Revista de Biología Tropical ISSN: 0034-7744 [email protected] Universidad de Costa Rica Costa Rica

Martín, Alberto; Díaz, Yusbelly; Miloslavich, Patricia; Escobar-Briones, Elva; Guerra-García, José Manuel; Ortiz, Manuel; Valencia, Bellineth; Giraldo, Alan; Klein, Eduardo Regional diversity of Amphipoda in the Caribbean Sea. Revista de Biología Tropical, vol. 61, núm. 4, diciembre, 2013, pp. 1681-1720 Universidad de Costa Rica San Pedro de Montes de Oca, Costa Rica

Available in: http://www.redalyc.org/articulo.oa?id=44930117012

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Alberto Martín1, Yusbelly Díaz1, Patricia Miloslavich1, Elva Escobar-Briones2, José Manuel Guerra-García3, Manuel Ortiz4, Bellineth Valencia5, Alan Giraldo5 & Eduardo Klein1 1. Departamento de Estudios Ambientales. Centro de Biodiversidad Marina. Universidad Simón Bolívar. Caracas, 89000. Venezuela; [email protected], [email protected], [email protected], [email protected] 2. Instituto de Ciencias del Mar y Limnología. Unidad Académica Ecología y Biodiversidad Acuática. Universidad Nacional Autónoma de México. Ciudad de México, 04510. México; [email protected] 3. Departamento de Zoología. Facultad de Biología. Universidad de Sevilla. Sevilla, 41012. España; [email protected] 4. Facultad de Estudios Superiores Iztacala. Laboratorio de Crustáceos. Universidad Nacional Autónoma de México. Los Reyes Iztacala, 54090. México; [email protected] 5. Departamento de Biología. Facultad de Ciencias Naturales y Exactas. Universidad del Valle. Cali, 10077. Colombia; [email protected], [email protected]

Received 21-xi-2012. Corrected 05-Iv-2013. Accepted 06-v-2013.

Abstract: Diversidad regional de Amphipoda en el Mar Caribe. The order Amphipoda is one of the most diverse within Peracarids, and comprises 6 950 described marine species. Amphipod research in the Caribbean Sea began in the late 1 800s, but has increased significantly since 1 980. In this study, we analized the amphipod biodiversity (Caprellidea, Gammaridea, Hyperiidea, and Ingolfiellidea) of the Caribbean Sea. For this, we compiled available data on species diversity of marine amphipods (data bases: WoRMS and OBIS and published species lists) into a comprehensive taxonomic list by country for the ecoregions of the Caribbean. Additionally, we anal- ized the relative contribution of each country to regional diversity and the rate of discovery of new species. The Caribbean amphipod fauna is composed of 535 species within 236 genera and 73 families for the higher taxon. The Western Caribbean ecoregion holds the largest diversity (282 species), while the Eastern Caribbean recorded the lowest one (73). Mexico and Venezuela recorded the largest number of species with 266 and 206, respectively. Twelve countries had less than 50 species. The richest suborder is the Gammaridea with 381 species followed by the suborder Hyperiidea with 116. From the total of 535 amphipod species reported for the Caribbean region, 218 have the Caribbean as the holotype locality, and 132 are endemic (about 25% of the total). Areas of higher diversity seem to be concentrated along the Mexican Caribbean, Cuba and the Northern coast of South America (Venezuela- Colombia); however, such pattern is most likely reflecting local collection efforts and taxonomic expertise rather than actual distribution. Knowledge of amphipod species is mostly limited to shallow, near-shore waters, with little information available on the deep sea fauna. Regional research priorities for this group should be focused on completing shallow water coastal inventories of species in Central America and the Greater and Lesser Antilles. In addition, sampling the deep sea ecosystems should follow along with other particular habitats such as anchialine cave systems. It is also neccessary to increase ecological research efforts, mainly in some speciose suborders, including the Caprellidea and Hyperiidea, known to exhibit high diversity in other tropical localities. Rev. Biol. Trop. 61 (4): 1681-1720. Epub 2013 December 01.

Key words: Amphipoda, diversity, Caribbean Sea, endemism.

The order Amphipoda is one of the most Winfield, Ortiz & Escobar-Briones, 2009). diverse and globally distributed within the They fulfill an important role in the trophic superorder Peracarida after order Isopoda. dynamics of many communities, as they break- Its members occur worldwide in terrestrial, down plant material and detritus, and also freshwater, subterranean and marine environ- function as grazers, filter feeders, suspension ments and in virtually all habitats (Foster, feeders, commensals, micropredators, scav- LeCroy, Heard & Vargas, 2009; LeCroy, Gasca, engers, or as prey for other carnivore species

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013 1681 (Bellan-Santini, 1998; Bellan-Santini & Ruffo, species. Jan Hendrik Stock explored exten- 1998; LeCroy et al., 2009). sively between 1973 and 1978 more than 25 Traditionally, the Amphipoda were divided islands in the Antilles in search of stygobionts, into four suborders: the Gammaridea Latreille, which resulted in a few new descriptions 1802; the Caprellidea Leach, 1814; the Hype- (Stock, 1976, 1977a,b, 1979). More recently, riidea Milne-Edwards, 1830; and the Ingolfiel- since the late 1960s, several authors have lidea Hansen, 1903 (Barnard, 1969; Bousfield, provided important publications on Caribbean 1973, 2001; Lincoln, 1979; Martin & Davis, amphipods, including descriptions of new spe- 2001). Despite, Myers & Lowry (2003) rees- cies, re-descriptions or additional information, tablished the suborder Corophiidea Leach, taxonomic lists and inventories, biogeographi- 1814, and placed the caprellideans sensu lato cal and ecological data, and illustrated keys for within it, we decided to use the traditional clas- specific taxa (see Table 1 for a comprehensive sification system because these categories will reference list by major topics). likely change with new phylogenetic studies. In this paper, we analyzed the amphipod Although there is a significant amount biodiversity (Caprellidea, Gammaridea, Hype- of scientific literature regarding Caribbean riidea, and Ingolfiellidea) of the Caribbean Sea. amphipod species in a variety of books, jour- Our goal was to carry out a spatial analysis of nals, and reports, many of these publications the data that allowed us to determine gaps in are restricted to one or a few species and many knowledge and to test if differences/similarities of them are old and therefore, not easily avail- in composition between ecoregions are due to able. Comprehensive works on amphipods of ecological and biogeographic factors or to geo- the region are relatively rare. graphic inequality in sampling efforts. We hope The first records and descriptions of amphi- that this paper will contribute to raise aware- pods in the Caribbean Sea, began with the col- ness about the value of taxonomic inventories lections of gammarideans and corophiideans of and to indicate how much and where scientific the H.M.S. Challenger between 1873-76, when sampling is needed, for a better understanding Stebbing described Ampelisca abyssicola from of the Caribbean amphipod biodiversity. Culebra Island, St. Thomas (Lesser Antilles) (Stebbing, 1888). Further descriptions included Material and methods Shoemakerella cubensis (as Lysianax cubensis) from Cuba (Stebbing, 1897), and Talorchestia Study area: The Caribbean Sea is a large fritzi (Stebbing, 1903), Chelorchestia costari- sea, closed off to the West and to the South by cana (Stebbing, 1906) and Pseudorchestoi- the Americas, and bordered to the North and dea biolleyi (Stebbing, 1908) (as Orchestoidea to the East by the islands chains of the Greater biolleyi) from Costa Rica. Other expeditions and Lesser Antilles (Spalding, 2004). It covers to the Caribbean, particularly to Puerto Rico about 2 754 000km2, has a volume of nearly and Cuba, provided new amphipod descrip- 6.5x106km3, borders over 13 500km of coast- tions (Shoemaker, 1933a,b, 1934, 1935, 1948). line, and is home to 12 island countries, nine Hyperiid amphipods are also poorly studied continental countries, and 12 island dependen- in the Caribbean, and the knowledge we have cies (to France, The Netherlands, U.K., and comes from some historical larger surveys U.S.). Towards the East and Northeast, the (Shoemaker, 1945, 1948; Fage, 1960; Lewis closely spaced chain of islands, banks, and sills & Fish, 1969; Shih, 1969; Moore & Sander, of the Antilles Islands Arc separate the Carib- 1977, 1979; Gasca & Shih, 2001, 2003; Gasca bean from the Atlantic Ocean, and act as a & Suarez-Morales, 2004; Gasca, 2009a,b) and sieve for the inflow of Atlantic water (Andrade mainly from the upper layers (0-200m) of the & Barton, 2000), whereas towards the North- water column. Globally, the ingolfiellideans west it is linked to the Gulf of Mexico by the have a very low biodiversity, with only 44 Yucatan Channel (Miloslavich, Diaz, Klein,

1682 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013 Table 1 Main topics developed in amphipod research in the Caribbean and bibliographical sources

CUADRO 1 Principales tópicos de investigación sobre anfípodos en el Mar Caribe con sus fuentes bibliográficas

Taxonomic Topic group References (Suborder) Species Caprellidea Ortiz, Guerra-García & Lalana (1998); Guerra-García (2003a); Guerra-García et al. (2006). description Gammaridea Stebbing (1888, 1897, 1903, 1906, 1908); Chevreux (1901); Shoemaker (1926, 1933a,b, 1934, 1948); Stephensen (1933, 1947, 1948); Ruffo (1950, 1954); Barnard (1954, 1960, 1961, 1962, 1964, 1972, 1973); Mateus & Mateus (1966); Holsinger & Peck (1968); Myers (1968, 1977, 1978, 1979); Holsinger (1977, 1990, 1992); Ortiz (1976a,b,c,d,e, 1980,1991); Just (1977, 1983, 1984); Stock (1977a,b, 1978, 1980, 1981, 1983, 1985); McKinney (1978, 1979, 1980); McKinney et al. (1978); Ortiz & Gómez (1979); Ortiz & Lalana (1980, 1989, 1994a,b, 1995, 2002a,b,c); Thomas (1983, 1997); Thomas & Barnard (1983, 1985, 1986, 1989, 1990, 1991a,b); Ortiz & Nazábal (1984a,b, 1988); Ortiz & Veledo (1985); Barnard & Agard (1986); Barnard & Thomas (1987, 1990); Vonk & Stock (1987); Vonk (1988, 1989, 1990, 1991); Ciavatti (1989); Bellan-Santini (1990, 1997); Ortiz, Lalana & López (1992); Ortiz, Lalana & Beltran (1993); LeCroy (1995); Ortiz & Pérez (1995); Lowry & Stoddart (1997); Ortiz & Lemaitre (1997); Jaume & Wagner (1998); Ortiz, Lalana & Lio (1999); Krapp-Schickel & Ruffo (2000, 2001); Martín, Ortiz & Atienza (2000, 2001); Ortiz, Lalana & Sánchez (2000); Ortiz, Martín & Atienza (2000); Ruffo et al. (2000); Berge et al. (2001); Lazo-Wasem & Gable (2001); Ortiz, Cházaro-Olvera & Winfield (2001); Ortiz, García-Debras & Lalana (2002, 2003); Ortiz, Lalana & Varela (2002, 2004, 2006, 2007a,b,c, 2008); Van der Ham & Vonk (2003); Thomas & Klebba (2006, 2007); Ortiz & Thomas (2007); Coleman (2011); Ortiz, Varela & Lalana (2011). Ingolfiellidea Stock (1976, 1979). Inventories, Caprellidea Díaz et al. (2005); Guerra-García et al. (2006). taxonomic lists Gammaridea Shoemaker (1935); McCain (1968); McCain & Steinberg (1970); McKinney (1977); Ortiz and reports (1978, 1979a,b, 1983); Galan (1983, 1984); Stoner & Lewis (1985); Lagarde (1987); Ortiz & Lalana (1993, 1996, 1998, 2002a, 2006); Ortiz & Lemaitre (1994); Villaroel & Graciani (1997); Díaz & Martín (2000a,b,c,d, 2001a,b, 2003, 2005); Martín, Atienza & Díaz (2000); Atienza & Martín (2001); Díaz (2001); Lazo-Wasem & Gable (2001); Martín (2001a,b, 2003, 2007); Ortiz, Lalana & Lio (2001); Escobar-Briones et al. (2002); Martín, Ortiz & Díaz (2002); Ayala & Martín (2003); Martín & Díaz (2003); Van der Ham & Vonk (2003); Varela et al. (2003); Jara (2007); Ortiz, Lalana & Varela (2007c, d,e, 2008); Ortiz, Martín & Díaz (2007); LeCroy et al. (2009); López et al. (2009). Hyperiidea Shoemaker (1945, 1948); Fage (1960); Lewis & Fish (1969); Shih (1969); Moore & Sander (1977, 1979); Ortiz, Lalana & Guevara (1990); Gasca & Shih (2001, 2003); Ortiz, Lalana, Varela & Leal (2003); Gasca & Suárez-Morales (2004); Ortiz, Lopeztegui, Lalana & Varela (2006); Gasca (2009a,b); LeCroy et al. (2009). Ingolfiellidea Stock (1977b, 1979); Vonk & Schram (2003). Biology, Caprellidea Guerra-García et al. (2006). ecology and Gammaridea Galan (1983, 1984); Sánchez (1985); Thomas (1997); Martín & Díaz (2003); Díaz & biogeography Martín (2005); Jara (2007). Hyperiidea Lewis & Fish (1969); Moore & Sander (1977, 1979). Ingolfiellidea Stock (1977b, 1979); Vonk & Schram (2003). Caprellidea Díaz et al. (2005); Guerra-García et al. (2006). Taxonomic Gammaridea Galan (1983,1984); LeCroy (1995); Thomas (1997); Ortiz, Alvarez & Winfield (2002); keys Ortiz, Martín, Winfield, Díaz & Atienza (2004).

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013 1683 Alvarado, Díaz, Gobin, Escobar-Briones, Cor- Using these estimates, we calculated the index tés, Weil, Bastidas, Robertson, Zapata, Martín, of relative contribution for each country where Kazandjan & Ortiz, 2010). the larger the number of species listed, the We compiled and analyzed the amphipod higher the alpha diversity value. Based on the species composition of the Caribbean region number of endemic species by country, the rel- from the Southeastern coast of the Yucatan evance of this country to the relative contribu- Peninsula in Mexico (Quintana Roo State) tion to the gamma diversity changed. to Belize, Guatemala, Honduras, Nicaragua, Additionally, in order to determine the Costa Rica, and Panama down to Colombia and rate of description of new species, we arranged Venezuela in South America, and also along the the number of species records per year and Greater Antilles (Cuba, the Cayman Islands, produced cumulative curves for all subor- Jamaica, Hispaniola, Navassa and Puerto ders. Based on the fact that the quality of Rico), the Lesser Antilles (St. Thomas, Tortola, taxonomic inventories depends on the avail- Anegada, Jost Van Dyke, Virgin Gorda, St. ability of identification guides and taxonomic Croix, Anguilla, St. Martin, St. Barthelemy, experts, our review also related these sources Saba, St. Eustatius, Barbuda, St. Kitts, Nevis, to each higher taxon. Antigua, Monserrat, Guadeloupe, Dominica, Martinique, St. Lucia, St. Vincent, Grenadines Results and Grenada), down to the Southern islands of Barbados, Tobago, Trinidad and the Nether- Diversity and distribution: There are lands Antilles (Bonaire, Curaçao, Aruba). 535 species identified within 236 genera and 73 families that have been reported for the Diversity inventories: To compile the Caribbean Sea. The most speciose taxon are available data on marine Amphipoda spe- the Gammaridea with 387 species (72.34%), cies diversity in the Caribbean, we used two followed by the Hyperiidea with 116 species approaches. The first approach summarized the (21.68%), the Caprellidea with 25 (4.67%) and number of marine amphipod species for Capre- the Ingolfiellidea with 7 (1.31%) species. A llidea, Gammaridea, Hyperiidea and Ingolfiel- complete taxonomic listing of these species and lidea using georeferenced species records from their distributions within the Caribbean is pre- the open-access databases (WoRMS and OBIS) sented in a supplementary table (Appendix 1). and from local, country, and regional check- Regarding the regional distribution of these lists. The second approach was to produce species, each taxon apparently exhibits its own revised species lists for the group by country spatial pattern of species richness, as no com- (where information was available). mon distribution pattern was evident (Table 1 and 2). The Western Caribbean ecoregion had Data analysis: The presence/absence data the largest diversity (282 species), followed by for each Amphipoda taxon was used to cal- the Southern Caribbean (230 species), while culate a Sørensen distance matrix. This sym- the Eastern Caribbean ecoregion had the low- metrical index excludes double absences in the est diversity (73 species). When examined calculation. Only countries with 10 or more by country, Mexico and Venezuela were the amphipod species were included in the matrix. most species-rich with 266 and 206 species To analyze the relative contribution of each respectively, followed by Cuba with 168 spe- country to the Caribbean regional diversity, we cies. The species number of the three countries used the contribution partition analysis pro- combined accounted for 49.72% of the total posed by Lu, Wagner & Chen (2007), using amphipod species reported for the Caribbean. species richness as a measure of diversity. The Twelve countries had less than 50 species gamma (regional) diversity is the sum of the (Table 2). When the richness was standardized alpha (local) and beta (interlocal) diversities. by coastline length, the average number of

1684 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013 species per 100 kilometers of country-coast- The Gammaridea were clearly the most line was 5±8species/100km, with Mexico hav- speciose taxon in all countries and ecoregions. ing the highest richness (29.4species/100km), The Southern Caribbean ecoregion (represent- and Nicaragua the lowest (0.2species/100km). ed by Venezuela, Aruba, Curaçao, Bonaire, When standardized by the Economic Exclusive Trinidad and Tobago) encompassed the largest Zone (EEZ) coverage, the average number of gammaridean diversity (188 species), while the species was 0.05±0.08species/100km2 of EEZ, Eastern Caribbean ecoregion (Lesser Antilles) with Mexico having the highest number of spe- had the lowest diversity (54 species) (Table cies (0.30species/100km2) of EEZ, followed by 2). When examined by country, Venezuela had Costa Rica (0.12species/100km2), and Nicara- the highest diversity with 169 species, fol- gua having the lowest (0.001species/100km2). lowed by Mexico and Cuba, with 140 and

Table 2 Number of Caribbean Amphipoda species per country within the five ecoregions

CUADRO 2 Riqueza de especies del orden Amphipoda por país en cada ecorregión del Mar Caribe

Amphipoda Suborders EEZ 2 )* Ecoregion 2 spp./100km Total species Total (km)* Coastline No. spp./100km of coastline (km EEZ Caprellidea Gammaridea Hyperiidea Ingolfiellidea

WESTERN CARIBBEAN 13 156 112 1 282 México (Yucatán) 13 140 112 1 266 905 29.4 88 874 0.30 Belize 0 37 0 0 37 516 7.2 35 351 0.10 Guatemala 0 1 0 0 1 150 0.7 1 642 0.06 Honduras 0 2 0 0 2 644 0.3 18 151 0.01 SOUTH-WESTERN CARIBBEAN 9 111 0 0 120 Nicaragua 0 1 0 0 1 503 0.2 87 930 0.001 Costa Rica 0 31 0 0 31 212 15 25 090 0.12 Panama 0 33 0 0 33 1 295 2.5 143 442 0.02 Colombia 9 67 0 0 67 1 760 3.8 472 891 0.01 SOUTHERN CARIBBEAN 12 188 24 6 230 ABC* 0 22 0 5 27 Venezuela 12 169 24 1 206 2 800 7.4 470 666 0.04 Trinidad & Tobago 0 8 0 0 8 362 2.2 74 199 0.01 GREATER ANTILLES 11 177 28 0 216 Cuba 10 131 27 0 168 3 755 4.5 350 751 0.05 Cayman Islands 0 4 0 0 4 160 2.5 Jamaica 0 5 3 0 8 1 022 0.8 258 137 0.003 Hispaniola 0 18 0 0 18 3 051 0.6 382 658 0.004 Puerto Rico 1 30 0 0 31 501 6.2 205 529 0.02 EASTERN CARIBBEAN 3 54 16 0 73 Lesser Antilles 3 54 16 0 73

(ABC*=Aruba, Bonaire and Curacao and*=Coastline information taken from The world fact book (http://www.cia.gov/ library/publications/the-world-factbook/geos) and Economic Exclusive Zone (EEZ) information was taken from www. seaaroundus.org/eez/eez.aspx)

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013 1685 131 species respectively, while Nicaragua and the Mexican Caribbean where pelagic samples Guatemala had the lowest with only one spe- have frequently been obtained (Table 2). Spe- cies each. Gammaridea are composed of 49 cies of this taxon were reported only in five of families (Table 3), of which the Melitidae and the 17 countries analyzed herein. In the Carib- Hadziidae were the most diverse, with 57 and bean Sea, the hyperiids are represented by 19 31 species, respectively, while 10 families were families (Table 3), of which the Scinidae holds only represented by one species (Appendix 1). 17 species, followed by the Lestrigonidae and The most eurytopic species within the region Platyscelidae with 14 species each (Appendix were Parahyale hawaiensis (eight countries), 1). The genus Scina is the most diverse with 16 Ampelisca lobata and P. fascigera (seven coun- species. The most eurytopic species is Lestrigo- tries); in contrast 208 of the 387 species of nus bengalensis, occurring in five countries, in Gammaridea were reported as unique to one contrast to 70 of the 116 species in this subor- country. The genus Ampelisca was the most der reported in one country. speciose, with 23 species. For Ingolfiellidea, only seven species The highest diversity was recorded in within one genus and one family were reported Caprellidea for the Western Caribbean ecore- for the Caribbean Sea (Table 3, Appendix 1). gion (13 species), the lowest occurred in the Of these species, six were recorded in the Eastern Caribbean (three species) (Table 2). Southern Caribbean ecoregion (Aruba, Cura- Mexico displayed the highest diversity val- çao, Bonaire and Venezuela) and the remaining ues, with 13 species, representing 52.0% of species in the Western Caribbean (Mexico, reported for the Caribbean, followed by Ven- Yucatan) (Table 2), and all were reported only ezuela with a total of 12 species and Cuba for one country. with 10 species. This taxon, composed of five families (Table 3), includes the Caprellidae Advances in regional knowledge: The and Pariambidae, the most diverse taxa, with first amphipod descriptions were made by ten and seven species respectively (Appendix Linnaeus in 1758 (Gammarus locusta, Gam- 1). In the Caribbean, the genus Caprella and marus pulex, and Cyamus ceti). Between 1800 Deutella were the most speciose with five spe- and 1850, an average of 6.28 species were cies reported, respectively. The most eurytopic described per year and then, between 1850 species were Paracaprella pusilla, Deutella and 1900, this number increased to 44.64 new incerta and Hemiaegina minuta, represented in descriptions per year, and to 51.04 between four countries, respectively. 1850 and 1900. Between 1900 and 1950, the The highest species richness (112 spe- number of descriptions almost doubled to cies) in the Hyperiidea occurred in the Western 97.70 species per year, and since 1950, about Caribbean ecoregion, mainly from Yucatan in 103.18 species are described per year (Fig. 1A). The accumulation curve of world species dis- Table 3 coveries (Fig. 1B) shows no sign of leveling Species, genera and families of Caribbean amphipods off, indicating that a full inventory of these taxa is still far from being completed despite CUADRO 3 Número de species, generos y familias de los anfípodos two centuries of efforts. In the Caribbean presentes en el Mar Caribe region, the description of species began in 1888 with Ampelisca abyssicola by Stebbing and Suborder Family Genus Species increased at a slow rate changing in trend with Caprellidea 5 2 25 a substantial accumulation of species in the Gammaridea 49 179 387 1980s (Fig. 2). As in the global graph, species Hyperiidea 18 44 116 description in the Caribbean region is far from Ingolfiellidea 1 1 7 reaching an asymptote, and at present, 218 spe- Total 73 236 535 cies of amphipods (40.75% of the Caribbean

1686 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013 total) have been described from this region. Of Venezuela was aggregated into one group these, 60.55% (132 species) are endemic to the (Fig. 3). The similarity between Mexico and Caribbean region, while the remaining 39.45% Venezuela is based on 119 common species (86 species) have been reported elsewhere, (22.24% of the total species in the region), mainly for the Gulf of Mexico and Florida. while the similarity between Cuba and Mexico is based in 99 common species (18.50% of the Similarity analysis: In the cluster tree total species in the region; Table 4). Mexico, generated with the Sorensen Distance Index, Cuba and Venezuela share species with other the amphipod fauna of Cuba, Mexico and countries in the region (14, 13 and 10 countries,

A Number of Species 0 50 100 150 200 250

1750 1800 1850 1900 1950 2000

Year

B Accumulated Number of Species Accumulated 0 2000 4000 6000 8000 10000

1750 1800 1850 1900 1950 2000 Year

Fig. 1. (A) Amphipoda species discoveries around the world; (B) Cumulative Amphipoda species discoveries around the world.

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013 1687 A Number of Species 0 5 10 15 1750 1800 1850 1900 1950 2000 Year

B Number of Species 0 50 100 150 200

1750 1800 1850 1900 1950 2000 Year

Fig. 2. (A) Amphipoda species discoveries in the Caribbean Sea; (B). Cumulative Amphipoda species discoveries in the Caribbean Sea. respectively). Countries with the highest alpha diversity values, i.e. Puerto Rico, Costa Rica, diversity were Mexico, Venezuela, and Cuba and Belize (Fig. 4). and those contributed proportionally to the gamma diversity. The Lesser Antilles contrib- Discussion uted to regional diversity, despite having fewer species than i.e. Colombia. Aruba, Curaçao The crustaceans, along with the mollusks, and Bonaire contributed more to the regional are the most speciose taxonomic groups among diversity than countries with higher alpha the marine fauna, with 44 950 reported species

1688 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013 (Bouchet, 2006). Despite the lack of a larval De Broyer, Lowry, Jazdzewski & Robert, 2007; stage, and therefore, with limited dispersal Zeidler & De Broyer, 2009), and Japan (544 capability, the order Amphipoda is a relative- species; Fujikura, Lindsay, Kitazato, Nishida ly species-rich group within the crustaceans, & Shirayama, 2010). In order to establish man- representing 15% of the total if we consider agement and conservation plans, hotspots of that the number of marine amphipod species endemism should be identified, the species in is 6 950 (Vader, 2005), or up to 26% if we need of protection prioritized, and their popu- consider that the number of species is 11 780 lation size estimated. This should also lead to (WoRMS, 2012). recommendations to upgrade their current con- In this sense, the 535 amphipod species servation status. There is currently little pub- reported in the Caribbean region represent a lished recognition that the diversity associated small contribution to global diversity. Despite with the islands exists at all and as a result, not this apparent low richness, large percentage of only its conservation has received little atten- unique species, represent an important contri- tion, but these species have often been affected bution to the endemic regional diversity. When by diverse, ongoing anthropogenic activities. comparing this amphipod diversity across geo- Gammaridea are the most diverse taxon graphical regions, the Caribbean has a larger among the Amphipoda, with at least 8 300 diversity than the Patagonian Shelf (98 species) species according to WoRMS (2012). In the and the Tropical West Atlantic (107 species) Caribbean, gammaridean amphipods were also (Miloslavich, Klein, Díaz, Hernández, Big- the most diverse group (387 species), repre- atti, Campos, Artigas, Castillo, Penchaszadeh, senting about 72% of the regional biodiver- Neill, Carranza, Retana, Díaz de Astarloa, sity. One of the most comprehensive reviews Lewis, Yorio, Piriz, Rodríguez, Yoneshigue- in marine gammaridean was carried out by Valentin, Gamboa & Martín, 2011), but lower Barnard & Karaman (1991) who provided than Australia (888 species; Butler, Rees, Bee- not only keys, diagnoses and species lists for sley & Bax, 2010), the South American Hum- the group, but also discussed their geographic boldt Current region (860 species; Miloslavich distribution based on thermal zones and depth, et al., 2011), the Southern Ocean (797 species; and some aspects of endemicity. According to these authors, in terms of generic diversity, the Gammaridea are mainly a cold water group, whereas tropical regions seem to be poor in tropical endemic genera. A higher richness in cool and temperate environments, contrasting HIS ABC with poor diversity in the tropics, is also a general trend in freshwater amphipods (Vain- PRC LAN

BEL ola, Witt, Grabowski, Bradbury, Jazdzewski CRC PAN

COL & Sket, 2008). In the tropics, even if many Distance new species are expected to be described, it is unlikely that any of these ones will be associ- ated with a new genus (Barnard & Karaman, 1991). In the Caribbean region, four new CUB

VEN genera have been described since Barnard’s MEX 0.4 0.5 0.6 0.7 0.8 0.9 1 and Karaman’s work in 1991: Ottenwalderia Fig. 3. Cluster tree based on species presence/absence (Jaume & Wagner, 1998), Miramarassa (Ortiz, matrix and generated with the Sørensen distance index. Lalana & Lio, 1999) Pleusiroides and Tantena PRC Puerto Rico, LAN Lesser Antilles, HIS Hispaniola, ABC Aruba, Bonaire and Curacao, CRC Costa Rica, PAN (Ortiz, Lalana & Varela, 2007a,b respectively). Panama, BEL Belize, COL Colombia, CUB Cuba, MEX Other important components of this taxon in Mexico, VEN Venezuela. the Caribbean are the subterranean and cave

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013 1689 taxa, whose present distribution and biogeo- Metaprotella hummelincki, Tritella chibcha, graphic pattern reflect their origin. In the case and Cubadeutella cavernicola), while about of marine and brackish species, it has been 40% of the species have an eurytopic dis- suggested that they may have derived from tribution either in the tropical and the tem- epigean marine or brackish water ancestors as perate marine ecosystems (Caprella andreae, they adapted to fluctuating sea levels, and even C. danilevskii, C. equilibra, C. penantis, C. some freshwater species may have derived scaura, Pseudaeginella biscaynensis, Hemiae- from marine and brackish species that were gina minuta, Paracaprella pusilla and Phtisica caught inland during regression of seawater marina; McCain & Steinberg, 1970). In the case (Holsinger, 1994; Biernbaum, 1996), in both of D. californica and Hemiproto wigleyi, these cases leading to speciation by isolation. species are distributed also in the East Pacific Caprellidea are a much less diverse taxon (Guerra-García, 2003a; Díaz, Guerra-García with 451 species worldwide and 25 species & Martín, 2005), while D. incerta is distrib- (4.72%) within 12 genera in the Caribbean uted along the East coast of the United States (Table 1). Of these, more than 30% could be and Canada, and Paracaprella digitimanus considered endemic for the region (Pseudae- has been also found on the Brazilian coast ginella colombiensis, Deutella caribensis, D. (Díaz et al., 2005). The Caribbean caprel- margaritae, D. mayeri, Mayerella redunca, lid diversity could be considered high when

Table 4 Species number of amphipods in common between countries in the Caribbean Sea

CUADRO 4 Número de especies de anfípodos communes entre cada uno de los países del Mar Caribe México Puerto Rico Hispaniola Jamaica Belize Honduras Guatemala Nicaragua Costa Rica Panamá Colombia Venezuela ABC* Lesser Antilles & Tobago Trinidad Islands Cayman

Cuba 99 6 2 3 8 0 0 0 13 6 38 87 5 16 2 1 México 13 3 4 21 2 1 0 17 10 41 119 6 35 3 1 Puerto Rico 2 0 2 1 0 0 3 1 5 8 2 9 1 0 Hispaniola 1 0 0 0 0 0 0 2 3 2 3 0 0 Jamaica 0 0 0 0 0 0 0 5 0 2 0 0 Belize 1 0 0 7 7 7 16 0 3 2 0 Honduras 0 0 1 1 1 1 0 0 0 0 Guatemala 0 1 1 1 0 0 0 0 0 Nicaragua 0 0 0 0 0 0 0 0 Costa Rica 7 12 13 3 4 2 0 Panamá 5 7 2 0 0 0 Colombia 45 7 6 2 1 Venezuela 8 27 3 3 ABC* 4 1 1 Lesser Antilles 3 2 Trinidad & Tobago 0

(ABC*=Aruba, Bonaire and Curacao).

1690 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013 Gamma Diversity HIS

ABC

CRC

PRC

PAN

BEL

LAN

COL

CUB

VEN

MEX

0 5 10 15 20 25 30 Relative Contribution (%)

Fig. 4. Contributions to the Amphipoda Caribbean gamma diversity by country. All countries are ordered by their alpha diversity value, from lowest to highest. HIS Hispaniola, PAN Panama, ABC Aruba, Bonaire and Curacao, BEL Belize, CRC Costa Rica, PRC Puerto Rico, LAN Lesser Antilles, COL Colombia, CUB Cuba, MEX Mexico, VEN Venezuela. compared to other tropical areas [Brazil: 12 The Hyperiidea have an overall number of species (Wakabara, Tararam, Valério-Berardo, valid species of 304 according to the WoRMS. Duleba & Pereira-Leite, 1991; Guerra-García, There are 222 species known to occur in the 2003b); Tanzania: 13 species (Guerra-García, Pacific Ocean and 175 species in the Atlantic 2001); Mauritius: seven species (Guerra-Gar- (Gasca, 2009a,b). This review recorded 116 cía, 2003c); India: 11 species (Guerra-García, species in the Caribbean region, representing Ganesh, Jaikumar & Raman, 2010); Phuket, 38.16% of the global hyperiid fauna. The few Thailand: 12 species (Guerra-García, 2004); inventories and species lists in the region have Indonesia: seven species (Krapp-Schickel been provided not only for shallow water spe- & Guerra-García, 2005); Philippines: three cies (Gasca & Shih, 2001, 2003), but also for species (Guerra-García, 2002); Papua New- the deep water masses (Gasca, 2009b). One Guinea: seven species (Guerra-García, 2003d); of the most complete studies on the Western Great Barrier Reef: 17 species (Guerra-García Atlantic Hyperiidea, although restricted to the & Lowry, 2009), but still significantly lower Southwestern Atlantic, reports 151 species in when compared to temperate waters (Thiel, that region (Vinogradov, 1999). In compari- Guerra-García, Lancellotti & Vásquez, 2003). son, the known species richness of hyperiid For example, 105 species have been reported amphipods from the Caribbean region is larger for Japan (Takeuchi, 1999) and 41 species for than in the other adjacent tropical areas, such the Mediterranean (Guerra-García & Takeuchi, as the Gulf of Mexico (100 species; LeCroy 2002; Sturaro & Guerra-García, 2012). et al., 2009), the Sargasso Sea (88 species;

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013 1691 Gasca, 2007), and Brazil (90 species; Montu, taxonomic expertise. Also, countries having a 1998; Lima & Valentin, 2001). We suggest that low reported biodiversity are probably unders- the number of hyperiid species inhabiting the ampled, especially if they are in close proxim- Caribbean region is higher, mainly because ity to highly diverse countries. For example, it the hyperiids can be divided roughly into is likely that the biodiversity of Belize, Colom- warm-water and cold-water species, and most bia, Aruba, Bonaire, Curaçao and Trinidad of the warm-water species are circumtropical and Tobago would resemble the biodiversity (including the tropical and subtropical belts). of their neighbouring countries Mexico and Most of the hyperiid specimens contained in Venezuela. Another important aspect to con- zooplankton collections have been identified sider is that a low number of species could also at the suborder level, which leads us to rec- be related to a small size of the coastal zone, ognize that new species and new records will to the lack of habitat diversity, and/or to the occur as the taxonomic effort to identify these level of degradation of the coast due to natu- samples increases. ral or anthropogenic disturbances (Dauvin & The ingolfiellidean amphipods are not Ruellet, 2009; De la Ossa-Carretero, Del-Pilar- very diverse in the Caribbean Sea (seven Ruso, Giménez-Casalduero, Sánchez-Lizaso species), but they still represent an important & Dauvin, 2012). In addition, the number of percentage (15.91%) of the global total of 44 species can be also related to the extent of the valid species. Such biodiversity is low, consid- shelf area, the hydrographic isolation of bays, ering the wide range of ecological conditions in coves, among others. which they occur, which varies from deep sea In conclusion, available information on soft muds, to high altitude freshwater rivers in amphipod taxonomy in the Caribbean region the continents, to subterranean fresh, brackish is heterogeneous, restricted for the most part and marine interstitial waters (Vonk & Sch- to Cuba, Mexico and Venezuela, and in these ram, 2003), but comparable to other families countries, mainly focused on Gammaridea of amphipods with partly overlapping habitat and Caprellidea. Given this heterogeneity in requirements such as hadziids (30 species) and the data, knowledge on the distributions of bogidiellids (seven species). Seven ingolfiel- the species is also limited. There is a need lidean amphipods are recorded for the Carib- to compare between habitats throughout the bean region, six of them are endemic and one, region, i.e. compare seagrass habitat diversity Ingolfiella fuscina is also recorded for the Gulf along a latitudinal gradient, or between depth of Mexico and Florida (Felder & Camp, 2009). zones, i.e. shelf, slope and abyssal marine More exploration and collecting efforts with- ecosystems. This would lead us to understand in anchialine cave systems in the Caribbean distribution patterns and recognize hotspots islands will very likely lead to the discovering that will allow for recommendations on man- of a larger number of species in this suborder. agement plans for each of them. On the other In general terms, Mexico, Venezuela and hand, knowledge of ecology and behaviour Cuba exhibit the highest species diversity, is virtually non existent for the majority of due in part to the higher intensity of research the species. More significantly, at this time of in these areas. In relation to species descrip- rapidly vanishing natural habitats, we have not tions and taxonomic publications, Cuba has yet estimated how many species are relatively taken the lead, particularly in coastal and conserved within the boundaries of national shallow waters. The least studied coasts in the parks and other protected areas. In addition to Caribbean are those of Guatemala, Honduras, climate change which is related to higher tem- Nicaragua, Trinidad and Tobago, Jamaica and peratures, sealevel rise, erosion of the coastal the Cayman Islands. The lack of knowledge habitats and the acidification and disappear- in these countries is probably due to limita- ance of important habitats, many intertidal tions in funding, research institutions, and/or zones are currently subject to disturbance from

1692 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013 industrialization and infrastructure develop- (Caprellidea, Gammaridea, Hyperiidea e Ingolfiellidea) en ment from the increasingly growing tourism el Mar Caribe, compilándose los registros de especies cita- das en las diversas ecorregiones del área, tanto en publica- industry. The fauna and flora of the inter- ciones como a través de bases de datos (WoRMS y OBIS). tidal zone are likely to be affected by such Se listan un total de 535 especies pertenecientes a 236 anthropogenic changes in environmental con- géneros y 73 familias. La ecorregión del Caribe Occidental ditions; therefore there is a need for monitor- presentó la mayor riqueza específica (282), mientras que ing programmes, which could help document el Caribe Oriental presentó el menor valor (73). México y Venezuela presentaron el mayor número de especies reg- such environmental changes. Thomas (1993) istradas (266 y 206, respectivamente), mientras que doce stressed that gammaridean amphipods are ideal países presentaron menos de 50. Los Gammaridea fueron el candidates for monitoring studies because they grupo más diverso con 387 especies, seguido de los Hype- are ecologically and trophically important, riidea (116 especies). Hasta el presente, se han descrito 218 numerically dominant, and have relative low especies nuevas en aguas del Caribe, de las cuales 132 son endémicas. Las áreas con mayor diversidad de anfípodos dispersal capabilities. However, the use of se concentraron a lo largo del caribe Mexicano, Cuba y these organisms as biomonitors in the tropics is la costa norte de Sur-América (Venezuela-Colombia); sin only possible if their taxonomic and ecological embargo, este patron refleja más bien un mayor esfuerzo en information is available. la recoleccion de estos organismos y una mayor experticia Besides increasing sampling efforts in taxonómica, que un patron específico de distribución del grupo. En general, el conocimiento del grupo se limita a coastal zones of poorly sampled countries las aguas someras, con muy poca información sobre zonas in Central America and the Lesser Antilles, profundas. En este sentido, las prioridades de investigación several ecosystems require additional sam- regional en este grupo, debe enfocar un mayor esfuerzo en pling efforts like the anchialine cave systems, la costa de Centro América y las Antillas Mayores y Meno- the continental margins, and the deep sea, res. Así mismo, es necesario aumentar las recolecciones en aguas profundas y otros ecosistemas particulares como inlcuding the escarpments, canyons, slopes, los anquialinos. Por último, es importante aumentar las seamounts and diapirs, the abyssal plains, and investigaciones taxonómicas y ecológicas de los taxones geologically active ecosystems such as vents que presentan mayor diversidad. and seeps. These collecting initiatives should be combined with an increase in taxonomic Palabras clave: Amphipoda, diversidad, riqueza especí- fica, Mar Caribe, endemismo. efforts, both in classical taxonomy and in the use of molecular tools like barcoding, for which regional capacity building is required. References All of these efforts should be carried out Andrade, C. A. & Barton, E. D. (2000). Eddy development through regional coordinated collaboration. and motion in the Caribbean Sea. Journal of Geo- physical Research, 105(C11), 26191-26201. Acknowledgments Atienza, A. & Martín, A. (2001). Tiburonella viscana (Amphipoda: Plastysnopidae) en las costas de Ven- The autors thank the countless students ezuela. Revista de Biología Tropical, 49(3), 1270. who have worked at the Laboratory of Peraca- Ayala, Y. & Martín, A. (2003). Ampliación del ámbito de Podocerus kleidus (Amphipoda: Podoceridae) al Mar ridean Crustaceans, as well as in the amphipod Caribe. Revista de Biología Tropical, 51(1), 267. collection of the Natural Science Museum at Barnard, J. L. (1954). Amphipoda of the family Ampelisci- Simon Bolivar University. dae collected by the Velero III in the Caribbean Sea. Allan Hancock Atlantic Expeditions Reports, 7, 1-13. RESUMEN Barnard, J. L. (1960). The amphipod family Phoxocephali- dae in the Eastern Pacific Ocean, with analysis of El orden Amphipoda es uno de los más diversos other species and notes for a revision of the family. dentro de los Peracáridos. Las investigaciones sobre este Allan Hancock Pacific Expeditions, 18, 174-368. orden en el Mar Caribe se iniciaron a finales de los años Barnard, J. L. (1961). Gammaridean Amphipoda from 1800, incrementándose significativamente a partir de 1980. depths of 400 to 6000 meters. Galathea Reports, 5, En este estudio se analizó la biodiversidad de los anfípodos 23-128.

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Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013 1697 Moore, E. A. & Sander, F. (1979). A comparative study of Ortiz, M. (1979b). Contribución al estudio de los anfípodos zooplankton from oceanic, shelf and harbor waters of (Gammaridea) del Mediterráneo americano. Inves- Jamaica. Biotropica, 11(3), 196-206. tigaciones Marinas, 45, 1-16. Myers, A. A. (1968). Two Aoridae (Amphipoda, Gam- Ortiz, M. (1980). Una nueva especie de anfípodo bentósico maridea) including a new species of Amphideuto- (Amphipoda, Gammaridea) de aguas cubanas. Revis- pus Barnard from Venezuelan waters. Crustaceana, ta de Investigaciones Marinas, 1, 92-103. 14(2), 127-130. Ortiz, M. (1983). Los anfípodos (Gammaridea) de las Myers, A. A. (1977). Studies on the genus Lembos Bate costas del mar Caribe de la República de Colombia. VI. Atlantic species: L. dentischium sp. nov., L. kun- Revista de Investigaciones Marinas, 4, 23-31. kelae sp. nov., L. rectangulatus sp. nov., L. unicornis Ortiz, M. (1991). Amphipod Crustacea. II. Family Bateidae. Bynum & Fox. Bollettino del Museo Civico di Storia Memoirs of Hourglass Cruises, VII, 1-31. Naturale di Verona, 4, 125-154. Ortiz, M., Alvarez, F. & Winfield, I. (2002). Caprellid Myers, A. A. (1978). Studies on the genus Lembos Bate Amphipods: Illustrated key for the genera and spe- VII. Atlantic species 4. L. setosus sp. nov., L. brun- cies from the Gulf of Mexico and the Caribbean neomaculatus Myers ssp. longicornis nov. and ssp. Sea. Mexico. Ediciones de la Universidad Nacional mackinneyi nov., L. foresti Mateus and Mateus, L. Autónoma de Mexico. longicarpus sp. nov. Bollettino del Museo Civico di Ortiz, M., Cházaro-Olvera, S. & Winfield, I. (2001). A Storia Naturale di Verona, 5, 183-209. new amphipod crustacean of the genus Haustorius (Gammaridea, Haustoriidae), from coast of Mexico. Myers, A. A. (1979). Studies on the genus Lembos Bate Avicennia, 14, 53-59. VIII. Atlantic species 5. L. tigrinus sp. nov., L. tem- pus sp. nov., L. spinicarpus (Pearse) comb. nov. with Ortiz, M., García-Debras, A. & Lalana, R. (2002). Una ssp. inermis nov., L. ovalipes sp. nov., L. unifasciatus nueva especie de anfípodo anquialina del género Melita (Gammaridea: Melitidae), de la Isla de Cuba. Myers ssp. reductus nov. Bollettino del Museo Civico Avicennia, 15, 43-52. di Storia Naturale di Verona, 6, 221-248. Ortiz, M., García-Debras, A. & Lalana, R. (2003). Una Myers, A. A. & Lowry, J. K. (2003). A phylogeny and a nueva especie de anfípodo anquialino del género new classification of the Corophiidea Leach, 1814 Socarnopsis (Amphipoda, Lysianassidae) del sistema (Amphipoda). Journal of Crustacean Biology, 23(2), cavernario de Playa Girón, Cuba. Avicennia, 16, 443-485. 71-77. Ortiz, M. (1976a). Un nuevo anfípodo de aguas cubanas Ortiz, M. & Gómez, O. (1979). Una nueva especie de (Amphipoda, Gammaridea, Phliantidae). Investiga- anfípodo (Amphipoda, Gammaridea) de aguas pro- ciones Marinas, 8(25), 21-35. fundas del sur de Cuba. Investigaciones Marinas, Ortiz, M. (1976b). Un nuevo género y una nueva especie 40, 23-30. de anfípodo de aguas cubanas (Amphipoda, Gam- Ortiz, M., Guerra-García, J. M. & Lalana, R. (1998). Cuba- maridea, Ampithoidae). Ciencias (La Habana). Inves- deutella cavernicola, a new genus and new species tigaciones Marinas, 8(27), 3-12. of Caprellidae (Crustacea: Amphipoda) from Cuba. Ortiz, M. (1976c). Un nuevo anfípodo (Amphipoda, Gam- Zootaxa, 2130, 60-68. maridea, Gammaridea) colectado del contenido Ortiz, M. & Lalana, R. (1980). Un nuevo anfípodo del estomacal del ronco amarillo Haemulun sciurus, género Leptocheirus (Amphipoda, Gammaridea) de de aguas cubanas. Investigaciones Marinas, 8(27), aguas cubanas. Revista de Investigaciones Marinas, 13-20. 1, 58-73. Ortiz, M. (1976d). Un nuevo anfípodo perforador de made- Ortiz, M. & Lalana, R. (1989). Una nueva especie de ra (Amphipoda, Gammaridea, Cheluridae) de aguas anfípodo del complejo Eriopisa (Amphipoda, Gam- cubanas. Investigaciones Marinas, 8(27), 21-26. maridea), de aguas cubanas. Revista de Investigacio- nes Marinas, 10, 233-237. Ortiz, M. (1976e). A new crustacean amphipod, Mallacoo- Ortiz, M. & Lalana, R. (1993). Adición a la lista de espe- ta carausui, from the Cuban waters. Revue Roumaine cies y bibliografía de los anfípodos (Crustacea, de Biologie, Série de Biologie Animale, 21(2), 93-95. Amphipoda) del Mediterráneo Americano. Revista de Ortiz, M. (1978). Invertebrados marinos bentónicos de Investigaciones Marinas, 14, 16-37. Cuba. I. Crustacea, Amphipoda, Gammaridea. Inves- Ortiz, M. & Lalana, R. (1994a). A new amphipod species tigaciones Marinas, 38, 3-10. of the genus Nasageneia (Gammaridea: Eusiridae) Ortiz, M. (1979a). Lista de especies y bibliografía de los from the south-western cuban marine waters. Travaux anfípodos (Crustacea: Amphipoda) del Mediterráneo Du Museum d’Histoire Naturelle “Grigore Antipa”, Americano. Investigaciones Marinas, 43, 1-40. 34, 285-292.

1698 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013 Ortiz, M. & Lalana, R. (1994b). Two new species of the Ortiz, M., Lalana, R. & Sánchez, A. (2000). Una nueva genus Elasmopus (Amphipoda, Gammaridea), from especie de anfípodo espongícola del género the Cuban marine waters. Travaux Du Museum Hoplopheonoides Shoemaker, 1956 (Gammaridea, d´Histoire Naturelle “Grigore Antipa”, 34, 293-302. Cyproideidae) de Cuba. Avicennia, 12/13, 63-68. Ortiz, M. & Lalana, R. (1995). Gitanopsis templadoi, una Ortiz, M., Lalana, R. & Varela, C. (2002). Descripción de nueva especie de anfípodo (Gammaridea, Amphilo- la hembra de Spathiopus cojimarensis (Amphipoda: chidae), de la plataforma suroccidental de Cuba. Ana- Melitidae). Solenodon, 2, 17-20. les del Instituto de Ciencias Marinas y Limnología, Ortiz, M., Lalana, R. & Varela, C. (2004). Una nueva especie 22, 121-124. de anfípodo marino del género Elasmopus (Amphipoda, Ortiz, M. & Lalana, R. (1996). Los anfípodos de la primera Melitidae) del Archipiélago cubano. Avicennia, 17, 35-40. expedición conjunta Cuba-US, a bordo del B/I “Ulis- Ortiz, M., Lalana, R. & Varela, C. (2006). A new species of the es”, a las aguas del Archipiélago Sabana-Camagüey, genus Elasmopus (Gammaridea, Melitidae) from Cuban Cuba, en 1994. Anales del Instituto de Biología, waters. Avicennia, 17, 35-40. Universidad Nacional Autonoma de México Serie Zoología, 67, 89-101. Ortiz, M., Lalana, R. & Varela, C. (2007a). Un género nuevo y una especie nueva de anfípodo espongícola (Gamma- Ortiz, M. & Lalana, R. (1998). Lista actualizada de los ridea: Eusiridae) de las aguas profundas del sur de Cuba. crustáceos no decápodos de Cuba. Revista de Inves- Solenodon, 6, 8-14. tigaciones Marinas, 19(2-3), 92-99. Ortiz, M. & Lalana, R. (2002a). Una nueva especie Ortiz, M., Lalana, R. & Varela, C. (2007b). Tantena, género de anfípodo del género Spathiopus (Gammaridea, nuevo y especie nueva de anfípodo marino (Lysianas- Melitidae), de la plataforma noroccidental de Cuba. sidae) y primera consignación de la familia Ochlesidae Avicennia, 15, 31-36. y del género Curidia, con la descripción de una especie nueva para Cuba (Amphipoda, Gammaridea). Solenodon, Ortiz, M. & Lalana, R. (2002b). Primer registro del género 6, 20-32. Neoischyrocerus (Amphipoda, Ischyroceridae) para el Mar Caribe y el archipiélago cubano, con la Ortiz, M., Lalana, R. & Varela, C. (2007c). Primer registro descripción de nueva especie de Cuba. Avicennia, del género Aristias (Amphipoda: Aristiidae) para el Mar 15, 37-51. Caribe, con la descripción de una especie nueva. Solen- odon, 6, 33-40. Ortiz, M. & Lalana, R. (2002c). Una nueva especie de anfípodo del género Bemlos (Amphipoda, Gamma- Ortiz, M., Lalana, R. & Varela, C. (2007d). Crustáceos no ridea, Aoridae), del archipiélago cubano. Revista de Decápodos - Filo Arthropoda, sub-filo Crustacea. Lista Investigaciones Marinas, 23(1), 69-76. de especies registradas en Cuba (diciembre de 2006). In R. Claro, (Ed.), La Biodiversidad marina de Cuba. (CD- Ortiz, M. & Lalana, R. (2006). Nuevos datos sobre crustá- ROM), La Habana: Instituto de Oceanología, Ministe- ceos peracáridos (Crustacea, Peracarida) de las costas del Caribe de Costa Rica. Cocuyo, 16, 16-18. rio de Ciencia, Tecnología y Medio Ambiente. Ortiz, M., Lalana, R. & Beltrán, J. (1993). Una nueva Ortiz, M., Lalana, R. & Varela, C. (2007e). First record especie de anfípodo hadzioideo (Amphipoda: Gam- of the genus Rhachotropis (Crustacea: Amphipoda: maridea) del Caribe de Nicaragua. Revista de Inves- Gammaridea: Eusiridae) for the cuban marine waters, tigaciones Marinas, 14, 103-109. with the description of a new species. Travaux Du Museum d’Histoire Naturelle “Grigore Antipa”, 50, Ortiz, M., Lalana, R. & Guevara, E. (1990). Platyscelus 25-30. ovoides (Amphipoda: Hyperiidae) en el contenido estomacal del bonito (Katsuwomus pelamis). Revista Ortiz, M., Lalana, R. & Varela, C. (2008). Primer registro de Investigaciones Marinas, 11, 174-175. de la familia Melphidippidae y del género Hornellia (Amphipoda: Gammaridea) para aguas cubanas, con Ortiz, M., Lalana, R. & Lio, V. (1999). Un nuevo género y la descripción de una especie nueva. Solenodon, 7, una especie de anfípodo marino (Amphipoda: Aristi- 7-13. idae), de Cuba. Avicennia, 10/11, 137-142. Ortiz, M., Lalana, R., Varela, C. & Leal, S. (2003). Crustá- Ortiz, M., Lalana, R. & Lio, V. (2001). Primera consig- ceos marinos planctónicos (Malacostraca), de Cuba. nación del género Protohadzia (Amphipoda, Gam- Revista de Investigaciones Marinas, 24, 77-79. maridea), para el archipiélago cubano. Avicennia, 14, 129-132. Ortiz, M. & Lemaitre, R. (1994). Crustáceos anfípodos Ortiz, M., Lalana, R. & López, M. (1992). Nueva espe- (Gammaridea) colectados en las costas del Caribe cie de anfípodo del género Bemlos (Gammaridea, colombiano, al sur de Cartagena. Anales de Investiga- Corophiidae), de Cuba. Anales del Instituto de Cien- ciones Marinas de Punta Betín, 23, 119-127. cias Marinas y Limnología, Universidad Nacional Ortiz, M. & Lemaitre, R. (1997). Seven new amphi- Autónoma de Mexico, 19, 163-166. pods (Crustacea: Peracarida: Gammaridea) from the

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013 1699 Caribbean coast of South America. Anales del Insti- Marcuzzi. Memorie del Museo Civico di Storia Natu- tuto de Investigaciones Marinas de Punta Betín, 26, rale di Verona, 4, 117- 125. 71-104. Ruffo, S., Krapp-Schickel, T. & Gable, M. F. (2000). The Ortiz, M., Lopeztegui, A., Lalana, R. & Varela, C. (2006). genus Maera (Crustacea: Amphipoda: Melitidae) Registros nuevos de anfípodos planctónicos (Crus- from Bermuda. Postilla, 221, 1-35. tacea, Amphipoda, Hyperiidea) de la costa Sur de la Sánchez, L. (1985). Algunos aspectos biológicos de Tal- Isla de la Juventud, suroeste de Cuba. Cocuyo, 16, 15. orchestia margaritae Stephensen, 1948 (Amphipoda: Ortiz, M., Martín, A. & Atienza, D. (2000). Una nueva Talitridae), de la Isla de Margarita, Venezuela. (Tesis especie de anfípodo del género Tiburonella (Crus- de licenciatura). Universidad de Oriente, Cumaná, tacea, Amphipoda, Platyischnopidae) de Venezuela. Venezuela. Acta Biologica Venezuelica, 20(2), 29-36. Shih, C-t. (1969). The systematic and biology of the family Ortiz, M., Martín, A. & Díaz, Y. J. (2007). Lista y referen- Phronimidae (Crustacea: Amphipoda). Dana Reports, cias de los crustáceos anfípodos (Amphipoda: Gam- 74, 1-100. maridea) del Atlántico occidental tropical. Revista de Shoemaker, C. R. (1926). Amphipods of the family Biología Tropical, 55(2), 479-498. Bateidae in the collection of the United States Muse- Ortiz, M., Martín, A., Winfield, I., Díaz, Y. & Atienza, D. um. Proceeding of the United States of Natural (2004). Anfípodos (Crustacea: Gammaridea). Clave Museum, 68(2626), 1-26. gráfica para la identificación de las familias, géneros y Shoemaker, C. R. (1933a). Two new genera and six new especies marinas y estuarinas del Atlántico occidental species of Amphipoda from Tortugas. Papers of the tropical. México: Universidad Nacional Autónoma Tortugas Laboratory. Carnegie Institution of Wash- de México, Facultad de Estudios Superiores Iztacala. ington, 28(435), 245-256. Ortiz, M. & Nazábal, J. (1984a). A new amphipod crus- Shoemaker, C. R. (1933b). Amphipoda from Florida and tacean of the genus Lembos (Gammaridea, Aoridae) the West Indies. American Museum Novitates, 598, from the Cuban marine waters. Travaux Du Museum 1-24. d´Histoire Naturelle “Grigore Antipa”, 26, 11-13. Shoemaker, C. R. (1934). Reports on the collections Ortiz, M. & Nazábal, J. (1984b). Corocubanus, un nuevo obtained by the first Johnson–Smithsonian deep-sea género de anfípodo (Amphipoda, Gammaridea, expedition to the Puerto Rican deep. Three new Corophiidae), de aguas Cubanas. Revista de Inves- amphipods. Smithsonian Miscelaneous Collections, tigaciones Marinas, 5, 3-21. 9(12), 1-6. Ortiz, M. & Nazábal, J. (1988). Una nueva especie de Shoemaker, C. R. (1935). The Amphipods of Porto Rico anfípodo del género Lembos (Crustacea, Amphi- and the Virgin Islands. New York Academy of Sci- poda), de aguas cubanas. Revista de Investigaciones ences Scientific Survey of Porto Rico and the Virgin Marinas, 9, 29-35. Islands, 15(2), 229-253. Ortiz, M. & Pérez, A. (1995). Una nueva especie de Shoemaker, C. R. (1945). The Amphipoda of the Bermuda anfípodo cavernícola Hadzioideo (Amphipoda, Gam- Oceanographic Expeditions 1929-1932. Zoologica, maridea) de Cuba. Graellsia, 51, 165-168. 30, 185-266. Ortiz, M. & Thomas, J. D. (2007). Cerapus orteai Shoemaker, C. R. (1948). The Amphipoda of the Smithso- (Corophioidea: Corophiidae) un nuevo anfípodo de nian-Roebling Expedition to Cuba in 1937. Smithson- la costa caribeña de Costa Rica. Avicennia, 19, 17-24. ian Miscelaneous Collections, 110(3), 1-15. Ortiz, M., Varela, C. & Lalana, R. (2011). Especie nueva de anfípodo del género Photis (Gammaridea: Phot- Spalding, M. D. (2004). A guide to the coral reefs of the idae) del Archipiélago cubano. Novitates Caribaea, Caribbean. Berkeley: University of California Press. 4, 10-16. Stebbing, T. R. R. (1888). Report on the Amphipoda col- Ortiz, M. & Veledo, T. (1985). Una nueva especie de lected by H.M.S. Challenger during the years 1873- anfípodo del género Garosyrrhoe (Synopiidae, Gam- 76. Zoology, 29, 1-1737. maridea) de aguas cubanas. Revista de Investigacio- Stebbing, T. R. R. (1897). Amphipoda from the Copen- nes Marinas, 6, 14-18. hagen Museum and other sources. Transactions of Ruffo, S. (1950). Studi sui crostacei anfipodi. XXII. Anfi- Linnean Society of London, Serie Zoology, 7, 25-45. podi del Venezuela raccolti dal dott. G. Marcuzzi. Stebbing, T. R. R. (1903). Amphipoda from Costa Rica. Memorie del Museo Civico di Storia Naturale di Proceedings of the United States Natural Museum, Verona, 2, 49-65. 26, 925-931. Ruffo, S. (1954). Studi sui crostacei anfipodi. XL. Stebbing, T. R. R. (1906). Amphipoda I. Gammaridea. Das Nuovi anfipodi raccolti nel Venezuelan dal Prof. G. Tierrich, 21, 806.

1700 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013 Stebbing, T. R. R. (1908). A new amphipod crustacean, Thiel, M., Guerra-García, J. M., Lancellotti, D. A. & Orchestoidea biolleyi, from Costa Rica. Proceedings Vásquez, N. (2003). The distribution of littoral capre- of the United States Natural Museum, 34, 241-244. llids (Crustacea: Amphipoda: Caprellidea) along the Stephensen, K. (1933). Amphipoda from the marine salines Pacific cost of continental Chile. Revista Chilena de of Bonaire and Curacao. Zoologische Ergebnisse Historia Natural, 76(2), 203-218. einer Reise nach Bonaire, Curacao und Aruba im Thomas, J. D. (1983). Curidia debrogania, a new genus Jahre 1930. Zoologische Jahrbucher, Systematik, 64, and species of amphipod (Crustacea: Ochlesidae) 437-446. from the Barrier Reef of Belize, Central America. Stephensen, K. (1948). Amphipods from Curacao, Bonaire, Proceedings of Biological Society of Washington, Aruba and Margarita. Studies on Fauna Curaçao, 96(1), 127-133. Aruba, Bonaire and the Venezuelan Islands, 11(3), Thomas, J. D. (1993). Biological monitoring and tropi- 1-20. cal biodiversity in marine environments: a critique Stock, J. H. (1976). A new member of the crustacean sub- with recommendations, and comments on the use of order Ingolfiellidea from Bonaire. Studies on Fauna amphipods as bioindicators. Journal of Natural His- Curaçao, 50, 56-75. tory, 27(4), 795-806. Stock, J. H. (1977a). The taxonomy and zoogeography of Thomas, J. D. (1997). Systematics, ecology and phylogeny the Hadziid amphipoda, with emphasis on the West of the Anamixidae (Crustacea: Amphipoda). Records Indian taxa. Studies on Fauna Curaçao, 55, 1-130. of Australian Museum, 49, 35-98. Stock, J. H. (1977b). The zoogeography of the crustacean Thomas, J. D. & Barnard, J. L. (1983). The Platyischnopi- suborder Ingolfiellidea with descriptions of new West dae of America (Crustacea: Amphipoda). Smithson- Indian taxa. Studies on Fauna Curaçao, 55, 131-146. ian Contributions to Zoology, 375, 1-33. Stock, J. H. (1978). Bogidiella martini, a new hypo- Thomas, J. D. & Barnard, J. L. (1985). Perioculodes gean amphipod from the island of Saint-Martin (West cerasinus, n.sp., the first record of the genus from Indies) and the zoogeography of the Bogidiellidae. the Caribbean Sea (Amphipoda: Oedicerotidae). Pro- International Journal of Speleology, 9(2), 103-113. ceedings of Biological Society of Washington, 98(1), Stock, J. H. (1979). New data on taxonomy and zooge- 98-106. ography of Ingolfiellid crustacea. Bijdragen tot de Thomas, J. D. & Barnard, J. L. (1986). New genera and Dierkunde, 49(1), 81-96. species of the Megaluropus group (Amphipoda, Stock, J. H. (1980). A new cave amphipod (Crustacea) Megaluropidae) from American Seas. Bulletin of from Curacao: Psammogammarus caesicolus n. sp. Marine Science, 38(3), 442-476. Bijdragen tot de Dierkunde, 50, 375-386. Thomas, J. D. & Barnard, J. L. (1989). Gammaropsis Stock, J. H. (1981). The taxonomy and zoogeography of arawakia, a new species of marine amphipoda (Crus- the family Bogidiellidae (Crustacea, Amphipoda), tacea) from Jamaica. Proceedings of Biological Soci- with emphasis on the west indian taxa. Bijdragen tot ety of Washington, 102(1), 89-94. de Dierkunde, 51, 345-374. Thomas, J. D. & Barnard, J. L. (1990). Gitana dominica, Stock, J. H. (1983). The stygobiont amphipoda of Jamaica. a new species from the Caribbean Sea (Amphipoda: Bijdragen tot de Dierkunde, 53, 267-286. Amphilochidae). Proceedings of Biological Society of Washington, 103(3), 617-623. Stock, J. H. (1985). Bogidiellidae (Amphipoda) from Haiti and some general rules on the occurrence of Crusta- Thomas, J. D. & Barnard, J. L. (1991a). Photis trapherus, cea Malacostraca in inland groundwaters of the West a new Elephantine species from the Caribbean Sea Indies. Stygologia, 1, 208-223. (Crustacea: Amphipoda). Proceedings of Biological Society of Washington, 104(1), 96-100. Stoner, A. W. & Lewis, F. G. (1985). The influence of quantitative and qualitative aspects of habitat com- Thomas, J. D. & Barnard, J. L. (1991b). Two new species plexity in tropical sea-grass meadows. Journal of of Netamelita from the Caribbean Sea (Crustacea: Experimental Marine Biology and Ecology, 94(1-3), Amphipoda). Proceedings of Biological Society of 19-40. Washington, 104(3), 583-592. Sturaro, N. & Guerra-García, J. M. (2012). A new species Thomas, J. D. & Klebba, K. N. (2006). Studies of com- of Caprella (Crustacea: Amphipoda) from the Medi- mensal leucothoid amphipods: two new sponge- terranean Sea. Helgolaender Marine Research, 66(1), inhabiting species from south Florida and the western 33-42. doi: 10.1007/s10152-011-0244-5. Caribbean. Journal of Crustacean Biology, 26(1), Takeuchi, I. (1999). Checklist and bibliography of the 13-22. Caprellidea (Crustacea: Amphipoda) from Japanese Thomas, J. D. & Klebba, K. N. (2007). New species waters. Otsuchi Marine Science, 24, 5-17. and host associations of commensal leucothoid

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013 1701 amphipods from coral reefs in Florida and Belize interstitial on Curaçao. Foundation for Scientific (Crustacea: Amphipoda). Zootaxa, 1494, 1-44. Research in Surinam and the Netherlands Antilles, Vader, W. (2005). How many Amphipod Species?. Poster Amsterdam, 123, 185-198. presented at XII International Amphipod Colloqui- Vonk, R. (1990). Thalassostygius exiguus n.g., n. sp., a new um. Cork, Ireland. marine interstitial melitid (Crustacea, Amphipoda) Vainola, R., Witt, J. D. S., Grabowski, M., Bradbury, J. from Curaçao and Klein Bonaire (Netherlands Antil- H., Jazdzewski, K. & Sket, B. (2008). Global diver- les). Stygologia, 5, 43-48. sity of amphipods (Amphipoda; Crustacea) in fresh- Vonk, R. (1991). Two marine interstitial Metaniphargus water. Hydrobiologia, 595, 241-255. doi: 10.1007/ species (Crustacea, Amphipoda) from Hawaii and the s10750-007-9020-6. Cayman Islands. Stygologia, 6, 111-118. Van der Ham, J. L. & Vonk, R. (2003). A phylogenetic Vonk, R. & Schram, F. R. (2003). Ingolfiellidea (Crusta- analysis of the Eriopisa complex (Crustacea: Amphi- cea, Malacostraca, Amphipoda): a phylogenetic and poda: Melitidae) and a new species from beach biogeographic analysis. Contributions to Zoology, interstitial in Venezuela. Journal of Natural History, 72(1), 39-72. 37(7), 779-796. Varela, C., Ortiz, M. & Lalana, R. (2003). Crustáceos (Per- Vonk, R. & Stock, J. H. (1987). Psammogammarus longi- acarida y Decapoda), de la costa sur de la Península dactilus n.sp., a new cave amphipod (Crustacea) and de Guanahabibes, Cuba. Revista de Investigaciones other stygobiont amphipods from Bonaire. Stygolo- Marinas, 24, 73-76. gia, 3, 241-251. Villarroel, E. J. & Graziani, C. A. (1997). Primer registro Wakabara, Y., Tararam, A. S., Valério-Berardo, M. T., de Caprella danilevskii Czerniavskii (Amphipoda: Duleba, W. & Pereira-Leite, F. P. (1991). Gammarid- Caprellidae) en Venezuela. Saber, Universidad de ean and caprellidean fauna from Brazil. Hydrobiolo- Oriente, 9, 102-103. gia, 223, 69-77. Vinogradov, G. (1999). Amphipoda. In D. Boltovskoy WoRMS. (2012). Amphipoda. In J. Lowry (Ed.), World (Ed.), Zooplankton of the Southwestern Atlantic. Amphipoda database. Recuperado de http://www. (1141-1240). Backhuys, Leiden. marinespecies.org/aphia.php?p=taxdetails&id=1135 Vonk, R. (1988). Psammomelita uncinata n.g., n.sp. (Crus- Zeidler, W. & De Broyer, C. (2009). Volume 3: Catalogue tacea, Amphipoda, Melitidae) from infralittoral sand of the Hyperiidean Amphipoda (Crustacea) of the interstices on Curaçao. Stygologia, 4, 166-176. Southern Ocean with distribution and ecological data. Vonk, R. (1989). Nuuanu curvata n.sp and Melita leio- Bulletin de L’Institut Royal des Sciences Naturelles telson n.sp. (Crustacea, Amphipoda) from beach de Belgique, 79(1), 1-96.

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X X X X X X X X X X X X X CUB X X X X X X X X X X APPENDI X 1 Species McCain, 1968 Ortiz, Guerra-García & Lalana, 2009 & Lalana, Ortiz, Guerra-García Czerniavskii, 1868 Czerniavskii, Mayer, 1890 Mayer, Mayer, 1903 Mayer, McCain, 1968 Say, 1818 Say, (Mayer, 1903) (Mayer, Guerra-García, 2003 Guerra-García, margaritae Deutella mayeri Stebbing, 1895 Deutella wigleyi Hemiproto antillensis Phtisica Slabber, 1769 marina Slabber, Phtisica McCain, 1968 redunca Mayerella 2006 & Müller, Krapp-Schickel chibcha Guerra-García, Tritella Pseudaeginella biscaynensis (McCain, 1968) Pseudaeginella 2006 & Müller, Krapp-Schickel columbiensis Guerra-García, Pseudaeginella cavernicola Cubadeutella californica Deutella 2006 & Müller, Krapp-Schickel caribensis Guerra-García, Deutella incerta Deutella 1890 Hemiaegina minuta Mayer, Pseudaeginella antiguae Barnard, 1932 Pseudaeginella Mayer, 1903 tenuis Mayer, Paracaprella Templeton, 1836 scaura Templeton, Caprella hummelincki Metaprotella 1971 digitimanus Quitete, Paracaprella 1890 pusilla Mayer, Paracaprella Caprella equilibra Caprella Leach, 1814 penantis Leach, Caprella Aeginella spinosa Boeck, 1861 Aeginella 1890 Mayer, andreae Caprella danilevskii Caprella Suborder / Family GUA, Guatemala, HON, Honduras, NIC, Nicaragua, CRC, Costa Rica, PAN, Panama, COL, Colombia, VEN, Venezuela, ABC, Netherlands Antilles, TYT, Trinidad & Tobago, LAN, Lesser Antilles LAN, Lesser Tobago, & Trinidad TYT, Antilles, ABC, Netherlands Venezuela, VEN, Panama, COL, Colombia, GUA, Guatemala, HON, Honduras, NIC, Nicaragua, CRC, Costa Rica, PAN, Taxonomic classification of the amphipod fauna from Caribbean countries or subregion.PRC, Puerto Rico, HIS, Hispaniola, JAM, Jamaica, CUB, Cuba, CAY, Cayman Islands, ME X , Mexico, BEL, Belize, classification of the amphipod fauna from Caribbean countries or subregion.PRC, Puerto Rico, HIS, Hispaniola, JAM, Jamaica, CUB, Cuba, CAY, Taxonomic Family Phtiscidae Vassilenko, 1968 Vassilenko, Phtiscidae Family Family Protellidae McCain, 1970 Protellidae Family Family Pariambidae Laubitz, 1993 Laubitz, Pariambidae Family Family Caprellinoididae Laubitz, 1993 Laubitz, Caprellinoididae Family Suborder Caprellidea Leach, 1814 Leach, Suborder Caprellidea 1814 Leach, Caprellidae Family

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X X X X X X X X X X X X X X X CUB X X X X X X X X X APPENDI X 1 ( Continued Species J.L. Barnard, 1954 J.L. Barnard, 1954 (Della Valle, 1893) Valle, (Della Martin, Ortiz & Atienza, 2001 Atienza, Martin, Ortiz & Goeke & Heard, 1984 Shoemaker, 1916 Shoemaker, Goeke & Heard, 1983 (McKinney, 1978) (McKinney, Stimpson, 1864 Holmes, 1908 Mills, 1967 (J. L. Barnard & Thomas, 1983) (J. L. Barnard & Apolochus pillaii Ampelisca spinipes Boeck, 1861 Ampelisca vadorum Mills, 1963 Ampelisca venetiensis Ampelisca verrilli Ampelisca cubensis (Ortiz & Gómez, 1979) Byblis blasensis J.L. Barnard, 1964 Byblisoides 1978) Apolochus casahoya (McKinney, Apolochus delacaya Apolochus neapolitanus Amphilochus ascidicola J.L. Barnard, 1954 romigi Ampelisca 1933 Shoemaker, schellenbergi Ampelisca Barnard & Agard, 1986 paria Barnard & Ampelisca pugetica Ampelisca Ampelisca parapanamensis J.L. Barnard, 1954 Ampelisca Ampelisca parapacifica Ampelisca Ampelisca mexicana J.L. Barnard, 1954 Ampelisca Ampelisca pacifica Ampelisca Ampelisca lobata Holmes, 1908 Ampelisca Ampelisca holmesi Pearse, 1908 Ampelisca J. L. Barnard & Thomas, 1989 burkei J. L. Barnard & Ampelisca cristata Holmes, 1908 Ampelisca cristoides J.L. Barnard, 1954 Ampelisca Ampelisca agassizi (Judd, 1896) Ampelisca Ampelisca abdita Mills, 1964 Ampelisca abyssicola Stebbing, 1888 Ampelisca bicarinata Ampelisca brevisimulata Ampelisca cristata microdentata Ampelisca Suborder / Family Family Ampeliscidae Costa, 1857 Ampeliscidae Family Family Amphilochidae Boeck, 1871 Amphilochidae Family Suborder Gammaridea Latreille, 1802 Latreille, Suborder Gammaridea Costa, 1857 Ampeliscidae Family

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X X X X X X X X X X X X X X X CUB X X X X X X X X X X X X X X X X X X APPENDI X 1 ( Continued Species Myers, 1968 (Milne-Edwards, 1830) Ortiz & Lalana, 1995 Ortiz & Lalana, (A. A. Myers, 1977) (A. (A. A. Myers, 1977) (A. Smith, 1873 Kunkel, 1910 Bemlos unicornis (Bynum & Fox, 1977) Globosolembos francanni (Reid, 1951) Globosolembos smithi (Holmes, 1905) Ampithoe valida Ampithoe Cymadusa compta (Smith, 1873) 1816 Cymadusa filosa Savigny, bacescui Ortiz, 1976 Pseudamphithoides incurvaria (Just, 1977) Pseudamphithoides 1938) Bemlos intermedius (Schellenberg, Bemlos kunkelae A. Myers, 1978) Bemlos longicornis (A. 2002 Ortiz & Lalana, Bemlos rolani 1992 & López, Bemlos sanmartini Ortiz, Lalana 1997 Bemlos scolosternum Ortiz & Lemaitre, A. Myers, 1979) Bemlos spinicarpus inermis (A. Bemlos spinicarpus (Pearse, 1912) Ampithoe pollex Ampithoe ramondi Audouin, Ampithoe 1826 Sunamphitoe pelagica Amphideutopus dolichocephalus 1988) (Ortiz & Nazábal, Bemlos barnardi Bemlos brunneomaculatus Myers, 1977 Bemlos dentischium (Mateus & Mateus, 1966) Bemlos foresti Ortiz & Lemaitre, 1997 hirsutimanus Ortiz & Lemaitre, Ampithoe longimana Smith, 1873 Ampithoe 1954 Ruffo, marcuzzii Ampithoe (Shoemaker, 1933) Hourstonius tortugae (Shoemaker, Thomas & Barnard, 1990 Gitana dominica Gitanopsis templadoi 1978) Hourstonius laguna (McKinney, Suborder / Family Family Aoridae Walker, 1908 Aoridae Family Walker, Family Ampithoidae Stebbing, 1899 Ampithoidae Family Family Amphilochidae Boeck, 1871 Amphilochidae Family

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X X X X X X X X X CUB X X X X X X X X X X X X X APPENDI X 1 ( Continued Species Myers, 1977 (A. A. Myers, 1977) (A. Stock, 1981 (A. A. Myers, 1979) (A. Bousfield, 1973 (Ortiz, 1991) Lowry & Stoddart, 1997 Ortiz & Lemaitre, 1997 Ortiz & Lemaitre, Ortiz & Lalana, 1996 lowryi Ortiz & Lalana, Biancolina cultrifera Actogidiella Plesiolembos ovalipes Plesiolembos rectangulatus Plesiolembos 1865 catharinensis Müller, Batea 1926) cuspidata (Shoemaker, Batea schotti Batea 1926 transversa Shoemaker, Batea martini (Stock, 1978) Antillogidiella hamatula Stock, 1985 Bogidiella perla Stock, 1981 Bogidiella prionura Stock, 1985 Bogidiella Stock, 1981 virginalis Bogidiella crassipes Stock, 1981 Marigidiella Paramicrodeutopus myersi (Bynum & Fox, 1977) Paramicrodeutopus laminosum (Pearse, 1912) Pedicorophium Boca campi Lowry & Stoddart, 1997 Miramarassa & Lio, 1999 sanchezi Ortiz, Lalana 2011 aldebaranae Coleman, Batea bousfieldi Batea campi (Ortiz, 1991) Batea 1926) carinata (Shoemaker, Batea Ortiz, Lalana & Varela, 2007 Varela, & Aristias bicornuta Ortiz, Lalana Aristias captiva Ortiz & Lalana, 1996 Ortiz & Lalana, heardi Rudilemboides naglei Rudilemboides hamatipes (Norman, 1869) Argissa Ortiz & Nazábal, 1984 Lembos mayensis Ortiz & Nazábal, Myers, 1979 reductus Lembos unifasciatus unifasciatus Lembos unifasciatus Stephensen, 1947 bonnieroides Grandidierella Suborder / Family Family Biancolinidae Barnard, 1972 Biancolinidae Family Family Bateidae Stebbing, 1906 Bateidae Family Family Aristiidae Lowry & Stoddart, Aristiidae Family 1997 Family Argissidae Walker, 1904 Argissidae Family Walker, Family Aoridae Walker, 1908 Aoridae Family Walker,

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X ME

)

X X X X X X X X X X X X X X X X CUB X X X X X X X X X X X X X APPENDI X 1 ( Continued Species (Giambiagi, 1926) (Giambiagi, Bousfield, 1973 (Milne Edwards, 1830) LeCroy, 1995 LeCroy, LeCroy, 1995 LeCroy, Stebbing, 1888 Heard & Perlmutter, 1977 Heard & Perlmutter, LeCroy, 1995 LeCroy, Colomastix bousfieldi 1995 Colomastix camura LeCroy, 1995 Colomastix falcirama LeCroy, LeCroy, 1995 Colomastix gibbosa LeCroy, Colomastix halichondriae 1995 LeCroy, Colomastix heardi Colomastix irciniae Colomastix janiceae Colomastix tridentata rioplatense Americorophium acutum (Chevreux, 1908) Apocorophium 1911) (Vanhöffen, lacustre Apocorophium 1934) louisianum (Shoemaker, Apocorophium 1934) simile (Shoemaker, Apocorophium barbadensis Just, 1983 Caribboecetes crassicornis Just, 1984 Caribboecetes intermedius Just, 1984 Caribboecetes (Calman, 1910) insulae (Calman, Tropichelura bonellii Crassicorophium 1934) baconi (Shoemaker, Laticorophium 1980 Ortiz & Lalana, Leptocheirus rhizophorae acherusicum (Costa, 1853) Monocorophium Ortiz & Lemaitre, 1997 justi Ortiz & Lemaitre, Caribboecetes pterycornis Just, 1984 Caribboecetes Cyphocaris anonyx Boeck, 1871 1934 Cyphocaris johnsoni Shoemaker, Monocorophium insidiosum (Crawford, 1937) Monocorophium Tropichelura gomezi Ortiz, 1976 Tropichelura Cebocaris grutesca J.L. Barnard, 1964 Stenopleura atlantica Stenopleura Suborder / Family Family Colomastigidae Stebbing, 1899 Colomastigidae Family Family Corophiidae Leach, 1814 Leach, Corophiidae Family Family Cyphocarididae Lowry & Cyphocarididae Family Stoddart, 1997 Family Cheluridae Allman, 1847 Cheluridae Family Family Cebocaridae Lowry & Stoddart, Cebocaridae Family 2011 Family Calliopiidae Sars, 1893 Calliopiidae Family

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CAY

TYT LAN

X

ABC VEN

X X X X X X X COL

X X X X PAN

X X CRC

X X

NIC

GUA

HON BEL

X X

JAM HIS

X X X PRC

X X X X

X ME

)

X X X X X X X X X CUB X X X X X X X X APPENDI X 1 ( Continued Species Ledoyer, 1986 Ledoyer, Ortiz, Lalana & Varela, 2007 Varela, & Ortiz, Lalana Holsinger, 1992 Holsinger, Ortiz & Lalana, 1994 Ortiz & Lalana, (Shoemaker, 1933) (Shoemaker, Stock, 1985 (A.O. Walker, 1905) Walker, (A.O. Bellan-Santini, 1997 Bellan-Santini, Apoweckelia serrata Stock, 1985 Apoweckelia Say, 1818 Say, Gammarus mucronatus Gammarus tigrinus Sexton, 1939 gurneei Holsinger & Peck, 1968 Alloweckelia 1992 Bahadzia bozonici Holsinger, 1998 Wagner, Bahadzia jaraguensis Jaume & Bahadzia latipalpus Bahadzia setodactylus Bahadzia yagerae Ortiz & Perez, 1995 Eusirogenes adad J.L. Barnard, 1964 Eusirogenes monoculoides (Haswell, 1880) Eusiroides Rhachotropis portoricana J.L. Barnard, 1964 Rhachotropis 2007 Varela, & wimvaderi Ortiz, Lalana Rhachotropis Barnard & Thomas, 1990 Ensayara jumane Barnard & obesus (Chevreux, 1905) Eurythenes yucatanensis McKinney et al., 1980 Eusiroides Nasageneia bacescui alcoladoi Pleusiroides 1948 bartschi Shoemaker, Pontogeneia 1934 lobata Shoemaker, Rhachotropis Sennaia bidactyla Lepechinella raua J.L. Barnard, 1973 Lepechinella minikoi Nototropis McKinney, 1980 McKinney, urocarinatus Atylus Ortiz, Lalana & Sanchez-Diaz, 2000 & Sanchez-Diaz, Hoplopheonoides shoemakeri Ortiz, Lalana Aberratylus aberrantis (J.L. Barnard, 1962) longleyi Tethygeneia Ortiz & Lemaitre, 1997 Nasageneia comisariensis Ortiz & Lemaitre, Nasageneia yucatanensis Shoemaker, 1956 Hoplopheonoides obesa Shoemaker, Suborder / Family Family Gammaridae Latreille, 1802 Latreille, Gammaridae Family Karaman, 1943 Hadziidae Family Family Eusiridae Stebbing, 1888 Eusiridae Family Family Endevouridae Lowry & Stoddart, Endevouridae Family 1997 Stoddart & Lowry, Eurytheneidae Family 2004 Family Dexaminidae Leach, 1814 Leach, Dexaminidae Family Family Cyproideidae Barnard, 1974 Cyproideidae Family

1708 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013 CAY

X TYT

X LAN

X X X X X ABC

X X X X VEN

X X X X COL

X PAN

X

CRC

NIC

GUA

HON

BEL JAM

X X X HIS

X X X X X X X X X PRC

X X

X ME

)

X X X X X X X CUB X X X X X APPENDI X 1 ( Continued Stock, 1977 Species Stock, 1985 Stock, 1983 Holsinger, 1977 Holsinger, Stock, 1977 Stock & Vermeulen, 1985 Vermeulen, Stock & Holsinger, 1977 Holsinger, (Shoemaker, 1933) (Shoemaker, Crangoweckelia mixta Stock, 1985 Crangoweckelia spinicauda Stock, 1985 Crangoweckelia cenoticola Mayaweckelia Mayaweckelia yucatanensis Mayaweckelia Metaniphargus anchihalinus Metaniphargus bousfieldi Metaniphargus 2001 Winfield, & Haustorius mexicanus Ortiz, Cházaro-Olvera Metaniphargus chaetodactylus Metaniphargus Stock, 1977 emarginata Saliweckelia Stock, 1985 crenatus Metaniphargus curasavicus Stephensen, 1933 Metaniphargus curasavicus orientalis Metaniphargus haitianus Stock, 1985 Metaniphargus Stock, 1983 hyporheicus Metaniphargus 1974) jamaicae (Holsinger, Metaniphargus juberthiei Metaniphargus longipalpus Stock, 1985 Metaniphargus palpator Stock, 1977 Metaniphargus pedunculatus Stock, 1985 Metaniphargus plumicauda Stock, 1985 Metaniphargus 1991 sabulonis Vonk, Metaniphargus grandis Stock, 1985 Pintaweckelia schoenerae (Fox, 1973) Protohadzia holsingeri Stock, 1977 Saliweckelia 1990 cernua Holsinger, Tuluweckelia & Gable, 2001 nisbetae Lazo-Wasem Parhyalella perieri (Lucas, 1849) Apohyale Parhyalella whelpleyi Parhyalella media (Dana, 1853) Apohyale 1950 Hyale pygmaea Ruffo, Parargissa galatheae americana J.L. Barnard, 1961 galatheae Parargissa Suborder / Family Family Hadziidae Karaman, 1943 Hadziidae Family Family Haustoriidae Stebbing, 1906 Haustoriidae Family Family Hadziidae Karaman, 1943 Hadziidae Family 1957 Bulycheva, Hyalellidae Family Family Hyalidae Bulycheva, 1957 Bulycheva, Hyalidae Family

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013 1709

CAY

TYT LAN

X X X X X X X ABC

X X VEN

X X X X X X X X X X X X X X X X X X X X X X X COL

X X X X X X X X

PAN CRC

X X X

NIC

GUA

HON BEL

X X X JAM

X X HIS

X X X PRC

X X

X ME

)

X X X X X X X X X X X X X X X X X X CUB X X X X X X X X X X X APPENDI X 1 ( Continued Species (Stebbing, 1888) Stebbing, 1899 (Stebbing, 1899) Ortiz & Lalana, 2002 Ortiz & Lalana, Conlan, 1995 Thomas & Heard, 1979 Ortiz & Nazábal, 1984 Ortiz & Nazábal, (Bate & Westwood, 1862) Westwood, & (Bate Barnard & Thomas, 1987 Barnard & Pearse, 1912 Walker, 1904 Walker, K.H. Barnard, 1916 Chevreux, 1901 McKinney, 1980 McKinney, Lowry & Thomas, 1991 Lowry & Shoemaker, 1945 Shoemaker, Ericthonius brasiliensis (Dana, 1853) Ericthonius Corocubanus guitarti Corocubanus rubricornis (Stimpson, 1853) Ericthonius tetradonta Microjassa vidali Neoischyrocerus Ortiz, Varela & Lalana, 2011 & Lalana, Varela Ortiz, Photis lecroyae Photis longicaudata Kalke & Holland, 1978 McKinney, Photis macromana Photis melanica 1945 Photis pugnator Shoemaker, Photis sarae Souza-Filho & Serejo, 2010 Thomas & J. L. Barnard, 1991 Photis trapherus Lowry, 1972 shoemakeri Lowry, Microprotopus Photis dentata Ortiz & Thomas, 2007 Cerapus orteai Ortiz & 1997 Cerapus thomasi Ortiz & Lemaitre, Cerapus benthophilus Cerapus cudjoe Wigley, 1966 raneyi Wigley, Microprotopus (Schellenberg, 1925) togoensis (Schellenberg, Latigammaropsis Thomas & Barnard, 1989 arawakia Gammaropsis atlantica Latigammaropsis Chevalia mexicana Chevalia Parhyale inyacka Parhyale diplodactyla Protohyale carpenteri Chevalia Parhyale hawaiensis (Dana, 1853) Parhyale Protohyale macrodactyla Protohyale aviculae Chevalia Parhyale fascigera Stebbing, 1897 Parhyale Audulla chelifera Audulla Suborder / Family Family Ischyroceridae Stebbing, 1899 Ischyroceridae Family Family Isaeidae Dana, 1853 Isaeidae Family Family Hyalidae Bulycheva, 1957 Bulycheva, Hyalidae Family

1710 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013

CAY

TYT LAN

X X X X ABC

X X VEN

X X X X X X X COL

X X PAN

X X X X X X X X X X X X X CRC

X X

NIC

GUA HON

X BEL

X X X X X X X X X X X

JAM

HIS PRC

X X

X ME

)

X X X X X X X X X X X CUB X X X X X X X X X APPENDI X 1 ( Continued Species Stephensen, 1933 McKinney, 1979 McKinney, Lowry & Stoddart, 1997 Thomas & Klebba, 2006 Ortiz & Lalana, 1996 Ortiz & Lalana, Lysianopsis alba Holmes, 1903 Lysianopsis 1942) (Shoemaker, madagascarensis Orchomene 1990 stocki Bellan-Santini, Orchomene 1997 serrata Bellan-Santini, Orenoqueia Lepiduristes lepidus (J.L. Barnard, 1964) Lysianassa hummelincki Lysianassa Eclecticus eclecticus Eclecticus (Stephensen, 1933) Bonassa bonairensis McKinney, 1979 quintana McKinney, Listriella 1933) Concarnes concavus (Shoemaker, Martin, Ortiz & Atienza, 2000 Atienza, Martin, Ortiz & andresi Listriella 1966 Wigley, barnardi Listriella 1979 carinata McKinney, Listriella diffusa J.L. Barnard, 1959 Listriella kensleyi Listriella Leucothoe panpulco J.L. Barnard, 1961 Leucothoe Bellan-Santini, 1997 ayrtonia Bellan-Santini, Leucothoe Thomas & Klebba, 2007 flammosa Leucothoe Thomas & Klebba, 2007 garifunae Leucothoe kensleyi Leucothoe Thomas & Ortiz, 1995 laurensi Leucothoe Thomas & Klebba, 2007 saron Leucothoe 1789) spinicarpa (Abildgaard, Leucothoe Thomas & Klebba, 2007 barana Leucothoe Anamixis cavatura Thomas, 1997 Anamixis hanseni Stebbing, 1897 Thomas & Klebba, 2007 wuriti Leucothoe Nepanamixis dianthus Thomas, 1997 bousfieldi Liljeborgia Thomas & Klebba, 2007 ubouhu Leucothoe Thomas & Klebba, 2006 ashleyae Leucothoe Anamixis vanga Thomas, 1997 Suborder / Family Family Lysiannassidae Dana, 1849 Lysiannassidae Family Family Leucothoidae Dana, 1852 Leucothoidae Family Family Leucothoidae Dana, 1852 Leucothoidae Family Family Liljeborgiidae Stebbing, 1899 Liljeborgiidae Family

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013 1711

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X X X X ABC

X VEN

X X X X X X X X X X X X X X X X X X X X COL

X X X X X X PAN

X CRC

X X

NIC

GUA

HON BEL

X X X

JAM HIS

X

PRC

X ME

)

X X X X X X X X X X CUB X X X X X X X X X X X X X APPENDI X 1 ( Continued Species (Say, 1818) (Say, Lowry & Stoddart, 1997 Jaume & Wagner, 1998 Wagner, Jaume & Chevreux, 1907 (Bate, 1862) (Bate, (Stock, 1980) (Bate, 1862) (Bate, Thomas & J. L. Barnard, 1985 Ortiz, Lalana & Varela, 2004 Varela, & Ortiz, Lalana (S. I. Smith, 1873) Elasmopus pocillimanus Elasmopus rapax Costa, 1853 1994 Elasmopus thomasi Ortiz & Lalana, Ortiz & Lalana, 1994 Ortiz & Lalana, Elasmopus lemaitrei Elasmopus levis Elasmopus pectenicrus Elasmopus spinidactylus Elasmopus bampo J.L. Barnard, 1979 Elasmopus brasiliensis (Dana, 1853) Elasmopus elieri Kalke & Holland, 1978 Eriopisa incisa McKinney, 2003 Vonk, der Ham & Van Eriopisa mochimae Hamimaera hamigera (Haswell, 1879) Thomas & J. L. Barnard, 1988 Elasmopus balkomanus Confodiopisa caesicola appendiculata Dulichiella Thomas & J. L. Barnard, 1985 Dumosus atari Ceradocus shoemakeri Fox, 1973 Oliveira, 1953 Anelasmopus kraui Oliveira, (Stimpson, 1855) Ceradocus (Denticeradocus) rubromaculatus 1948 Shoemaker, Ceradocus (Denticeradocus) sheardi Socarnopsis catacumba (Clark & Barnard, 1985) 2003 Socarnopsis karamani Ortiz, Garcia-Debras & Lalana, apalachicola Tryphosella 1997 uristesi Bellan-Santini, Tryphosella 1980) myersi (McKinney, Gibberosus Thomas & Barnard, 1986 Resupinus spinicaudatus Anamaera hixoni Ottenwalderia kymbalion Ottenwalderia carabicus J.L. Barnard, 1964 Paracentromedon Shoemakerella cubensis (Stebbing, 1897) Shoemakerella Suborder / Family Family Melitidae Bousfield, 1973 Melitidae Family Family Megaluropidae Thomas & Megaluropidae Family Barnard, 1986 Bousfield, 1973 Melitidae Family Family Lysiannassidae Dana, 1849 Lysiannassidae Family

1712 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013 CAY

X X X

TYT LAN

X X X X ABC

X X X X X X VEN

X X X X X X X X X X X X X X COL

X X X X

PAN

CRC NIC

X

GUA

HON BEL

X

JAM

HIS

PRC

X ME

)

X X X X X X X X X X CUB X X X X X X X X X X X APPENDI X 1 ( Continued Species Ortiz, Lalana & Beltran, 1993 & Beltran, Ortiz, Lalana Vonk & Stock, 1987 Vonk (Schellenberg, 1938) (Schellenberg, (A. Costa, 1851) Krapp-Schickel & Ruffo, 2000 & Ruffo, Krapp-Schickel Vonk, 1988 Vonk, (Schellenberg, 1938) (Schellenberg, J.L. Barnard, 1969 Krapp-Schickel & Ruffo, 2000 & Ruffo, Krapp-Schickel Thomas & Barnard, 1991 Ortiz, 1976 Vonk, 1989 Vonk, Smith, 1873 Quadrimaera pieteri pacifica Quadrimaera prope Quadrimaera quadrimana (Dana, 1852) 2000) & Ruffo, Quadrimaera sarae (Krapp-Schickel 1938) Quadrimaera serrata (Schellenberg, 1933) Quadrivisio lutzi (Shoemaker, Psammogammarus scopulorum Stock, 1983 Psammomelita uncinata Quadrimaera cristianae Quadrimaera inaequipes Krapp & Gable, 2000) Quadrimaera miranda (Ruffo, Psammogammarus bluefieldensis Psammogammarus longidactylus Quadrimaera pacifica Watling, 1981 Jerbarnia americana Watling, 2000 & Jarrett, Maera jerrica Krapp-Schickel 2001 & Ruffo, Maeracoota galani Krapp-Schickel rathbunae (Pearse, 1908) Maeropsis carausui Ortiz, 1976 Mallacoota 2002 guanaense Ortiz, Garcia-Debras & Lalana, Melita leiotelson Melita longisetosa Sheridan, 1980 Melita nitida Melita palmata (Montagu, 1804) Melita persona G. Karaman, 1987 Melita Kunkel, 1910 planaterga Melita Meximaera diffidentia Kalke & Holland, 1978 McKinney, barnardi Netamelita tabaci Netamelita 1989 Nuuanu curvata Vonk, Nuuanu muelleri Suborder / Family Family Melitidae Bousfield, 1973 Melitidae Family Family Melitidae Bousfield, 1973 Melitidae Family

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013 1713

CAY TYT

X LAN

X ABC

X VEN

X X X X X X X X X X X COL

X X X PAN

X X X CRC

X X X X X X

NIC

GUA

HON BEL

X X X X X X

JAM

HIS PRC

X X

X ME

)

X X X X X X X X CUB X X X X X X X X X X X APPENDI X 1 ( Continued Species Thomas & Barnard, 1986 Thomas & Barnard, 1986 Myers, 1968 J. L. Barnard & Thomas, 1987 J. L. Barnard & J.L. Barnard, 1980 (Ortiz, 1976) (Bynum & Fox, 1977) Bellan-Santini, 1997 Bellan-Santini, Chevreux, 1888 (Shoemaker, 1933) (Shoemaker, Microphoxus minimus J.L. Barnard, 1960 Microphoxus Metharpinia oripacifica Metharpinia Harpinia excavata Eobrolgus spinosus Holmes, 1905 Eobrolgus Harpinia intermedia Pariphinotus seticoxa Parpano cebus J.L. Barnard, 1964 Parpano composturus J.L. Barnard, 1964 oculatus (Holmes, 1908) Heterophoxus 1933) floridana (Shoemaker, Metharpinia (Shoemaker, 1933) Pariphinotus seclusus (Shoemaker, Shoemaker, 1934 longimana Shoemaker, Westwoodilla Monoculodes cf. carinatus (Bate, 1857) Thomas & J. L. Barnard, 1985 cerasinus Perioculodes Hartmanodes nyei Ortiz, Lalana & Varela, 2007 Varela, & Curidia monicae Ortiz, Lalana Thomas, 1983 Curidia debrogania (Patoides) synparis (J.L. Barnard, 1964) Aceroides Americhelidium americanum (Bousfield, 1973) Americhelidium Myers, 1968 Pseudomegamphopus barnardi Pseudomegamphopus excavatus Ruffomaera williamsi 2002 Ortiz & Lalana, Spathiopus cojimarensis 1990 Thalassostygius exiguus Vonk, 1989 Ortiz & Lalana, guanarocana Victoriopisa atlantica Hornellia (Metaceradocus) Thomas, 1987 Barnard & Neomegamphopus heardi Thomas, 1987 Neomegamphopus hiatus J. L. Barnard & Neomegamphopus kalanii Ortiz, Lalana & Varela, 2008 Varela, & habanensis Ortiz, Lalana Hornellia (Metaceradocus) tequestae Hornellia (Metaceradocus) Suborder / Family Family Phoxocephalidae Sars, 1891 Phoxocephalidae Family Family Pardaliscidae Boeck, 1871 Pardaliscidae Family Family Phliantidae Stebbing, 1899 Phliantidae Family Family Ochlesidae Stebbing, 1910 Ochlesidae Family 1865 Liljeborg, Oedicerotidae Family Family Melitidae Bousfield, 1973 Melitidae Family Stebbing, 1899 Melphidippidae Family Myers, 1981 Neomegamphopidae Family Myers, 1981 Neomegamphopidae Family

1714 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013

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X LAN

X X X X X X X

ABC VEN

X X X X X X X X X X X X X X X X COL

X X X X X X PAN

X X X CRC

X X X

NIC

GUA HON

X BEL

X X X X X

JAM

HIS PRC

X X

X ME

)

X X X X X X X X CUB X X X X X X X X APPENDI X 1 ( Continued Species (J.L. Barnard, 1964) McKinney, Kalke & Holland, 1978 McKinney, Bovallius, 1886 Bovallius, (Just, 1981) Baldinger & Gable, 2002 Baldinger Thomas & J. L. Barnard, 1992 Dana, 1852 J.L. Barnard, 1962 Syrrhoe papyracea Stebbing, 1888 Syrrhoe biocellata Tiron Latacunga comanita J.L. Barnard, 1972 Ortiz & Veledo, 1988 Veledo, luquei Ortiz & Garosyrrhoe torpens (J.L. Barnard, 1962) Ileraustroe bellairsi Metatiron (J. L. Barnard, 1972) tropakis Metatiron Synopia scheeleana Synopia ultramarina Dana, 1853 Verge, Vader & Galan, 2001 Vader Verge, calypsonis Stegocephaloides texensis Parametopella 1904 gallensis Walker, Stenothoe valida Stenothoe Podocerus cristatus Thomson, 1879 Podocerus fissipes Serejo, 1995 Podocerus jareckii Ortiz & Lemaitre, 1997 Ortiz & Lemaitre, Seba robusta Bruzeliopsis turba J.L. Barnard, 1964 bigarra (J.L. Barnard, 1962) Garosyrrhoe Podocerus kleidus & Gable, 1994 Podocerus lazowasemi Baldinger americana Lowry & Stoddart, 1997 Aroui 1980 McKinney, Seba tropica Bruzelia pericu J.L. Barnard, 1972 Podocerus brasiliensis (Dana, 1853) Eudevenopus metagracilis 2000 Atienza, Ortiz, Martin & morrocoyensis Tiburonella viscana (J.L. Barnard, 1964) Tiburonella Bellan-Santini, 1997 barbada Bellan-Santini, Proharpinia 1938) epistomus (Shoemaker, Rhepoxynius Thomas & J. L. Barnard, 1983 Eudevenopus honduranus Suborder / Family Family Synopiidae Dana, 1853 Synopiidae Family Family Stegocephalidae Dana, 1853 Stegocephalidae Family Boeck, 1871 Stenothoidae Family Family Sebidae Walker, 1908 Walker, Sebidae Family Family Scopelocheiridae Lowry & Scopelocheiridae Family Stoddart, 1997 Dana, 1853 Synopiidae Family Family Podoceridae Leach, 1814 Leach, Podoceridae Family Family Phoxocephalidae Sars, 1891 Phoxocephalidae Family Barnard & Platyischnopidae Family Drummond, 1979

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013 1715

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X X X X X X X X X

ABC VEN

X X X X X COL

X X PAN

X X CRC

X X

NIC

GUA

HON

BEL JAM

X

HIS PRC

X X X

X ME

)

X X X X X X X X X X X X X X CUB X X X X X APPENDI X 1 ( Continued Species (Stebbing, 1908) Bousfield, 1984 (Claus, 1879) (Bovallius, 1885) (Bovallius, (Stephensen, 1918) Bovallius, 1887 Bovallius, Tethorchestia antillensis Tethorchestia Vemana lizata J.L. Barnard, 1964 Vemana Cystisoma latipes Bellan-Santini, 1997 carinatus Bellan-Santini, Stephonyx 1997 incertus Bellan-Santini, Stephonyx J.L. Barnard, 1964 compressa Vemana 1861 Brachyscelus crusculum Bate, Brachyscelus globiceps Stephensen, 1925 Brachyscelus macrocephalus Brachyscelus rapacoides Stephensen, 1925 1887 Bovallius, Thamneus rostratus Stephonyx biscayensis (Chevreux, 1908) Stephonyx Orchestia grillus (Bosc, 1802) Orchestia 1845) platensis (Krøyer, Platorchestia biolleyi Pseudorchestoidea 1950 Ruffo, marcuzzi Talorchestia Stephensen, 1948 margaritae Talorchestia sulensoni (Stebbing, 1899) Talorchestia 1989 karukarae Ciavatti, Tethorchestia 1930) uhleri (Shoemaker, Uhlorchestia Talorchestia fritzi Stebbing, 1903 Talorchestia Bovallius, 1885 Lanceola sayana Bovallius, Iulopis loveni Tryphana malmii Boeck, 1871 Tryphana Dairella californica Dairella (Bovallius, 1886) Cystisoma longipes (Bovallius, Ciavatti, 1989 guadalupensis Ciavatti, Floresorchestia Chelorchestia costaricana (Stebbing, 1906) Chelorchestia Suborder / Family Family Cystisomatidae Willemoes-Suhm, Family Willemoes-Suhm, Cystisomatidae 1875 Family Vitjazianidae Birstein & Birstein Vitjazianidae Family 1955 Vinogradov, Milne Edwards, 1830 Suborder Hyperiidea Stephensen, 1923 Brachyscelidae Family Stephensen, 1923 Brachyscelidae Family Family Uristidae Hurley, 1963 Hurley, Uristidae Family Family Lanceolidae Bovallius, 1887 Lanceolidae Family Family Iulopididae Zeidler, 2004 Zeidler, Iulopididae Family Family Lycaeidae Claus, 1879 Lycaeidae Family Family Dairellidae Bovallius, 1887 Bovallius, Dairellidae Family Family Talitridae Rafinesque, 1815 Talitridae Family

1716 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013

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X X X X X X X X X X

COL

PAN

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X X

HIS

PRC

X ME

)

X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X CUB X X X X X X X APPENDI X 1 ( Continued Species (Stephensen, 1924) Claus, 1879 Bowman, 1973 Bowman, 1973 Stephensen, 1925 Stebbing, 1888 Chevreux, 1900 Hyperionyx macrodactylus Lestrigonus bengalensis Giles, 1887 Hyperietta vosseleri (Stebbing, 1904) Hyperietta Hyperioides longipes Chevreux, 1900 Hyperietta luzoni (Stebbing, 1888) Hyperietta parviceps Bowman, 1973 Hyperietta stebbingi Hyperietta stephenseni Hyperietta (Bovallius, 1889) Lestrigonus crucipes (Bovallius, Lestrigonus latissimus (Bovallius, 1889) 1901) (Vosseler, Lestrigonus macrophthalmus (Streets, 1878) Cranocephalus scleroticus 1887 Bovallius, Glossocephalus milneedwardsi (Claus, 1871) tenuirostris Leptocotis 1887 Oxycephalus clausi Bovallius, Oxycephalus piscator Milne Edwards, 1830 Rhabdosoma minor Fage, 1954 (Woltereck, 1904) Mimonectes gaussi (Woltereck, Lycaea vincentii Lycaea zamboangae (Stebbing, 1888) Lycaeopsis Lycaea bovallii Lycaea bovallioides Lycaea pachypoda (Claus, 1879) Lycaea pauli Stebbing, 1888 Lycaea pulex Marion, 1874 Lycaea antennarius (Claus, 1871) Simorhynchotus Lestrigonus schizogeneios (Stebbing, 1888) spinifera Claus, 1879 Phronimopsis fusca (Dana, 1853) Themistella 1925 bajensis Shoemaker, Lycaea Lycaeopsis themistoides Lycaeopsis Suborder / Family Family Lestrigonidae Zeidler, 2004 Zeidler, Lestrigonidae Family Family Oxycephalidae Dana, 1853 Oxycephalidae Family Family Mimonectidae Bovallius, 1887 Bovallius, Mimonectidae Family Family Lycaeidae Claus, 1879 Lycaeidae Family Family Lycaeidae Claus, 1879 Lycaeidae Family Family Lycaeopsidae Chevreux, 1913 Lycaeopsidae Family

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HIS

PRC

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)

X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X CUB X X X X X X APPENDI X 1 ( Continued Species Milne Edwards, 1830 (Claus, 1862) Claus, 1879 Stebbing, 1888 Bate, 1862 Bate, Vosseler, 1901 Vosseler, Streets, 1887 Bowman, 1978 (Claus, 1879) Bovallius, 1887 Bovallius, Stebbing, 1888 Phronima colletti Phronima 1901 curvipes Vosseler, Phronima pacifica Phronima (Claus, 1879) sphaeroma Thyropus sedentaria (Forskal, 1775) Phronima Streetsia challengeri Streetsia mindanaonis (Stebbing, 1888) Streetsia porcella Streetsia 1887) steenstrupi (Bovallius, Streetsia crassipes Claus, 1879 Paraphronima gracilis Claus, 1879 Paraphronima Claus, 1879 Parascelus edwardsi Parascelus typhoides Schizoscelus ornatus Claus, 1879 Anchylomera blossevillei Guérin-Méneville, 1836 solitaria Guérin-Méneville, Phronima stebbingi Phronima elongata Phronimella semilunata Risso, 1882 Phrosina Rhabdosoma whitei Primno abyssalis Bowman, 1968 Primno brevidens 1986 Primno evansi Sheader, Primno johnsoni Bowman, 1978 Primno latreillei 1836 Guérin-Méneville, Primno macropa Amphithyrus bispinosus Claus, 1879 Amphithyrus glaber Spandl, 1924 1982 Amphithyrus muratus Volkov, Amphithyrus sculpturatus Claus, 1879 Hemityphis tenuimanus Claus, 1879 Suborder / Family Family Phronimidae Rafinesque, 1815 Rafinesque, Phronimidae Family Family Parascelidae Bate, 1862 Bate, Parascelidae Family Family Paraphronimidae Bovallius, 1887 Bovallius, Paraphronimidae Family 1862 Bate, Parascelidae Family Family Phrosinidae Dana, 1853 Family Family Oxycephalidae Dana, 1853 Oxycephalidae Family Family Platyscelidae Bate, 1862 Bate, Platyscelidae Family

1718 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013

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X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X CUB X X X X X X X APPENDI X 1 ( Continued Species Stephensen, 1925 Chevreux, 1914 Stebbing, 1888 (Bovallius, 1887) (Bovallius, M. Vinogradov, 1964 Vinogradov, M. Scina crassicornis (Fabricius, 1775) Scina curvidactyla Scina damasi Pirlot, 1929 1998 australis Zeidler, Scina hurleyi Scina indica 1926 Scina langhansi Wagler, 1885) (Bovallius, Scina marginata 1926 Scina nana Wagler, Scina pacifica Scina similis Stebbing, 1895 Scina stenopus Stebbing, 1895 Acanthoscina acanthodes (Stebbing, 1895) Acanthoscina (Sars, 1882) Scina borealis Paratyphis maculatus Claus, 1879 Paratyphis parvus Claus, 1887 Stebbing, 1888 Paratyphis promontori Paratyphis spinosus Spandl, 1924 armatus (Claus, 1879) Platyscelus crustulatus (Claus, 1879) Platyscelus ovoides (Risso, 1816) Platyscelus serratulus Stebbing, 1888 Platyscelus Claus, 1879 forcipatus Tetrathyrus intermedia Stebbing, 1888 Eupronoe laticarpa Eupronoe maculata Claus, 1879 Eupronoe minuta Claus, 1879 Eupronoe gracilis Claus, 1879 Paralycaea hoylei Paralycaea crustulum Claus, 1879 Parapronoe parva Claus, 1879 Parapronoe 1836 capito Guérin-Méneville, Pronoe Suborder / Family Family Scinidae Stebbing, 1888 Scinidae Family Family Platyscelidae Bate, 1862 Bate, Platyscelidae Family 1862 Bate, Platyscelidae Family Pronoidae Dana, 1853 Family

Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013 1719

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X X X X X X X X X X X X X CUB X X X X APPENDI X 1 ( Continued Species Stock, 1979 Stock, 1977 Stock, 1976 Behning & Woltereck, 1912 Woltereck, Behning & Bovallius, 1887 Bovallius, Bovallius, 1887 armata Bovallius, Vibilia australis Stebbing, 1888 Vibilia Tattersall, 1906 Tattersall, Scina submarginata 1885) (Bovallius, Scina tullbergi 1906 Scina vosseleri Tattersall, Thurston, 1976 Scina wagleri atlantis 1912 Woltereck, chuni Behning & Vibilia 1887 gibbosa Bovallius, Vibilia Lucas, 1845 jeangerardi Vibilia Stebbing, 1888 propinqua Vibilia stebbingi Vibilia Ingolfiella fontinalis Ingolfiella fuscina Dojiri & Sieg, 1987 Ingolfiella grandispina Stock, 1979 Ingolfiella Stock, 1979 margaritae Ingolfiella putealis Ingolfiella quadridentata Ingolfiella tabularis Stock, 1977 Ingolfiella Vibilia viatrix Vibilia Suborder / Family Family Vibiliidae Dana, 1853 Vibiliidae Family Family Scinidae Stebbing, 1888 Scinidae Family Suborder Ingolfiellidea Hansen, 1903 Suborder Ingolfiellidea Hansen, 1903 Ingolfiellidae Family

1720 Rev. Biol. Trop. (Int. J. Trop. Biol. ISSN-0034-7744) Vol. 61 (4): 1681-1720, December 2013