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Updated Checklist and Identification of Areas of Endemism of Benthic Amphipods (Caprellidea and Gammaridea) from Offshore Habitats in the SW Gulf of Mexico*

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Updated Checklist and Identification of Areas of Endemism of Benthic Amphipods (Caprellidea and Gammaridea) from Offshore Habitats in the SW Gulf of Mexico* sm70n1099-2034 1/3/06 16:29 Página 99 SCI. MAR., 70 (1): 99-108 SCIENTIA MARINA 2006 Updated checklist and identification of areas of endemism of benthic amphipods (Caprellidea and Gammaridea) from offshore habitats in the SW Gulf of Mexico* IGNACIO WINFIELD 1, ELVA ESCOBAR-BRIONES 2 and JUAN J. MORRONE 3 1 Laboratorio de Ecología de Crustáceos, Facultad de Estudios Superiores Iztacala. Posgrado en Ciencias del Mar y Limnología-UNAM, A. P. 314, Tlalnepantla, C.P.54090, Estado de México, México. E-mail [email protected] 2 Laboratorio de Biodiversidad y Macroecología, Instituto de Ciencias del Mar y Limnología-UNAM, A. P. 70-305, 04510, México, D.F., México. 3 Museo de Zoología “Alfonso L. Herrera”, Departamento de Biología Evolutiva, Facultad de Ciencias-UNAM, A. P. 70-379, 04510, México, D.F., México. SUMMARY: An updated checklist of the benthic amphipods associated with soft bottoms in the SW Gulf of Mexico has been made and the areas of endemism have been identified. Amphipods were collected in a depth interval of 25 to 3700 m during six cruises from 1994 to 1999. A total number of 56 species were identified, 11 belonging to the suborder Caprellidea and 45 to the suborder Gammaridea. Thirteen species extended their geographical distribution within the Gulf of Mexico and four species were considered new records in the Gulf of Mexico. A biogeographic analysis based on the shared species and the parsimony criterion helped identify three areas of endemism in the SW Gulf of Mexico. They were named: area A, a small area on the continental shelf of the Bay of Campeche; area B, the continental slope of the Bay of Campeche; and area C, the lower continental slope of the Mexican ridges extending into the western Sigsbee abyssal plain. These areas are defined in terms of their complex topography that may help isolate habitats that promote endemicity. Keywords: Amphipoda, biogeography, continental shelf, continental slope, abyssal plain. RESUMEN: LISTADO ACTUALIZADO E IDENTIFICACIÓN DE ÁREAS DE ENDEMISMO DE LOS ANFÍPODOS BENTÓNICOS (CAPRELLIDEA Y GAMMARIDEA) DE HÁBITATS MARINOS EN EL SO DEL GOLFO DE MÉXICO. – Se presenta una lista actualizada de los anfípo- dos bentónicos asociados a fondos suaves en el SO del Golfo de México y las áreas de endemismo reconocidas. Los anfí- podos fueron colectados en un intervalo de profundidad de los 25 a 3700 m en seis cruceros oceanográficos de 1994 a 1999. Un total de 56 especies fueron identificadas, 11 pertenecientes al suborden Caprellidea y 45 al suborden Gammaridea. Trece especies ampliaron su distribución geográfica en el Golfo de México y cuatro especies fueron registros nuevos para el Golfo de México. Un análisis biogeográfico basado sobre las especies compartidas de acuerdo con el criterio de parsimonia ayudó a identificar tres áreas de endemismo en el sector SO del Golfo de México, nombradas como área A, un área pequeña sobre la plataforma continental de la Bahía de Campeche; área B, el talud continental de la Bahía de Campeche; y área C, del talud continental inferior de las Cordilleras Mexicanas hasta el oeste de la planicie abisal de Sigsbee. Estas áreas fueron definidas en términos de su topografía compleja que permite aislar hábitats promoviendo el endemismo. Palabras clave: Amphipoda, biogeografía, plataforma continental, talud continental, planicie abisal. *Received December 13, 2004. Accepted July 22, 2005. BENTHIC AMPHIPODS AND AREAS OF ENDEMISM 99 sm70n1099-2034 1/3/06 16:29 Página 100 INTRODUCTION The first reports of amphipods from offshore habitats were documented during the last decades of the 19th century by Sars, Chevreux, and Stebbing (Thurston, 2000). A large number of amphipods have been recorded subsequently in diverse benth- ic habitats (Barnard, 1961, 1962, 1964). Endemicity to the generic level was analyzed by Barnard and Karaman (1991) and Bellan-Santini (1990) within different biogeographic regions. At a worldwide scale, the surveys by Marquez and Bellan-Santini (1993), Cartes and Sorbe (1999), Bachelet et al. (2003), and Guerra-García (2003) have contributed with new species to the checklist, showing the restricted geographic distribution of species of this group. Our knowledge of the amphipod fauna has sig- nificantly increased in the last years; however, it remains limited for offshore habitats in most of the ocean regions. The benthic amphipod fauna of the Gulf of Mexico has been extensively documented in the coastal ecosystems including the continental shelf (Myers, 1981; Ortiz, 1991; Thomas, 1993; LeCroy, 1995, 2000, 2001). Our objective was to update the checklist of ben- thic amphipod species (Caprellidea and FIG. 1. – Markers show the location of sampling stations of the six cruises where amphipods were sorted for this study. Abbreviations Gammaridea) occurring between 25 and 3700 m used: SBI: Sigsbee I, SBII: Sigsbee II, ENI–II: ENOS I and II, depth, and to recognize areas of endemism of OGXII-XVI: OGMEX XII and XVI. amphipod fauna in the offshore habitats of the south-western region of the Gulf of Mexico. western sector of the Gulf, a canyon and knolls in the Bay of Campeche, and an escarpment along the northern edge of the Campeche Bank. The MATERIAL AND METHODS Sigsbee abyssal plain occupies most of the central portion of the basin and is characterized by thin The Gulf of Mexico is characterized by a com- turbidite layers (Bryant et al., 1991). The surface plex geomorphology (Roberts et al., 1999) with var- circulation is dominated by the Loop Current and ious canyons, escarpments, salt diapirs and massive interaction with eddies (Welsh and Inoue, 2000), carbonated deposits (Bryant et al., 1991). The semi- in general the water column is thermally stratified enclosed basin is linked with the Caribbean Sea by and displays space and time variability along the the Yucatan Channel and with the Western Tropical continental shelf (Monreal-Gómez and Salas de Atlantic Ocean through the Straits of Florida and the León, 1997). different straits and sills through the Caribbean Sea. The amphipod species were sorted out by siev- The seafloor is an accumulation of terrigenous clas- ing, with a 250 µm sieve, superficial sediment tic sediments, the sedimentary regime is the result of samples that were collected with GOMEX (0.06 the influence of the Mississippi, Panuco, m2) and USNEL (0.25 m2) boxcorers on six cruis- Papaloapan and Grijalva-Usumacinta rivers (McKee es: SIGSBEE.1 (SBI), SIGSBEE.2 (SBII), and Baskaran, 1999). ENOS.1 (ENI), ENOS.2 (ENII), OGMEX.12 The continental shelf represents approximately (OGXII), and OGMEX.16 (OGXVI). These cruis- 30% of the total area (Rabalais et al., 1999) with a es were in the south-western (SW) region of the variable extension (Bouma, 1972). The continental GM in the polygon delimited by 18º38.0’ and slope is characterized by the Mexican ridges in the 25º27.0’N - 92º19.0’ and 97º12.0’W on board the 100 I. WINFIELD et al. sm70n1099-2034 5/3/06 21:09 Página 101 R/V Justo Sierra-UNAM from 1994 to 1999. The RESULTS bathymetric range (25 to 3700 m) encompassed soft bottom habitats of the continental shelf, con- Order AMPHIPODA Latreille, 1816 tinental slope and abyssal plain. Sampling sites Suborder CAPRELLIDEA Leach, 1814 were graphically posted with the SURFER V.7 Superfamily CAPRELLOIDEA Laubitz, 1993 software. Caprellidea and Gammaridea Family CAPRELLIDAE White, 1847 amphipods were preserved in ethanol solution, 1. Caprella sp. and identified according to McCain (1968), Myers (1981), Lowry (1984), Barnard and This species is a common component, it occurs Karaman (1991), Ortiz (1994), Bousfield and in soft bottoms of the Mexican ridges (SBI cruise) Hoover (1997), Lowry and Stoddart (1997), and the Bay of Campeche (OGXVI cruise) within a LeCroy (2000, 2001), and Ortiz et al. (2002). The depth range of 550 to 2120 m. classification and the superfamily arrangement followed Bousfield and Shih (1994) and 2. Caprella danilevskii Czerniavski, 1868 Bousfield (2001). Incomplete or damaged amphipods were identified to the lowest taxonom- This species is a dominant component, it occurs ic level possible. Each species herein reported in muddy sediments of the Mexican ridges (SBI includes the geographical occurrence, the infor- cruise) within a depth range of 550 to 2120 m. mation for depth and sediment type and the cruis- es on which they were collected. The specimens 3. Caprella equilibra Say, 1818 were characterized by their frequency and abun- dance of occurrence in the samples and herein This species is a dominant component, it occurs named as dominant, common and rare faunal in muddy sediments of the Mexican ridges and components. The specimens are deposited at the Sigsbee abyssal plain (SBI cruise) within a depth Colección Nacional de Crustáceos (CNCR- range of 1231 to 3700 m. UNAM) and counted with a reference number to the database and collection. 4. Caprella penantis Leach, 1814 Endemicity in this study was defined as local areas with high survival of species (allochthonous This species is a dominant component, it endemics) (Pielou, 1992). The Parsimony occurs in muddy sediments of the Sigsbee abyssal Analysis of Endemicity (PAE) was used to identi- plain (SBI cruise) within a depth range of 2200 to fy the areas of endemism in the SW region of GM. 3700 m. This method classifies the study sites according to their shared taxa, following the parsimony criteri- Family PARIAMBIDAE Laubitz, 1993 on (Rosen, 1988; Morrone, 2004). To obtain the 5. Deutella sp. cladograms and the strict consensus cladogram WINCLADA V 0.9.99 beta (Nixon, 1999) cladis- This species is a rare component, it occurs in soft tic software was used. The data matrix (which can bottoms of the Bay of Campeche (OGXVI cruise) at be requested from the first author) included 40 897 m depth. species (columns) that were identified as 0 to 39, collected in 15 sites (rows) that were identified as 6.
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