Benthic amphipods (Amphipoda: Gammaridea and Corophiidea) from the Mexican southeast sector of the Gulf of Mexico: checklist, new records and zoogeographic comments

Pedro L Ardisson; Carlos E.  Paz Ríos

The southeast region of the Gulf of Mexico is considered to be biologically important, because it is a connection and transition zone between the Caribbean and the Gulf of Mexico, harboring great marine biodiversity. Nevertheless, benthic amphipods have been poorly studied in the Mexican southeast sector of the Gulf of Mexico with few studies listing species. The aim of this study is to provide an update checklist of species for the Mexican southeast sector (based on literature review and records from the present study) as well as a brief zoogeographical analysis for the Gulf of Mexico amphipod fauna, putting them in context with the fauna on the tropical western Atlantic. Fifty-five species were listed for the Mexican southeast sector; 36 of them showed a geographical extension to the Yucatan continental shelf representing 23 new records for the Mexican southeast sector, nine for the southeast region and four for the Gulf of Mexico. Based on the zoogeographical analysis, there is support of the application of Carolinian and Caribbean zoogeographic provinces to amphipods in the Gulf of Mexico

Zootaxa 3635 (2): 137–173 www.mapress.com / zootaxa / Copyright © 2013 Magnolia Press ISSN 1175-5326 (print edition) Article ZOOTAXA ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3635.2.4 http://zoobank.org/urn:lsid:zoobank.org:pub:A64F4DF7-B31C-45FE-A9B3-BF933609D9DC Benthic amphipods (Amphipoda: Gammaridea and Corophiidea) from the Mexican southeast sector of the Gulf of Mexico: checklist, new records and zoogeographic comments CARLOS E. PAZ-RÍOS1 & PEDRO-LUIS ARDISSON2 Departamento de Recursos del Mar, Cinvestav, Carretera antigua a Progreso, km 6, Apdo. Postal 73, Cordemex 97310 Merida, Yucatan, Mexico. E-mail: 1cpaz@mda.cinvestav.mx, 2ardisson@mda.cinvestav.mx Abstract The southeast region of the Gulf of Mexico is considered to be biologically important, because it is a connection and transition zone between the Caribbean and the Gulf of Mexico, harboring great marine biodiversity. Nevertheless, benthic amphipods have been poorly studied in the Mexican southeast sector of the Gulf of Mexico with few studies listing species. The aim of this study is to provide an update checklist of species for the Mexican southeast sector (based on literature review and records from the present study) as well as a brief zoogeographical analysis for the Gulf of Mexico amphipod fauna, putting them in context with the fauna on the tropical western Atlantic. Fifty-five species were listed for the Mexican southeast sector; 36 of them showed a geographical extension to the Yucatan continental shelf representing 23 new records for the Mexican southeast sector, nine for the southeast region and four for the Gulf of Mexico. Based on the zoogeographical analysis, there is support of the application of Carolinian and Caribbean zoogeographic provinces to amphipods in the Gulf of Mexico. Key words: Yucatan, Crustacea, Peracarida, marine, biodiversity, biogeography Introduction Benthic amphipods inhabit a variety of substrata, including hard and soft bottoms, submerged aquatic vegetation and other benthic organisms (e.g. sponges, corals, anemones, tunicates). The high morphological diversity of amphipods dwelling on sandy sediments suggests a great adaptive radiation (Bousfield 1970). Benthic amphipods obtain a great part of their food from the sediment or water column, displaying major feeding modes such as detritivory, filter-feeding, herbivory, predatory, scavenging and sand-cleaning (Bousfield 1973; Biernbaum 1979). This ecological diversity has conferred them the ability to colonize all marine habitats and a broad interval of depth, from intertidal to abyssal zone (Thomas 1993; Escobar-Briones & Winfield 2003; LeCroy et al. 2009). Biologically, the reproductive strategy in amphipods is to brood their eggs in a marsupium, releasing them as miniature versions of adults (direct development), and in some cases providing parental care (Thiel 1999). Thereby species show a trade-off condition to achieve reproductive success (Sainte-Marie 1991; William et al. 2001). Amphipods are important for bioturbation because they promote the interchange of gases (mainly oxygen and nitrogen) and contribute high rates of secondary production for energy flow to higher trophic levels (Marques & Bellan-Santini 1993; Duffy & Hay 2000; Soliman & Rowe 2008). These ecological and biological traits have positioned amphipods as one key link between the sediment and the water column. Despite their importance, there are many marine regions where amphipods are often ignored, leaving a gap of information which provides an erroneous assessment of the regional biodiversity. One approximation to solve this crisis is to make checklists of species based on reliable identification or taxonomic revision. This is the first step in establishing a framework of how many and which species are in which habitats (alpha, beta and gamma diversity). According to Hendrickx and Harvey (1999), a checklist of species provides information about the complexity of the biological communities and the live resources available, helping to recognize and to delimit the protected areas and to identify potential anthropogenic impacts. Accepted by G. Karaman: 11 Feb. 2013; published: 26 Mar. 2013 137 Regarding amphipod fauna in the Gulf of Mexico (GM), the study of gammarideans sensu lato has been constant and productive. It can be noted in the number of species, for instance, of the 418 species (excluding anchialine, freshwater and troglobitic) documented in the tropical western Atlantic (TWA) (Ortiz et al. 2007), 131 of them (31%) have been reported in the GM (LeCroy et al. 2009). In addition, based on the composition and abundance of benthic macrofauna, amphipods in the northeastern region of the GM represent the second most important taxa, just below the polychaetes (Brooks et al. 2006). However, excluding the relatively well-known amphipod fauna of the Florida Keys and Northwestern Cuba, this taxon has been poorly studied in the southeast region of the GM. For instance, to date only two studies recording benthic amphipods in the Mexican southeast sector have been published (Ortiz 1979; Oliva-Rivera & Jiménez-Cueto 1997), becoming the sector with the lowest number of species in the GM (Escobar-Briones & Winfield 2003). Likewise, only two studies describing the distribution and abundance patterns of species have been published (Paz-Ríos 2008; Rodríguez-Pliego et al. 2011). More recently, in adjacent waters toward the southeastern region Escobar-Briones and Jiménez-Guadarrama (2010) described the distribution and abundance patterns of peracarids (including amphipods) on the Campeche Bank, although their analysis was at the family level. Simões et al. (2010) provided a list of crustaceans for the coast of Yucatan State, including amphipods, but this list is partially based on unconfirmed material from biological collections. Therefore, for updating and providing new information on the amphipod fauna in southeastern GM, the aims of this study are 1) to provide an annotated checklist of species for the Mexican southeast sector of the GM, 2) to report geographical extensions for those species and 3) to provide a brief analysis of zoogeographical affinity of the GM amphipod fauna inside the TWA amphipod fauna. Methods Study area. Sampling was carried out in the Yucatan continental shelf at 20–40 m depth, in the southeast region of the GM (Fig 1). The Yucatan continental shelf includes the submerged portion of a shallow limestone plateau, which is expanded 220 km toward the northeast (Logan et al. 1969). These same authors also state that the inner continental shelf has a gentle slope from south to north and is bounded on the west, north, and east by a precipitous continental slope that plunges from the submerged plateau margin to the abyssal depths of the GM and Caribbean. The water column on the continental shelf forms part of a superficial layer (100–150 m) which is well-known as a layer of mixture (Pica & Pineda 1991). The dominant current comes from a branch of the North Atlantic current, which penetrates through the Yucatan strait to the GM (DiMarco et al. 2005). Substratum is mainly composed of carbonated sand of autochthonous origin with more than 75% of carbonates, which reflects the presence of coral reefs on the shelf (Logan et al. 1969; Balsam & Beeson 2003). Sample collection and processing. Benthic samples were collected on board the R/V Onjuko during an oceanographic cruise, “Yucatan 2004-A” (April 5–16, 2004) carried out by the Department of Oceanography of the Mexican Navy. Eighteen stations were positioned (Fig 1A, Table 1) and each station was sampled in duplicate using a Petite Ponar grab (0.23 m-2), excepting the stations 2, 14 and 18 due to adverse meteorological conditions. Both sediment and fauna were analyzed. Sediment texture was analyzed following Bale and Kenny (2005) and is characterized in Table 1. Faunal samples were anaesthetised with MgCl2, and then fixed with 10% formaldehyde solution buffered with seawater before storage. The samples were washed through 500 µm size mesh and transferred to 70% ethanol. Species were sorted and identified to the lower taxonomical level. All species are deposited in the Colección de Invertebrados Bentónicos de Yucatán (CYMX), Cinvestav; each species herein reported has a catalog number for the collection (collected and identified by C.E. Paz-Ríos). The classification of Gammaridea is based on Martin and Davis (2001). The classification of Corophiidea is based on Myers and Lowry (2003); members of family Caprellidae and Cyamidae were excluded. Data analysis. An annotated checklist of amphipods on the Mexican southeast sector is provided. It constitutes a compilation based on literature and records from identified species in the present study; doubtful records and personal communication were excluded. The checklist includes material examined (if it is the case), distribution, habitat and remarks (if necessary). 138 · Zootaxa 3635 (2) © 2013 Magnolia Press PAZ-RÍOS & ARDISSON FIGURE 1. Localization of the study area in the Gulf of Mexico. (A) Sampling stations on the northern Yucatan continental shelf; (○) sample not collected due to adverse meteorological conditions. (B) Nested spatial scales in the Gulf of Mexico for recording the geographical extension of species; dashed line represents the Mexico's exclusive economic zones. BENTHIC AMPHIPODS IN SOUTHEASTERN GULF OF MEXICO Zootaxa 3635 (2) © 2013 Magnolia Press · 139 TABLE 1. Summary information for sampling stations on the northern Yucatan continental shelf, southeastern Gulf of Mexico. The sampling stations 12, 14 and 18 were not collected due to adverse meteorological conditions. Station Coordinate Date Depth (m) Sediment texture 1 21°47’N 87°30’W 9 April 2004 19.6 Medium grain size, poorly sorted 2 22°02’N 87°30’W 9 April 2004 31.4 Fine grain size, poorly sorted 3 22°17’N 87°29’W 9 April 2004 43.8 Medium grain size, poorly sorted 4 22°17’N 88°00’W 10 April 2004 40.9 Medium grain size, poorly sorted 5 22°06’N 88°59’W 10April 2004 31.4 Coarse grain size, moderately sorted 6 21°53’N 88°00’W 10 April 2004 19.3 Medium grain size, moderately sorted 7 21°52’N 88°30’W 10 April 2004 20.9 Medium grain size, poorly sorted 8 22°03’N 88°30´W 10 April 2004 30.1 Fine grain size, very poorly sorted 9 22°17’N 88°30’W 10 April 2004 42.1 Fine grain size, moderately sorted 10 22°06’N 89°00’W 11 April 2004 42.2 Fine grain size, poorly sorted 11 21°59’N 89°00’W 11 April 2004 35.3 Medium grain size, poorly sorted 12 21°49’N 89°00’W 11 April 2004 24.9 - 13 21°28’N 90°24’W 14 April 2004 14.2 Medium grain size, moderately sorted 14 21°41’N 90°18’W 14 April 2004 33.9 - 15 21°53’N 90°17’W 15 April 2004 42.9 Fine grain size, moderately sorted 16 22°01’N 89°36’W 15 April 2004 40.5 Fine grain size, moderately sorted 17 21°49’N 89°35’W 15 April 2004 33.4 Medium grain size, poorly sorted 18 21°28’N 89°36’W 15 April 2004 22.5 - From identified species, the most abundant and widely distributed species were determined by the absolute abundance (Ind) and relative frequency (%). To record the geographic extension of species three nested spatial scales were established (Fig 1B): 1) Mexican southeast sector, 2) southeast region and 3) GM. To explore the zoogeographical affinity of the GM fauna with the TWA fauna, a hierarchical cluster analysis was applied to one species composition matrix of presence/absence. A matrix of species (excluding anchialine, freshwater and troglobitic) for the TWA fauna was derived from compilation by Ortiz et al. (2007) for six regions: Bahamas, Barbados, Bermuda, Caribbean, Florida and GM. Likewise, a matrix of species for the GM fauna was derived from compilation by LeCroy et al. (2009) for four regions: Northeast, Southeast, Southwest and Northwest. Thus, a total of 418 and 232 species were listed for the TWA and the GM, respectively. After that, another matrix was generated from the two above compilations to carry out the analysis; to avoid redundant information and ambiguous interpretations in analysis, of those regions proposed by Ortiz et al. (2007), the Florida region was excluded and interchanged by the four regions of the GM employed by LeCroy et al. (2009). Thus, this last matrix included 381 shared species among eight regions (Appendix 1). The Sorensen coefficient as similarity index and the average linkage clustering were used to do the hierarchical cluster analysis due to the differences in sample size and sampling effort at the different compilations. The statistical software used for analysis was PRIMER v6 (Clarke & Gorley 2006). Results Faunal composition Based on literature review and records from the present study, the amphipod fauna for the Mexican southeast sector is made up so far by 55 species. The suborder Gammaridea is represented by 16 families, 26 genera and 34 species, whereas the suborder Corophiidea is represented by nine families, 14 genera and 21 species. Families with the highest numbers of species were Melitidae (8), Photidae (6), Ampeliscidae (5), Aoridae (4) and Lysianassidae (4). These families represented 49% of all the fauna. 140 · Zootaxa 3635 (2) © 2013 Magnolia Press PAZ-RÍOS & ARDISSON A total abundance of 267 individuals was counted from the collected samples in the Yucatan continental shelf, identifying an assemblage of 39 species. The most abundant and widely distributed species in the area were Eudevenopus honduranus Thomas & Barnard, 1983 (60 Ind, 46%), Gammaropsis sp. B sensu LeCroy, 2000 (32 Ind, 54%), Acuminodeutopus naglei (Bousfield, 1973) (27 Ind, 46%) and Americhelidium americanum (Bousfield, 1973) (26 Ind, 46%). Annotated checklist of amphipods on the Mexican southeastern sector From the compilation of literature and records of the present study, so far 55 species have been recorded (listed below) in the Mexican southeastern sector. Nineteen of these had been already recorded in the sector and the other 36 were added in the present study. From identified species in the present study, 36 showed a geographical extension to the Yucatan continental shelf. Twenty-three species are the first records for the Mexican southeast sector (two of which are also the first records for the southwest region), nine species are the first records for the southeast region, and four species are the first records for the GM. These species are indicated by a symbol ( ) preceding their names. ► Order Amphipoda Latreille, 1816 Suborder Gammaridea Latreille, 1802 Family Ampeliscidae Costa, 1857 ►Ampelisca agassizi (Judd, 1896) Material examined. CYMX-68-PY, 1 male, station 3; CYMX-69-PY, 1 female, station 9. General distribution. Western Atlantic; eastern Pacific (Mills 1965, 1967; Ortiz et al. 2007; García-Madrigal 2008; LeCroy et al. 2009). GM distribution. Louisiana; Mobile Bay (Goeke & Heard 1984); Perdido Key (Rakocinski et al. 1996); Tamaulipas and Veracruz continental shelf (Escobar-Briones & Winfield 2003); Biloxi Bay (Pederson & Peterson 2002); Bay of Campeche (Winfield et al. 2006). Habitat. Coarse sand or in fine to medium sand mixed with shell hash; at depths of 2 to 450 m (LeCroy 2002). Remarks. First record for the southeast region. ►Ampelisca brevisimulata Barnard, 1954 Material examined. CYMX-67-PY, 1 male, station 4. General distribution. Western Atlantic; eastern Pacific (Barnard 1954a, b; Mills 1965; Ortiz et al. 2007; García-Madrigal 2008; LeCroy et al. 2009). GM distribution. Mexican southwestern continental shelf (Escobar-Briones & Winfield 2003); Veracruz continental shelf (Winfield & Ortiz 2011). Habitat. Sand to silt clay sediments; at depths of 11 to 172 m (García-Madrigal 2008). Remarks. First record for the southeast region. Ampelisca cristata Holmes, 1908 General distribution. Western Atlantic; eastern Pacific (Barnard 1954b; Mills 1967; Ortiz et al. 2007; GarcíaMadrigal 2008; LeCroy et al. 2009). GM distribution. Off Port Charlotte; Yucatan continental shelf (Ortiz 1979); Florida Bay (LeCroy 2002). Habitat. Sand sediments; at depths of 2 to 152 m (Ortiz et al. 2007; García-Madrigal 2008; LeCroy et al. 2009). BENTHIC AMPHIPODS IN SOUTHEASTERN GULF OF MEXICO Zootaxa 3635 (2) © 2013 Magnolia Press · 141 Ampelisca lobata Holmes, 1908 General distribution. Western Atlantic; eastern Pacific (Barnard 1954b; Mills 1967; Ortiz et al. 2007; GarcíaMadrigal 2008; LeCroy et al. 2009). GM distribution. central West Florida shelf; Yucatan continental shelf (Ortiz 1979); Río Soto la Marina Lagoon (Cházaro-Olvera et al. 2002); Tamaulipas continental shelf (Winfield et al. 2006); Veracruz coral reef system (Winfield et al. 2007a); Northwestern Cuba (Ortiz & Lalana 2010). Habitat. Soft bottoms; from intertidal depths to 450 m (Ortiz et al. 2007; García-Madrigal 2008; LeCroy et al. 2009). Ampelisca vadorum Mills, 1963 Material examined. CYMX-74-PY, 2 females, station 3; CYMX-73-PY, 1 female, station 10. General distribution. Western Atlantic (Mills 1963, 1965; Bousfield 1973; Nelson 1995; Ortiz et al. 2007; LeCroy et al. 2009). GM distribution. Key Largo (Shoemaker 1933a); western Florida (Mills 1963); Gulf Coast of the USA (Mills 1965); Yucatan continental shelf (Ortiz 1979); Terminos Lagoon (Ledoyer 1986); off the Papaloapan river (Escobar-Briones & Winfield 2003); Bay of Campeche (Winfield et al. 2006). Habitat. Sand bottoms, often mixed with shell; from intertidal depths to 70 m (Bousfield 1973). Family Amphilochidae Boeck, 1871 ►Gitana dominica Thomas & Barnard, 1990 Material examined. CYMX-71-PY, 1 female, 1 unsexed, station 2; CYMX-70-PY, 1 male, station 5; CYMX72-PY, 1 female, station 10. General distribution. Caribbean (Thomas & Barnard 1990; Ortiz et al. 2007; Winfield & Escobar-Briones 2007). GM distribution. Previously unreported. Habitat. Sand among reef outcrop; at depths of 4 to 112 m (Thomas & Barnard 1990; Winfield & EscobarBriones 2007). Remarks. First record for the GM. LeCroy (2002) reported to Gitana cf. dominica, one species very similar to G. dominica which is presented at the northeastern Florida. The difference between records is in the serration of the ventral margins of coxae 2–4, which is less on G. dominica than G. cf. dominica (LeCroy 2002). Family Argissidae Walker, 1904 ►Argissa hamatipes (Norman, 1869) Material examined. CYMX-89-PY, 1 female, station 1; CYMX-90-PY, 1 male, 1 unsexed, station 2; CYMX92-PY, 1 female, station 3; CYMX-91-PY, 1 female, station 5; CYMX-93-PY, 1 female, station 11. General distribution. Arctic Ocean; Northeastern and Northwestern Atlantic; Northeastern Pacific; Southern Africa (LeCroy 2002; LeCroy et al. 2009). GM distribution. Southern Texas (McKinney 1977); Perdido Key (Rakocinski et al. 1996); Cape Romano; Tampa Bay; Apalachee Bay; lower Florida Keys (LeCroy 2002); Louisiana continental shelf (Dubois et al. 2009). Habitat. Pelagic, but mainly epibenthic on hard and soft bottoms; at depths of 2 to 1720 m (LeCroy 2002). Remarks. First record for the Mexican southeast sector. Family Bateidae Stebbing, 1906 ►Batea cuspidate (Shoemaker, 1926) Material examined. CYMX-63-PY, 1 male, station 7. General distribution. Central western Atlantic (Ortiz et al. 2007; LeCroy et al. 2009; Coleman 2011). 142 · Zootaxa 3635 (2) © 2013 Magnolia Press PAZ-RÍOS & ARDISSON GM distribution. Western Florida (Shoemaker 1926; Ortiz 1991); off Key Largo (Shoemaker 1933a); Faro Roncali; Playa Las Tumbas (Varela et al. 2003); lower Florida Keys; Florida Bay; Cape Sable; Port Richey; Homosassa Island (LeCroy 2004); Bay of Campeche (Winfield et al. 2006); Northwestern Cuba (Ortiz & Lalana 2010). Habitat. Thalassia sp. or Halodule sp. grassbeds, algae on sandy, shelly or muddy bottoms, live bottom areas near corals and sponges with patches of sand or shell covered with green macroalga Caulerpa sp.; at depths of 1 to 49 m (Ortiz 1991; LeCroy 2004). Remarks. First record for the Mexican southeast sector. Family Hyalidae Bulycheva, 1957 Protohyale macrodactyla (Stebbing, 1899) General distribution. Western Atlantic; Madagascar; western Africa (McKinney 1977; Ortiz et al. 2007; LeCroy et al. 2009). GM distribution. Yalahau Lagoon (Oliva-Rivera & Jiménez-Cueto 1997); Río Soto la Marina Lagoon; Camaronera Lagoon; Terminos Lagoon (Cházaro-Olvera et al. 2002). Habitat. Free-swimming on algae and hard substrates; from intertidal depths to shallow subtidal zone (Bousfield & Hendrycks 2002; LeCroy et al. 2009). Family Leucothoidae Dana, 1852 ►Anamixis cavatura Thomas, 1997 Material examined. CYMX-66-PY, 2 males, station 7. General distribution. Tropical western Atlantic (Thomas 1997; LeCroy 2007; Ortiz et al. 2007; LeCroy et al. 2009; White 2011). GM distribution. From the Mississippi Delta to Tampa; Florida Keys (Pearse 1912; Shoemaker 1933b; Thomas 1997; White 2011); Northwestern Cuba (Varela et al. 2003; Ortiz & Lalana 2010). Habitat. Coral rubble; internal cavities of small asconoid sponges and colonial ascidians; at depths of 1 to 5 m (Thomas 1997; White 2011). Remarks. First record for the Mexican southeast sector. This record extended its maximum depth from 5 m (White 2011) to 20.9 m. Escobar-Briones and Winfield (2003) reported to this species “in the Bank of Campeche in the southern Gulf”; however, no record of that occurrence had been established. Family Liljeborgiidae Stebbing, 1899 ►Liljeborgia bousfieldi McKinney, 1979 Material examined. CYMX-31-PY, 1 male, station 7. General distribution. Central western Atlantic (McKinney 1979; Oliva-Rivera & Jiménez-Cueto 1992; Ortiz et al. 2007; LeCroy et al. 2009). GM distribution. Florida Keys (Thomas 1993; LeCroy 2011). Habitat. Among algae and on coral reefs; at depths of 0.5 to 3m (McKinney 1979; Thomas 1993). Remarks. First record for the Mexican southeast sector. This record extended its maximum depth from 3 m (McKinney 1979) to 20.9 m. ►Listriella barnardi Wigley, 1966 Material examined. CYMX-15-PY, 1 female, station 1; CYMX-16-PY, 1 female, station 2; CYMX-14-PY, 2 males, 1 female, station 3. General distribution. Western Atlantic (McKinney 1979; Thomas 1993; Nelson 1995; LeCroy et al. 2009; LeCroy 2011). BENTHIC AMPHIPODS IN SOUTHEASTERN GULF OF MEXICO Zootaxa 3635 (2) © 2013 Magnolia Press · 143 GM distribution. North Texas coast (McKinney 1979); Perdido Key (Rakocinski et al. 1996); Florida Keys (Thomas 1993); Louisiana continental shelf (Dubois et al. 2009). Habitat. Patches of Zostera sp. and Thalassia sp. grassbeds (Wigley 1966; Lewis 1984); common on soft bottoms (McKinney 1979); at depths of 1 to 22 m (Thomas 1993; LeCroy 2011). They also are inhabitants of polychaetes tubes (Bousfield 1973). Remarks. First record for the Mexican southeast sector. This record extended its maximum depth from 22 m (LeCroy 2011) to 43.8 m. Escobar-Briones et al. (2002) listed this species for Mexico; however, no record of that occurrence had been established. LeCroy (2011) reviewed the morphology of this species for the Florida and GM material, identifying several morphs over a wide range of locations, treating them as Listriella cf. barnardi. She determined that no important morphological differences on those materials for considering them as another species. Family Lysianassidae Dana, 1849 ►Hippomedon pensacola Lowry & Stoddart, 1997 Material examined. CYMX-7-PY, 1 female, station 4. General distribution. South Atlantic Bight; GM (LeCroy 2007; LeCroy et al. 2009). GM distribution. Santa Rosa Island (Lowry & Stoddart 1997); Apalachee Bay (LeCroy 2007). Habitat. Infaunal in sandy beaches or muddy substrates; at depths of 5 to 34 m (Lowry & Stoddart 1997; LeCroy et al. 2009). Remarks. First record for the southeast region. This record extended its maximum depth from 34 m (LeCroy 2007) to 40.9 m. ►Hippomedon sp. B sensu LeCroy, 2007 Material examined. CYMX-9-PY, 1 female, station 1; CYMX-10-PY, 2 females, station 3; CYMX-8-PY, 1 unsexed, station 8; CYMX-11-PY, 1 unsexed, station 10. General distribution. South Carolina to southeastern Florida; GM (LeCroy 2007). GM distribution. Cape Sable to Cape Romano; Tampa Bay (LeCroy 2007). Habitat. In sandy bottoms; at depths of 8 to 17 m (LeCroy 2007). Remarks. First record for the southeast region. This record extended its maximum depth from 17 m (LeCroy 2007) to 43.8 m. The main difference between this species and H. pensacola, the only other member of the genus know from GM is, the lack of a notch on epimeron 3 and the presence of an expanded distal flange on the dorsolateral margin of the peduncle of uropod 2 (LeCroy 2007). ►Orchomenella thomasi Lowry & Stoddart, 1997 Material examined. CYMX-4-PY, 1 male, station 5. General distribution. GM (LeCroy et al. 2009). GM distribution. Sanibel Island (Lowry & Stoddart 1997); Louisiana coast (LeCroy 2007). Habitat. Fine to medium sand with crushed shell, dead bryozoans or calcareous algae present; at depths of 10 to 73 m (Lowry & Stoddart 1997). Remarks. First record for the southeast region. ►Shoemakerella cubensis (Stebbing, 1897) Material examined. CYMX-5-PY, 1 male, 1 female, station 7. General distribution. GM; Caribbean (Lowry & Stoddart 1997; Ortiz et al. 2007; LeCroy et al. 2009). GM distribution. Dry Tortugas to Panama City; Mississippi Delta (Thomas 1993; Lowry & Stoddart 1997); Mexican southwestern continental shelf (Escobar-Briones & Winfield 2003); Veracruz coral reef system (Winfield et al. 2007a); Northwestern Cuba (Ortiz & Lalana 2010); Tuxpan river (Winfield & Ortiz 2011). Habitat. In grassbeds, rubble, mixed algae and sandy bottoms; at depths of 1 to 69 m (Lowry & Stoddart 1997; LeCroy 2007). Remarks. First record for the Mexican southeast sector. 144 · Zootaxa 3635 (2) © 2013 Magnolia Press PAZ-RÍOS & ARDISSON Family Megaluropidae Thomas & Barnard, 1986 ►Gibberosus myersi (McKinney, 1980) Material examined. CYMX-94-PY, 3 males, 1 unsexed, station 7. General distribution. Western Atlantic; eastern Pacific (Thomas & Barnard 1986a; García-Madrigal 2008; LeCroy et al. 2009). GM distribution. Looe Key Reef (Thomas & Barnard 1986a); Perdido Key (Rakocinski et al. 1996); lower Florida Keys to Cape Romano; Tampa Bay (LeCroy 2007). Habitat. In or near grassbeds, and from fine to medium sand often mixed with shell; at depths of 1 to 29 m (Thomas & Barnard 1986a; LeCroy 2007). This species feeds by perching upside down, filtering suspended material from the water column (Thomas & Barnard 1986a). Remarks. First record for the Mexican southeast sector. Family Melitidae Bousfield, 1973 Ceradocus sheardi Shoemaker, 1948 General distribution. Tropical western Atlantic (Ortiz et al. 2007; Krapp-Schickel & Vader 2009; LeCroy et al. 2009). GM distribution. Dry Tortugas (Shoemaker 1948); Yucatan continental shelf (Ortiz 1979); Northwestern Cuba (Varela et al. 2003; Ortiz & Lalana 2010); central West Florida shelf (Ortiz 1979; Krapp-Schickel & Vader 2009); Fish Haven off Louisiana (LeCroy et al. 2009). Habitat. Coral rubble and sandy bottoms in the lagoon and fore-reef environment; at depths of 1 to 73 m (Thomas 1993; Krapp-Schickel & Vader 2009). ►Ceradocus shoemakeri Fox, 1973 Material examined. CYMX-3-PY, 1 male, station 7; CYMX-2-PY, 1 female, station 13. General distribution. Tropical western Atlantic (Fox 1973; Martín & Díaz 2003; Ortiz et al. 2007; LeCroy et al. 2009). GM distribution. Dry Tortugas; Perdido Key; Apalachee Bay (LeCroy 2000); central West Florida shelf (Krapp-Schickel & Vader 2009); Fish Haven off Louisiana (LeCroy et al. 2009); Northwestern Cuba (Ortiz & Lalana 2010). Habitat. Sandy bottoms, coral rubble, algae, sponges, and also occasionally found in grassbeds of Thallassia testudinum (LeCroy 2000); at depths of 0.5 to 73 m (Krapp-Schickel & Vader 2009). Remarks. First record for the southwest region and Mexican southeast sector. Dulichiella appendiculata (Say, 1818) General distribution. Northwestern Atlantic (Lowry & Springthorpe 2007). GM distribution. Central West Florida shelf (Ortiz 1979); Florida Keys; Florida Bay to Louisiana (Sheridan 1980; LeCroy 2000; Lowry & Springthorpe 2007); Yalahau Lagoon (Oliva-Rivera & Jiménez-Cueto 1997); Northwestern Cuba (Ortiz & Lalana 2010). Habitat. Marine epibenthic, polyhaline on sponges, hydroids and bryozoans; at depths of <1 to 80 m (Lowry & Springthorpe 2007; LeCroy et al. 2009). Elasmopus levis (Smith, 1873) General distribution. Northwestern Atlantic (Ortiz et al. 2007; LeCroy et al. 2009). GM distribution. San Antonio Bay; Corpus Christi Bay; Laguna Madre of Texas; Port Isabel (McKinney 1977); Terminos Lagoon (Ledoyer 1986); Perdido Key (Rakocinski et al. 1996); Yalahau Lagoon (Oliva-Rivera & Jiménez-Cueto 1997); Veracruz coral reef system (Winfield et al. 2007a); central West Florida shelf (Vader & Krapp-Schickel 2012). Habitat. Grassbeds, algae and sandy bottoms; at depths of 0 to 18 m (LeCroy 2000; LeCroy et al. 2009; Vader & Krapp-Schickel 2012). BENTHIC AMPHIPODS IN SOUTHEASTERN GULF OF MEXICO Zootaxa 3635 (2) © 2013 Magnolia Press · 145 Elasmopus rapax Costa, 1853 General distribution. Mediterranean Sea; Red Sea; eastern Atlantic; western Atlantic; Indo-Pacific (Hughes & Lowry 2010). GM distribution. Pigeon Key; Florida Bay (Pearse 1912); Key West (Pearse 1912; LeCroy 2000); Plantation Key (Shoemaker 1933a); north Texas offshore; Texas offshore reefs; Port Isabel; Lobos Reef (McKinney 1977); central West Florida shelf (Ortiz 1979); Yalahau lagoon (Oliva-Rivera & Jiménez-Cueto 1997). Habitat. This species is found on a broad range of habitats (e. g. jetties, piers, rocks, dead coral, mangrove roots, algae, sponges, sand bottom, mud bottom); occasionally it is also found on the carapace of the Loggerhead sea turtle; at depths of 1 to 50 m (LeCroy 2000; LeCroy et al. 2009). Jerbarnia americana Watling, 1981 General distribution. Northwestern Atlantic (Watling 1981; LeCroy et al. 2009). GM distribution. Yucatan continental shelf (Ortiz 1979; Watling 1981); off Sanibel Island (Krapp-Schickel & Vader 2009). Habitat. Soft bottoms; at depths of 35 to 179 m (LeCroy et al. 2009). ►Netamelita tabaci Thomas & Barnard, 1991 Material examined. CYMX-1-PY, 1 male, 1 unsexed, station 1; CYMX-58-PY, 2 males, station 5; CYMX-59PY, 1 unsexed, station 8. General distribution. Caribbean (Thomas & Barnard 1991; Ortiz et al. 2007). GM distribution. Previously unreported. Habitat. Deep fore-reef sediments, with coralgal mud and Halimeda sp. flakes (Thomas & Barnard 1991). Remarks. First record for the GM. This record extended its maximum depth from 29 m (Thomas & Barnard 1991) to 31.4 m. Quadrimaera quadrimana (Dana, 1853) General distribution. Tropical western Atlantic; Indo-Pacific (Krapp-Schickel & Ruffo 2000; Ruffo et al. 2000; LeCroy et al. 2009). GM distribution. Florida Keys (Thomas 1993); Yalahau Lagoon (Oliva-Rivera & Jiménez-Cueto 1997); Northwestern Cuba (Ortiz 2001; Ortiz & Lalana 2010). Habitat. Coral rubble and rocks in the lagoon and fore-reef environment; at depths of 1 to 5 m (Thomas 1993; LeCroy et al. 2009). Remarks. As LeCroy (2000) pointed out, records of this species in the western Atlantic region prior to 2000 should be rechecked to ensure that they do not represent one of the closely similar species described by Ruffo et al. (2000) and Krapp-Schickel and Ruffo (2000) from the quadrimana complex. For instance, Q. quadrimana was recorded from southern Gulf of Mexico (Terminos Lagoon) as Maera quadrimana by Ledoyer (1986), but actually it corresponds to Quadrimaera miranda (Ruffo, Krapp & Gable, 2000). Family Melphidippidae Stebbing, 1899 ►Hornellia tequestae Thomas & Barnard, 1986 Material examined. CYMX-28-PY, 3 males, 3 females, 5 unsexed, station 1. General distribution. Southeastern Florida; GM (LeCroy 2007; LeCroy et al. 2009). GM distribution. Looe Key Reef (Thomas & Barnard 1986b); lower Florida Keys to Cape Romano (LeCroy 2007). Habitat. Algal turf on dead coral, coral rubble and sandy shell bottoms in the fore-reef environment; at depths of 2 to 45 m (Thomas & Barnard 1986b; Thomas 1993; LeCroy 2007). Remarks. First record for the Mexican southeast sector. 146 · Zootaxa 3635 (2) © 2013 Magnolia Press PAZ-RÍOS & ARDISSON Family Oedicerotidae Lilljeborg, 1865 ►Americhelidium americanum (Bousfield, 1973) Material examined. CYMX-52-PY, 1 female, station 1; CYMX-54-PY, 8 females, station 2; CYMX-56-PY, 2 females, station 3; CYMX-57-PY, 5 females, station 10; CYMX-55-PY, 3 males, 1 female, station 11; CYMX-53PY, 6 females, station 15. General distribution. Western Atlantic (Bousfield 1973; LeCroy 2000; Ortiz et al. 2007; LeCroy et al. 2009). GM distribution. Texas coast (McKinney 1977); Florida Keys (Thomas 1993); Perdido Key (Rakocinski et al. 1996); Veracruz continental shelf (Winfield et al. 2006). Habitat. Burrowing species in medium to coarse well-sorted sediments from forereef, lagoon, and rubble zone habitats; at depths of 1 to 76 m (Bousfield 1973; Thomas 1993; Winfield et al. 2006). Remarks. First record for the Mexican southeast sector. ►Hartmanodes nyei (Shoemaker, 1933) Material examined. CYMX-29-PY, 1 unsexed, station 10; CYMX-30-PY, 1 female, station 15. General distribution. Western Atlantic; eastern Pacific (LeCroy 2000; Ortiz et al. 2007; García-Madrigal 2008; LeCroy et al. 2009). GM distribution. Key West; off Key Largo (Shoemaker 1933a); Corpus Christi Bay (McKinney 1977); Florida Keys (Thomas 1993); Perdido Key (Rakocinski et al. 1996); Cojímar Bay (Ortiz 2001); Camaronera Lagoon (Cházaro-Olvera et al. 2002); Northwestern Cuba (Ortiz & Lalana 2010). Habitat. Fine to medium sand in back reef, lagoon or open beach habitats; at depths of 1 to 7 m (Thomas 1993, Rakocinski et al. 1996). Remarks. First record for the Mexican southeast sector. This record extended its maximum depth from 7 (Rakocinski et al. 1996) to 42.9 m. Family Phoxocephalidae Sars, 1891 Metharpinia floridana (Shoemaker, 1933) Material examined. CYMX-19-PY, 1 unsexed, station 3; CYMX-18-PY, 2 females, station 7; CYMX-20-PY, 1 unsexed, station 8; CYMX-17-PY, 2 females, 4 unsexed, station 9. General distribution. Western Atlantic (Shoemaker 1933a; Barnard 1980; Ortiz et al. 2007; LeCroy et al. 2009; LeCroy 2011). GM distribution. Florida Keys (Shoemaker 1933a; Barnard 1980; Thomas 1993); Yucatan continental shelf (Ortiz 1979); Perdido Key (Rakocinski et al. 1996); Camaronera Lagoon (Cházaro-Olvera et al. 2002); Bank of Campeche (Escobar-Briones & Winfield 2003); Bay of Campeche (Winfield et al. 2006); Veracruz continental shelf (Winfield & Ortiz 2011). Habitat. In plankton and sandy bottoms (Rakocinski et al. 1996; Thomas 1993; LeCroy 2011); at depths of 0 to 60 m (LeCroy 2011), but it has been also recorded to depth of 3360 m (Winfield et al. 2006). Family Platyischnopidae Barnard & Drummond, 1979 ►Eudevenopus honduranus Thomas & Barnard, 1983 Material examined. CYMX-22-PY, 24 females, 11 unsexed, station 1; CYMX-21-PY, 7 females, station 2; CYMX-24-PY, 1 male, 5 females, station 8; CYMX-23-PY, 2 females, station 9; CYMX-25-PY, 1 female, station 10; CYMX-26-PY, 7 females, 2 unsexed, station 11. General distribution. Western Atlantic; eastern Pacific (Thomas & Barnard 1983; Ortiz et al. 2007; GarcíaMadrigal 2008; LeCroy et al. 2009; LeCroy 2011). GM distribution. Off Panama City (Thomas & Barnard 1983); Florida Keys (Thomas 1993); Perdido Key (Rakocisnki et al. 1996); Louisiana continental shelf (Dubois et al. 2009). Habitat. Sandy and muddy bottoms where they are fast swimmers, rapid burrowers and effective predators; at depths of 1 to 40 m (Thomas & Barnard 1983). BENTHIC AMPHIPODS IN SOUTHEASTERN GULF OF MEXICO Zootaxa 3635 (2) © 2013 Magnolia Press · 147 Remarks. First record for the Mexican southeast sector. This record extended its maximum depth a few meters from 40 (Thomas & Barnard 1983) to 42.2 m. Escobar-Briones et al. (2002) included this species for Mexico, however, no record of that occurrence had been established. On the Yucatan continental shelf this species has been documented as dominant on sandy bottoms, decreasing its abundance with depth (Paz-Ríos 2008). Family Stenothoidae Boeck, 1871 ►Parametopella texensis McKinney, Kalke & Holland, 1978 Material examined. CYMX-12-PY, 2 unsexed, station 3. General distribution. GM (LeCroy et al. 2009; LeCroy 2011). GM distribution. Texas coast shelf (McKinney et al. 1978); Biloxi Bay (Pederson & Peterson 2002); Camaronera Lagoon (Cházaro-Olvera et al. 2002). Habitat. Sandy bottoms mixed with silt/clay; at depth of 1 to 18 (McKinney et al. 1978). Remarks. First record for the southeast region. This record extended its maximum depth from 18 (McKinney et al. 1978) to 43.8 m. Family Synopiidae Dana, 1853 ►Metatiron bellairsi (Just, 1981) Material examined. CYMX-35-PY, 1 male, 1 female, station 5; CYMX-36-PY, 1 female, station 7; CYMX-34PY, 2 females, station 11. General distribution. Central western Atlantic (Just 1981; Barnard & Thomas 1989; Ortiz et al. 2007; LeCroy 2011). GM distribution. Florida Bay; southwest Florida shelf (LeCroy 2011). Habitat. Coral sand with detritus and in or near sparsely grassbeds; at depths of 2.5 to 34 (Just 1981; Barnard & Thomas 1989; LeCroy 2011). Detailed biology and behavior of this species can be found in Just (1981). Remarks. First record for the southeast region. This record extended its maximum depth by one meter from 34 (Barnard & Thomas 1989) to 35.3 m. Escobar-Briones et al. (2002) included this species for Mexico, however, no record of that occurrence had been established. This species is very similar to the other species of the genus reported in the GM, Metatiron triocellatus (Goeke, 1982); in fact Thomas (1993) pointed out that “they are closely related that could be the same species”. LeCroy (2011) examined the morphology for both species and established that they are distinct species. ►Metatiron tropakis (Barnard, 1972) Material examined. CYMX-27-PY, 1 female, station 1. General distribution. Western Atlantic; eastern Pacific (Barnard 1972; LeCroy et al. 2009; LeCroy 2011). GM distribution. Tampa Bay (Barnard 1972); offshore of West Bay; Lavaca Bay (McKinney 1977); Perdido Key (Racocinski et al. 1996); Louisiana continental shelf (Dubois et al. 2009). Habitat. In plankton and sandy bottoms; at depths of 3 to 157 m (Barnard 1972). Remarks. First record for the southeast region. Escobar-Briones et al. (2002) included this species for Mexico, however, no record of that occurrence had been established. This species can be easily distinguished from the other two sympatric species M. bellairsi and M. triocellatus, because it has an accessory flagellum with four or five articles; accessory eye composed of two rather than three ommatidia; and presences of dorsal spines on the telson. ►Synopia ultramarina Dana, 1853 Material examined. CYMX-6-PY, 1 male, station 9. General distribution. Cosmopolitan in warm-temperate, subtropical and tropical regions (Hughes 2009; LeCroy et al. 2009). GM distribution. Off Key Largo (Shoemaker 1933a); Looe Key Reef (Barnard & Thomas 1989); off southwest Florida (LeCroy 2011). 148 · Zootaxa 3635 (2) © 2013 Magnolia Press PAZ-RÍOS & ARDISSON Habitat. Sandy bottoms or coral rubble near coral reefs; it is also a nektonic species which has often been collected in light traps; at depths of 0 to 18 m (Barnard & Thomas 1989; Thomas 1993; Hughes 2009; LeCroy et al. 2009). Remarks. First record for the Mexican southeast sector. This record extended its maximum depth from 18 (Hughes 2009) to 42.1 m. Although this species is widely distributed around the world, few records have been documented in the GM, reflecting a poor sampling effort in the area. Barnard and Thomas (1989) pointed out that species of this genus are rarely reported in modern times because they are found mainly in neritic waters or in epipelagic waters and that if species were collected, it is unlikely that they will be identified by an expert. Suborder Corophiidea Leach, 1814 Infraorder Corophiida Leach, 1814 Superfamily Aoroidea Stebbing, 1899 Family Aoridae Stebbing, 1899 ►Bemlos mackinneyi (Myers, 1978) Material examined. CYMX-65-PY, 1 male, 2 females, station 2. General distribution. GM (LeCroy et al. 2009). GM distribution. Galveston (Myers 1978); central West Florida shelf; Dauphin Island (Myers 1981); Perdido Key (Rakocinski et al. 1996); lower Florida Keys (LeCroy 2002). Habitat. Sand or shell hash bottoms; at depths of 5 to 6 m (LeCroy 2002). Remarks. First record for the Mexican southeast sector. This record extended its maximum depth from 6 m (Myers 1981) to 31.4 m. ►Bemlos spinicarpus inermis (Myers, 1979) Material examined. CYMX-87-PY, 4 females, station 7; CYMX-85-PY, 1 female, station 10; CYMX-88-PY, 2 females, station 13. General distribution. Tropical western Atlantic (Ortiz et al. 2007; LeCroy et al. 2009). GM distribution. Central West Florida shelf (Myers 1981). Habitat. Sand and shell hash bottoms; also in live bottom (e.g. growth of sponges, corals, bryozoans, algae); at depths of 6 to 73 m (Myers 1981; LeCroy 2002). Remarks. First record for the southeast region. The main characters to identify females of this species in the GM after the key provided by LeCroy (2002) are: i) palmar margin convex and angle without tooth on propodus of gnathopod 1, ii) propodus subequal to length of carpus in gnathopod 1, iii) inner ramus longer than peduncle in uropod 3 and iv) distoventral process on peduncle of uropod 3 relatively long. Sub-species was established here on base to the reported distribution, habitat and depth range; nevertheless, to determine the actual sub-species it is necessary to examine male specimens. Grandidierella bonnieroides Stephensen, 1948 General distribution. Cosmopolitan in tropical and temperate regions (Myers 1970, 2009; LeCroy et al. 2009). GM distribution. Port St. Joe; Lake Pontchartrain; Baffin Bay (Myers 1970); Barataria Bay (Thomas 1976); Terminos Lagoon (Ledoyer 1986); Yalahau Lagoon (Oliva-Rivera & Jiménez-Cueto 1997); Alvarado Lagoon (Winfield et al. 2001); Río Soto la Marina Lagoon (Cházaro-Olvera et al. 2002); western coast of Florida; Florida Keys (LeCroy 2002); Northwestern Cuba (Ortiz 2001; Ortiz & Lalana 2010). Habitat. This species is widespread in the GM, occurring frequently in grassbeds or oyster reefs; at depths of 0 to 8 m (Heard 1982; LeCroy 2002). ►Lembos unifasciatus reductus Myers, 1979 Material examined. CYMX-86-PY, 1 male, station 7. General distribution. Tropical western Atlantic (Ortiz et al. 2007; LeCroy et al. 2009). BENTHIC AMPHIPODS IN SOUTHEASTERN GULF OF MEXICO Zootaxa 3635 (2) © 2013 Magnolia Press · 149 GM distribution. Central West Florida shelf (Myers 1981); lower Florida Keys to Cape Romano; Apalachee Bay (LeCroy 2002). Habitat. Sand and shell bottoms; at depths of 2 to 73 m (Myers 1981; LeCroy 2002). Remarks. First record for the Mexican southeast sector. Family Unciolidae Myers & Lowry, 2003 ►Acuminodeutopus naglei (Bousfield, 1973) Material examined. CYMX-84-PY, 1 female, station 4; CYMX-81-PY, 2 females, station 8; CYMX-83-PY, 3 females, station 10; CYMX-82-PY, 11 females, station 11; CYMX-79-PY, 2 females, station 13; CYMX-80-PY, 8 females, station 15. General distribution. Western Atlantic (Bousfield 1973; Ortiz et al. 2007; Winfield & Escobar-Briones 2007; LeCroy et al. 2009). GM distribution. Captiva Island; central West Florida shelf; west of Egmont Key; St. Andrew Bay; Dauphin Island (Myers 1981); Perdido Key (Rakocinski et al. 1996); Florida Keys to Cape Sable (LeCroy 2002). Habitat. Grassbeds and soft sediments with shell debris; from intertidal depths to 55 m (Bousfield 1973; Myers 1981). Remarks. First record for the southwest region and Mexican southeast sector. Superfamily Chevalioidea Myers & Lowry, 2003 Family Chevaliidae Myers & Lowry, 2003 Chevalia aviculae Walker, 1904 General distribution. Circumtropical (Barnard & Thomas 1987b; LeCroy et al. 2009). GM distribution. Central West Florida shelf; Yucatan continental shelf (Ortiz 1979); Florida Keys (Thomas 1993); Northwestern Cuba (Ortiz & Lalana 2010). Habitat. Sandy bottoms; coral rubble and coral overhangs in the fore-reef environment; at depths of 2 to 56 m (Ortiz 1979; Thomas 1993; LeCroy et al. 2009). Remarks. Barnard and Thomas (1987b) showed that a C. aviculae species complex exists. Because of this, LeCroy (2000) proposed that material previously reported as C. aviculae from the western Atlantic needs to be reexamined to determine its status. ►Chevalia carpenteri Barnard & Thomas, 1987 Material examined. CYMX-60-PY, 1 male, station 5. General distribution. Central western Atlantic (LeCroy et al. 2009). GM distribution. Looe Key Reef (Barnard & Thomas 1987b). Habitat. Coral rubble with associated algae and coral overhangs in the fore-reef environment; at depths of 1 to 5 m (Barnard & Thomas 1987b; LeCroy 2000). Remarks. First record for the Mexican southeast sector. This record extended its maximum depth from 5 (Barnard & Thomas 1987b) to 31.4 m. Superfamily Corophioidea Leach, 1814 Family Ampithoidae Boeck, 1871 Ampithoe ramondi Audouin, 1826 General distribution. Cosmopolitan in warm-temperate and tropical regions (LeCroy et al. 2009). GM distribution. Florida Keys (Shoemaker 1933b; Thomas 1993); Yucatan continental shelf (Ortiz 1979); Río Soto la Marina Lagoon (Cházaro-Olvera et al. 2002); Northwestern Cuba (Ortiz 2001; Ortiz & Lalana 2010); Veracruz coral reef system (Winfield et al. 2007a; Winfield et al. 2010). 150 · Zootaxa 3635 (2) © 2013 Magnolia Press PAZ-RÍOS & ARDISSON Habitat. Sandy bottoms, grassbeds, algae, polichaete colonies, ascidians and fouling communities; at depths of <1 to 35 m (Thomas 1993; Ortiz et al. 2007; LeCroy et al. 2009). Cymadusa filosa Savigny, 1816 General distribution. Cosmopolitan in tropical regions (Peart 2004; LeCroy et al. 2009). GM distribution. Yalahau Lagoon (Oliva-Rivera & Jiménez-Cueto 1997); Northwestern Cuba (Ortiz 2001; Ortiz & Lalana 2010). Habitat. Grassbeds and as a tube-building species on algae; at depths of 0 to 5 m (Appadoo & Myers 2003; Ortiz et al. 2007; LeCroy et al. 2009). Remarks. Peart (2004) showed that a C. filosa species complex exists; records of this species by Pears (1912) as Grubia compta and Shoemaker (1935) as Grubia filosa from the Florida Keys are synonymized with Cymadusa setosa (Haswell, 1879). Further material from the southern GM needs to be reexamined to determine its actual distribution in the Gulf basin. Family Corophiidae Leach, 1814 Apocorophium louisianum (Shoemaker, 1934) General distribution. Eastern Florida; GM (LeCroy 2004; LeCroy et al. 2009). GM distribution. Lake St. Catherine; Sarasota Bay (Shoemaker 1947); Barataria Bay (Thomas 1976); Terminos Lagoon (Ledoyer 1986; Cházaro-Olvera et al. 2002); Yalahau Lagoon (Oliva-Rivera & Jiménez-Cueto 1997); Biloxi Bay (Shoemaker 1947; Pederson & Peterson 2002); Alvarado Lagoon (Winfield et al. 2001). Habitat. This species is widespread in the GM, occurring in grassbeds, sandy or muddy bottoms, and as a tubebuilding species on wood, shell, leaves and submerged grasses; at depths of 1 to 15 m (Heard 1982; LeCroy 2004). Infraorder Caprellida Leach, 1814 Superfamily Caprelloidea Leach, 1814 Family Podoceridae Leach, 1814 ►Podocerus kleidus Thomas & Barnard, 1992 Material examined. CYMX-13-PY, 1 male, station 7. General distribution. Central western Atlantic (Ortiz et al. 2007; LeCroy et al. 2009). GM distribution. Cudjoe Key (Thomas & Barnard 1992); Veracruz coral reef system (Winfield et al. 2007a); Northwestern Cuba (Ortiz & Lalana 2010); southwest Florida (LeCroy 2011). Habitat. Clinging to alga Gracilaria sp. (Thomas & Barnard 1992) and in tubes attached to erect substrates, e.g. algae and gorgonians in areas of current flow and wave action (Thomas 1993); at depths of 1 to 13 m (Winfield et al. 2007a; LeCroy 2011). Remarks. First record for the Mexican southeast sector. This record extends its maximum depth from 13 (Winfield et al. 2007a) to 31.4 m. Superfamily Neomegamphopoidea Myers, 1981 Family Neomegamphopidae Myers, 1981 ►Neomegamphopus kalanii Barnard & Thomas, 1987 Material examined. CYMX-64-PY, 1 male, 4 females, station 5. General distribution. Southeastern Florida (Barnard & Thomas 1987a; Ortiz et al. 2007; LeCroy 2011). GM distribution. Previously unreported. Habitat. Sandy bottoms; at depths of 33 to 124 m (Barnard & Thomas 1987a). BENTHIC AMPHIPODS IN SOUTHEASTERN GULF OF MEXICO Zootaxa 3635 (2) © 2013 Magnolia Press · 151 Remarks. First record for the GM. Another species of this genus is found in the GM (Florida), Neomegamphopus hiatus Barnard & Thomas, 1987. Both species are very similar; Barnard and Thomas (1987a) documented that “at first we believed specimens of N. kalanii might be untransformed juvenile of N. hiatus”, but “no stage of transformation between the two species can be demonstrated”. To date those species are separated (based on males) because N. kalanii has a carpus longer than wide, palm beaded, a carpal tooth not gaping, and a remarkable propodus expanded proximally. Superfamily Photoidea Boeck, 1871 Family Ischyroceridae Stebbing, 1899 ►Caribboecetes sp. Material examined. CYMX-61-PY, 1 male, station 1; CYMX-62-PY, 2 females, station 6. General distribution. Genus on the central western Atlantic, and Mexican Pacific (Just 1984; Ortiz & Lemaitre 1997; LeCroy 2007). GM distribution. Genus previously unreported. Habitat. Material collected in medium size and poorly/moderately sorted sand to depth of 19 m. Members of this genus inhabit grass beds, sandy bottoms, and a variety of portable dwellings (e.g. gastropod shells, polychaete tubes, claw tips from decapod molts scaphopod shells); at depths of 0.5 to 54 m (Just 1983, 1984). Remarks. First record for the GM. This species is similar to Caribboecetes barbadensis Just, 1983 from Barbados and Caribboecetes jenikarpae Just, 1984 from Mexican Pacific. Caribboecetes sp. differs from C. barbadensis by bearing dorsolateral setae on peduncular article 5 of antenna 2 in a single row and by coxal plate 7 and article 2 of pereopod 7 with setose lateral surface. It differs from C. jenikarpae by displaying on article 2 of pereopod 2 a strongly triangular anterolateral margin. Caribboecetes sp. presumably represents an undescribed species. Ericthonius brasiliensis (Dana, 1853) Material examined. CYMX-33PY, 1 female, station 5. General distribution. Cosmopolitan in temperate, subtropical and tropical regions (Ortiz et al. 2007; GarcíaMadrigal 2008; LeCroy et al. 2009; Gable et al. 2010). GM distribution. Central and northern Texas shelf; Port Isabel (McKinney 1977); central West Florida shelf (Ortiz 1979); Florida Keys (Thomas 1993); Perdido Key (Rakocinski et al. 1996); Yalahau Lagoon (Oliva-Rivera & Jiménez-Cueto 1997); Cojímar Bay (Ortiz 2001); Biloxi Bay (Pederson & Peterson 2002); Camaronera Lagoon (Cházaro-Olvera et al. 2002); Laguna Madre of Tamaulipas (Barba & Sánchez 2005); Sigsbee abyssal plain (Winfield et al. 2006); Veracruz coral reef system (Winfield et al. 2007a, 2010); Northwestern Cuba (Ortiz & Lalana 2010). Habitat. This is a tube-dwelling fouling species and is often found in diverse soft and hard bottoms (e.g. rocks, mangroves, oyster shells, sponges, sand, seagrasses), generally in shallow waters (1–10 m) (LeCroy 2007). It also has been collected in deep waters (200–3690 m) (Winfield et al. 2006; LeCroy et al. 2009). ►Ericthonius sp. A sensu LeCroy, 2007 Material examined. CYMX-32-PY, 1 male, 1 female, station 7. General distribution. Southeastern and southwestern Florida (LeCroy 2007). GM distribution. Lower Florida Keys to Cape Romano (LeCroy 2007). Habitat. Grassbeds, from sandy shell to silty mud mixed with shell hash; at depths of 1 to 15 m (LeCroy 2007). Remarks. First record for the Mexican southeast sector. This record extended its maximum depth from 15 (LeCroy 2007) to 20.9 m. This species is different from all western Atlantic species of Ericthonius in the morphology of peraeopod 5 of the adult male. The basis is expanded posterodistally to form a wing-like process and an expanded posterodistal margin on the merus; females are more easily distinguished by the presence of long marginal setae on the anterior lobe of coxa 5 and the setae lacking on the margin of the basis of peraeopod 5 (LeCroy 2007). 152 · Zootaxa 3635 (2) © 2013 Magnolia Press PAZ-RÍOS & ARDISSON Family Photidae Boeck, 1871 Gammaropsis atlantica Stebbing, 1888 General distribution. Cosmopolitan in tropical regions (LeCroy 2000; LeCroy et al. 2009). GM distribution. Yucatan continental shelf (Ortiz 1979); Florida Keys (Thomas 1993); Camaronera Lagoon (Cházaro-Olvera et al. 2002); Northwestern Cuba (Ortiz & Lalana 2010). Habitat. Sandy bottoms, coral Acropora sp., alga Halimeda sp. and coral rubble; at depths of <1 to 40 m (Thomas 1993; LeCroy 2000; Ortiz et al. 2007). ►Gammaropsis sp. B sensu LeCroy, 2000 Material examined. CYMX-41-PY, 1 male, 1 unsexed, station 2; CYMX-38-PY, 2 females, 3 unsexed, station 3; CYMX-42-PY, 2 males, 1 female, station 4; CYMX-43-PY, 3 females, 5 unsexed, station 5; CYMX-40-PY, 2 females, station 9; CYMX-44-PY, 1 male, 1 female, station 10; CYMX-39-PY, 1 male, 6 females, 3 unsexed, station 11. General distribution. Southwestern Florida (LeCroy 2000). GM distribution. Lower Florida Keys to Cape Romano (LeCroy 2000). Habitat. Material collected in fine-medium sand; at depths of 31.4 to 43.8 m. Remarks. First record for the Mexican southeast sector. This species is different from all Gammaropsis species in the TWA region by article 2 of mandibular palp relatively slender, much longer than article 3 and whit this last article without subdistal anterior notch; rounded ocular lobe; gnathopod 2 of male densely setose with palmar angle on propodus defined by small cusp and dactyl with setae on extensor margin; pereopod 3–4 with propodus linear and longer than carpus; basis of pereopod 5 narrow, with approximately length twice width; and with ramus twice length of peduncle (peduncle of uropod 3 not elongate). ►Photis macromana Mckinney, Kalke & Holland, 1978 Material examined. CYMX-37-PY, 1 male, 1 female, station 5. General distribution. Central western Atlantic (McKinney et al. 1978; Ortiz et al. 2007; LeCroy et al. 2009). GM distribution. Florida Bay to Texas (McKinney et al. 1978; LeCroy 2000); Veracruz coral reef system (Winfield et al. 2007a). Habitat. Sandy bottoms; at depths of 3 to 42 m (McKinney et al. 1978; LeCroy 2000). Remarks. First record for the Mexican southeast sector. LeCroy (2000) determined that male adult of P. macromana is very similar to male subadult of Photis pugnator Shoemaker, 1945 in gnathopod 2. On base to the key provided by the same author, currently those species can be separate by differences on coxa 3–5, as well as on inner and outer ramus of uropod 3. LeCroy (2000) suggests taxonomical analyses to determine whether or not the two species should be synonymized. ►Photis melanica (McKinney, 1980) Material examined. CYMX-47-PY, 1, male, 1 female, 4 unsexed, station 2; CYMX-48-PY, 1 female, station 10; CYMX-45-PY, 2 females, station 11. General distribution. Central western Atlantic (McKinney 1980; Ortiz et al. 2007; LeCroy et al. 2009). GM distribution. Texas coast (McKinney 1980); Florida Bay; Tampa Bay (LeCroy 2000). Habitat. Fine sand or sandy mud bottoms with some shell hash present; at depths of 1 to 6 m (LeCroy 2000; Martín & Díaz 2003). Remarks. First record for the Mexican southeast sector. This record extended its maximum depth from 6 (LeCroy 2000) to 31.4 m. Photis pugnator Shoemaker, 1945 General distribution. Northwestern Atlantic (Shoemaker 1945; LeCroy 2000). GM distribution. Cedar Key; Tampa Bay; Sarasota Bay; Dry Tortuga (Shoemaker 1945); 7.5 Fathom Reef (McKinney 1977, 1980); central West Florida shelf; Yucatan continental shelf (Ortiz 1979); Veracruz coral reef system (Winfield et al. 2010). Habitat. Hard and sandy bottoms, and in floating Sargassum sp. mats; at depths of 3 to 54 m (McKinney 1980; LeCroy 2000; LeCroy et al. 2009). BENTHIC AMPHIPODS IN SOUTHEASTERN GULF OF MEXICO Zootaxa 3635 (2) © 2013 Magnolia Press · 153 ►Photis sp. C sensu LeCroy, 2000 Material examined. CYMX-94-PY, 1 male, station 2; CYMX-51-PY, 1 male, 1 female, station 3; CYMX-46PY, 2 unsexed, station 7; CYMX-49-PY, 1 male, station 8; CYMX-50-PY, 1 female, station 9. General distribution. Eastern Florida; GM (LeCroy 2000). GM distribution. Florida Keys to Perdido Key (LeCroy 2000). Habitat. Widespread species in fine and well sorted sand and in grassbeds; at depths of 1 to 8 m (LeCroy 2000). Remarks. First record for the Mexican southeast sector. This record extended its maximum depth from 8 (LeCroy 2000) to 43.8 m. Two distinctive characteristics in the coxa 1 separate to this species from the other species in the TWA region: a produced anteroventral angle, forming blunt process and a ventral margin with an anterior gap in row of marginal setae. Zoogeographical affinities Affinity among the eight regions is displayed in four clusters with 40% of similarity. Three of these clusters represented three independent regions (Bahamas, Bermuda and Barbados), whereas the fourth cluster represented five related regions (NW, NE, SW, SE and Caribbean) (Fig 2B). Geographically, independent regions collectively corresponded to fauna from the outer fringe of the Antillean arc, while related regions corresponded to fauna from the GM and Caribbean (Fig 2A). Fauna in the cluster with related regions displayed a gradual affinity southward in geographical distribution, from the GM to the Caribbean. Northern regions (NE and NW) established the highest similarity (57%) with the SW region also related (49%), while the SE region and Caribbean were also highly related (50%). Zoogeographically, the similarity among regions resembles the two major provinces in the TWA: Carolinian province and Caribbean province. Northern regions corresponded to fauna from the Carolinian province, the SW region corresponded to turnover fauna from the Carolinian province to the Caribbean province, and the SE region and Caribbean corresponded to fauna from the Caribbean province. Discussion Species richness. The study of amphipods in the GM is conducted by three countries: Cuba, Mexico and USA. The most updated compilation has listed the greatest number of species for the southeast region (Fig 3); this region harbors 66% of the amphipod diversity in the GM. Of the countries mentioned earlier, Cuba and USA have contributed significantly with descriptions of new species and records. Thomas (1993) listed 52 species from the Florida Keys alone, pointing out that “these species represent approximately one-third of the total number of amphipod species found in the area”. On the other hand, Ortiz and Lalana (2010) recently listed 99 species from the Cuban Gulf. In the Mexican Gulf, amphipods from the southwest sector have been more heavily studied than those in the southeast sector. This is reflected in the great number of species (66) recognized in the former sector. Studies carried out in the southwest sector include those that focus on ecological roles (Corona et al. 2000; Winfield et al. 2007b), checklists of species (Cházaro-Olvera et al. 2002; Winfield et al. 2006, 2007a; Winfield & Ortiz 2011) and description of new species (Winfield & Álvarez 2009; Winfield & Ortiz 2009, 2010, 2012). The few studies in the southeast sector focus on the ecological distribution (Paz-Ríos 2008; Rodríguez-Pliego et al. 2011) and checklists of species (Ortiz 1979; Oliva-Rivera & Jiménez-Cueto 1997; Simões et al. 2010). Nevertheless, due to the great heterogeneity of habitat in the southeast sector (e.g. beaches, coastal lagoons, grassbeds, reefs), Escobar-Briones and Jiménez-Guadarrama (2010) have suggested that the Campeche Bank on the southeast sector is like a “hotspot” of biological diversity for peracarids associated with carbonated bottoms. Therefore, the reported number of amphipod species here stated will increase with the exploration of new habitats (e.g. Alacranes Reef) and description of new species (e.g. Caribboecetes sp.). Four species previously reported in the region were excluded from the checklist of amphipods for the Mexican southeast sector. Three of these species [Linguimaera hamigera (Haswell, 1879), Eudevenopus cf. metagracilis 154 · Zootaxa 3635 (2) © 2013 Magnolia Press PAZ-RÍOS & ARDISSON (Barnard, 1964) and Pseudohaustorius carolinensis Bousfield, 1965] are listed in Ortiz (1979) and one [Bemlos waipio (Barnard, 1970) as Lembos processifer Pirlot, 1938], is listed in Oliva-Rivera and Jiménez-Cueto (1997). These species were excluded based on the atypical habitats and distribution ranges reported. In addition, these species have not been listed for the GM in the most updated checklists of amphipods in the region by LeCroy (2002, 2011), Ortiz et al. (2007), LeCroy et al. (2009) and Ortiz and Lalana (2010). FIGURE 2. Zoogeographical affinity of amphipod fauna in the tropical western Atlantic. (A) Regions in the tropical western Atlantic after Ortiz et al. (2007) and LeCroy et al. (2009); gray lines represent the borders of regions in the GM and gray arrows represent the borders of the Carolinian and Caribbean provinces. (B) Cluster diagram of amphipod assemblage among regions of the tropical western Atlantic. BENTHIC AMPHIPODS IN SOUTHEASTERN GULF OF MEXICO Zootaxa 3635 (2) © 2013 Magnolia Press · 155 FIGURE 3. Number of amphipod species (Gammaridea and Corophiidea) in regions of the Gulf of Mexico. NE, Northeast; SE, Southeast; SW, Southwest; NW, Northwest. Data based on LeCroy et al. (2009); family Caprellidae and Cyamidae excluded. Another species previously recorded from the GM (see LeCroy et al. 2009), but excluded from this checklist was Melita nitida Smith, 1873. This species was listed by Oliva-Rivera and Jiménez-Cueto (1997) from the Yalahau Lagoon, but based on specimens recently examined from that lagoon; M. nitida actually represents Melita planaterga Kunkel, 1910. Zoogeographic comments. The two zoogeographical provinces mostly recognized and widely accepted for the benthic macrofauna of the TWA are the Carolinian and Caribbean provinces (Hedgpeth 1953; Briggs 1974; Barnwell & Thurman 1984; Kensley & Schotte 1989; Rabalais et al. 1999; Engle & Summers 2000; SalazarVallejo 2000; Raz-Guzman et al. 2004; Neigel 2009; Briggs & Bowen 2012). The Carolinian province is located in the northern GM and has been considered part of the warm-temperate province, which includes the northwestern Atlantic coast from Cape Hatteras (North Carolina) to Cape Canaveral (Florida). The Caribbean province is located in the southern GM and has been considered part of the tropical province, which extends to the Orinoco Delta River. The approximate boundaries between Carolinian and Caribbean provinces in the GM are Cabo Rojo (Veracruz) on the southwestern end and Cape Romano (Florida) on the northeastern end. The hierarchical cluster analysis displayed an overlap between the amphipod fauna and the zoogeographical provinces described above. In this case the use of multivariate statistical methods (e.g. similarity index and average linkage clustering) for establishing zoogeographic provinces was not diminished, supporting its efficacy. The use of these statistical methods has been widely accepted among researchers who have explored the zoogeographical boundaries in different benthic taxa in the western Atlantic; including mysids (Escobar-Briones & Soto 1991), gastropods (Diaz 1995), macroinfauna (Engle & Summers 2000), decapods (Wickstec & Packard 2005), cumaceans (Watling 2009), amphipods, echinoderms, mollusks, polychaetes, sponges and stony corals (Miloslavich et al. 2010). Winfield et al. (2007c) listed the biotic components dominant for amphipods in the southwestern GM as Endemic (29%), Caribbean (25%) and Carolinian (25%). In this respect, although the native species are the major component, the shared affinity between Caribbean and Carolinian provinces is also denoting a faunal turnover, as is expected in this region. Thus, the distributional patterns found for southern GM amphipods by Winfield et al. (2007c) and the present study suggest a gradual zoogeographical change from the Carolinian-Caribbean province (southwestern region) to the Caribbean province (southeastern region). 156 · Zootaxa 3635 (2) © 2013 Magnolia Press PAZ-RÍOS & ARDISSON Based on the analyzed data matrix in the present study, extreme levels of endemism were found in the fauna from regions of the Antillean arc with the highest levels in Barbados (100%) and Bermuda (69%), characteristic to distant islands, and with the lowest level in Bahamas (16%), characteristic to a connected archipelago. Intermediate levels of endemism but denoting a strong biotic isolation were presented in the fauna from the Caribbean and GM with 47% and 25%, respectively. Particularly, the level of endemism in the GM was estimated at 27% by LeCroy et al. (2009), pointing out that this level will diminish with a better knowledge of species distribution. In the present study the endemicity in the GM diminished slightly due to the calculation, as it had been predicted above; it was estimated with 59 endemic species in the present study, instead of 62 endemic species in LeCroy et al. (2009). With respect to the endemism in the GM regions, there is a geographic latitudinal and longitudinal difference, likely related to a thermal biogeographic barrier and a sedimentary biogeographic barrier, respectively. Latitudinally, northern regions with a warm-temperate regime (NW= 5%, NE= 21%) presented higher levels of endemism than southern regions with a tropical regime (SW= 1.5%, SE= 15%). Longitudinally, eastern regions with carbonated sediments (NE= 21%, SE= 15%) presented higher levels of endemism than western regions with terrigenous sediments (NW= 5%, SW= 1.5%). Furthermore, the southwest region showed the lowest level of endemism, suitable to a region where presumably come together both Carolinian and Caribbean provinces and there is a high faunal turnover (dissimilarity) from the northwest region to the southwest region (57%). To explain both distributions in the GM provinces we should focus on historical and ecological biogeography. The former is concerned with historical meteorological/geological events which actually formed the Gulf basin (e.g. continental drift, cycles of glaciations, emergence of the Central American Isthmus, emergence of the Florida peninsula, redirection of the North Equatorial Current northward), which have been treated by Mills (1965), Briggs (1970), McKinney (1977) and Neigel (2009). The latter investigates and describes current environmental traits in the Gulf basin (e.g. currents, depth, sediment, temperature, seabed topography, biotic interactions), which have been treated by Hedgpeth (1953), Rabalais et al. (1999), Wicksten and Packard (2005), Winfield et al. (2006) and Richards et al. (2007). In summary, amphipod distribution in the GM may have originated via historical events in the tropical Atlantic (Myers & Lowry 2009). Thus, although the influence of the east Pacific (via Central American Isthmus) on the Caribbean (and after on the GM) was high, the influence from the Atlantic was stronger (McKinney 1977; Myers & Lowry 2009). This is currently supported by the amphipod zoogeography of Winfield et al. (2007c), where the biotic components with less dominance in the southwestern GM are Euritopic (16%) and Oregonian (5%). 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(2006) Update checklist and identification of areas of endemism of benthic amphipods (Caprellidae and Gammaridea) from offshore habitats in the SW Gulf of Mexico. Scientia Marina, 70, 99–108. Winfield, I., Cházaro-Olvera, S., Horta-Puga, G., Lozano-Aburto, M.A. & Arenas-Fuentes, V. (2010) Macrocrustáceos incrustantes en el Parque Nacional Sistema Arrecifal Veracruzano: biodiversidad, abundancia y distribución. Revista Mexicana de Biodiversidad, 80, S165–S175. 164 · Zootaxa 3635 (2) © 2013 Magnolia Press PAZ-RÍOS & ARDISSON APPENDIX 1. List of amphipod species in alphabetical order by geographic region used in the zoogeographic analysis. NE, Northeast; SE, Southeast; SW, Southwest; NW, Northwest. Data based on Ortiz et al. (2007) and LeCroy et al. (2009). Species Bermuda Bahamas Caribbean Barbados NE SE Acanthohaustorius intermedius 1 Acanthohaustorius pansus 1 Acanthohaustorius shoemakeri 1 Acanthohaustorius uncinus 1 1 SW NW 1 1 1 Actogidiella cultrifera 1 Acuminodeutopus heardi 1 Acuminodeutopus naglei 1 1 1 Americhelidium americanum 1 1 1 Americorchestia barbarae 1 Americorchestia heardi 1 Americorchestia salomani 1 Americorophium ellisi 1 Americorophium rioplatense 1 Ameroculodes edwardsi 1 Ameroculodes miltoni 1 Ampelisca abdita 1 Ampelisca abyssicola 1 Ampelisca agassizi 1 1 Ampelisca bicarinata 1 1 Ampelisca brevisimulata 1 Ampelisca burkeri 1 Ampelisca cristata cristata 1 Ampelisca cristata microdentata 1 Ampelisca cristoides 1 Ampelisca holmesi Ampelisca lobata 1 Ampelisca parapacifica 1 Ampelisca parapanamensis 1 Ampelisca paria 1 Ampelisca pugetica 1 Ampelisca romigi 1 Ampelisca schellenbergi 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Ampelisca vadorum 1 1 1 1 Ampelisca venetiensis 1 Ampelisca verrilli 1 Amphideutopus dolichocephalus 1 Amphilochus ascidicola 1 Ampithoe hirsutimanus 1 Ampithoe longimana 1 1 1 Ampithoe marcuzzii 1 1 Ampithoe ramondi 1 1 1 Ampithoe pollex 1 1 1 1 1 1 1 1 ......continued on the next page BENTHIC AMPHIPODS IN SOUTHEASTERN GULF OF MEXICO Zootaxa 3635 (2) © 2013 Magnolia Press · 165 APPENDIX 1. (Continued) Species Bermuda Bahamas Ampithoe valida Caribbean 1 Barbados NE SE 1 Anamaera hixoni NW 1 1 Anamixis cavatura 1 1 Anamixis vanga 1 1 Anelasmopus kraui 1 Apocorophium acutum 1 1 Apocorophium lacustre 1 1 1 Apocorophium louisianum 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Apocorophium simile 1 Apohyale media 1 1 Apohyale perieri 1 1 Apohyale wakabarae 1 Apolochus casahoya 1 Apolochus delacaya Apolochus neapolitanus 1 1 Apolochus pillaii 1 1 Argissa hamatipes 1 Aristias captiva 1 Aroui americana 1 Aruga holmesi 1 Audulla chelifera SW 1 1 1 1 Batea campi 1 1 Batea carinata 1 1 Batea catharinensis 1 1 1 Batea cuspidata 1 1 1 Batea schotti 1 Batea transversa 1 Bemlos barnardi 1 Bemlos foresti 1 1 Bemlos kunkelae 1 1 Bemlos longicornis 1 1 Bemlos mackinneyi 1 1 1 1 1 Bemlos minimus 1 Bemlos sanmartini 1 1 1 Bemlos setosus Bemlos spinicapus inermis Bemlos spinicarpus spincarpus 1 1 1 Bemlos scolosternum 1 1 Bemlos habanensis Bemlos mayensis 1 1 Batea bousfieldi Bemlos dentischium 1 1 1 1 1 1 ......continued on the next page 166 · Zootaxa 3635 (2) © 2013 Magnolia Press PAZ-RÍOS & ARDISSON APPENDIX 1. (Continued) Species Bermuda Bahamas Caribbean Barbados NE Bemlos tempus 1 Bemlos tigrinus 1 Bemlos unicornis 1 Biancolina brassicacephala Biancolina lowryi 1 SE SW 1 1 1 1 1 Boca campi 1 Boca elvae 1 Boca megachela 1 1 Bogidiella perla 1 Bogidiella virginalis 1 Bruzelia pericu 1 Bruzeliopsis turba 1 Byblis cubensis 1 Byblisoides blasensis 1 Caribboecetes barbadensis 1 Caribboecetes crassicornis 1 Caribboecetes intermedius 1 Caribboecetes justi 1 Caribboecetes pterycornis 1 Caribseba tropica 1 Cebocaris grutesca 1 Ceradocus rubromaculatus 1 Ceradocus sheardi 1 1 1 1 1 1 Ceradocus shoemakeri NW 1 Cerapus benthophilus 1 1 Cerapus cudjoe 1 Cerapus thomasi 1 Cerapus tubularis 1 Chelorchestia costaricana 1 Chelorchestia forceps 1 1 1 1 1 1 1 1 1 1 1 Chevalia carpenteri 1 1 Chevalia mexicana 1 1 1 Colomastix bousfieldi 1 1 1 Colomastix camura 1 1 Colomastix cornuticauda 1 Colomastix denticornis 1 1 1 1 1 Colomastix falcirama 1 1 1 Colomastix gibbosa 1 1 1 Colomastix halichondriae 1 1 1 Colomastix heardi 1 1 Colomastix irciniae 1 1 1 1 1 1 1 Chevalia aviculae Colomastix janiceae 1 1 1 ......continued on the next page BENTHIC AMPHIPODS IN SOUTHEASTERN GULF OF MEXICO Zootaxa 3635 (2) © 2013 Magnolia Press · 167 APPENDIX 1. (Continued) Species Bermuda Bahamas Caribbean Barbados NE SE Colomastix pusilla 1 Colomastix tridenta 1 1 1 Concarnes concavus 1 1 1 Confodiopisa scopulorum 1 Corocubanus guitarti SW NW 1 1 1 Curidia debrogania 1 Cymadusa compta 1 1 1 1 Cymadusa filosa 1 1 1 1 Cyphocaris johnsoni 1 1 1 1 1 Cyphocaris tunicola 1 1 1 Dissiminassa homosassa 1 1 Deflexiodes intermedius 1 Dulichiella appendiculata 1 1 1 Dumosus atari 1 1 1 Eclecticus eclecticus 1 Elasmopus balkomanus 1 1 1 Elasmopus bampo 1 Elasmopus brasiliensis 1 Elasmopus lemaitre 1 Elasmopus levis 1 1 1 Elasmopus pectenicrus 1 1 1 Elasmopus pocillimanus 1 1 1 1 1 Elasmopus rapax 1 1 1 1 1 Elasmopus spinidactylus 1 Elasmopus thomasi 1 Ensayara entrichoma 1 1 Ensayara jumane 1 Eobrolgus spinosus 1 Ericthonius brasiliensis 1 Ericthonius rubricornis 1 Eriopisa incisa 1 Eudevenopus honduranus 1 Eudevenopus metagracilis 1 Eurythenes obesus 1 1 1 1 1 1 1 1 1 Eusirogenes adad 1 Eusiroides monoculoides 1 Eusiroides yucatanensis 1 Floresorchestia guadalupensis 1 Gammarella curvata 1 Gammaropsis arawakia 1 Gammaropsis atlantica 1 Gammaropsis sutherlandi Gammaropsis togoensis 1 1 1 1 1 1 1 ......continued on the next page 168 · Zootaxa 3635 (2) © 2013 Magnolia Press PAZ-RÍOS & ARDISSON APPENDIX 1. (Continued) Species Bermuda Bahamas Caribbean NE SE SW NW Gammarus mucronatus 1 1 1 1 Gammarus tigrinus 1 Garosyrrhoe bigarra 1 Garosyrrhoe luquei 1 Gibberosus myersi 1 Gitana dominica 1 Gitanopsis petulans Barbados 1 1 1 Gitanopsis templadoi 1 Globosolembos francanni 1 Globosolembos smithi 1 1 1 1 1 Grandidierella bonnieroides 1 1 1 1 1 Haploops setosa 1 Harpinia excavata 1 1 1 1 1 Harpinia intermedia 1 Hartmanodes nyei 1 Haustorius jayneae 1 Haustorius mexicanus 1 Heterophoxus oculatus 1 Hippomedon pensacola 1 Hoplopheonoides obesa 1 Hoplopheonoides shoemakeri 1 Hornellia atlanticus 1 Hornellia tequestae 1 Hourstonius laguna 1 1 1 Hourstonius tortugae 1 1 1 Hyale diplodactyla 1 Hyale pygmaea 1 Jassa marmorata 1 1 Jerbania americana 1 Idunella sketi 1 1 1 Iphimedia zora 1 Latacunga comanita 1 1 Laticorophium baconi 1 Lembos unifasciatus reductus 1 1 1 1 Lembos websteri 1 1 1 1 Lepechinella aberrantis 1 Lepechinella raua 1 Lepidactylus triarticulatus 1 Lepiduristes lepidus 1 Leptocheirus rhizophorae 1 Leucothoe ashleyae 1 1 Ileraustroe torpens Lembos unifasciatus unifasciatus 1 1 1 1 1 1 ......continued on the next page BENTHIC AMPHIPODS IN SOUTHEASTERN GULF OF MEXICO Zootaxa 3635 (2) © 2013 Magnolia Press · 169 APPENDIX 1. (Continued) Species Bermuda Bahamas Caribbean Leucothoe ayrtonia Barbados NE SE SW NW 1 1 1 Leucothoe kensleyi 1 1 Leucothoe laurensi 1 Leucothoe panpulco 1 Leucothoe spinicarpa 1 Liljeborgia bousfieldi 1 Liocuna caeca 1 1 1 1 Listriella andresi 1 Listriella bahia 1 Listriella barnardi 1 1 Listriella carinata 1 Listriella Kensleyi 1 Listriella quintana 1 Lysianopsis alba 1 Lysianopsis hummelincki 1 Lysianopsis ozona 1 Maera diffidentia 1 1 1 1 1 1 1 1 1 Maera lucinae Maera tinkerensis 1 1 1 Maera williamsi 1 Maeracoota galani 1 Maeropsis rathbunae 1 Mallacoota carausui 1 Marigidiella crassipes 1 Melita dentata 1 Melita elongata 1 1 Melita guanaense 1 1 1 Melita intermedia 1 1 Melita leiotelson 1 Melita longisetosa 1 1 1 1 Melita nitida 1 1 1 1 1 Melita persona 1 1 1 1 1 1 Melita planaterga Melita stocki 1 1 Metaniphargus sabulonis 1 Metatiron bellairsi 1 Metatiron triocellatus Metatiron tropakis 1 1 Metharpinia floridana 1 1 Microjassa bahamensis 1 1 1 Microjassa floridensis 1 Microjassa tetradonta 1 Microphoxus minimus 1 1 ......continued on the next page 170 · Zootaxa 3635 (2) © 2013 Magnolia Press PAZ-RÍOS & ARDISSON APPENDIX 1. (Continued) Species Bermuda Bahamas Caribbean Barbados NE SE 1 Microprotopus raneyi 1 1 Microprotopus shoemakeri 1 1 Miramarassa sanchezi SW 1 1 1 Monocorophium acherusicum 1 Monocorophium insidiosum 1 1 1 Nasagenia bacescui 1 Nasagenia comisariensis 1 Nasagenia yucatanensis 1 Neoischyrocerus vidali 1 Neomegamphopus heardi 1 Neomegamphopus hiatus 1 Neomegamphopus roosevelti 1 Nepanamixis dianthus 1 Netamelita barnardi 1 1 Nototropis minikoi 1 1 1 1 1 1 1 Nototropis urocarinatus 1 1 1 1 1 Oediceroides synparis 1 1 Netamelita brocha Netamelita tabaci 1 1 Monocorophium tuberculatum 1 1 1 1 Orchestia grillus 1 Orchomene stocki 1 1 Orchomenella perdido 1 Orchomenella thomasi 1 Orenoquia serrata 1 Paracentromedon carabicus 1 Parahaustorius longimerus 1 Parahaustorius obliquus 1 Parametopella cypris 1 Parametopella texensis 1 Paramicrodeutopus myersi 1 Parargissa galatheae 1 Parhyale fascigera 1 1 Parhyale hawaiensis 1 1 Parhyale inyacka 1 Parhyalella batesoni NW 1 1 1 1 1 1 1 1 Parhyalella whelpleyi 1 1 Pariphinotus seclusus 1 1 Pariphinotus seticoxa Parpano cebus 1 Parpano composturus 1 Pedicorophium laminosum 1 Perioculodes cerasinus 1 1 1 1 1 1 1 ......continued on the next page BENTHIC AMPHIPODS IN SOUTHEASTERN GULF OF MEXICO Zootaxa 3635 (2) © 2013 Magnolia Press · 171 APPENDIX 1. (Continued) Species Bermuda Bahamas Photis dentata Caribbean Barbados NE SE SW 1 1 1 NW 1 Photis longicaudata Photis macromanus 1 1 1 1 Photis melanica 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 Photis pugnator Photis reinhardi 1 Photis trapherus 1 Platorchestia platensis 1 Podobothrus bermudensis 1 1 1 1 1 1 1 Plesiolembos ovalipes Plesiolembos rectangulatus 1 1 Podocerus brasiliensis Podocerus chelonophilus 1 Podocerus cristatus 1 Podocerus kleidus 1 Podocerus lazowasemi 1 Podocerus tachyrheo 1 Pontogeneia bartschi 1 1 Proharpinia barbada 1 Protohadzia schoenerae 1 1 Protohaustorius bousfieldi 1 1 Protohyale macrodactyla 1 Psammogammarus bluefieldensis 1 Psammomelita uncinata 1 1 1 Pseudamphithoides bacescui 1 Pseudoamphitoides incurvaria 1 1 1 Pseudohaustorius americanus 1 Pseudomegamphopus excavatus 1 1 Quadrimaera ariel 1 Quadrimaera caliban 1 Quadrimaera ceres 1 Quadrimaera cristianae 1 1 Quadrimaera inaequipes 1 Quadrimaera miranda 1 1 Quadrimaera pacifica 1 1 Quadrimaera pieteri 1 Quadrimaera quadrimana 1 1 Quadrimaera sarae 1 1 Rachotropis lobata 1 Rachotropis portoricana 1 Resupinus spinicaudatus 1 Rimakoroga floridana Seba robusta 1 1 1 1 ......continued on the next page 172 · Zootaxa 3635 (2) © 2013 Magnolia Press PAZ-RÍOS & ARDISSON APPENDIX 1. (Continued) Species Bermuda Bahamas Caribbean Sennaia bidactyla NE SE SW 1 1 1 1 1 Socarnopsis catacumba 1 Spathiopus cojimarensis 1 Spathiopus looensis 1 Stegocephaloides calypsonis 1 Stenothoe gallensis 1 Stenothoe georgiana 1 1 1 1 Stenothoe symbiotica 1 1 1 1 1 Stephonyx biscayensis 1 Stephonyx carinatus 1 1 Stephonyx incertus 1 Sunamphitoe pelagica 1 Synopia scheeleana 1 Synopia ultramarina NW 1 Shoemakerella cubensis Shoemakerella lowryi Barbados 1 1 1 Syrrhoe papyracea 1 Tabatzius muelleri 1 Talorchestia fritzi 1 Talorchestia marcuzzi 1 Talorchestia margaritae 1 1 1 Tethygeneia longleyi 1 Thalassostygius exiguus 1 Thetorchestia antillensis 1 Thetorchestia karukerae 1 Thetorchestia sulensoni 1 Tiburonella morrocoyensis 1 Tiburonella viscana 1 Tropichelura gomezi 1 Tropichelura insulae 1 Tryphosella apalachicola 1 Tryphosella uristesi 1 1 1 1 Uhlorchestia uhleri 1 Unciola irrorata 1 Unciola serrata 1 Vemana compressa 1 Vemana lizata 1 Victoriopisa guanarocana 1 Westwoodilla longimana 1 BENTHIC AMPHIPODS IN SOUTHEASTERN GULF OF MEXICO 1 Zootaxa 3635 (2) © 2013 Magnolia Press · 173

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