Marine Biodiversity Records, page 1 of 16. # Marine Biological Association of the United Kingdom, 2013 doi:10.1017/S175526721300064X; Vol. 6; e90; 2013 Published online Records and observations of amphipods (Amphipoda: Gammaridea and Corophiidea) from fouling assemblages in the Alacranes Reef, southern Gulf of Mexico carlos e. paz-ri’os1, nuno simo~es2 and pedro-luis ardisson1 1 Departamento de Recursos del Mar, Cinvestav, Carretera antigua a Progreso, km 6, Apartado Postal 73, Cordemex 97310 Merida, Yucatan, Mexico, 2Unidad Multidisciplinaria de Docencia e Investigacion, Facultad de Ciencias, Universidad Nacional Autonoma de Mexico, Puerto de Abrigo s/n, Sisal, Yucatan, Mexico The present study documents for the first time the occurrence of amphipods in the Alacranes Reef, Gulf of Mexico. Six macrofouling assemblages were sampled from two types of man-made structures (wood docks and navigation buoys). After sorting the material, 17 species were identified. An annotated checklist for all species is provided, documenting new records for 11 species (Ampithoe marcuzzii, Apolochus cf. delacaya, Audulla chelifera, Cymadusa compta, Leucothoe cf. wuriti, Neomegamphopus hiatus, Parhyale hawaiensis, Podocerus fissipes, Quadrimaera prope pacifica, Stenothoe gallensis and Stenothoe valida). Keywords: macrofouling, Peracarida, invasive species, hotspot, biodiversity, taxonomy, Yucatan Submitted 6 February 2013; accepted 13 June 2013 INTRODUCTION The Gulf of Mexico is a large marine ecosystem harbouring a great diversity of habitats (Kumpf et al., 1999), including oyster reefs, salt marshes, seagrass beds, estuaries, lagoons, mangroves, coral reefs, and soft and hard bottoms. Among these habitats, the coral reefs are remarkable for their complexity and high associated biodiversity. Coral reefs are common on the continental shelf throughout the Gulf of Mexico, although they also occur in deep water (Cordes et al., 2008). According to Tunnell et al. (2007a), the greatest coral reef area in the Gulf of Mexico is localized on the southern region. The main coral reefs are in the Florida Keys, north-western Cuba, the Campeche Bank, the Veracruz reef system and the Tuxpan reef system (Figure 1A). Regarding the above mentioned area, amphipods associated with coral reef environments have been reported for the Tuxpan reef system (McKinney, 1977), Florida Keys (Thomas, 1993), north-western Cuba (Ortiz & Lalana, 2010) and Veracruz reef system (Winfield & Ortiz, 2011). There are three studies recording amphipods associated with the typical carbonated bottom habitats of the Campeche Bank (Ortiz, 1979; Escobar-Briones & Jiménez-Guadarrama, 2010; Paz-Rı́os & Ardisson, 2013), but only one of these studies has sampled near coral reef environments (Escobar-Briones & Jiménez-Guadarrama, 2010). In this last study amphipods were identified only to family level; hence, there is currently Corresponding author: C.E. Paz-Rı́os Email: cpaz@mda.cinvestav.mx no checklist of amphipods to the species level associated with coral reef environments in the Campeche Bank. The Alacranes Reef is the largest coral reef area within the Gulf of Mexico and it has been categorized as a National Park since 1994. Its benthic fauna has been studied and it is listed in the Park’s management plan by the National Commission on Natural Protected Areas (CONANP, 2006) and in the checklist of the biota associated with the southern Gulf of Mexico coral reefs by Tunnell et al. (2007b). Nevertheless, these lists do not include several crustacean taxa, such as amphipods. The present study therefore provides the first checklist of amphipod species associated with the Alacranes Reef environment. This study constitutes the first part of an ongoing investigation that explores the species diversity of amphipods on artificial and natural substrates in the Alacranes Reef. The present checklist is made up of species collected on man-made structures (artificial substrates). Man-made structures have been classified as artificial reefs as they offer an extra substrate for the settlement, colonization and development of fouling organisms (Seaman & Jensen, 2000; Lin & Shao, 2002). The present study thereby contributes by documenting the species richness of fouling amphipods in the Alacranes Reef. Studies of fouling assemblages in the Gulf of Mexico have been mostly focused on the USA coastal area (Northern Gulf of Mexico), including but not limited to the following studies: Gunter & Geyer (1955); Pequegnat & Pequegnat (1968); Culpepper (1969); Lewbel et al. (1987); Martin & Bortone (1997). Few publications on fouling assemblages exist for the southern Gulf of Mexico; macrofouling species have been documented in four studies to date: Pech et al. (2002); Winfield et al. (2007, 2010, 2011). 1 2 carlos e. paz-ri’ os et al. Fig. 1. Area of study in the Gulf of Mexico: (A) coral reefs in the southern Gulf of Mexico; (B) area of study in the Alacranes Reef; (C) sampling sites on Isla Pérez. To increase our knowledge about the Alacranes Reef biodiversity, the aim of the present study is to document the occurrence of fouling amphipods on man-made structures (wood docks and navigation buoys) by means of an annotated checklist. The identified species composition will also contribute to a species occurrence baseline to be used in future comparisons of species composition collected from natural substrates. In turn, this will lead to the possible identification of potential invasive species, similar to the study of Winfield et al. (2011) for the Veracruz reef system. MATERIALS AND METHODS The Alacranes Reef forms part of North America’s marine ecoregion 14 (southern Gulf of Mexico) situated in the Campeche/Yucatanean outer neritic region (Wilkinson et al., 2009) (Figure 1A). It is a platform reef with a semi-elliptic shape, covering an area larger than 650 km2 (Bonet, 1967; Jordán-Dahlgren & Rodrı́guez-Martı́nez, 2003) (Figure 1B). Fouling amphipods were collected from six man-made structures: two wood docks (D) in Isla Pérez (the only permanently inhabited island in the Alacranes Reef) and four navigation buoys (B) near the same island (Table 1; Figure 1C). On each dock and buoy algae from an unmeasured area was recovered from the surface. The material was fixed in 90% alcohol in the field. To separate the species from the algae, the material was washed with freshwater and then preserved in 70% alcohol for sorting and identification in the laboratory. Species classification followed LeCroy et al. (2009); members of the family Caprellidae were excluded. Specimens are deposited in the regional collection Crustáceos de Yucatán (YUC-CC-255-11), UNAM-Sisal. All species are recorded in fouling amphipods in alacranes reef Table 1. Sampled man-made structures in Alacranes Reef. (B) Navigation buoy; (D) wood dock. Sample Latitude (N) ′ B1 B2 B3 B4 D1 D2 ′′ 22823 00 22822′ 55′′ 22822′ 36′′ 22822′ 35′′ 22822′ 57′′ 22823′ 00′′ Longitude (W) ′ ′′ 89840 35 89′ 40′ 24′′ 89840′ 23′′ 89840′ 17′′ 89840′ 55′′ 89841′ 02′′ an annotated checklist, with personal observations and/or figures. Species of taxonomic interest were illustrated to corroborate the species identity with a geographical extension or to document intraspecific morphological variation. The geographical range extension of species was recorded within two geographical scales: (i) the Mexican south-east sector; and (ii) the Gulf of Mexico. Species with geographical range extensions are indicated with a symbol (Q) preceding the name. RESULTS Date Dominant algae 20 April 2010 20 April 2010 20 April 2010 18 April 2010 17 October 2011 17 October 2011 Padina; Udotea; Ulva Padina; Udotea; Ulva Padina; Udotea; Ulva Padina; Udotea; Ulva Cladophora Acanthophora; Enteromorpha Family AMPHILOCHIDAE Boeck, 1871 Q Apolochus cf. delacaya (McKinney, 1978) (Figure 2) material examined 90 females (29 ovigerous), 58 unsexed, B1, YUC-CC-255-11-2. 2 females (1 ovigerous), 4 unsexed, B2, YUC-CC-255-11-3. 3 females (2 ovigerous), B3, YUC-CC-255-11-4. 19 females (4 ovigerous), 4 unsexed, B4, YUC-CC-255-11-5. distribution A total of 2204 individual amphipods were collected, counted and identified. Species composition is represented by two suborders, 11 families, 15 genera, and 17 species (Table 2). Two species are new records for the Gulf of Mexico (Podocerus fissipes Serejo, 1995 and Quadrimaera prope pacifica [Schellenberg, 1938]) and nine species are new records for the Mexican south-east sector (Ampithoe marcuzzii Ruffo, 1954, Apolochus cf. delacaya [McKinney, 1978], Audulla chelifera Chevreux, 1901, Cymadusa compta [Smith, 1873], Leucothoe cf. wuriti Thomas & Klebba, 2007, Neomegamphopus hiatus Barnard & Thomas, 1987, Parhyale hawaiensis [Dana, 1853], Stenothoe gallensis Walker, 1904 and Stenothoe valida Dana, 1853). systematics Order AMPHIPODA Latreille, 1816 Suborder GAMMARIDEA Latreille, 1802 General distribution: Gulf of Mexico (LeCroy et al., 2009). Gulf of Mexico records: Lobos Reef (McKinney, 1978); Florida Keys (Barnard & Thomas, 1983). habitat Apolochus delacaya has been found in the groove and buttress of coral reefs in shallow waters (5 m) (McKinney, 1978). Material examined of Apolochus cf. delacaya was collected on macrofouling communities with Padina, Udotea and Ulva as dominant algae from buoys anchored in the subtidal zone. remarks This is the first record for the Mexican south-east sector. There are three other recorded Apolochus species in the Gulf of Mexico (LeCroy et al., 2009): Apolochus casahoya (McKinney, 1978), Apolochus neapolitanus (Della Valle, 1893) and Apolochus pillaii (Barnard & Thomas, 1983). Apolochus cf. delacaya is distinguished from A. casahoya Table 2. Species composition and abundance of fouling amphipods in Alacranes Reef. (B) Navigation buoy; (D) wood dock. Suborder Family Species Sample Abundance Gammaridea Amphilochidae Bateidae Hyalidae Apolochus cf. delacaya Batea cuspidata Parhyale hawaiensis Protohyale macrodactyla Anamixis cavatura Leucothoe cf. wuriti Elasmopus rapax Quadrimaera prope pacifica Stenothoe gallensis Stenothoe valida Ampithoe marcuzzii Ampithoe ramondi Cymadusa compta Ericthonius sp. D Neomegamphopus hiatus Audulla chelifera Podocerus fissipes B1, B1, D2 B3 B1, B1, B1, B1, B1, B1, B1, B1, D2 B1, B2 B2, B1, 180 4 10 20 3 39 529 159 247 24 231 470 1 92 1 131 63 Leucothoidae Melitidae Stenothoidae Corophidea Ampithoidae Ischyroceridae Neomegamphopidae Photidae Podoceridae B2, B3, B4 B4 B3 B2, B4 B3, B4 B2, B3, B2, B3, B2, B3, B2, B3, B2, B3, B4 B4 B4 B4 B4, D1, D2 B2, B3, B4 B3 B2, B3, B4 3 4 carlos e. paz-ri’ os et al. records: western Florida; Florida Keys (Ortiz, 1991; LeCroy, 2004); western Cuba (Varela et al., 2003); Bay of Campeche (Winfield et al., 2006); Campeche Bank (Paz-Rı́os & Ardisson, 2013). habitat Found in Thalassia or Halodule grassbeds and among algae (Caulerpa and Halimeda) on sandy or muddy bottoms; depth range from 0 to 52 m (LeCroy, 2004; Winfield et al., 2006; Coleman, 2010). Family HYALIDAE Bulycheva, 1957 Q Parhyale hawaiensis (Dana, 1853) material examined 2 males, 5 females, 3 unsexed, D2, YUC-CC-255-11-18. distribution General distribution: cosmopolitan in tropical and warm temperate seas (Serejo, 1999; Bousfield & Hendrycks, 2002; LeCroy, 2007). Gulf of Mexico records: western Florida (Shoemaker, 1956; LeCroy, 2007); Barataria Bay area (Thomas, 1976); Galveston; Port Isabel (McKinney, 1977); Biloxi Bay (Pederson & Peterson, 2002). habitat Fig. 2. Apolochus cf. delacaya (McKinney, 1978), male: (A) mandible; (B) gnathopod 2; (C) uropods 1 –3. Scale bars: A, 0.1 mm; B, C, 0.1 mm. and A. neapolitanus by having four submarginal spines on the anterolateral surface of the propodus of gnathopod 2 (1 –2 in A. casahoya; without spines in A. neapolitanus). It differs from A. pillaii in having spines on the medial margin of uropod 1 outer ramus (without spines in A. pillaii). Apolochus cf. delacaya from the Alacranes Reef is well aligned with the description of Apolochus delacaya from the Lobos Reef by McKinney (1978), but differs in having spines on the medial margin of the peduncle of uropod 3 (without spines in A. delacaya). Family BATEIDAE Stebbing, 1906 Batea cuspidata (Shoemaker, 1926) material examined 3 females, B1, YUC-CC-255-11-7. 1 female, B4, YUC-CC255-11-8. From the intertidal or shallow subtidal zone; in algae and other fouling growth on hard substrates (e.g. piers, jetties, oyster beds, bryozoans, mangroves and coral rock); this species has also been found in the root system of Spartina marshes and intertidally on wet sand under shells (LeCroy, 2007). Parhyale hawaiensis has recently been recorded in a continental water body called Cenote Aerolito on Cozumel Island by Trujillo-Pisanty et al. (2010). The Yucatan Peninsula, Mexico, is a karstic zone with the main characteristic being cenotes. Cenotes (sinkholes) are the most peculiar aquatic ecosystem of the peninsula and are formed by dissolution of the carbonate rock in the karstic platform (Schmitter-Soto et al., 2002). Most cenotes are classified as anchialine since they are characterized by a salinity gradient due to their connection with the sea (Mejı́a-Ortı́z et al., 2006). remarks This is the first record for the Mexican south-east sector. Identified specimens were aligned with the descriptions and illustrations of Shoemaker (1956) and Serejo (1999) with one variation that appears to be developmental. The examined material has one and zero spines on the outer ramus of uropods 1–2, respectively (compared to two spines on each uropod). The largest specimen examined in this study was 3.6 mm, whereas the largest specimens examined by Shoemaker (1956) and Serejo (1999) were 12 mm and 10.2 mm, respectively. Protohyale macrodactyla (Stebbing, 1899) (Figure 3) distribution General distribution: eastern Florida; Caribbean; Gulf of Mexico (LeCroy, 2004; Coleman, 2010). Gulf of Mexico material examined 7 males, 13 females (2 ovigerous), B3, YUC-CC-255-11-25. fouling amphipods in alacranes reef half of the gnathopod 2 palm depend on the developmental stage of specimens. Therefore, this characteristic is confirmed and should be considered part of the diagnosis for Protohyale macrodactyla. The largest specimen examined in this study was 5 mm, whereas the largest specimens examined by Stebbing (1899), Schellenberg (1939) and Serejo (1999) were 4 mm, 10 mm and 7.2 mm, respectively. Family LEUCOTHOIDAE Dana, 1852 Anamixis cavatura Thomas, 1997 material examined 1 male, B1, YUC-CC-255-11-26. 2 unsexed, B3, YUC-CC255-11-27. distribution General distribution: Gulf of Mexico; south-eastern Florida; Bahamas; Caribbean (Thomas, 1997; LeCroy, 2007; White, 2011a). Gulf of Mexico records: from the Mississippi Delta to Tampa (Pearse, 1912); Florida Keys (Shoemaker, 1933b; Thomas, 1997); western Cuba (Varela et al., 2003); Campeche Bank (Paz-Rı́os & Ardisson, 2013). habitat Shallow subtidal (1 – 20.9 m), among sandy bottoms and coral rubble; endocommensal associates of asconoid sponges and colonial ascidians (White, 2011a; Paz-Rı́os & Ardisson, 2013). Q Leucothoe cf. wuriti Thomas & Klebba, 2007 (Figure 4) Fig. 3. Protohyale macrodactyla (Stebbing, 1899), male: (A) gnathopod 2 of juvenile 1; (B) gnathopod 2 of juvenile 2; (C) gnathopod 2 of adult 1; (D) gnathopod 2 of adult 2; (E) coxa 3 of adult 2; (F) pereopod 7 of adult 2; (G) uropod 1 of adult 2. Scale bar: A– G, 0.2 mm. distribution General distribution: Caribbean; Gulf of Mexico; Brazil; Madagascar; western Africa (Serejo, 1999; Ortiz et al., 2007; LeCroy et al., 2009). Gulf of Mexico records: Laguna Yalahau (Oliva-Rivera & Jiménez-Cueto, 1997); Laguna Rı́o Soto la Marina; Laguna Camaronera; Laguna Terminos (Cházaro-Olvera et al., 2002). habitat From the low intertidal to shallow subtidal zone; freeswimming on algae and hard substrates (Bousfield & Hendrycks, 2002; LeCroy et al., 2009). remarks Serejo (1999) redescribed this species with Brazilian specimens. Based on her observations and Schellenberg’s (1939) conclusions, gnathopod 2 palm is nearly straight without visible swellings on the distal half, as Stebbing (1899) had described and illustrated. The material examined in the present study clearly shows that these swellings on the distal Fig. 4. Leucothoe cf. wuriti Thomas & Klebba, 2007, male: (A) head and antennae; (B) gnathopod 1; (C) gnathopod 2; (D) epimera 1 –3. Scale bar: A– D, 0.5 mm. 5 6 carlos e. paz-ri’ os et al. material examined 10 males, 13 females (1 ovigerous), B1, YUC-CC-255-11-32. 1 male, 3 females, B2, YUC-CC-255-11-33. 6 males, 5 females (1 ovigerous), 1 unsexed, B4, YUC-CC-255-11-34. distribution General distribution: Gulf of Mexico; south-eastern Florida; Caribbean (LeCroy, 2011; White, 2011a, b). Gulf of Mexico records: Florida Keys (Thomas & Klebba, 2007). habitat Leucothoe wuriti has been collected in the shallow subtidal (2 –15 m) of coral reefs and mangrove lagoons as an endocommensal of ascidians (Thomas & Klebba, 2007; White, 2011b). Material examined of Leucothoe cf. wuriti was collected on macrofouling communities with Padina, Udotea and Ulva as dominant algae from buoys anchored in the subtidal zone. remarks This is the first record for the Mexican south-eastern sector. To date there are 11 recorded nominal species of leucothoids in the Gulf of Mexico (LeCroy et al., 2009; Winfield & Álvarez, 2009; Winfield et al., 2009b; LeCroy, 2011; White, 2011a; Ortiz & Winfield, 2012): Leucothoe ashleyae Thomas & Klebba, 2006, Leucothoe barana Thomas & Klebba, 2007, Leucothoe flammosa Thomas & Klebba, 2007, Leucothoe hendrickxi Winfield & Álvarez, 2009, Leucothoe hortapugai Winfield et al. 2009b, Leucothoe kensleyi Thomas & Klebba, 2006, Leucothoe laurensis Thomas & Ortiz, 1995, Leucothoe luquei Ortiz & Winfield, 2012; Leucothoe ortizi Winfield & Álvarez, 2009, Leucothoe ubouhu Thomas & Klebba, 2007, and Leucothoe wuriti. Due to the great leucothoid host diversity, specialized underwater collecting techniques are necessary in order to isolate commensal amphipods (Thomas & Klebba, 2007). However, these authors also recognize that collections of leucothoid amphipods are often the result of incidental capture, as happened in the present study from fouling communities dominated by algae. Leucothoe cf. wuriti from the Alacranes Reef is aligned with the description of Leucothoe wuriti in Thomas & Klebba (2007) and White (2011b), as well as the remarks in LeCroy (2011). However, it differs in having epimeron 1 with three ventral setae (2 in L. wuriti), and epimeron 3 with two ventral setae (bare in L. wuriti). Family MELITIDAE Bousfield, 1973 Elasmopus rapax (Costa, 1853) (Figure 5) material examined 128 males, 182 females (77 ovigerous), 10 unsexed, B1, YUCCC-255-11-46. 48 males, 109 females (26 ovigerous), B3, YUC-CC-255-11-47. 14 males, 38 females (10 ovigerous), B4, YUC-CC-255-11-48. distribution General distribution: Mediterranean Sea; Red Sea; eastern Atlantic; western Atlantic; Indo-Pacific (Hughes & Lowry, 2010). Gulf of Mexico records: Pigeon Key; Florida Bay Fig. 5. Elasmopus rapax (Costa, 1853), male morphotype A: (A) gnathopod 2, inner side; (B) basis of pereopod 7; (C) epimera 1 –3; (D) telson; male morphotype B: (E) gnathopod 2, inner side; (F) basis of pereopod 7; (G) epimera 1 – 3; (H) telson. Scale bars: A– C, E – G, 0.5 mm; D, H, 0.1 mm. (Pearse, 1912); Key West (Pearse, 1912; LeCroy, 2000); Plantation Key (Shoemaker, 1933a); offshore Texas; Lobos Reef (McKinney, 1977); Yalahau lagoon (Oliva-Rivera & Jiménez-Cueto, 1997). habitat This species is found in a broad range of habitats (e.g. jetties, piers, rocks, dead coral, mangrove roots, algae, sponges; sand bottoms and mud bottoms); occasionally it is found on the carapace of the loggerhead sea turtle (LeCroy, 2000). remarks McKinney (1977) documented that specimens of Elasmopus rapax from the Caribbean and Gulf of Mexico are highly variable, particularly the juvenile forms. Karaman (1982) noted the presence of two male morphotypes in the Mediterranean: morphotype A (with long setae on epimera 1– 3, long posterior setae on the basis of peraeopods 5– 7, and telson with very short robust setae) and morphotype B (with robust setae on epimera 1– 3, very short posterior setae on the basis of peraeopods 5–7, and telson with long robust setae). LeCroy (2000) determined that the Karaman’s (1982) morphotype A appears to be similar to E. rapax from the Gulf of Mexico. Recently, Hughes & Lowry (2010) redescribed E. rapax and considered morphotype B (with propodus palm of gnathopod 2 with two teeth) to be the juvenile male form in comparison to morphotype A (with propodus palm of gnathopod 2 with three teeth). They also pointed out that the great morphological variation of this species fouling amphipods in alacranes reef depends on the marine region it is collected from. In the examined material from the Alacranes Reef both morphotypes A and B are present. However, there is a variation in morphotype B, having a telson with short lobes and long robust setae. There is also a transitional form with the above mentioned characteristics for morphotype B, but with propodus palm of gnathopod 2 with three teeth as in morphotype A. Material examined in the present study is well aligned with the description of Q. prope pacifica except for the presence of six long setae (instead of 5) on the posterior margin of basis in gnathopod 1, and eight spines (instead of 7) on the ventral margin of epimeron 3. As Krapp-Schickel & Ruffo (2000) have noted, there is morphological variability in this species. Family STENOTHOIDAE Boeck, 1871 Q Stenothoe gallensis Walker, 1904 (Figure 7) Q Quadrimaera prope pacifica (Schellenberg, 1938) (Figure 6) material examined material examined 28 males, 40 females (5 ovigerous), B1, YUC-CC-255-11-60. 14 males, 21 females (2 ovigerous), B2, YUC-CC-255-11-61. 19 males, 29 females (6 ovigerous), B3, YUC-CC-255-11-62. 3 males, 5 females, B4, YUC-CC-255-11-63. 5 males, 46 females (10 ovigerous), B1, YUC-CC-255-11-78. 2 males, 17 females (3 ovigerous), B2, YUC-CC-255-11-79. 4 males, 47 females (11 ovigerous), 2 unsexed, B3, YUC-CC-255-11-80. 9 males, 115 females, B4, YUC-CC-255-11-81. distribution General distribution: Caribbean (Krapp-Schickel & Ruffo, 2000). Gulf of Mexico records: previously unreported. habitat Marine, epibenthic, from the intertidal to shallow subtidal zone (0 –3 m), on rocky and sandy bottoms (Krapp-Schickel & Ruffo, 2000). distribution General distribution: cosmopolitan in tropical and temperate seas (LeCroy, 2011). Gulf of Mexico records: Panama City Beach (Culpepper, 1969); Lobos Reef (McKinney, 1977); off Louisiana (Lewbel et al., 1987); Florida Keys (Thomas, 1993); Sigsbee abyssal plain (Winfield et al., 2006); Veracruz reef system (Winfield et al., 2007); Perdido Key (LeCroy, 2011). remarks This is the first record for the Gulf of Mexico. Krapp-Schickel & Ruffo (2000) described Quadrimaera prope pacifica (Caribbean), a species very close to Quadrimaera pacifica (Indo-Pacific). A redefinition of this species using topotypic material is required (type locality of Makin, Kiribati), due to the need for confirmation of a triangular spine on the inner margin of the dactylus of gnathopod 2 displayed on the original illustration by Schellenberg (Krapp-Schickel & Ruffo, 2000; Ruffo et al., 2000; Krapp-Schickel, 2009). habitat Marine, epibenthic from the low intertidal to shallow subtidal zone (0.5 – 44 m); hard and soft bottoms, roots of mangroves, algae, ascidians, bryozoans, hydroids, plankton (Fox & Bynum, 1975; Thomas, 1993; Ortiz et al., 2007; LeCroy, 2011). This species is non-indigenous to the Gulf of Mexico, but is commonly found in fouling communities on pilings, floats, ship bottoms, buoys, rocks and mangrove prop roots (Fox & Bynum, 1975; LeCroy et al., 2009; LeCroy, 2011). Fig. 6. Quadrimaera prope pacifica Schellenberg, 1938, male: (A) head; (B) gnathopod 1; (C) gnathopod 2; (D) pereopod 7; (E) epimera 2– 3; (F) telson. Scale bars: A– E, 0.5 mm; F, 0.2 mm. 7 8 carlos e. paz-ri’ os et al. Fig. 8. Stenothoe valida Dana, 1853, male: (A) gnathopod 2; (C) pereopod 7; (D) uropod 1; (E) uropod 3; female: (B) gnathopod 2. Scale bars: A– C, 0.5 mm; D, E, 0.1 mm. Fig. 7. Stenothoe gallensis Walker, 1904, male: (A) gnathopod 2; (C) pereopod 7; (D) uropod 1; (E) uropod 3; female: (B) gnathopod 2. Scale bars: A– C, 0.5 mm; D, E, 0.1 mm. remarks This is the first record for the Mexican south-eastern sector. Stenothoe gallensis was recorded in the Sigsbee abyssal plain, south-western Gulf of Mexico, to 3635 m depth by Winfield et al. (2006), but this record has been overlooked by two checklists of species in the region (LeCroy et al., 2009; LeCroy, 2011), likely because that occurrence reports an atypical habitat for the species. Material examined is aligned with the regional diagnosis by LeCroy (2011) for the Florida specimens. The main characteristics used to distinguish between males of S. gallensis from Stenothoe valida are on gnathopod 2 (merus, posterior margin crenulate; propodus, distal palmar tooth relatively small), merus of pereopod 7 (not broadly expanded with posterodistal lobe not reaching more than halfway to distal margin of carpus), uropod 1 (peduncle with distoventral spur) and uropod 3 (terminal article of ramus expanded proximally and dorsal margin with crenulations). The main characteristic used to distinguish between females of S. gallensis from S. valida is on gnathopod 2 (propodus, posterior margin without distal palmar teeth). Q Stenothoe valida Dana, 1853 (Figure 8) material examined 1 male, 6 females (2 ovigerous), B1, YUC-CC-255-11-82. 3 males, 6 females, B2, YUC-CC-255-11-83. 2 males, 2 females, B3, YUC-CC-255-11-84. 1 male, 3 females, B4, YUC-CC-255-11-85. distribution General distribution: nearly cosmopolitan in tropical and warm temperate waters (LeCroy, 2011). Gulf of Mexico records: Dry Tortugas (Pearse, 1932). habitat Marine, epibenthic, from the low intertidal to shallow subtidal zone (1 –33 m); hard and soft bottoms, algae, Thalassia grassbeds, hydroids, empty shells of barnacle Megabalanus, sponges, commensal of hydrocoral Millepora complanata Lamarck, 1816 (Pearse, 1932; Lewis, 1992; LeCroy, 2011). remarks This is the first record for the Mexican south-eastern sector. Stenothoe valida is a species commonly found in fouling communities on pilings, ship bottoms and buoys (LeCroy, 2011). Material examined is aligned with the regional diagnosis by LeCroy (2011) for the Florida specimens. The main characteristics used to distinguish between males of S. valida from Stenothoe gallensis are on gnathopod 2 (merus, posterior margin entire, not crenulate; propodus, distal palmar tooth relatively large), merus of pereopod 7 (broadly expanded with posterodistal lobe reaching distal margin of carpus), fouling amphipods in alacranes reef uropod 1 (peduncle without distoventral spur), and uropod 3 (terminal article of ramus lanceolate, not expanded proximally and dorsal margin without crenulations). The main characteristic used to distinguish between females of S. valida from S. gallensis is on gnathopod 2 (propodus, posterior margin straight with 1–3 small distal palmar teeth). Suborder COROPHIIDEA Leach, 1814 Family AMPITHOIDAE Stebbing, 1899 Q Ampithoe marcuzzii Ruffo, 1954 (Figure 9) material examined 21 males, 75 females (17 ovigerous), 5 unsexed, B1, YUC-CC-255-11-98. 8 males, 29 females (8 ovigerous), B2, YUC-CC-255-11-99. 5 males, 31 females (6 ovigerous), B3, YUC-CC-255-11-100. 16 males, 41 females (15 ovigerous), B4, YUC-CC-255-11-101. habitat From the intertidal zone to depths of 6 m; hard and soft bottoms, coral rubble, pilings, algae, grassbeds, ascidians, sponges, bryozoans and colonies of polychaetes (LeCroy, 2002; Martı́n & Dı́az, 2003). remarks This is the first record for the Mexican south-eastern sector. There is a very similar form from Florida described by LeCroy (2002) as Ampithoe cf. marcuzzii, which was listed as a Gulf of Mexico species by LeCroy et al. (2009). LeCroy (2002) inferred that the discrepancy between the Florida and Caribbean material could be a developmental difference; the palmar angle of gnathopod 1 is somewhat more rounded and the spine on the ventral margin of the inner ramus of uropod 3 is not present in sub-adult males of A. cf. marcuzzii. The material examined is aligned with the original illustrations by Ruffo (1954); the adult males have a moderately sinuous palmar angle of gnathopod 1 and a spine on the ventral margin of the inner ramus of uropod 3. Ampithoe ramondi Audouin, 1826 distribution material examined General distribution: eastern Florida; Gulf of Mexico; Caribbean (Ruffo, 1954; Ortiz & Lemaitre, 1994; Ortiz & Lalana, 1996; LeCroy, 2002; Martı́n & Dı́az, 2003; Foster et al., 2009). Gulf of Mexico records: Pigeon Key (LeCroy, 2002). 51 males, 130 females (35 ovigerous), 7 unsexed, B1, YUC-CC-255-11-102. 14 males, 78 females (26 ovigerous), 10 unsexed, B2, YUC-CC-255-11-103. 18 males, 42 females (11 ovigerous), 3 unsexed, B3, YUC-CC-255-11-104. 25 males, 28 females (8 ovigerous), B4, YUC-CC-255-11-105. 1 ovigerous female, D1, YUC-CC-255-11-106. 5 males, 16 females, 42 unsexed, D2, YUC-CC-255-11-107. distribution General distribution: cosmopolitan in tropical and subtropical seas (LeCroy, 2002). Gulf of Mexico records: Dry Tortugas (as Ampithoe divisura Shoemaker, 1933b); Lobos Reef (McKinney, 1977); Continental Shelf of Yucatan (Ortiz, 1979); Florida Keys (Thomas, 1993); Laguna Rı́o Soto la Marina (Cházaro-Olvera et al., 2002); Cojı́mar Bay (Ortiz, 2001); Veracruz reef system (Winfield et al., 2007, 2010). habitat Fouling growth, algae, grassbeds, ascidians, sponges, bryozoans, sandy bottoms; associates of polychaete colonies, and plankton tows over Thalassia (Serejo, 1998; LeCroy, 2002; Ortiz et al., 2007). Q Cymadusa compta (Smith, 1873) material examined 1 female, D1, YUC-CC-255-11-108. distribution Fig. 9. Ampithoe marcuzzii Ruffo, 1954, male: (A) gnathopod 1 of juvenile; (B) gnathopod 1 of subadult; (C) gnathopod 2 of adult; (E) uropod 1 of adult; (F) uropod 3 of adult; female: (D) gnathopod 2. Scale bars: A– D, 0.5 mm; E, 0.2 mm; F, 0.2 mm. General distribution: Bermuda; east coast of the United States; Gulf of Mexico; Caribbean (Bousfield, 1973; LeCroy, 2002; Ortiz et al., 2007; LeCroy et al., 2009; Gable et al., 2010; Ortiz & Lalana, 2010). Gulf of Mexico records: West Bay; Matagorda Bay; St Antonio Bay; Corpus Christi Bay; Port Aransas; St Luis Pass; Port Mansfield; Port Isabel; Laguna Madre (McKinney, 1977); Apalachicola Bay (Sheridan, 1980); Laguna Terminos (Ledoyer, 1986; Corona et al., 2000; Cházaro-Olvera et al., 2002); Florida Keys (Thomas, 9 10 carlos e. paz-ri’ os et al. 1993); Laguna Rı́o Soto la Marina (Cházaro-Olvera et al., 2002); St Andrew Bay (Foster et al., 2004); Habana-Matanzas region (Ortiz & Lalana, 2010). habitat From the intertidal zone to depths of 10 m; in a wide range of salinities (5– 33 ppt) on hard and soft bottoms, algae and grassbeds (LeCroy, 2002; Ortiz et al., 2007). remarks This is the first record for the Mexican south-eastern sector. Cymadusa compta is one of the most widespread shallowwater amphipod species in the Gulf of Mexico (McKinney, 1977). Nevertheless, there are still regions in the southern sector where it has not been reported, denoting the scarce knowledge about the taxa in that sector. Family ISCHYROCERIDAE Stebbing, 1899 Ericthonius sp. D (Figure 10) material examined 27 males, 28 females (2 ovigerous), B1, YUC-CC-255-11-118. 4 males, 7 females (1 ovigerous), B2, YUC-CC-255-11-119. 1 male, B3, YUC-CC-255-11-120. 14 males, 11 females, B4, YUC-CC-255-11-121. distribution General distribution: Gulf of Mexico. Gulf of Mexico records: this species has been collected so far only from the Alacranes Reef. habitat Members of the genus are tube-dwelling fouling species, often found on hard substrates like rocks, pilings, oil rings, mangroves and oyster shells; sandy bottoms, algae, grassbeds, sponges, hydroids (LeCroy, 2007). Ericthonius is generally a shallow-water genus (1 –10 m) (Bousfield, 1973; Thomas, 1993), although it has been recorded in the Gulf of Mexico from the continental slope and abyssal plain to depths of 3690 m by Winfield et al. (2006). Ericthonius sp. D was collected on macrofouling communities with Padina, Udotea and Ulva as dominant algae from buoys anchored in the subtidal zone. remarks This species forms part of Group 1 of Ericthonius according to Myers & McGrath (1984) as having stridulating ridges on coxa 2 in males, coxa 2 widely separated from 1 and 3, and male gnathopod 2 carpus with two posterodistal teeth. There are three other species sharing these characteristics in the Atlantic: Ericthonius brasiliensis (Dana, 1853), Ericthonius difformis Milne-Edwards, 1830 and Ericthonius punctatus (Bate, 1857). Ericthonius sp. D differs (males) from these species in having short antennae (elongate in E. punctatus), lacking a posterior swelling on the basis of gnathopod 1 (with swelling in E. brasiliensis) and having a major accessory tooth on the carpus of gnathopod 2 (small or absent accessory tooth in E. difformis). LeCroy (2007) described briefly three possible new species from Florida and the east coast of the United States: Ericthonius sp. A, B and C. Ericthonius sp. D differs from Ericthonius sp. A and C by a posterodistal margin not produced, lacking wing-shaped process on the basis of pereopod 5. It differs from Ericthonius sp. B in having small eyes. The Fig. 10. Ericthonius sp. D, male: (A) habitus; (B) carpus, propodus and dactylus of gnathopod 1; (C) gnathopod 2; (D) coxa, basis and ischium of pereopod 5; (E) epimeron 3; (F) uropod 3. Scale bars: A, 0.5 mm; C, 0.2 mm; B, D– F, 0.1 mm. fouling amphipods in alacranes reef material examined in the present study is still under examination because it is suspected to be an undescribed species. At the moment, it closely resembles the Indo-Pacific species Ericthonius parabrasiliensis Just, 2009, but more observations are required prior to final identification. (longer than wide in N. kalanii), a gaping carpal tooth (not gaping in N. kalanii); propodus not expanded or lobate proximally (expanded or lobate proximally in N. kalanii) and an entire palm (beaded or minutely serrate in N. kalanii). Family PHOTIDAE Boeck, 1871 Q Audulla chelifera Chevreux, 1901 Family NEOMEGAMPHOPIDAE Myers, 1981 Q Neomegamphopus hiatus Barnard & Thomas, 1987 material examined material examined 1 male, B2, YUC-CC-255-11-122. 1 male, B2, YUC-CC-255-11-123. 61 males, 69 females (13 ovigerous), B3, YUC-CC-255-11-124. distribution distribution General distribution: eastern Florida; Gulf of Mexico; Caribbean (Barnard & Thomas, 1987; LeCroy et al., 2009; LeCroy, 2011). Gulf of Mexico records: Florida Keys; northwestern Florida (Barnard & Thomas, 1987; Thomas, 1993); Apalachee Bay (Munguia, 2004); St Joe Bay (Munguia et al., 2007). General distribution: Bermuda; Gulf of Mexico; Caribbean; tropical Indo-Pacific (Thomas & Barnard, 1987; Lazo-Wasem & Gable, 1987; LeCroy, 2000; Ortiz et al., 2007; LeCroy et al., 2009). Gulf of Mexico records: southern Florida Keys (LeCroy, 2000). habitat Marine, epibenthic, from the low intertidal to shallow subtidal zone (1 –49 m); sandy bottoms, coral rubble, algae (Halimeda), grassbeds (Syringodium and Thalassia), associates of pen shell (Atrina) living on grassbeds (Barnard & Thomas, 1987; Munguia et al., 2007). remarks This is the first record for the Mexican south-eastern sector. For a long time, Neomegamphopus hiatus was the only reported species of neomegamphopid in the Gulf of Mexico. Recently, Paz-Rı́os & Ardisson (2013) recorded another Neomegamphopus species in the Gulf of Mexico (Yucatan continental shelf), Neomegamphopus kalanii Barnard & Thomas, 1987. These species are separated (based on males) because antenna 1 of N. hiatus has an accessory flagellum longer than article 1 of primary flagellum (subequal in N. kalanii); carpus of gnathopod 1 almost as wide as long habitat Marine, epibenthic, on shallow waters (0 – 5 m) and high energy areas from coral reefs; algae (Halimeda, Caulerpa, Turbinaria, Ulva and Sargassum) (Thomas & Barnard, 1987; LeCroy, 2000; Ortiz et al., 2007). remarks This is the first record for the Mexican south-eastern sector. Audulla Chevreux, 1901 is a monotypic genus. Its single species, Audulla chelifera, is characterized by a flattened flagellum (expanded and paddle-shaped) of the male antenna 2, and the chelate male gnathopod 2. Currently, it is reported in Gable et al. (2010) for Bermuda as Gammaropsis lina Lazo-Wasem & Gable (1987) with a redescribed type specimen from Kunkel (1910). Family PODOCERIDAE Leach, 1814 Q Podocerus fissipes Serejo, 1995 (Figure 11) Fig. 11. Podocerus fissipes Serejo, 1995, male: (A) habitus; (B) uropods 1 – 3. Scale bars: A, 1 mm; B, 0.2 mm. 11 12 carlos e. paz-ri’ os et al. material examined 2 males, 17 females (1 ovigerous), B1, YUC-CC-255-11-125. 3 males, 2 females (1 ovigerous), B2, YUC-CC-255-11-126. 1 male, 1 female, B3, YUC-CC-255-11-127. 23 males, 14 females, B4, YUC-CC-255-11-128. distribution General distribution: south-eastern Florida; Caribbean; southeastern Brazil; (Serejo, 1995; Baldinger & Gable, 2002; LeCroy, 2011). Gulf of Mexico records: previously unreported. habitat Podocerus fissipes is a marine epibenthic species in shallow waters (0 – 5 m); associated with the sponges Dysidea fragilis (Montagu, 1818) and Mycale microsigmatosa Arndt, 1927 in Brazil (Serejo, 1995, 1998; Ribeiro et al., 2003). The genus is frequently found in fouling communities on hard substrates, clinging to algae, barnacles, sponges, hydroids, gorgonians and other fouling growth in relatively high energy habitats (LeCroy, 2011). remarks This is the first record for the Gulf of Mexico. Given the fouling lifestyle of members of this genus and artefacts caused by a lack of sampling, Podocerus fissipes may eventually be found in more localities in the north-western Atlantic (LeCroy, 2011). So far there are four recorded species of Podocerus in the Gulf of Mexico (including the new record in the present study): Podocerus brasiliensis (Dana, 1853), Podocerus chelonophilus (Chevreux & Guerne, 1888), P. fissipes and Podocerus kleidus Thomas & Barnard, 1992. Podocerus fissipes is easily distinguished from the other Gulf of Mexico Podocerus species by the presence of long, stout, dorsal setae on pereonites 5–6 and pleonites 1–2; one small seta at anterodistal corner of coxal plates 1– 4; and by the presence of two long apical spines on the telson. Only one adult male presented three apical spines on the telson. DISCUSSION Amphipods associated with coral reefs have been studied around the world. Currently the amphipod fauna of the Great Barrier Reef is the richest fauna yet known from any tropical reef area with 256 species (Lowry & Myers, 2009), as a result of an intensive sampling effort. On the other hand, in the tropical western Atlantic (mainly in the Caribbean and Gulf of Mexico) most of the species are known from sporadic records and/or new single species descriptions and coral reef species have not been the subject of detailed systematic study (Thomas, 1993; Baldinger, 2000). Among studies which have reported coral reef species in the Mexican Atlantic are McKinney (1977, 1978, 1979, 1980a, b), Oliva-Rivera (2003), Winfield & Escobar-Briones (2007), Winfield et al. (2007, 2010) and Winfield & Ortiz (2011). The recorded amphipod species from the Mexican Gulf reefs have been from the Tuxpan –Veracruz reef system in the south-western sector. The present study is the first checklist of fouling amphipods associated with coral reef environments from the Campeche Bank in the south-eastern sector. Recent studies in the south-western sector have focused on the space – time variations of fouling assemblages on artificial panels established in the Veracruz reef system (Winfield et al., 2007, 2010) and species descriptions (Winfield & Álvarez, 2009; Winfield & Ortiz, 2009, 2010; Winfield et al., 2009a, b). Regarding the studies of fouling assemblages, the accumulated species richness from artificial panels in Winfield et al. (2007, 2010) reached 23 nominal species of Gammaridea sensu lato, whereas the accumulated species richness from man-made structures (navigation buoys and wood docks) in the present study reached 16 nominal species of Gammaridea sensu lato. The difference between the species number from the present study and previous studies may be related to the sampling effort. There are three navigation buoys and eight mooring buoys in the present study area that were not sampled. More species are expected to be found if these buoys are sampled and if the regional weather seasonality is examined (cold fronts, rainy and dry seasons). An updated checklist of the marine invasive amphipod species from Mexico (particularly from the Veracruz reef systems) was provided by Winfield et al. (2011). This list includes 11 species, but as those authors suggested, this is likely a subestimation of the current number of invasive amphipods, considering the structural complexity and environmental heterogeneity of the different marine ecosystems and the importance of the traffic of national and international ships. For instance, three possible invasive species (Elasmopus rapax, Stenothoe gallensis and Stenothoe valida) are listed in the present study, supporting the possibility of subestimation. These species were qualified as strong candidates for introduced species in Hawaii by means of shipping (Barnard, 1970; Coles et al., 1999; Carlton & Eldredge, 2009). To date, these species have been characterized as invasive species in other coastal areas. Elasmopus rapax has been determined as an invasive species on the Australian coast (Hughes & Lowry, 2010), S. gallensis as an invasive species on the Mediterranean coast (Streftaris et al., 2005; Bakir et al., 2010) and S. valida as a invasive species on the Californian coast (Carlton & Hodder, 1995; Ruiz et al., 2000). However, before establishing an invasive status for these species in the Alacranes Reef, a detailed risk assessment that includes criteria (among others) of Chapman & Carlton (1991) for introduced species (e.g. type locality, distribution, habitat, natural history and dispersal mechanism) will be necessary. ACKNOWLEDGEMENTS We thank Jazmı́n Deneb Ortigosa (UNAM, Mexico) and Julio Canton Avila (Pronatura, Mexico) for their help with the sample collection from the Alacranes Reef; and we extend special thanks to the Oceanography Research Station in Progreso, Mexican Navy for the transportation logistics on several occasions to the Alacranes Reef. The authors also thank the anonymous referees who helped to improve the manuscript. REFERENCES Arndt W. (1927) Kalk- und Kieselschwämme von Curaçao. Bijdragen tot de Dierkunde 25, 133–158. fouling amphipods in alacranes reef Audouin V. (1826) Explication sommaire des planches de crustaces ded l’Egypte et de la Syrie, publiées par Jules-César Savigny, membre de l’Institut: offrant un exposé des caracteres naturels des genres, avec la distinction des especes. Description de I’Egypte, Histoires Naturelle 1, 77–98. Bakir K., Sezgin M. and Katağan T. (2010) Alien amphipods on the Turkish coasts. Zoologica Baetica 21, 191 –196. species. Journal of the Marine Biological Association of the United Kingdom 91, 455–469. Coles S.L., DeFelice R.C., Eldredge L.G. and Carlton J.T. (1999) Historical and recent introductions of non-indigenous marine species into Pearl Harbor, Oahu, Hawaiian Islands. Marine Biology 135, 147 –158. Baldinger A.J. (2000) Amphipoda of the Caribbean region and the Gulf of Mexico. Polskie Archiwum Hydrobiologii 47, 697–702. CONANP (2006) Programa de conservación y manejo Parque Nacional Arrecife Alacranes. 1st edition. México: Comisión Nacional de Áreas Naturales Protegidas. Baldinger A.J. and Gable M.F. (2002) The genus Podocerus (Crustacea: Amphipoda: Podoceridae) from Guana Island, British Virgin Islands. Postilla 226, 1–21. Cordes E.E., McGinley M.P., Podowski E.L., Becker E.L., Lessard-Pilon S., Viada S.T. and Fisher C.R. (2008) Coral communities of the deep Gulf of Mexico. Deep-Sea Research I 55, 777–787. Barnard J.L. (1970) Sublittoral Gammaridea (Amphipoda) of the Hawaiian Islands. Smithsonian Contributions to Zoology 34, 1 –286. Corona A., Soto L.A. and Sánchez A.J. (2000) Epibenthic amphipod abundance and predation efficiency of the pink shrimp Farfantepenaeus duorarum (Burkenroad, 1939) in habitats with different physical complexity in a tropical estuarine system. Journal of Experimental Marine Biology and Ecology 253, 33–48. Barnard J.L. and Thomas J.D. (1983) A new species of Amphilochus from the gorgonian Pterogorgia anceps in the Caribbean Sea. In John P.A. (ed.) Selected papers on Crustacean. Trivandrum: The Aquarium, pp. 179–187. Barnard J.L. and Thomas J.D. (1987) New species of Neomegamphopus from tropical America (Crustacea: marine Amphipoda). Proceedings of the Biological Society of Washington 100, 147–163. Costa A. (1853) Relazione sulla memoria del Dottor Achille Costa, di ricerche su’ crostacei amfipodi del regno di Napoli. Rendiconto della Società Reale Borbonica, Accademia delle Scienze, new series 2, 167–178. Bate C.S. (1857) A synopsis of the British edriophthalmous Crustacea. Annals and Magazine of Natural History, series 2 19, 135 –152. Culpepper T.J. (1969) A taxonomic and ecological study of selected benthonic and gammarid crustaceans from the northeastern Gulf of Mexico. PhD thesis. Texas A&M University, Texas, USA. Boeck A. (1871) Crustacea Amphipoda borealia et arctica. Forhandlinger i Videnskabs-Selskabet i Christiania 1870, 81–280. Dana J.D. (1852) On the classification of the Crustacea Choristopoda or Tetradecapoda. American Journal of Science and Arts (2) 14, 297–316. Bonet F. (1967) Biogeologia subsuperficial del Arrecife Alacranes, Yucatán. Universidad Nacional Autónoma de México, Instituto de Geologı́a 80, 1 –92. Dana J.D. (1853) Crustacea. Part II. United States Exploring Expedition 14, 689–1618. Bousfield E.L. (1973) Shallow-water gammaridean Amphipoda of New England. 1st edition. Ithaca, NY: Cornell University Press. Bousfield E.L. and Hendrycks E.A. (2002) The talitroidean amphipod family Hyalidae revised, with emphasis on the North Pacific fauna: systematics and distributional ecology. Amphipacifica 3, 17–134. Bulycheva A.I. (1957) [Beach-fleas of the seas of the USSR and adjacent waters (Amphipoda–Talitroidea)]. Akademiya Nauk SSSR, Opredeliteli po Faune SSSR 65, 1 –185. Carlton J.T. and Eldredge L. (2009) Marine bioinvasions of Hawai’i: the introduced and cryptogenic marine and estuarine animals and plants of the Hawaiian archipelago. Bishop Museum Bulletins in Cultural and Environmental Studies 4, 1 –202. Della Valle A. (1893) Gammarini del Golfo di Napoli. Fauna und Flora des Golfes von Neapel 20, 1 –948. Escobar-Briones E. and Jiménez-Guadarrama E.L. (2010) Macrocrustáceos (Peracarida, Decapoda) de fondos carbonatados del sector occidental del Banco de Campeche en el sur del Golfo de México. Revista Mexicana de Biodiversidad 81, S63–S72. Foster J.M., Heard R.W. and Knott D.M. (2004) Northern range extensions for Caprella scaura Templeton, 1836 (Crustacea: Amphipoda: Caprellidae) on the Florida gulf coast and in South Carolina. Gulf and Caribbean Research 16, 65–69. Foster J.M., LeCroy S.E., Heard R.W. and Vargas R. (2009) Gammaridean Amphipods. In Wehrtmann I.S. and Cortés J. (eds) Marine biodiversity of Costa Rica, Central America. Berlin: Springer Verlag, pp. 265–274. Carlton J.T. and Hodder J. (1995) Biogeography and dispersal of coastal marine organisms: experimental studies on a replica of a 16th-century sailing vessel. Marine Biology 121, 721–730. Fox R.S. and Bynum K.H. (1975) The amphipod crustaceans of North Carolina estuarine waters. Chesapeake Science 16, 223 –237. Chapman J.W. and Carlton J.T. (1991) A test of criteria for introduced species: the global invasion by the isopod Synidotea laevidorsalis (Miers, 1881). Journal of Crustacean Biology 11, 386–400. Gable M.F., Lazo-Wasem E.A. and Baldinger A.J. (2010) The Amphipoda of Bermuda—a century of taxonomy. Zoologica Baetica 21, 131–141. Cházaro-Olvera S., Winfield I., Ortiz M. and Álvarez F. (2002) Peracarid crustaceans from three inlets in the southwestern Gulf of Mexico: new records and range extensions. Zootaxa 123, 1 –16. Gunter G. and Geyer R.A. (1955) Studies on fouling organisms of the northwest Gulf of Mexico. Publications of the Institute of Marine Science 4, 37–67. Chevreux E. (1901) Crustacés amphipodes: Mission Scientifique de M. Ch. Alluaud aux Îles Seychelles, (Mars, Avril, Mai 1892). Mémoires de la Société Zoologique de France 14, 388–438. Hughes L.E. and Lowry J.K. (2010) Establishing a neotype for Elasmopus rapax Costa, 1853 and its presence as an invasive species in temperate Australian waters. Journal of Crustacean Biology 30, 699–709. Chevreux E. and de Guerne J. (1888) Sur un amphipode nouveau (Cyrtophium chelonophilum) commensal de Thalassocheles caretta Linné. Comptes Rendus Hebdomadaires des Séances de l’Académie des Sciences 106, 625–628. Jordán-Dahlgren E. and Rodrı́guez-Martı́nez R.E. (2003) The Atlantic coral reefs of Mexico. In Cortés J. (ed.) Latin American coral reefs. Amsterdam: Elsevier Press, pp. 131–158. Coleman C.O. (2010) Batea aldebaranae sp. nov. (Crustacea: Amphipoda: Bateidae) from the coast of Belize with redescriptions of two related Just J. (2009) Ischyroceridae. In Lowry J.K. and Myers A.A. (eds) Amphipoda (Crustacea: Peracarida) of the Great Barrier Reef, Australia. Zootaxa 2260, 463–486. 13 14 carlos e. paz-ri’ os et al. Karaman G.S. (1982) Family Gammaridae. In Ruffo S. (ed.) The Amphipoda of the Mediterranean, Part 1. Gammaridea (Acanthonotozomatidae to Gammaridae). Mémoires de l’Institut Océanographique 13, 245–364. Krapp-Schickel T. (2009) Maeridae, the Ceradocus group. In Lowry J.K. and Myers A.A. (eds) Amphipoda (Crustacea: Peracarida) of the Great Barrier Reef, Australia. Zootaxa 2260, 598–642. Krapp-Schickel T. and Ruffo S. (2000) The Maera quadrimana-complex (Crustacea, Amphipoda, Melitidae) demands a new concept: Quadrimaera n. gen. (with description of three new species from Western Atlantic). Bollettino del Museo Civico di Storia Naturale di Verona 24, 193 –214. Kumpf H., Steidinger K. and Sherman K. (1999) The Gulf of Mexico Large Marine Ecosystem: assessment, sustainability, and management. Malden, MA: Blackwell Science. and Camp D.K. (eds) Gulf of Mexico: origins, waters and biota, Volume 1, Biodiversity. College Station, TX: Texas A&M University Press, pp. 941 –972. Ledoyer M. (1986) Faune mobile des herbiers de phanérogames marines (Halodule et Thalassia) de la Laguna de Términos (Mexique, Campeche). II. Les gammariens (Crustacea). Anales del Instituto de Ciencias del Mar y Limnologı́a, Universidad Nacional Autónoma de México 13, 171–200. Lewbel G.S., Howard R.L. and Gallaway B.J. (1987) Zonation of dominant fouling organism on northern Gulf of Mexico petroleum platforms. Marine Environmental Research 21, 199–224. Lewis J.B. (1992) Abundance, distribution and behavior of a commensal amphipod Stenothoe Valida Dana on the hydrocoral Millepora complanata Lamarck. Bulletin of Marine Science 51, 245–249. Kunkel B.W. (1910) The Amphipoda of Bermuda. Transactions of the Connecticut Academy of Arts and Sciences 16, 1 –126. Lin H.-J. and Shao K.-T. (2002) The development of subtidal fouling assemblages on artificial structures in Keelung Harbor, northern Taiwan. Zoological Studies 41, 170 –182. Lamarck J.B.P.A. de M. de (1816) Histoire naturelle des animaux sans vertèbres; Volume 2. Verdière: Libraire, quai des Augustins, No. 27. Lowry J.K. and Myers A.A. (2009) Amphipoda (Crustacea: Peracarida) of the Great Barrier Reef, Australia. Zootaxa 2260, 16. Latreille P.A. (1802) Histoire naturelle générale et particulière des Crustacés et des Insectes. Paris: F. Dufart. Martı́n A. and Dı́az Y.J. (2003) La fauna de anfı́podos (Crustacea: Amphipoda) en las aguas costeras de la región oriental de Venezuela. Boletı́n del Instituto Español de Oceanografı́a 19, 327–344. Latreille P.A. (1816) Amphipoda. In Nouveau Dictionaire d’histoire naturelle, appliquée aux Arts, à l’Agriculture, à l’Économie rurale et domestique, à la Médecine, etc. Par une société de Naturalistes et d’Agriculteurs. Paris, pp. 467–469. Martin T.R. and Bortone S.A. (1997) Development of an epifaunal assemblage on an estuarine artificial reef. Gulf of Mexico Science 15, 55–70. Lazo-Wasem E.A. and Gable M.F. (1987) A review of recently discovered type specimens of Bermuda Amphipoda (Crustacea: Peracarida) described by B.W. Kunkel (1882–1969). Proceedings of the Biological Society of Washington 100, 321–336. McKinney L.D. (1977) The origin and distribution of shallow water gammaridean Amphipoda in the Gulf of Mexico and Caribbean Sea with notes on their ecology. PhD thesis. Texas A&M University, Texas, USA. Leach W.F. (1814) Crustaceology. The Edinburgh Encyclopaedia 7, 402–403. McKinney L.D. (1978) Amphilochidae (Crustacea: Amphipoda) from the western Gulf of Mexico and Caribbean Sea. Gulf Research Reports 6, 137–143. LeCroy S.E. (2000) An Illustrated Identification Guide to the Nearshore Marine and Estuarine Gammaridean Amphipoda of Florida, Volume 1, Families Gammaridae, Hadziidae, Isaeidae, Melitidae and Oedicerotidae. Florida Department of Environmental Protection, Tallahassee, Contract No. WM724, 195 pp. LeCroy S.E. (2002) An Illustrated Identification Guide to the Nearshore Marine and Estuarine Gammaridean Amphipoda of Florida, Volume 2, Families Ampeliscidae, Amphilochidae, Amphithoidae, Aoridae, Argissidae and Haustoriidae. Florida Department of Environmental Protection, Tallahassee, Contract No. WM724, 214 pp. LeCroy S.E. (2004) An Illustrated Identification Guide to the Nearshore Marine and Estuarine Gammaridean Amphipoda of Florida, Volume 3, Families Bateidae, Biancolinidae, Cheluridae, Colomastigidae, Corophidae, Cyproideidae and Dexaminidae. Florida Department of Environmental Protection, Tallahassee, Contract No. WM724, 91 pp. LeCroy S.E. (2007) An Illustrated Identification Guide to the Nearshore Marine and Estuarine Gammaridean Amphipoda of Florida, Volume 4, Families Anamixidae, Eusiridae, Hyalellidae, Hyalidae, Iphimediidae, Ischyroceridae, Lysianassidae, Megaluropidae and Melphidippidae. Florida Department of Environmental Protection, Tallahassee, Contract No. WM880, 111 pp. LeCroy S.E. (2011) An Illustrated Identification Guide to the Nearshore Marine and Estuarine Gammaridean Amphipoda of Florida, Volume 5, Families Leucothoidae, Liljeborgiidae, Neomegamphopidae, Ochlesidae, Phliantidae, Phoxocephalidae, Platyischnopidae, Pleustidae, Podoceridae, Pontoporeiidae, Sebidae, Stenothoidae, Synopiidae and Talitridae. Florida Department of Environmental Protection, Tallahassee, Contract No. WM949, 209 pp. LeCroy S.E., Gasca R., Winfield I., Ortiz M. and Escobar-Briones E. (2009) Amphipoda (Crustacea) of the Gulf of Mexico. In Felder D.L. McKinney L.D. (1979) Liljeborgiid amphipods from the Gulf of Mexico and Caribbean Sea. Bulletin of Marine Science 29, 140 –159. McKinney L.D. (1980a) Four new and unusual amphipods from the Gulf of Mexico and Caribbean Sea. Proceedings of the Biological Society of Washington 93, 83–103. McKinney L.D. (1980b) The genus Photis (Crustacea: Amphipoda) from the Texas coast with the description of a new species, Photis melanicus. Contributions in Marine Science 23, 57–61. Mejı́a-Ortı́z L.M., Yañez G. and López-Mejı́a M. (2006) Fauna of five anchialine caves in Cozumel Island, México. National Association for Cave Diving Journal 39, 11–15. Milne-Edwards H. (1830) Extrait de recherches pour servir à l’histoire naturelle des crustacés amphipodes. Annales des Sciences Naturelles 20, 353–399. Montagu G. (1818) An essay on sponges, with descriptions of all the species that have been discovered on the coast of Great Britain. Memoirs of the Wernerian Natural History Society 2, 67–122. Munguia P. (2004) Successional patterns on pen shell communities at local and regional scales. Journal of Animal Ecology 73, 64–74. Munguia P., Mackie C. and Levitan D.R. (2007) The influence of stagedependent dispersal on the population dynamics of three amphipod species. Oecologia 153, 533–541. Myers A.A. (1981) Amphipod Crustacea I. Family Aoridae. Memoirs of the Hourglass Cruises 5, 1–73. Myers A.A. and McGrath D. (1984) A revision of the north-east Atlantic species of Ericthonius (Crustacea: Amphipoda). Journal of the Marine Biological Association of the United Kingdom 64, 379–400. fouling amphipods in alacranes reef Oliva-Rivera J.J. (2003) The amphipod fauna of Banco Chinchorro, Quintana Roo, Mexico with ecological notes. Bulletin of Marine Science 73, 77–89. Ruffo S. (1954) Studi sui crostacei anfipodi. XL. Nuovi anfipodi raccolti nel Venezuelan dal Prof. G. Marcuzzi. Memorie del Museo Civico di Storia Naturale di Verona 4, 117 –125. Oliva-Rivera J.J. and Jiménez-Cueto M.S. (1997) Composición, distribución y abundancia de los crustáceos peracáridos de la laguna de Yalahau, Quintana Roo. AvaCient 23, 26–31. Ruffo S., Krapp T. and Gable M.F. (2000) The genus Maera (Crustacea: Amphipoda: Melitidae) from Bermuda. Postilla 221, 3–35. Ortiz M. (1979) Contribución al estudio de los anfı́podos (Gammaridea) del Mediterráneo Americano. Revista de Investigaciones Marinas Serie 8, 1 –16. Ortiz M. (1991) Amphipod Crustacea 2, Family Bateidae. Memoirs of the Hourglass Cruises 8, 1–31. Ortiz M. (2001) Lista de invertebrados marinos, estuarinos y semiterrestres de la playa de Cojı́mar en la costa norte de la Provincia Ciudad de la Habana. Revista de Investigaciones Marinas 22, 93–102. Ruiz G.M., Fofonoff P.W., Carlton J.T., Wonham M.J. and Hines A.N. (2000) Invasion of coastal marine communities in North America: apparent patterns, processes, and biases. Annual Review of Ecology, Evolution and Systematics 31, 481–531. Schellenberg A. (1938) Litorale Amphipoden des tropischen Pazifiks nach Sammlungen von Prof. Bock (Stockholm), Prof. Dahl (Berlin) und Prof. Pietschmann (Wein). Kungliga Svenska Vetenskapsakademiens Handlingar, series 3 16, 1–105. Schellenberg A. (1939) Amphipoden des Kongo-Mündungsgebietes. Revue de Zoologie et de Botanique Africaines 32, 122–138. Ortiz M. and Lalana R. (1996) Los anfı́podos de la primera expedición conjunta Cuba–USA, a bordo del B/I Ulises, a las aguas del Archipélago Sabana-Camagüey, Cuba, en 1994. Anales del Instituto de Biologı́a. Universidad Nacional Autónoma de México, Serie Zoologı́a 67, 89–101. Schmitter-Soto J.J., Comı́n F.A., Escobar-Briones E., Herrera-Silveira J., Alcocer J., Suárez-Morales E., Elı́as-Gutiérrez M., Dı́az-Arce V., Marı́n L.E. and Steinich B. (2002) Hydrogeochemical and biological characteristics of cenotes in the Yucatan Peninsula (SE Mexico). Hydrobiologia 467, 215 –228. Ortiz M. and Lalana R. (2010) Distribución de los anfı́podos (Crustacea, Malacostraca, Peracarida) de los subórdenes Gammaridea, Caprellidea e Hyperiidea, presentes en el archipiélago cubano. Revista de Investigaciones Marinas 31, 75–90. Seaman W. and Jensen A.C. (2000) Purposes and practices of artificial reef evaluation. In Seaman W. (ed.) Artificial reef evaluation with application to natural marine habitats. Boca Raton, FL: CRC Press LLC, pp. 2 –19. Ortiz M. and Lemaitre R. (1994) Crustáceos anfı́podos (Gammaridea) colectados en las costas del Caribe colombiano, al sur de Cartagena. Anales del Instituto de Investigaciones Marinas de Punta de Betı́n 23, 119–127. Serejo C.S. (1995) Podocerus fissipes n. sp., a new species of sponge-dwelling amphipod (Amphipoda, Podoceridae) from the Brazilian coast. Nauplius 3, 49–57. Ortiz M. and Winfield I. (2012) A new commensal species of the genus Leucothoe Leach, 1814 (Amphipoda: Leucothoidae) from the Cuban Archipelago. Travaux du Muséum National d’Histoire Naturelle ‘Grigore Antipa’ 55, 17–26. Ortiz M., Martı́n A. and Dı́az Y.J. (2007) Lista y referencias de los crustáceos anfı́podos (Amphipoda: Gammaridea) del Atlántico occidental tropical. Revista de Biologı́a Tropical 55, 479–528. Paz-Rı́os C.E. and Ardisson P.-L. (2013) Benthic amphipods (Amphipoda: Gammaridea and Corophiidea) from the Mexican southeast sector of the Gulf of Mexico: checklist, new records and zoogeographic comments. Zootaxa 3635, 137–173. Pearse A.S. (1912) Notes on certain amphipods from the Gulf of Mexico, with description of new genera and new species. Proceedings of the United States National Museum 43, 369 –379. Pearse A.S. (1932) Inhabitants of certain sponges at Dry Tortugas. Papers from the Tortugas Laboratory. 7. Carnegie Institute of Washington Publication 435, 119–122. Pech D., Ardisson P.-L. and Bourget E. (2002) Settlement of a tropical marine epibenthic assemblage on artificial panels: influence of substratum heterogeneity and complexity scales. Estuarine, Coastal and Shelf Science 55, 743 –750. Serejo C.S. (1998) Gammaridean and caprellidean fauna (Crustacea) associated with the sponge Dysidea fragilis Johnston at Arraial do Cabo, Rio de Janeiro, Brazil. Bulletin of Marine Science 63, 363–385. Serejo C.S. (1999) Taxonomy and distribution of the family Hyalidae (Amphipoda, Talitroidea) on the Brazilian Coast. In Schram F.R. and von Vaupel Klein J.C. (eds) Proceedings of the Fourth International Crustacean Congress, Amsterdam, The Netherlands 20–24 July 1998. Crustaceans and the Biodiversity Crisis. Leiden, The Netherlands: Brill, pp. 591 –616. Sheridan P.F. (1980) Three new special [sic] of Melita (Crustacea: Amphipoda), with notes on the amphipod fauna of the Apalachicola estuary of northwest Florida. Northeast Gulf Science 3, 60–73. Shoemaker C.R. (1926) Amphipods of the family Bateidae in the collection of the United States National Museum. Proceedings of the United States National Museum 68, 1–26. Shoemaker C.R. (1933a) Amphipoda from Florida and the West Indies. American Museum Novitates 598, 1 –24. Shoemaker C.R. (1933b) Two new genera and six new species of Amphipoda from Tortugas. Papers from the Tortugas Laboratory 28, 247–256. Shoemaker C.R. (1956) Observations on the amphipod genus Parhyale. Proceedings of the United States National Museum 106, 345 –358. Pederson E.J. and Peterson M.S. (2002) Bryozoans as ephemeral estuarine habitat and a larval transport mechanism for mobile benthos and young fishes in the north-central Gulf of Mexico. Marine Biology 140, 935–947. Smith S.I. (1873) Crustacea, ex. Isopoda. Report upon the invertebrate animals of Vineyard Sound and adjacent waters. U.S. Commission on Fish and Fisheries, Part 1, Report on the Condition of the Sea Fisheries of the South Coast of New England in 1871 and 1872, 483 pp. Pequegnat W.E. and Pequegnat L.H. (1968) Ecological aspect of marine fouling in the northeastern Gulf of Mexico. Texas A&M University. Department of Oceanography, Texas, Reference 68-22T, 80 pp. Stebbing T.R.R. (1899) Amphipoda from the Copenhagen Museum and other sources. Part II. Transactions of the Linnean Society of London (2, Zoology) 8, 395–432. Ribeiro S.M., Omena E.P. and Muricy G. (2003) Macrofauna associated to Mycale macrostigmatosa (Porifera: Demospongiae) in Rio de Janeiro State, SE Brazil. Estuarine, Coastal and Shelf Science 57, 951–959. Stebbing T.R.R. (1906) Amphipoda I. Gammaridea. Das Tierreich 21, 1–806. Streftaris N., Zenetos A. and Papathanassiou E. (2005) Globalisation in marine ecosystems—the story of non indigenous marine species across 15 16 carlos e. paz-ri’ os et al. Europe Seas. Oceanography and Marine Biology: an Annual Review 43, 419–453. Thomas J.D. (1976) A survey of gammarid amphipods of the Barataria Bay, Louisiana region. Contribution in Marine Science 20, 87–100. Thomas J.D. (1993) Identification manual for the marine Amphipoda (Gammaridea): I. Common coral reef and rock bottom amphipods of South Florida. Florida Department of Environmental Protection, Tallahassee, Contract No. SP290, 83 pp. Thomas J.D. (1997) Systematics, ecology and phylogeny of the Anamixidae (Crustacea: Amphipoda). Records of the Australian Museum 49, 35–98. Thomas J.D. and Barnard J.L. (1987) The Indo-Pacific Audulla chelifera reported from the Caribbean Sea (Crustacea: Amphipoda). Proceedings of the Biological Society of Washington 100, 364–370. Thomas J.D. and Barnard J.L. (1992) Podocerus kleidus, new species from the Florida Keys (Crustacea, Amphipoda, Dulichiidae). Bulletin of Marine Science 51, 309–314. Thomas J.D. and Klebba K.N. (2006) Studies of commensal leucothoid amphipods, two new sponge-inhabiting species from south Florida and the western Caribbean. Journal of Crustacean Biology 26, 13–22. Winfield I. and Álvarez F. (2009) Two new species of amphipods (Peracarida, Amphipoda, Leucothoidae) from the Veracruz Coral Reef System, SW Gulf of Mexico. Crustaceana 82, 11–25. Winfield I. and Escobar-Briones E. (2007) Anfı́podos (Crustacea: Gammaridea) del sector norte del Mar Caribe: listado faunı́stico, registros nuevos y distribución espacial. Revista Mexicana de Biodiversidad 78, 51–61. Winfield I. and Ortiz M. (2009) Lysianopsis adelae, new species of amphipod (Gammaridea: Lysianassidae) from the Veracruz Coral Reef System, SW Gulf of Mexico. Zootaxa 2205, 53–61. Winfield I. and Ortiz M. (2010) Colomastigids (Amphipoda: Gammaridea: Colomastigidae) from the Veracruz Coral Reef System, SW Gulf of Mexico, with a description of two new species associated with sponges. Scientia Marina 74, 773–782. Winfield I. and Ortiz M. (2011) Crustáceos con bolsa incubadora (Crustacea: Malacostraca: Peracarida). In Hernández-Ortiz V. (ed.) La biodiversidad en Veracruz: estudio de estado. Volume II, Diversidad de especies: conocimiento actual. Cap. Invertebrados. México: CONABIO, pp. 277 –286. Thomas J.D. and Klebba K.N. (2007) New species and host associations of commensal leucothoid amphipods from coral reefs in Florida and Belize (Crustacea, Amphipoda). Zootaxa 1494, 1–44. Winfield I., Abarca-Arenas L.G. and Cházaro-Olvera S.I. (2007) Crustacean macrofoulers in the Veracruz coral reef system, SW Gulf of Mexico: checklist, spatial distribution and diversity. Cahiers de Biologie Marine 48, 287–295. Thomas J.D. and Ortiz M. (1995) Leucothoe laurensi, a new species of leucothoid amphipod from Cuban waters (Crustacea, Amphipoda, Leucothoidae). Proceedings of the Biological Society of Washington 108, 613 –616. Winfield I., Escobar-Briones E. and Morrone J.J. (2006) Updated 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. Trujillo-Pisanty D., Chávez-Solı́s E.M., Solı́s-Weiss V. and Hermoso-Salazár M. (2010) Peracarids (Crustacea: Malacostraca) from cenote Aerolito, Cozumel, Mexican Caribbean. Cahier de Biologie Marine 51, 177–180. Tunnell J.W. Jr, Chavez E.A. and Withers K. (2007a) Coral reefs of the southern Gulf of Mexico. College Station, TX: Texas A&M University Press. Tunnell J.W. Jr, Barrera N., Beaver C.R., Davidson J., Gourley J.E., Moretzsohn F., Nañez-James S., Pearce J.J. and Vega M.E. (2007b) Checklist of the biota associated with southern Gulf of Mexico coral reefs and coral reef islands. Available through: GulfBase at http:// www.gulfbase.org/checklist/ (accessed 3 August 2012). Varela C., Ortiz M. and Lalana R. (2003) Crustáceos (Peracarida y Decapoda) de la costa sur de Guanahacabibes, Cuba. Revista de Investigaciones Marinas 24, 73–76. Walker A.O. (1904) Report on the Amphipoda collected by Professor Herdman, at Ceylon, in 1902. Report to the Government of Ceylon on the Pearl Oyster Fisheries of the Gulf of Mannar, Supplementary Report, no. 17, 71 pp. White K.N. (2011a) A taxonomic review of the Leucothoidae (Crustacea: Amphipoda). Zootaxa 3078, 1–113. White K.N. (2011b) Caribbean Leucothoidae (Crustacea: Amphipoda) of Panama. Gulf and Caribbean Research 23, 23–35. Wilkinson T., Wiken E., Bezaury-Creel J., Hourigan T., Agardy T., Herrmann H., Janishevski L., Madden C., Morgan L. and Padilla M. (2009) Marine ecoregions of North America. Montreal: Commission for Environmental Cooperation. Winfield I., Ortiz M. and Cházaro-Olvera S. (2009a) A new sponge-inhabiting amphipod species (Crustacea, Gammaridea, Sebidae) from the Veracruz Coral Reef System, southwestern Gulf of Mexico. Organisms, Diversity & Evolution 9, 72.e1–72.e8. Winfield I., Ortiz M. and Cházaro-Olvera S. (2009b) Especie nueva de anfı́podo comensal (Amphipoda: Gammaridea: Leucothoidae) del Parque Nacional Sistema Arrecifal Veracruzano, SO del golfo de México. Revista Mexicana de Biodiversidad 80, 315–321. Winfield I., Cházaro-Olvera S., Horta-Puga G., Lozano-Aburto M.A. and Arenas-Fuentes V. (2010) Macrocrustáceos incrustantes en el Parque Nacional Sistema Arrecifal Veracruzano: biodiversidad, abundancia y distribución. Revista Mexicana de Biodiversidad 80, S165– S175. and Winfield I., Cházaro-Olvera S., Ortiz M. and Palomo-Aguayo U. (2011) Lista actualizada de las especies de anfı́podos (Peracarida: Gammaridea y Corophiidea) marinos invasores en México. Revista de Biologı́a Marina y Oceanografı́a 46, 349–361. Correspondence should be addressed to: C.E. Paz-Rı́os Departamento de Recursos del Mar Cinvestav, Carretera antigua a Progreso, km 6 Apartado Postal 73, Cordemex 97310 Merida, Yucatan, Mexico email: cpaz@mda.cinvestav.mx