on this list of threatened endemic species we conducted additional bibliographic searches to locate published molecular phylogenies and additional conservation information that included these Endangered/Critically Endangered taxa [sensu IUCN (1980, 2001)]. As primary data base/bibliographic resources for these searches we used the ISI Web of Knowledge (Thomson Reuters, 2011), the GenBank NIH Genetic Sequence Data Base (Benson et al., 2011), the Kew Bibliographic Data Base (Royal Botanic Gardens – Kew, 2011), and relevant conservation reviews, monographs and taxonomic treatments (e.g., Hágsater & Dumont, 1996; Oldfield, 1997; Donaldson, 2003; Zona et al., 2007; Jestrow et al., 2010). These studies provided us with details concerning the phylogenetic position of the Caribbean Islands endemics. We looked into statistical support for relevant nodes, taxonomic sampling of these studies, number of DNA independent markers that were utilized to generate the phylogenies, and the actual phylogenetic placement of the threatened species. Results of Review Based on the list of threatened species that we prepared for this study we estimate that there are over 1,300 seed-plant species that are either Endangered or Critically Endangered [sensu IUCN (1980, 2001)] on the Caribbean Islands. This represents approximately 16 % of the endemic flora of vascular plants. However, we believe this to be a “conservative” value because our review did not include ferns and the available red lists do not cover the complete Caribbean flora. For instance, 30 % of the Cuban flora still needs to be assessed (Leiva & González-Torres, 2010). We anticipate that this figure will increase once there are completed IUCN red lists for: (1) Haiti, the Bahamas, most of the Lesser Antilles and (2) the component of the endemic flora not included for the other islands. The three genera with the highest number of threatened species are Eugenia (Myrtaceae) (54 species), Lepanthes (Orchidaceae) (27 species), and Pilea (Urticaceae) (31 species). Including studies from our research teams, there are molecular topologies available for 112 of the threatened species (ca. 1 % of the total endemic flora) [Appendices S1-S3 (supplementary online material)]. However, molecular studies for four species [i.e., Acidocroton trichophyllus (Euphorbiaceae), Brunfelsia portoricensis (Solanaceae), Synapsis ilicifolia (Bignoniaceae), and Zanthoxylum ekmanii (Rutaceae)] were not informative for our research as these studies had the endemic species as outgroups or to test particular phylogenetic methods or to investigate the utility of DNA barcode markers in floristic studies (Appendix S3). Molecular phylogenies for 57 of the threatened species are based on a single linked DNA region, or have low statistical support for the relevant nodes. With few exceptions Author's personal copy Caribbean Island Threatened Seed Plants 533 (i.e., McDowell & Bremer, 1998; Lavin et al., 2003; Steinmann et al., 2007; BécquerGranados et al., 2008; Michelangeli et al., 2008; Roncal et al., 2008; Salazar & Nixon, 2008; Schaefer et al., 2008; Van Ee et al., 2008; Graham, 2010) most of the reviewed studies did not focus on molecular systematics of Caribbean groups but on developing phylogenies for particular genera or suprageneric taxa. Molecular Phylogenies: Biogeographical Patterns Our review of studies from other research groups found that a large proportion of the threatened endemics have sister relationships with other West Indies endemics (53 species in 33 genera: Acacia (Fabaceae), Calycogonium (Melastomataceae), Calyptranthes (Myrtaceae), Catesbaea (Rubiaceae), Cestrum (Solanaceae), Cinnamodendron (Canellaceae), Coccoloba (Polygonaceae), Croton (Euphorbiaceae), Elekmania (Asteraceae), Euphorbia (Euphorbiaceae), Exostema (Rubiaceae), Gaussia (Arecaceae), Gesneria (Gesneriaceae), Ginoria (Lythraceae), Guettarda (Rubiaceae), Harnackia (Asteraceae), Henriettea (Melastomataceae), Isidorea (Rubiaceae), Lescaillea (Asteraceae), Matelea (Apocynaceae), Mecranium (Melastomataceae), Miconia (Melastomataceae), Pachyanthus (Melastomataceae), Pereskia (Cactaceae), Phialanthus (Rubiaceae), Phyllomelia (Rubiaceae), Poitea (Fabaceae), Portlandia (Rubiaceae), Rhytidophyllum (Gesneriaceae), Rondeletia (Rubiaceae), Scrophularia (Scrophulariaceae), Solanum (Solanaceae), and Tolumnia (Orchidaceae), see Appendix S3). A few of the threatened endemics are nested inside clades restricted to South America or sister to lineages from this region (10 species in 6 genera: Crescentia (Bignoniaceae), Doerpfeldia (Rhamnaceae), Elekmania, Gesneria, Rhytidophyllum, and Styrax (Styracaceae), see Appendix S3). The rest of the threatened species had sister relationships with species/clades with a widespread distribution or restricted to North or Central America. Our review did not find any species that was sister to an Old World clade; however, the clade that includes of the Cuban monotypic genus Doerpfeldia and the South American genus Ampelozizyphus (Rhamnaceae) (two species) was sister to a genus from Madagascar, Bathiorhamnus (Rhamnaceae) (seven species) (Richardson et al., 2000) (see below). The second case that showed a connection with the Old World was found in Ginoria, the ITS phylogeny supports Tetrataxis (Lythraceae) (unispecific genus endemic to Mauritius Island, Indian Ocean) as part of a clade that has five of the species of Ginoria. The threatened G. jimenezii is sister to this clade (Graham, 2010). Molecular Phylogenies: Sister Relationships Our review was concordant with the results obtained in our laboratories and showed that three of the threatened species are sister to clades with a large number of species or genera. They included the recently described endemic and monotypic genus Linnaeosicyos (Cucurbitaceae), an Hispaniolan taxon that is sister to the Sicyueae clade (Cucurbitaceae), a group composed of 125 species in 16 genera (Schaefer et al., 2008). The Cuban unispecific genus Lepturidium (Poaceae) follows a similar pattern as it is sister to the Mulhlenbergiinae (Poaceae) [~176 species in a single genus, Muhlenbergia (Poaceae)] (Peterson et al., 2010). The third example is provided by Leptogonum (Polygonaceae), a monotypic Hispaniolan genus (Burke et al., 2010) that is sister to the six-tepaled clade of the Author's personal copy 534 N. Oleas et al. subfamily Eriogonoideae (Polygonaceae) (Burke et al., 2010). This group includes nine genera (~300 species). Six of the threatened species belong to monophyletic groups that have a small number of taxa but are sister to species/genus-rich clades. For instance the Endangered (EN) Magnolia minor (Magnoliaceae) together with four other species of this genus form a monophyletic assemblage that is sister to the rest of the genus. Three species-poor palm genera with threatened endemics appear to follow the same pattern. A phylogenetic analyses based on morphological and molecular data supports Gaussia (five species, three of them endemic to the Caribbean Islands) as sister to Chamaedorea (77 species) (Cuenca et al., 2009). Likewise a molecular systematic study for the subfamily Ceroxyloideae shows Pseudophoenix (four species, three of them endemic to the Caribbean Islands) as sister to the tribes Ceroxyleae and Phytelepheae (Trénel et al., 2007). These two tribes encompass seven genera with 39 species. Within the subfamily Coryophoideae, Colpothrinax (three species, one of them endemic to Cuba) is sister to a clade that includes of nine genera (36 species). Within the Rhamnaceae, the Cuban genus Doerpfeldia, Ampelozizyphus, and Bathiorhamnus (see above) form a lineage that is sister to a clade composed of four tribes with 22 genera (Richardson et al., 2000). The last example is provided by Domingoa (Orchidaceae) (three species endemic in the Caribbean Islands, including the Endangered (EN) D. nodosa), as this genus together with Homalopetalum (Orchidaceae) (seven species, genus found in Mexico, Central and South America, and Greater Antilles) form a monophyletic group (with ~12 species) that is sister to a clade that has a great portion of the taxonomic diversity of the tribe Laeliinae (Orchidaceae), including large genera such as Cattleya (ca. 40 species) and Epidendrum (ca. 1,500 species) (Van Den Berg et al., 2009). Two other examples concern threatened species that are sister to clades with few continental species but that have unique morphological features. For instance Tillandsia arizajuliae (Bromeliaceae) is sister to a clade that has the pseudobulbous taxa of this genus (12 species, none of them endemic to the Caribbean Islands) (Chew et al., 2010). A second case concerns Acacia anegadensis and A. roigii. These two Critically Endangered species form a monophyletic group that is sister to a clade with 13 species (Gómez-Acevedo et al., 2010). Ten of these species form a unique monophyletic group known as the “Myrmecophillous group.” This group has those New World species that maintain a symbiosis with several species of ants. The Myrmecophillous species display morphological modifications associated with ant mutualism, such as domatia, extrafloral nectaries, and leaflet tips that provide food for the ants. Among the threatened species belonging to endemic genera, the unispecific genus Phyllacanthus (Rubiaceae) is sister to a clade with four genera, two of them, Isidorea (Rubiaceae) and Portlandia, endemic to the Caribbean Islands (Motley et al., 2005). Also within the Rubiaceae, the endemic genus Phialanthus (18 species, two species included in the study) is part of a clade that include two additional endemic genera: Ceratopyxis (monotypic) and Schmidtottia (16 species, one included in the study) (Motley et al., 2005). Molecular Phylogenies: Taxonomic Implications Molecular phylogenies have had an impact on the current taxonomic placement of some of these threatened species. For instance, based on DNA and morphological Author's personal copy Caribbean Island Threatened Seed Plants 535 evidence the Critically Endangered species Trichosanthes amara (Cucurbitaceae) was transferred to Linnaeosicyos (Schaefer et al., 2008, see above). Likewise molecular and micro/macromorphological data led Jestrow et al. (2010) to describe Garciadelia (see Introduction above) to accommodate Leucocroton leprosus, a Critically Endangered species from Haiti known from only two herbarium collections from the early 20th century. Jestrow et al. (2010) described three additional species in this genus; all of them are threatened. Results from molecular systematic studies have resulted in taxonomic changes that have been relevant in three additional genera. In the Euphorbiaceae, DNA phylogenies clearly show that the Cuban genus Moacroton (Euphorbiaceae) is paraphyletic with four species of Croton from the mainland (Van Ee et al., 2008); therefore, the Critically Endangered M. trigonocarpus is currently named C. trigonocarpus. Molecular phylogenies for subtribe Laeliinae (see above) reveal the Caribbean genus Domingoa as paraphyletic with Nageliella (Orchidaceae), the most recent taxonomic treatment for this group merges both genera in a single one: Domingoa (Pridgeon et al., 2005). Within the Rubiaceae, Rova et al. (2009) reported Rondeletia (two threatened species) as paraphyletic with the Caribbean Island genus Stevensia (Rubiaceae) (11 species), this resulted in the recent merging of the latter within Rondeletia (Borhidi, 2010). There are molecular phylogenetic data for 14 threatened species belonging to the Melastomataceae. These species are placed in six genera within the tribes Miconieae or Henrietteeae. Only two of these genera (i.e., Henriettea and Mecranium) appear to be monophyletic (Michelangeli et al., 2008; Penneys et al., 2010). In contrast, Calycogonium, Miconia, Pachyanthus, and Tetrazygia are polyphyletic (Michelangeli et al., 2008; Bécquer-Granados et al., 2008) and it is likely that some of the Caribbean endemics currently in these four genera will need new taxonomic placements (Michelangeli, pers. comm). Most of these genera have a large number of species and are taxonomically “difficult” to resolve. Cinnamodendron provides another example of a polyphyletic genus with a Critically Endangered species (i.e., C. cubense). Molecular systematic studies conducted by Salazar and Nixon (2008) show that Caribbean Island species of Cinnamodendron will need to be placed in a different genus. The generitype, C. axillare, is a Brazilian endemic that belongs to a “South American” clade that does not have any of the Caribbean Island endemics. Matelea (Cactaceae) [generitype: M. palustris (Farr et al., 1979)] and Pereskia [generitype: P. aculeata (Leuenberger, 1986)] follow similar phylogenetic patterns. Caribbean island endemics belonging to these polyphyletic genera are not placed in the clades that have their generitypes. In contrast, West Indian threatened endemics in the polyphyletic genera Elekmania [generitype: E. barahonensis (Nordenstam, 2006)], Guettarda [generitype: G. speciosa (Jarvis, 2007)], Mollugo (Molluginaceae) [generitype: M. verticillata (Jarvis, 2007)] belong to the clades that have the types of these genera. The phylogenetic patterns of three other genera with endemic species appear to have taxonomic implications and we believe that will need to be addressed in the future. The placement of the Critically Endangered species Phyllomelia coronata within the Cuban genus Mazaea (Rubiaceae), makes the latter paraphyletic (Manns & Bremer, 2010). Likewise, Basiphyllaea (Orchidaceae) [with the Endangered (EN) species B. carabiaiana] is paraphyletic with the North American genus Hexalectris (Orchidaceae) (Sosa, 2007). Ginoria (with three threatened species) is also Author's personal copy 536 N. Oleas et al. paraphyletic, as the clade that has all of the species of this genus also includes Tetrataxis (Graham, 2010). Discussion Our review finds that a small proportion of the endemic threatened flora of the Caribbean islands has been the subject of molecular systematic. The general finding was that most rare West Indian seed-plants have evolved from lineages largely or entirely confined to the Caribbean Basin. However, in this study we have also shown that several of the endemic species have unique phylogenetic patterns. These species have a high priority for conservation not only because of their Endangered or Critically Endangered status [sensu IUCN (1980, 2001)] but because of their phylogenetic placements. Few of the molecular studies of the threatened species support links between the Caribbean Islands and the Old World. However; in a previous review we reported that at least three endemic genera, Narvalina (Asteraceae), Selleophytum (Asteraceae), and Siemensia (Rubiaceae) are sister to different clades from Polynesia and the South Pacific (Namoff et al., 2010b) suggesting phytogeographical links with these remote regions. The Tropical Sea Way connected the Caribbean Sea and the Pacific Ocean until the late Pliocene (ca. 4 mya; Coates, 1997). This dispersal route was enhanced by the Circumtropical Current, which ran from the coasts of Brazil to the Pacific (Collins et al., 1996). Once the Panama Isthmus was formed, this geographical connection disappeared and this current became an important part of the Gulf Stream. The Caribbean Islands have had a complex environmental history and have been severely affected by eustatic/isostatic sea-level changes, climatic changes, volcanism, and migration of lithospheric plate boundaries (Iturralde-Vinent & MacPhee, 1999; Iturralde-Vinent, 2006). It is suggested that prior to the middle Eocene (~ 49 mya) the islands went through several waves of emergence and submergence, but it is only after this geological epoch that the Caribbean Islands started a period of “sustained emergence” (Graham, 2003a, 2003b). According to this model, the middle Eocene is considered as the starting point for the current fauna and flora of the Antilles (Iturralde-Vinent & MacPhee, 1999; Graham, 2003a, 2003b). Thus, the flora of the Caribbean Islands is relatively “recent” when compared with that of South and North America. Despite this relative young age, our review shows that several of the threatened species are part of lineages that are sister to groups with a large number of taxa and clades. They represent the only surviving elements of these early divergent lineages. A full review concerning to what extent this phylogenetic pattern is followed by other endemics is outside the scope of this study; however, a review focusing on the endemic genera (Francisco-Ortega et al., 2007) found that Verhuellia, a genus with three species endemic in Cuba and Hispaniola, is sister to the rest of the Piperaceae (Wanke et al., 2006; Samain et al., 2008). The significance of this finding is that this family is one of the early-branching species-poor groups of angiosperms (as defined by Soltis et al., 2005). Other endemic genera follow similar patterns, including the endemic legumes Arcoa, and Pictetia as they are the only representatives of early divergent lineages in the Umtiza and the Ormocarpum clade, respectively (Lavin et al., 2001; Wojciechowski et al., 2004; Bruneau et al., 2008; Lavin & Beyra Matos, 2008). Likewise, two Antillean orchid genera, Dilomilis and Neocogniauxia form a clade sister to the subtribe Pleurothallidinae Author's personal copy Caribbean Island Threatened Seed Plants 537 (Van Den Berg et al., 2009). Similarly the Caribbean genus Ekmanianthe (Bignoniaceae) is sister to Tabebuia (Bignoniaceae) (Grose & Olmstead, 2007) and the Caribbean genus Bellonia (Gesneriaceae) is sister to the tribe Gesnerieae (Martén-Rodríguez et al., 2010). Our review has shown that the results obtained by us regarding the unique phylogenetic placement of some of the threatened endemic species of the Caribbean Islands are also found in other studies. They stress the conservation relevance of the Caribbean Island Biodiversity Hotspot because of its: (1) high number of endemics, (2) high rate of habitat loss, and (3) unique branches of the tree of life. Acknowledgments We dedicate this paper to Dr. Victoria Sosa in recognition for her contributions to the advancement of molecular systematics in Latin America. Nora Oleas thanks Fairchild Tropical Garden and Florida International University for their support to attend the “10th Congreso Latinoamericano de Botánica”. We are grateful to Dr. Fabián Michelangeli and Prof. Richard Olsmtead for their insights concerning taxonomy of the Melastomataceae and Bignoniaceae, respectively. Thanks are due Dr. Larry Noblick and Dr. Scott Zona for their help on the biogeography of Caribbean palms, and to Dr. Johan Rova for helping in tracking taxonomic literature pertinent to Rondeletia. Dr. Brian Boom and an anonymous referee critically reviewed the manuscript. This is contribution 209 from the Tropical Biology Program of Florida International University. Literature Cited Andrus, N., A. Tye, G. Nesom, D. Bogler, C. Lewis, R. Noyes, P. Jaramillo & J. Francisco-Ortega. 2009. 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