Check List ‘23. ae ie 7% vv, ANNOTATED LIST OF SPECIES Check List 14 (2): 479-494 BS https://doi.org/10.15560/14.2.479 PENSUFT. Updated checklist of estuarine caridean shrimps (Decapoda: Caridea) from the southern region of Laguna Madre, Tamaulipas, Mexico, with new records and a key for taxonomic identification Hiram Herrera-Barquin,' Antonio Leija-Tristan,' Susana Favela-Lara’ 1 Laboratorio de Ecologia Pesquera, Laboratorio de Ecologia Molecular, Departamento de Ecologia, Facultad de Ciencias Bioldgicas, Universidad Autonoma de Nuevo Leon, Av. Pedro de Alba s/n cruz con Av. Manuel L. Barragan, 66451, San Nicolas de los Garza, Nuevo Leon, Mexico. 2 Laboratorio de Ecologia Molecular, Departamento de Ecologia, Facultad de Ciencias Biolégicas, Universidad Autonoma de Nuevo Leon, Av. Pedro de Alba s/n cruz con Av. Manuel L. Barragan, 66451, San Nicolas de los Garza, Nuevo Leon, Mexico. Corresponding author: Hiram Herrera-Barquin, hiram.division@gmail.com Abstract We provide an updated list of the caridean shrimp species from the southern region of the Laguna Madre, Tamauli- pas, Mexico, along with a key for taxonomic identification. The survey was conducted in 3 sites during 3 temporal seasons. A total of 2,989 specimens were collected belonging to 12 species, 6 genera, and to the following 4 families: Alpheidae, Hippolytidae, Palaemonidae, and Processidae. Hippolytidae was the most abundant family, followed by Palaemonidae, Alpheidae, and Processidae. The hippolytid Hippolyte obliquimanus Dana, 1852, the palaemonids Palaemon floridanus Chace, 1942, and P. northropi (Rankin, 1898), and the alpheid A/pheus cf. packardii Kingsley, 1880 represent new records for the Laguna Madre and selected areas of the Gulf of Mexico. Key words Estuary; crustacean; hypersaline; new record; Neartic; Neotropical. Academic editor: Gianna Innocenti | Received 21 February 2018 | Accepted 30 March 2018 | Published 27 April 2018 Citation: Herrera-Barquin H, Leija-Tristan A, Favela-Lara S (2018) Updated checklist of estuarine caridean shrimps (Decapoda: Caridea) from the southern region of Laguna Madre, Tamaulipas, Mexico, with new records and a key for taxonomic identification. Check List 14 (2): 479-494. https://doi.org/10.15560/14.2.479 Introduction tions and contributing to the maintenance of submerged aquatic vegetation (SAV) habitats (Barba-Macias 2012). The Decapoda Latreille, 1802 represent one of the most ; The Laguna Madre of Tamaulipas 1s the largest coastal diverse orders within the crustaceans. With almost 3,500 ; species described, caridean shrimps constitute the most '@800n in Mexico (Contreras and Castafieda 2004), and diverse group of shrimp-like crustaceans (De Grave and together with Laguna Madre of Texas, form the largest Fransen 2011). They have ecological relevance due to hypersaline system in the world (Tunnell and Judd 2002). their presence in a great variety of habitats (Bauer 2004), _ It 1s placed under the influence of 2 biogeographic regions In coastal estuaries, they represent a numerically abun- and 2 marine provinces, making it a dynamic, wide eco- dant component, playing an important role as links that tone (Escobar-Briones 2004, Aubriot et al. 2005) that transfer energy to higher levels of the food web (Almeida supports a rich variety of organisms from both freshwater et al. 2013), recycling nutrients through fecal deposi- and marine environments. It also has an estuarine biota Copyright Herrera-Barquin et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 480 98°48"W 97°48'W 24°33'N 26°54'N 26°54'N 25°54'N 25°54'N 24°30'N 24°54'N 24°54'N 24°27'N 23°54'N 23°54'N 22°54'N 22°54'N 24°24'N } 20 LJ 20 40 60 80km Sa 99°48'W 98°48'W 97°48'W 97°50'W 97°50'W Check List 14 (2) 97°47'W 97°44'W 97°41'W 24°33'N 24°30'N 24°27'N i Sampling points 24°24'N 97°47'W 97°44'W Figure 1. Sampling points considered for this work in the southern region of Laguna Madre, Tamaulipas, Mexico. (Contreras and Castafieda 2004) which use the great extent of habitats for shelter, feeding, and as nursing areas. Among the faunistic groups occurring in Laguna Madre of Tamaulipas, crustaceans account for a total of 96 species (Leija-Tristan et al. 2000); regarding caridean shrimp community Leija-Tristan et al. (2000) found 14 species in the Mexican basin, and most of the subsequent surveys recorded 9 (Barba-Macias 1999, Rodriguez- Almaraz et al. 2000, Barba-Macias et al. 2005) and 7 species (Barba-Macias 2012). All of these studies found the Alpheidae, Hippolytidae, and Palaemonidae to be the 3 main families, with occasional occurrences of Proces- sidae, and underlined the strong association between this decapod group and SAV habitats. In Laguna Madre, SAV is the most important primary producer (Rendon Von-Osten and Garcia-Guzman 1995), and greatly determines the abundance and richness of estuarine benthic biota (Barba-Macias et al. 2005). This occurs due to the spatial complexity provided by these vegetated areas, compared with bare substrates which, instead, do not serve either as foraging or sheltering grounds against predators (Llanso et al. 1998). Knowledge about caridean shrimps in the study area has been less updated than that regarding the structure and ecology of edible crustaceans. Furthermore, the taxonomy of some groups is complex or under revision (Roman-Contreras and Martinez-Mayén 2010, Vera- Caripe et al. 2012, Almeida et al. 2012), resulting in scattered knowledge. Our main objective is to provide an updated list of the caridean shrimps occurring in the southern area of Laguna Madre of Tamaulipas, as well as a key for taxonomic identification of the species. Methods Study area. The Laguna Madre is situated in the coastal plain of the Gulf of Mexico, in the states of Tamaulipas, Mexico and Texas, USA (Rendon-von Osten and Garcia- Guzman 1995). On the Mexican side, it has a shoreline of approximately 160 km long from Rio Bravo delta in the north to Rio Soto la Marina at south, covering an area of 272,844.6 ha (Carrera 2004). The climate in the southern region is hot and semiarid; the mean annual temperature is 24.3 °C (17 °C in January and 29.6 °C in June); the mean annual precipitation is 748 mm, with the rainy season ranging from June to October (Tunnel and Judd 2002). The hydrology of the system is characterized by a mean salinity of 41.2%o0 (33.5—-63), a temperature of 25.9 °C (Contreras and Castafieda 2004), an average dissolved oxygen of 4.5-6.3 mg/L, and a pH of 8.4-9.0 (Leija-Tristan 2005). The poor drainage from the land into the lagoon pro- vides several components involved in the natural bio- geochemical cycles, but also transports substances such as pesticides, heavy metals, agricultural fertilizers and domestic waste, derived mainly from anthropogenic activities around the basin (Bello-Pineda et al. 2009). Our work was carried out in the southern region of Laguna Madre, at 3 sampling stations (Table 1; Fig. 1), adjacent to the inlet Boca de Catan and the neighboring Table 1. Collection sites in the southern region of Laguna Madre, Tamaulipas, Mexico. Station Latitude (N) Longitude (W) 12ST) 24°29.1000' 097°41.4500’ 2 (S2) 24°29.1833' 097°41.9667' 3 ($3) 24°29.1000' 097°41.4500’ Herrera-Barquin et al. | Caridean shrimps from Laguna Madre, Tamaulipas 481 town of Punta de Piedra, Tamaulipas (Fig. 1). Station 1 (S1) is located nearby an inlet, it is influenced by daily tidal regimes and presence of oyster reefs. Station 2 (S2) is adjacent to a deep tidal channel, over a flooded sedi- ment flat covered with seagrass. Station 3 (S3) is located 2 km west to Punta Piedra town, at the edge of an internal basin called Bahia de Catan. This area was selected for its fisheries activities and the presence of seagrass meadows, which are habitats for the caridean shrimps. Sampling methods. The collections were made in 3 seasons—March (dry season) and August 2014 (rainy season), and February 2017 (northern winds season)}— at each of the 3 sites, in order to cover a whole year of seasonal variations. At each site, the sampling procedure consisted of 5 separate points along 2 linear transects of 23 m each Over Seagrass-covered substrates. Caridean shrimps were collected by sieving over the seagrass leaves, removing incidental sediments and vegetation, selecting the speci- mens with tweezers. Specimens were stored in a jar with salt water and later preserved in 90% isopropyl alcohol. Additionally, seagrass samples were randomly collected within the same sampling area, stored in plastic bags and preserved in ice for processing in the laboratory. The species were identified according to Holthuis (1950, 1952), Chace (1972), Williams (1984), Abele and Kim (1986) and Hernandez et al. (2005). A ZEISS Stemi DV4 stereoscopic microscope and a LEICA EZ4HD with added camera were used for specimen observation and photography, respectively. The families, as well as the species and their syn- onyms are listed following the criteria of De Grave and Fransen (2011), and the taxa within the families are ordered alphabetically. Additional information, such as distribution or morphological observations is summa- rized from the above-mentioned literature. The collected specimens were deposited in the Carcinological Collec- tion of the Biological Sciences Faculty of Autonomous University of Nuevo Leén (UANL-FCB-C). Vouchers for each record are presented in the Results. Seagrass were processed by washing them to remove the sediment from the leaves. The seagrass species were determined by observing vegetative structures such as shoots and leaves (Dawes et al. 2004). Results A total of 2,989 shrimp specimens were identified belong- ing to 4 families, 6 genera and 12 species. The seagrass samples identification showed an integrated community of 3 species: Syringodium filiforme Kutz., Halodule wrightii Asch. and Thalassia testudinum Banks & Sol. ex K.D. Koenig. Superfamily Palaemonoidea Rafinesque, 1815 Family Palaemonidae Rafinesque, 1815 Leander tenuicornis (Say, 1818) Material examined (16 specimens). 5 August 2014 (UANL-FCB-C17-8139), S2 (24°29.1833' N, 097° 41.9667' W), among seagrass of Syringodium filiforme and Halodule wrightii, 3 specimens (29, 1¢); 5 August 2014 (UANL-FCB-C17-8149), S3 (24°29.2667' N 097°45.9833' W), same habitat, 13 specimens (89, 5<). p) Distribution. Western Atlantic Ocean: Canada, USA (Massachusetts, Virginia, North Carolina, South Caro- lina, Florida, Louisiana and Texas), Bermudas, Mexico, Panama, Bahamas, Cuba, Jamaica, Puerto Rico, Virgin Islands, Colombia, Venezuela, Brazil, Malvinas Islands. Oriental Atlantic Ocean: Azores Archipelago. Mediter- ranean: Spain, France, Italy and Libya. Pacific and Indian oceans: Red Sea, India, Japan, Papua New Guinea, Aus- tralia and New Zealand (Ferreira et al. 2010). Previous records from Laguna Madre. Leija-Tristan et al. (2000); Rodriguez et al. (2000). Remarks. The morphology of our specimens agrees with the description provided by Abele and Kim (1986). Specimens were more easily identified by observing the stylocerite reaching to distal third of basal antennular segment and the deep rostrum of females, rather than the shallow one showed by males. Palaemon floridanus Chace, 1942 Figure 2A, B Material examined (46 specimens). Among seagrass meadows. 5 August 2014 (UANL-FCB-C17-8143), S2 (24°29.1833' N, 097°41.9667' W), 11 specimens (99, 24); 5 August 2014 (UANL-FCB-C17-8147), S3 (24°29.2667' N, 097°45.9833' W), same habitat, 35 specimens (162, 19); 18 February 2017 (UANL-FCB- C17-8161), S3 (same previous point), 1 specimen (<3). Distribution. Florida, Panama, Belize, Texas, Mexico (Laguna Madre, Tamaulipas [this study, new record]) (Chace 1972; Holthuis 1952: Coen et al. 1981; Strenth and Chace 1995; Baeza and Fuentes 2012). Previous records from Laguna Madre: None. New record. Remarks. This species is closely related with Palaemon northropi (Rankin, 1898), from which it was differenti- ated by Chace (1942) by the shape and dentition of the rostrum, the second legs, which have distinctly longer fingers, and the more slender and longer dactyli of the third pereopods (Holthuis 1952). Nevertheless, the most consistent identification character is the shape of the rostrum, which is slender and possesses more teeth on the ventral margin than in P. northropi (Holthuis 1952) (Fig. 2A). Our specimens typically showed 5 teeth on the ventral margin of rostrum. 482 Check List 14 (2) Figure 2. A, B. Palaemon floridanus, male (UANL-FCB-C17-8147): (A) anterior region lateral view; (B) mandible lateral view. C, D. Palaemon northropi, male (UANL-FCB-C17-8142): (C) anterior region lateral view; (D) mandible lateral view. Scale bars = 1 mm. (ip: incisor process; mp: mandibular process; p: palp). Palaemon northropi (Rankin, 1898) Figure 2C, D Material examined (8 specimens). Among seagrass beds. 5 August 2014 (UANL-FCB-C17-8142), S2 (24° 29.1833’ N, 097°41.9667' W), 5 specimens (29, 34); 5 August 2014 (UANL-FCB-C17-8146), S3 (24°29.2667' N, 097°45.9833' W), 3 specimens (19, 2<3). Distribution. Eastern American coastal regions from Bermuda to Brazil (Holthuis 1952; Ferreira et al. 2010); Mexico: Bahia de Chetumal (Castellanos-Osorio 2009), Laguna de Términos, Campeche; Bahia de la Ascencion, Quintana Roo (Chace 1972, Roman-Contreras 1988, Wicksten 2005a); Laguna Madre, Tamaulipas (this study [new record]). Previous records from Laguna Madre. None. New record. Remarks. This species can be confused with Palaemon floridanus, from which it differs by bearing 3 or 4 ventral teeth in the rostrum (Fig. 2C). Palaemon mundusnovus De Grave & Ashelby, 2013 Figure 3A, B Material examined (150 specimens). Among seagrass. 8 March 2014 (UANL-FCB-C17-8119), S1 (24°29.1000' N, 097°41.4500' W), 49 specimens (59, 44); 8 March 2014 (UANL-FCB-C17-8123), S2 (24°29.1833' N, 097°41.9667' W), 1 specimen ('); 9 March 2014 (UANL- FCB-C17-8130), S3 (24°29.2667' N, 097°45.9833' W), 14 specimens (59, 93’); 4 August 2014 (UANL-FCB- C17-8133), S1 (same point), 86 specimens (309, 57); 18 February 2017 (UANL-FCB-C17-8162), S3 (same point), 43 specimens (299, 14). Distribution. Vineyard Sound, Massachusetts, to Port Aransas, Texas (Holthuis 1952); Laguna de Mecoacan, Tabasco; Laguna de Términos, Campeche; Bahia de la Ascension, Quintana Roo; Isla Arenas, Yucatan, México (Chace 1972, Williams 1984, Roman-Contreras 1988, Dominguez et al. 2003, Barba-Macias et al. 2005, Wick- sten 2005a, Barba-Macias 2012). Previous records from Laguna Madre. Hildebrand (1958), Barba-Macias (1999), Leija-Tristan et al. (2000), Rodriguez-Almaraz et al. (2000), Sheridan and Minello (2003, Laguna Madre of Texas), Barba-Macias et al. (2005), Barba-Macias (2012). Remarks. The rostrum bears 4 or 5 ventral teeth, seldom 3; our material typically showed 4 ventral teeth, ranging Herrera-Barquin et al. | Caridean shrimps from Laguna Madre, Tamaulipas 483 A Figure 3. A, B. Palaemon mundusnovus, female (UANL-FCB-C17-8133) : (A) anterior region lateral view; (B) left chela lateral view. C, D. Palae- mon pugio, male (UANL-FCB-C17-8134): (€) anterior region lateral view; (D) left chela lateral view. E, F. Palaemon vulgaris, female (UANL-FCB- C17-8122): (E) anterior region lateral view; (F) left chela lateral view. Scale bars = 1 mm. (ft: finger tooth; mft: movable finger tooth). from 3 to 5 (Fig. 3A), and agreed with descriptions given by Abele and Kim (1986), and also Holthuis (1952), particularly in the number of ventral teeth of rostrum, the dorsal teeth behind the margin of orbit, and also the presence of a blunt and tiny tooth on the dactylus of second pereopod. This species is reported as Palaemon- etes intermedius in all the literature, but following the transfer of P. intermedius (Holthuis, 1949) to the genus Palaemon, the name became the junior homonym of Palaemon intermedius Stimpson, 1860. Thus, De Grave and Ashelby (2013) proposed the replacement name now in use. All members of the genus Palaemonetes were transferred to Palaemon by these authors based on morphological, cladistic and genetic evidence (Pereira 1997, Ashelby et al. 2012, De Grave and Ashelby 2013), concluding that the presence/absence of mandibular palp can no longer be used as the sole character that separates both genera. Furthermore, larval development studies comparing 3 coastal Palaemonetes species with 3 species of Palaemon concluded that the differences within the genera are more pronounced that between genera, which suggests a close evolutionary relationship (Knowlton and Vargo 2004). A84 Palaemon pugio (Holthuis, 1949) Figure 3C, D Material examined (22 specimens). Among seagrass beds. (UANL-FCB-C17-8120), S1 (24°29.1000' N, 097° 41.4500’ W), 1 specimen (2); 8 March 2014 (UANL- FCB-C17-8125), S2 (24°29.1833' N, 097°41.9667' W), 1 specimen (3); 4 August 2014 (UANL-FCB-C 17-8134), Sl (same point), 17 specimens (62, 114); 5 August 2014 (UANL-FCB-C17-8145), S3(24°29.2667' N, 097° 45.9833’ W), 3 specimens (2); 18 February 2017 (UANL- FCB-C17-8163), S3 (same point), 11 specimens (79, 4). Distribution. Quebec; near Yarmouth, Nova Scotia; Newcastle and East Brunswick, Maine; northern Massa- chusetts to Texas; intermittent from Verte River, 3 miles to the west of St Modeste (Holthuis 1952, Williams 1974, 1984, Abele and Kim 1986); islands and coastal lagoons from Veracruz to Campeche; Puerto Progreso, Yucatan; Akumal, Quintana Roo (Zarur 1962, Roman-Contreras 1988, Rodriguez et al. 2000, Dominguez et al. 2003, Barba-Macias et al. 2005, Barba-Macias 2012). Previous records from Laguna Madre. Hildebrand (1958, 1969), Roman Contreras (1988), Barba-Macias (1999), Rodriguez et al. (2000), Leija-Tristan et al. (2000), Sheridan and Minello (2003), Barba-Macias et al. (2005), Barba-Macias (2012). Remarks. This species is recognized by having an unarmed stretch in both sides of the rostrum before a dagger-shaped tip, and 2—5, generally 3, ventral teeth. The fingers of second pereiopod are without teeth on their cutting edges (Abele and Kim 1986) (Fig. 3C, D). Our material agreed with the previous features. Palaemon vulgaris Say, 1818 Figure 3E, F Material examined (187 specimens). In seagrass beds, tidal zones and presence of oyster reefs. 8 March 2014 (UANL-FCB-C17-8118), S1 (24°29.1000' N, 097° 41.4500’ W), 1 specimen (3); 8 March 2014 (UANL- FCB-C17-8122), S2 (24°29.1833' N, 097°41.9667' W), 96 specimens (629, 34), 4 August 2014 (UANL-FCB- C17-8135), S1 (same point), 15 specimens (9); 5 August 2014 (UANL-FCB-C17-8141), S2 (same point), 75 specimens (402, 354); 17 February 2017 (UANL-FCB- C17-8154), S1(same point), 104 specimens (459, 59); 17 February 2017 (UANL-FCB-C17-8157), S2 (same point), 64 specimens (309, 34). Distribution. Southern Gulf of St Lawrence, south- ward to Cameron County, Texas; Laguna de Tamiahua, Veracruz; Laguna Mecoacan, Tabasco; Rio Champoton, Laguna de Términos, Campeche; near Progreso, Yucatan; Puerto Morelos, Punta Hualapich and Akumal, Quintana Roo (Holthuis 1952, Williams 1984, Abele and Kim 1986, Roman-Contreras 1988, Barba-Macias et al. 2005, Barba- Macias 2012). Check List 14 (2) Previous records from Laguna Madre. Barba-Macias (1999), Sheridan and Minello (2003), Sheridan (2004, Laguna Madre of Texas), Barba-Macias et al. (2005), Barba-Macias (2012). Remarks. The rostrum has 2 teeth on dorsal series behind posterior margin of orbit, 3—5 ventral teeth, the dactylus of the second pereopod is armed with 2 teeth and the immovable finger with 1 tooth on its cutting edge. Our material agreed with descriptions given by Holthuis (1952), Williams (1984), and Abele and Kim (1986). This species was also part of the re-appraisal of the genus Palaemonetes made by De Grave and Ashelby (2013). Superfamily Alpheoidea Rafinesque, 1815 Family Alpheidae Rafinesque, 1815 Alpheus heterochaelis Say, 1818 Figure 4A—C Material examined (9 specimens). In seagrass beds. 8 March 2014 (UANL-FCB-C19-8127), S2 (24°29.1833' N, 097°41.9667' W), 5 specimens (49, 1); 4 August 2014 (UANL-FCB-C19-8136), S$1(24°29.1000' N, 097°41.4500' W), 3 specimens (12, 2); 5 August 2014 (UANL-FCB-C19-8144), S3 (24°29.2667' N, 097°45.9833' W), 1 specimen (4); 17 February 2017 (UANL-FCB-C19-8151), S1 (same point), 5 specimens (42, 13); 18 February 2017 (UANL-FCB-C19-8158), S3 (same point), 2 specimens (@). Distribution. Lower portion of Chesapeake Bay (USA) to South and West, through the Gulf of Mexico, Cuba, Curacao, Bermuda, Suriname and Brazil (Verrill 1922, Chace 1972, Williams 1984, McClure 2005); in Mexico, from Veracruz to Quintana Roo (Roman-Contreras 1988, Markham et al. 1990, Hermoso-Salazar and Martinez- Guzman 1991, Hernandez et al. 1996, Rodriguez et al. 2000). Previous records from Laguna Madre. Barba-Macias (1999); Rodriguez et al. (2000); Leija-Tristan et al. (2000); Sheridan and Minello (2003); Sheridan (2004); Barba-Macias et al. (2005). Remarks. Our material agreed with descriptions by Chace (1972) and McClure (2005) regarding male pro- podus of major first chelae, showing deep notches on both ventral and dorsal margins and the merus of first pereopods unarmed distoventrally (Figure 4B, C). Alpheus cf. packardii Kingsley, 1880 Figure 4D—F Material examined (25 specimens) Among seagrass meadows, in shallow waters adjacent to deeper areas. 8 March 2014 (UANL-FCB-C19-8126), S2 (24°29.1833' N, 097°41.9667' W), 14 specimens (72, 73); 9 March 2014 (UANL-FCB-C19-8131), S3 (24°29.2667' N, 097° 45.9833' W), 11 specimens (49, 7’). Herrera-Barquin et al. | Caridean shrimps from Laguna Madre, Tamaulipas 485 cp | | i es wath a mvs pr i nial all | Figure 4. A-C. Alpheus heterochaelis: (A) female (UANL-FCB-C19-8151), anterior margin lateral view; (B) male (UANL-FCB-C19-8144), first left pereiopod ventral view ; (C) female (UANL-FCB-C19-8127), major chela lateral view. D-F. Alpheus cf. packardii: (D) female (UANL-FCB- C19-8126), anterior margin lateral view; (E) first left pereiopod ventral view; (F) female (UANL-FCB-C19-8126), major chela lateral view. Scale bars = 1 mm. (cp: carpus; dt: dactylus; ot: overhanging tooth; pr: propodus; cvn: chela ventral notch). Distribution. Western Atlantic: Bermuda; North Carolina to Florida; Gulf of Mexico; throughout the Caribbean Sea; Brazil (Anker et al. 2016). In Mexico: Bahia de la Ascension, Bahia del Espiritu Santo, Arrecife Mahahual (Roman-Contreras and Martinez-Mayén 2010); Isla Verde, Artrecife Hornos, Isla de Sacrificios, Isla de Enmedio, Veracruz (Hermoso-Salazar and Arvizu-Coyotzi 2015); Laguna Madre, Tamaulipas (this study [new record]). Previous records from Laguna Madre. None. New record. Remarks. Kingsley (1878) described A/pheus normanni based on material from the Pacific coast of Panama. Two years later, he described A. packardii based on specimens from Key West, Florida (Kingsley 1880). Chace (1937) considered both as identical morphologically, and placed A. packardii as a synonym of A. normanni. Several authors continued treating these species as synonyms or only reported A. normanni for the western Atlantic (Wil- liams 1965, Chace 1972, Christoffersen 1979, Williams 1984, Abele and Kim 1986, McClure 2005). Neverthe- less, Kim and Abele (1988) compared material from 486 the eastern Pacific and western Atlantic (Florida), and found consistent morphological differences in the form of the male minor chelae, being more elongated in the Pacific specimens (5.8 times as long as broad); thus, they removed A. packardii from the synonymy of A. normanni. The minor chelae of the males we examined ranges from 3.17-4.67 times as long as broad, which overlaps the range of 3.64—4.38 described by Roman- Contreras and Martinez-Mayén (2010) as an argument to conclude their material belongs to A. packardii species complex. Moreover, our material is morphologically similar to specimens identified also as A. cf. packardii or A. packardii (Soledade and Almeida 2013, Giraldes and Freire 2015, Anker et al. 2016, see figures). Additionally, according to Christoffersen (1998), the western Atlantic Specimens previously named as A. normanni must be attributed to A. packardii. Regardless the above, both species are part of the transisthmian A. normanni—A. packardii species complex that is currently being revised (Anker et al. 2016, Anker and Santos unpubl.). Species complexes are relatively common within highly species-rich caridean taxa like the genus Alpheus, which comprises many species with very similar morphology, even in the presence of high genetic or protein divergence (Knowlton et al. 1993, McClure and Greenbaum 1994, Knowlton and Weight 1998, Mathews et al. 2002). This has led to nomenclatural confusion, and the taxonomic identity of the western Atlantic material identified as A. normanni will need to be carefully re- assessed since it is quite possible that it refers to more than 1 species (Anker et al. 2016). The presence of several cryptic taxa within the genus A/pheus from eastern Pacific and western Atlantic (Williams et al. 2001), the molecular, morphological, coloration patterns and distributional evidence show- ing that A. normanni and A. packardii are different, as well as the existence of at least 5 undescribed cryptic species belonging to normanni—packardii complex (3 in western Atlantic and 2 in the eastern Pacific) (Almeida Figure 5. Hippolyte obliquimanus, male (UANL-FCB-C20-8137), anterior region with cephalic appendages, lateral view. Scale bar = 1 mm. (fas: first antennular peduncle segment; dls: distolateral spines; ant: antennule; rc: rostral carina). Check List 14 (2) et al. 2007, Roman-Contreras and Martinez-Mayen 2010, Vera-Caripe et al. 2012, A. Anker pers. com.) makes the taxonomy unsettled for both, but the establishment of sev- eral separate species is the likely outcome. Thus, in this work we identified the collected specimens as Alpheus cf. packardii for the aforesaid considerations, evidence and taxonomic issues, and because recent studies have used the name A. packardii for records from the Gulf of Mexico (Roman-Contreras and Martinez-Mayén 2010, Hermoso-Salazar and Arvizu-Coyotzi 2015), the Carib- bean Sea (Vera-Caripe et al. 2012) and Brazil (Souza et al. 2011; Santos et al. 2012, Soledade and Almeida 2013 and all previous records mentioned there in, Giraldes and Freire 2015, Anker et al. 2016). Family Hippolytidae Spence-Bate, 1888 Hippolyte obliquimanus Dana, 1852 Figure 5 Material examined. | specimen, heavily damaged, 4. On seagrass beds. 5 August 2014 (UANL-FCB-C20-8137), S2 (24°29.1833'N, 097°41.9667' W). Distribution. North Carolina, south to Florida, through the Caribbean to Brazil (d’?Udekem d’Acoz 1997); in Mexico from Laguna Madre, Tamaulipas (this study [new record]) to Quintana Roo (Escobar 1984, Roman- Contreras 1988, Hernandez et al. 1996, Rodriguez et al. 2000, Roman-Contreras and Martinez-Mayén 2010, Hermoso-Salazar and Arvizu-Coyotzi 2015). Previous records from Laguna Madre. None. New record. Remarks. Previous records for the Gulf of Mexico in USA are inexistent, except those from Florida. Hippolyte curacaoensis 1s considered a junior synonym of H. obliquimanus (d’Udekem d’Acoz 1997). It differs from its closest relative reported for the area, H. zostericola, by having a rostrum usually armed with 3 or 4 strong teeth on dorsal margin, a strong lateral carina in proximal third of length, and the basal segment of antennular peduncle armed with 1-3 strong distolateral spines (Figs. 5, 6A, B) (Abele and Kim 1986; d’Udekem d’Acoz 1997). All these previous features were observed in the collected material. Hippolyte zostericola (Smith, 1873) Figure 6A, B Material examined (1825 specimens). On seagrass meadows of Syringodium filiforme and Halodule wrightii. 8 March 2014 (UANL-FCB-C20-8117), S1(24°29.1000' N, 097°41.4500' W), 294 specimens (2409, 54); 8 March 2014 (UANL-FCB-C20-8124), S2 (24°29.1833' N, 097°41.9667' W), 312 specimens (30792, 5); 9 March 2014 (UANL-FCB-C20-8129), S3 (24°29.2667' N, 097°45.9833' W), 64 specimens (Q°); 5 August 2014 Herrera-Barquin et al. | Caridean shrimps from Laguna Madre, Tamaulipas 487 fas Am - Figure 6. Hippolyte zostericola, female (UANL-FCB-C20-8124): (A) rostrum, lateral view; (B) antennular peduncle, dorsal view. Scale bars = 1 mm. (r: rostrum; fas: first antennular peduncle segment). (UANL-FCB-C20-8140), S2 (same point), 21 specimens (2); 5 August 2014 (UANL-FCB-C20-8148), S3 (same point), 8 specimens (79, 14); 17 February 2017 (UANL- FCB-C20-8152), S1 (same point), 248 specimens (1709, 783), 17 February 2017 (UANL-FCB-C20-8155), S2 (same point), 856 specimens (5999, 255); 18 Febru- ary 2017 (UANL-FCB-C20-8159), S3 (same point), 22 specimens (219, 13). Distribution. From Massachusetts, USA, and Bermuda, south to Florida, Trinidad and Curagao (Chace 1972, Markham and McDermott 1980, Zupo and Nelson 1999). Through Gulf of Mexico from Redfish Bay Texas to Quintana Roo (Chace 1972, Wicksten 2005b, Roman- Contreras 1988, Markham et al. 1990, Roman-Contreras and Romero-Rodriguez 2005, Roman-Contreras and Martinez-Mayén 2010). Also present for eastern Pacific in San Antonio, Robles municipality, Tumaco, western Colombia (Wicksten 1989, Lemaitre and Alvarez-Leon 1992). Previous records from Laguna Madre. Hildebrand (1958), Barba-Macias (1999), Lower Laguna Madre, Texas (Sheridan and Minello 2003), Laguna Madre of Texas, Corpus Christi Bay (Sheridan 2004), Barba- Macias et al. (2005), Barba-Macias (2012). Tozeuma carolinense Kingsley, 1878 Material examined (415 specimens). In seagrass beds. 8 March 2014 (UANL-FCB-C20-8121), S1(24°29.1000' N, 097°41.4500' W), 2 specimens (2 ); 8 March 2014 (UANL- FCB-C20-8128), S2 (24°29.1833’ N, 097° 41.9667’ W), 35 specimens (149, 21); 9 March 2014 (UANL-FCB- C20-8132), S3 (24°29.2667' N, 097° 45.9833’ W), 67 specimens (339, 34); 5 August 2015 (UANL-FCB- C20-8138), S2 (same point), 73 specimens (449, 29); 5 August 2014 (UANL-FCB-C20-8150), S3 (same point), 238 specimens (1939; 45); 17 February 2017 (UANL- FCB-C20-8156), S2 (same point), 4 specimens (29, 2); 18 February 2017 (UANL-FCB-C20-8160), S3 (same point), 14 specimens (129, 2¢). Distribution. From Massachusetts, USA, and Bermuda, to Sao Paulo, Brazil including Gulf of Mexico, Cuba, Isla Santa Lucia and Curacao (Chace 1972, Markham and McDermott 1980, Williams 1984, Hernandez et al. 1996, Martinez-Iglesias et al. 1996, Christoffersen 1998); in Mexico from Tamaulipas (Laguna Madre, see references below) to Quintana Roo (Chace 1972, Roman-Contreras 1988, Markham et al. 1990, Hernandez et al. 1996). Previous records from Laguna Madre. Hildebrand (1958); Barba-Macias (1999); Rodriguez et al. (2000); as Tozeuma carolinensis by Leija-Tristan et al. (2000); in Laguna Madre of Texas by Sheridan and Minello (2003); Corpus Christi Bay and adjacent zones (Sheridan 2004); Barba-Macias et al. (2005); Barba-Macias (2012). Remarks. Our material agreed with the description in Abele and Kim (1986). This species 1s recognized by its elongated and dorsally unarmed rostrum. Superfamily Processoidea Ortmann, 1896 Family Processidae Ortmann, 1896 Ambidexter symmetricus Manning & Chace, 1971 Material examined. 8 March 2014 (UANL-FCB-C23- 8164), S1 (24°29.1000' N, 097°41.4500' W), 1 specimen (2); in seagrass meadows of Syringodium filiforme and Halodule wrightii. Distribution. Gulf of Mexico, to Trinidad (Chace 1972); western Atlantic from Florida to Brazil (Christoffersen 1998); Tamaulipas, Mexico; Louisiana, Florida, Puerto Rico, Trinidad (Abele 1972). Previous records from Laguna Madre. Barba-Macias (1999); Laguna Madre of Texas (Sheridan and Minello 2003); Barba-Macias et al. (2005). Remarks. The features observed agreed with those included in Abele and Kim (1986). 488 Key for the taxonomic identification of caridean shrimps associated with seagrass beds from the southern region of Laguna Madre (Adapted from Chace 1972, Williams 1984, Abele and Kim 1986) 1 ire Oh 3! 4! 5! 6! T' Carpus of second pair of pereopods entire (Palae- TO LNICHAG a eye sae PS conan ceux umn oe to wae arene | ae 2 Carpus of second pair of pereopods subdivided ..... 7 Carapace without branchiostegal groove ventral to antennal spine; endopod of the first pleopod of male with an appendix interna (genus Leander E. Desma- rest, 1849); lateral extension of anterior margin of basal antennular segment concave or straight; sty- locerite may reach to distal third of basal antennular segment; rostrum shallow in mature males, but very deep in mature females; fingers of second pereopod Ui Med ee.4 Bees: Leander tenuicornis (Say, 1818) Carapace with branchiostegal groove; endopod of the first pleopod of male entire, without appendix interna (genus Palaemon Weber, 1795) .........0.0ccccccceetcecees 3 Mandible usually with palp, formed by 2-3 articles; rostrum curved upwards (Fig. 2).....00....ccccee 4 Mandible usually without palp; rostrum straight or lessicuiveddpwards:ChIe S)P este csc0t 2 hate 5 Rostrum high, ventral margin with 3 or 4 teeth ......... Palaemon northropi (Rankin, 1898) Rostrum slender, ventral margin with 5 or 7 teeth .... Bi onl al soe A Palaemon floridanus Chace, 1942 Rostrum with 2 dorsal teeth behind posterior margin of orbit, teeth reaching to tip, 3—5 ventral teeth; dac- tylus of second pereopod with 2 teeth, immovable finger with 1 tooth on cutting edge... Br eee lly asia Palaemon vulgaris Say, 1818 Rostrum with only 1 dorsal tooth behind posterior margin of orbit; dactylus of second pereopod may have a tiny blunt tooth or may be unarmed, fixed fin- ger without tooth on cutting edge ........0.0. 6 Rostrum with dorsal teeth reaching to often bifurcate tip; 4 or 5, seldom 3, ventral teeth; dactylus of second pereopod with tiny and sometimes blunt tooth ......... ..Palaemon mundusnovus De Grave & Ashelby, 2013 Both margins of rostrum with unarmed stretch before dagger-shaped tip; 2-5, generally 3, ventral teeth; fingers of second pereopod without teeth on cutting COSC rea tier es ll Palaemon pugio (Holthuis, 1949) Chelae of first pair of pereopods distinct, at least on one side; first pair of pereopods both chelate; rostrum dentate or unarmed, not with single subdistal dorsal COOL I Ree Mine. 8 A ot BB tat Sine th ore ate Bn eee 8 Usually right first pereopod chelate, the other ending in simple claw-like dactyl; if both chelate, rostrum with subdistal dorsal tooth (Processidae); first pereo- pods similar, both chelate and lacking exopods; second pEercopods SVIMME ICAL: er ccs. .cmssedeer ti vawndenvds .... Ambidexter symmetricus Manning & Chace, 1971 Check List 14 (2) 8 Fingertips of first pair of chelae usually dark colored; first pair of chelipeds short and rather heavy but not swollen; eyes free, never extremely elongate (Hippo- NVA GI suid sete ea, Re Be oe, 22, Oe ee, 9 8’ Fingertips of first pair of chelae not dark colored; eyes never extremely elongate; first pair of pereopods distinctly stronger than second, often consisting in major and minor chelae (Alpheidae); eyes covered by carapace, epipods present on at least first 2 pairs of pereopods (genus Alpheus Fabricius, 1798) .......... 1] 9 Body short, total length 14 mm in males, 15.5 mm in females; rostrum shorter or longer than carapace, but clearly less than 1.3 times its length; supraorbital tooth present; with 1-3 (usually 2) dorsal and 14 ventral teeth (genus Hippolyte Leach, 1814)......... 10 9’ Body elongated, maximum length of males 40 mm, females 50 mm; rostrum noticeably longer than cara- pace, 1.4 times to nearly twice its length; supraorbital tooth absent; rostrum unarmed dorsally and with up LOMPOSVCTIAl TECHN. NS ka AAP Tae mee SATE Mfg, Wrensdeateee 10 Rostrum armed with 3 or 4 strong teeth on dorsal mar- gin and with a strong lateral carina in proximal third of length; rostrum reaching beyond end of antennular peduncle in both sexes; basal segment of antennular peduncle armed with 1-3 strong distolateral spines ETS Oy) sxanntaaaee Hippolyte obliquimanus Dana, 1852 10’ Rostrum usually armed with 2 (rarely 1 or 3) strong teeth in proximal half of dorsal margin and without distinct lateral carina; rostrum reaching beyond anten- nular peduncle in females and nearly to distal margin of second antennular segment in males; first segment of antennular peduncle unarmed (Fig. 6A, B)........... Pris ree Se Ee Hippolyte zostericola (Smith, 1873) 11 Major first chelae superiorly and inferiorly notched; merus of fisrt pereiopods unarmed distoventrally; major chela with distal ends of propodus and dac- tyl rounded; upper margin deeply notched forming SaddlelikerdepressiOns sr.07 eM ni totes ees sate evenly tare UL 8 ym Alpheus heterochaelis Say, 1818 Major first chelae not notched inferiorly; merus armed distoventrally with one large spine and 2—4 smaller spines; major chela with distal ends of propodus and dactyl narrowly-rounded:; upper margin of propodus deeply notched with depression ending distally in acute-overhanging tooth (Fig. 4D, F) ..0...0..0. pegs, Mina ds 520s Alpheus cf. packardii Kingsley, 1880 ~ 11 Discussion In recent years, several aspects of Laguna Madre of Tam- aulipas have been analyzed, with major focus on species with great importance to local fisheries, like peneaid shrimps, fishes, and oysters (Rendon-von Osten and Garcia-Guzman 1995). Although no population of cari- dean shrimp species has been commercially exploited in Herrera-Barquin et al. | Caridean shrimps from Laguna Madre, Tamaulipas 489 the lagoon, these shrimps have been studied due to their important ecological relevance. As a result, not only their diversity, richness, and habitat-associated distribution patterns have been documented, as well as their biomass and variations of density (Barba-Macias 1999, 2012, Barba-Macias et al. 2005), but also species lists (Rodri- guez et al. 2000, Leija-Tristan et al. 2000) and restoration assessments of submerged aquatic vegetation-covered areas have been done (Sheridan and Minello 2003, Sheri- dan 2004, King and Sheridan 2006). This study reports 12 caridean shrimp taxa, showing a higher species richness than most previous studies in the system (Barba-Macias 1999, 2012, Rodriguez et al. 2000, Barba-Macias et al. 2005), but not as many species (14) as recorded by Leia-Tristan et al. (2000). The observed richness is due to the influence of tropical and subtropical waters masses from Caribbean—Carolinian marine prov- inces, which explains the convergence of different types of fauna (Barba-Macias et al. 2012). The family Hippolytidae was the most abundant, with high numbers of individuals of Hippolyte zostericola and Tozeuma carolinense, followed by Palaemonidae with abundant records of Palaemon vulgaris, P mundusnovus, P. floridanus and P. pugio. Finally, Alpheidae and Pro- cessidae were the least abundant families. Commonly, alpheid shrimp richness is higher in tropical regions (Anker et al. 2006), whereas palaemonid and hippolytid shrimps are proportionally more abundant in temperate shallow water environments, especially those with SAV substrates (Lopez de la Rosa et al. 2002, Glancy et al. 2003). Estuarine fauna is distinguished by its tolerance to salinity variations, displaying strategies like high growth rates, rapid colonization, and high but variable abundances under suitable environmental conditions (Rendon-von Osten and Garcia-Guzman 1995, Kennish and Paerl 2010). Hippolyte zostericola was the most abundant species with 1,825 specimens, being higher in February > March > August, and at S2 > S1 > S3, maybe in response to the raised temperatures and salinities registered in August, as well as to a less seagrass structural coverage observed. Abundance of this shrimp has been correlated with water depth, and seagrass coverage and morphology as part of an adaptive response to improve avoidance of visual predators (Howard 1984, Zupo and Nelson 1999); fur- thermore, Zupo and Nelson (1999) also found that the positive correlation of H. zostericola abundance and water depth was consistent with negative phototropism, as part of the same predator-avoidance mechanism. Along with this, a potentially important predatory interaction has been noted between H. zostericola and Palaemon mundusnovus, indicated by the significant inverse correlation of abundance of both species, and being demonstrated in laboratory experiments (Zupo and Nelson 1999). In this work, maximum abundance of H. zostericola was observed at S2, whereas P. mundusno- vus was virtually absent (only 1 specimen found), which could be associated with the aforementioned predator— prey interaction, as well with more suitable conditions of water depth and seagrass coverage observed at the site. Nevertheless, this species is a common and dominant ele- ment in estuarine environments along Gulf of Mexico, due to its wide physiological tolerance to salinity changes (Barba-Macias et al. 2005), and further studies should be performed to better understand its ecological behavior in the area. Another dominant hippolytid shrimp was Tozeuma carolinense, with 431 collected specimens. This spe- cies, as well as Hippolyte zostericola, is less represented under mesohaline water conditions (Sanchez et al. 1996) and strongly associated with SAV meadows, commonly being attached to seagrass and macroalgae leaves, feed- ing on epiphytes and sheltering from predators (Kneib 1988). Both species were absent at S1 during August and February, maybe as a result of an increase in pre- dation by fishes foraging near the inlet, which changes their diets throughout rainy season, primarily feeding on macrocrustaceans (Barba-Macias 1999). However, T: carolinense was poorly represented at this station in March as well, whereas H. zostericola remained abun- dant. Other factors like tidal variations, less observed density of Syringodium filiforme in March and February, sampling effort, or the more euhaline—mesohaline waters occurring adjacent to Boca de Catan inlet could be influ- encing this phenomenon. Lesser abundances for Alpheidae might be explained by the geographical distribution of the family, highly diverse and inhabiting a great variety of marine and estuarine habitats in tropical waters (Anker et al. 2006), suggesting less richness and abundance towards Laguna Madre region. Lower representation could also be related to sampling exclusion, derived from the use of a selective epifaunal sampling method over SAV, excluding habitats such coral and rocky substrates where alpheid shrimps are commonly found, given their infaunal burrowing habits in those areas (Bauer 1985, Roman-Contreras and Martinez-Mayén 2010, Barba-Macias 2012). The only record of a processid shrimp was for Ambidexter sym- metricus at S1, but it was absent in the inner zones of the lagoon, which have euhaline-hypersaline waters and extensive SAV meadows. . However, this shrimp has been recognized as a rare or occasional element in Laguna Madre (Barba-Macias 1999, Barba-Macias et al. 2005), and more associated with inlet areas, euhaline conditions, less variable salinity and temperature, deeper zones, and bare sandy substrates (without SAV) (Barba-Macias 1999). Furthermore, it has been noted that processid shrimps are mostly nocturnal and that A. symmetricus is difficult to collect due to its cryptic habits (Pachelle et al. 2016, Santana-Moreno et al. 2016), so its presence could be accidental. We provide 4 new records for the study area: Palae- mon floridanus, P. northropi, Alpheus cf. packardii and Hippolyte obliquimanus. The last species has been mostly recorded in estuaries along southern Gulf of Mex- ico (Escobar 1984, Hernandez et al. 1996, Rodriguez et 490 al. 2000, Roman-Contreras 1988, Roman-Contreras and Martinez-Mayen 2010, Hermoso-Salazar and Arvizu- Coyotzi 2015), and in estuarine zones of Florida (d’Udekem d’Acoz 1997), but absent for the rest of the U.S. Gulf coast. The northernmost records in Mexico are known for Laguna de Tamiahua, Veracruz, and coral reefs of Veracruz state (Rodriguez et al. 2000). Hippolyte obliquimanus was collected at S2 in August, over seagrass meadows primarily composed by Syringo- dium filiforme and Halodule wrightii and represented by only | damaged specimen (Fig. 5). Nevertheless, it was possible to observe its distinct morphological features (see species remarks in results section). This species was absent in the other sampling stations despite the presence of SAV. It prefers shallow water environments, among Sargassum sp. algae (Mantelatto et al. 1999) and sea- grass meadows (Thalassia testudinum and S. filiforme) (d’Udekem d’Acoz 1997), on muddy and sandy bot- toms (Chace 1972); these same habitats are common in Laguna Madre of Tamaulipas (Estavillo and Aguayo 1985, Rendon-von Osten and Garcia-Guzman 1995). This only record was thought to be due to its occasional presence associated with seasonality and abundance of Sargassum sp. The density of Hippolyte obliquimanus is higher in autumn-winter periods in association with these algae (Terossi and Mantelatto 2010), which also influence variations in abundance of many caridean shrimp spe- cies (Leite and Turra 2003). The decrease in abundance of Sargassum sp. promotes the migration of associated caridean shrimps to deeper zones, which typically occurs during spring-summer in tropical regions, as a result of the rising temperatures (Terossi and Mantelatto 2010). In Laguna Madre, Sargassum sp. enters during spring, drifts through the system, including the sampling zone (Barba- Macias 1999), and leaves in early summer (Breuer 1962) as temperature increases. This dynamic might explain the observed abundance of H. obliquimanus, which could have entered to the system during spring among Sargas- sum, decreasing in number towards August as a result of a decreased Sargassum abundance and rising tem- peratures, thus searching for areas with SAV and deeper/ cooler waters. Reproductive traits could also be involved in the occurrence of Hippolyte obliquimanus. Clarke (1987) suggested that the reproductive output of some hippoly- tid shrimps decreases with increasing latitude. This was confirmed by Terossi et al. (2010), who observed lower mean reproductive output in populations from Brazil (23° S) than those from Costa Rica (09° N). Moreover, Woodward and White (1981) suggested that lower tem- peratures allow for a higher energy investment in embryo production, compared to animals living in warmer waters (Diaz 1980). However, Terossi and Mantelatto (2010) recommended that considerations should be given to local and regional environmental conditions that might contribute significantly to the plasticity of the reproduc- tive biology of decapods. Although we collected only 1 specimen of Hippolyte obliquimanus, a nearby occurrence Check List 14 (2) from Laguna de Tamiahua, Veracruz, makes it likely that there are well-established populations in Laguna Madre and possibly along the rest of the U.S. Gulf coastline; we attributed the scarcity of records to a lack of sampling effort. Palaemonidae showed the highest species richness with 6 species. Within this family, the genus Palaemon comprised 5 species, decreasing in abundance as follows: Palaemon vulgaris, P. mundusnovus, P. floridanus, P. pugio and P. northropi. Palaemonid shrimps are abundant in tropical and temperate estuaries over the world, serv- ing as detritivores, small invertebrate predators and an important food source for several fish and bird species (Bauer 2004). Here we report 2 new records for Laguna Madre and estuarine environments of Tamaulipas: Palaemon floridanus and P. northropi. Palaemon floridanus has no previous records along the Mexican Gulf, with just 1 nearby record from Padre Island and Laguna Madre of Texas (Strenth and Chace 1995). Its range includes the Gulf of Mexico, Caribbean Sea and adjacent zones (Chace 1942, Coen et al. 1981). On the other hand, P. northropi has been observed in Laguna de Términos, Campeche, and several points along Quintana Roo (Chace 1972, Roman-Contreras 1988, Hernandez et al. 2005), over a wide area from Bermuda to Uruguay (Holthuis 1952). Given their distribution range, the scarcity of published records around the study area is interesting, and might be due to a lack of surveys dealing with this genus in North America (Knowlton and Vargo 2004), which is supported by the notable difference in time between this study and the only local record of the genus (Palaemon floridanus) made by Strenth and Chace (1995) (19 years apart). Palaemon floridanus also has been associated with red algae like Digenia simplex, Laurencia poitei and some species of Gracilaria (Hooks et al. 1976). Con- sidering that most of the studies in the region addressed the relation of caridean shrimps with seagrass substrates (Barba-Macias 1999, 2012, Sheridan and Minello 2003, Sheridan 2004, Barba-Macias et al. 2005, King and Sheridan 2006), it is possible that habitat preferences are influencing the recorded occurrences; however, studies have not been carried out to clarify this relationship. Coen et al. (1981) found post-larvae and juvenile specimens among litter of Thalassia testudinum and Syringodium filiforme, species also occurring in Laguna Madre, posing the need for regular samplings for a better understanding of the frequency of this shrimp in the ecosystem. In their study, Strenth and Chace (1995) mentioned that it is com- mon during spring and summer months, and ovigerous females are present from March to September, reach- ing their maximum size in January and February. Our material of Palaemon floridanus is from August, with ovigerous females present, which suggests that suitable conditions allow for their growth early in the year and for development of their eggs later in spring and summer; this supports the idea of a well-settled species at least in Laguna Madre, and probably in other coastal lagoons or Herrera-Barquin et al. | Caridean shrimps from Laguna Madre, Tamaulipas 491 estuarine environments along Mexican Gulf coastline. Palaemon northropi had no previous records in the region until this work, which, in addition to the lack of studies, could be due to a inappropriate sampling method, as well as this species’ preference for more marine habitats (Anger and Moreira 1998, Pralon and Negreiros- Fransozo 2006) and not being commonly associated with SAV habitats, as are other caridean shrimp species. Another new record for Laguna Madre is Alpheus cf. packardii. This shrimp ranges from North Carolina, Ber- muda, Gulf of Mexico, Caribbean Sea, and Brazil (Anker et al. 2016) and with records in the Gulf of Mexico from Key West, Florida and Texas (Christoffersen 1979); Mexi- can records includes coral reefs off Veracruz (Hernandez et al. 1996, Hermoso-Salazar and Arvizu-Coyotzi 2015), but it is absent from the rest of the Gulf coast both in Mexico and the U.S. Our material consists of few speci- mens, which we attribute to an inappropriate sampling method considering their habitat preferences (Bauer 1985, Roman-Martinez 2010, Barba-Macias 2012). In Laguna Madre, the genus A/pheus has been related to SAV, but showing low abundance, density and biomass, and representing a rare or occasional element (Barba- Macias 1999, Barba-Macias et al. 2005). Moreover, the distribution patterns of alpheid shrimps are difficult to study since alpheids tend to form cryptic species com- plex (Anker 2001, Almeida et al. 2007, 2012), which pose future challenges as nomenclature and geographic distribution of the genus Alpheus are expected (Anker et al. 2007). Our work contributes to the knowledge of the caridean shrimp fauna in the southern region of Laguna Madre, adding new species to the list for the area and extending species’ ranges in the Gulf of Mexico. It will be necessary to design sampling methods fitted to the biology of each species in this group, to better describe their structure, ecology, and dynamics, Future studies on caridean shrimp might be useful for potential conserva- tion and maintenance of habitat integrity, and thus, giving support to many other ecologically and commercially rel- evant organisms. Our contribution also sets a precedent to aim efforts towards the clarification of geographic distribution, taxonomy, and even systematic status of this important and diverse decapod group. Authors’ Contributions HHB and ALT collected the specimens; HHB identified the seagrass material; HHB, ALT, and SFL wrote the text; HHB and ALT identified the caridean shrimps. References Abele LG (1972) A review of the genus Ambidexter (Crustacea: Decap- oda: Processidae) in Panama. Bulletin of Marine Science 22 (2): 365-380. Abele LG, Kim W (1986) An illustrated Guide to the Marine Decapod Crustaceans of Florida. Department of Biological Science, Florida State University, 784 pp. Almeida AO, Guerrazzi MC, Coelho PA (2007) Stomatopod and deca- pod crustaceans from Camamu Bay, state of Bahia, Brazil. Zootaxa 1553: 145. http:// doi.org/10.11646/zootaxa.1553.1.1 Almeida AO, Boehs G, Araujo-Silva CS, Bezerra LEA (2012) Shallow- water caridean shrimps from southern Bahia, Brazil, including the first record of Synalpheus ul (Rios & Duffy, 2007) (Alpheidae) in the southwestern Atlantic Ocean. Zootaxa 3347: 135. http://do1. org/10.11646/zootaxa.3905.1.3 Almeida AO, Costa-Souza AC, Cunha AM, Souza-Santos P, Oliveira MV, Oliveira-Soledade G (2013) Estuarine caridean shrimps (Crus- tacea: Decapoda) from I/héus, Bahia, Brazil: updated checklist and a key for their identification. Check List 9 (6): 1396-1405. https:// doi.org/10.15560/9.6.1396 Anger K, Moreira GS (1998) Morphometric and reproductive traits of tropical Caridean shrimps. Journal of Crustacean Biology 18 (4): 823-838. http://doi.org/10.1163/193724098 X00674 Anker A (2001) Two new species of snapping shrimps from the Indo- Pacific, with remarks on colour patterns and sibling species in Alpheidae (Crustacea: Caridea). The Raffles Bulletin of Zoology 49: 57-72. Anker A, Ahyong ST, Noel PY, Palmer AR (2006) Morphological phy- logeny of alpheid shrimps: parallel preadaptation and the origin of a key morphological innovation, the snapping claw. Evolution 60 (12): 2607-2528. http://doi.org/10.1111/j.0014-3820.2006.tb01886.x Anker A, Hurt C, Knowlton N (2007) Three transisthmian snapping shrimps (Crustacea: Decapoda: Alpheidae: A/pheus) associated with innkeeper worms (Echiura: Thalassematidae) in Panama. Zootaxa 1626: 1-23. Anker A, Tavares M, Mendoga JB (2016) Alpheid shrimps (Decapoda: Caridea) of the Trinidade & Martin Vaz Archipelago, off Brazil, with new records, description of a new species of Synalpheus and remarks on zoogeographical patterns in the oceanic islands of the tropical southern Atlantic. Zootaxa 4138 (1): 001-058. http://doi. org/10.11646/zootaxa.4138.1.1 Ashelby CW, Page TJ, De Grave S, Hughes JM, Johnson ML (2012) Regional scale speciation reveals multiple invasions of freshwa- ter in Palaemoninae (Decapoda). Zoologica Scripta 41: 293-306. https://doi.org/10.1111/j.1463-6409.2012.00535.x Aubriot L, Conde D, Bonilla S, Hein V, Britos A (2005) Vulnerabilidad de una laguna costera en una Reserva de Bidsfera: indicios recien- tes de eutrofizacion. In: Vila I, Pizarro J (Eds) Taller Internacional de Eutrofizacion de Lagos y Embalses. CYTED XVIIB. Patagonia Impresiones, Santiago, 65-85. Baeza JA, Fuentes MS (2012) Phylogeography of the shrimp Palaemon floridanus (Crustacea: Caridea: Palaemonidae): a partial test of meta-population genetic structure in the wider Caribbean. Marine Ecology 34: 381-393. http://doi.org/10.1111/maec.12038 Barba-Macias E (1999) Variacion de la densidad y la biomasa de peces juveniles y decapodos epibénticos de la region central de Laguna Madre, Tamaulipas. Hidrobioldgica 9: 103-116. Barba-Macias E (2012) Faunistic analysis of the caridean shrimps inhabiting seagrasses along the NW coast of the Gulf of Mexico and Caribbean Sea. Revista de Biologia Tropical 60 (3): 1161-1175. http://doi.org/10.15517/rbt.v6013.1765 Barba-Macias E, Raz-Guzman A, Sanchez AJ (2005) Distribu- tion patterns of estuarine caridean shrimps in the southwest- ern Gulf of México. Crustaceana 78 (6): 709-726. http:// doi. org/10.1163/156854005774353502 Bauer RT (1985) Diel and seasonal variation in species composition and abundance of caridean shrimps (Crustacea, Decapoda) from seagrass meadows on the north coast of Puerto Rico. Bulletin of Marine Science 36: 150-162. Bauer RT (2004) Remarkable shrimps: adaptations and natural history of the Carideans. University of Oklahoma Press, Norman, 282 pp. Bello-Pineda J, GO6mez-Mendoza L, Magafia-Rueda V, Graizbord B, Rodriguez-Herrero PH, Cervantes-Abrego M (2009) Descripcion detallada de los sitios piloto. In: Buenfil-Friedman J (Ed) Adap- tacidn a los Impactos del Cambio Climatico en los Humedales 492 Costeros del Golfo de México. Secretaria de Medio Ambiente y Recursos Naturales, Instituto Nacional de Ecologia, Ciudad de México, 389-546. Breuer JP (1962) An ecological survey of the lower Laguna Madre of Texas, 1953-1959. Publications of the Institute of Marine Science, University of Texas 8: 153-83. Carrera E (2004) Ficha Informativa de los Humedales de Ramsar. Ramsar Sites Information Database https://rsis.ramsar.org/ris/1362. Accessed on: 2016-9-5. Castellanos-Osorio I (2009) Estudios de zooplancton: logros y retos. In: J Espinoza-Avalos, Islebe GA, Hernandez-Arana HA (Eds.) El sistema ecoldgico de la bahia de Chetumal/Corozal: costa occiden- tal del Mar Caribe. ECOSUR, Chetumal, 61-68. Chace FA (1937) The Templeton Crocker Expedition, Part 7: caridean decapod Crustacea from the Gulf of California and the west coast of Lower California. Zoologica 22 (2): 109-138. Chace FA (1942) Six new species of decapod and stomatopod Crustacea from the Gulf of Mexico. Proceedings of the New England Zoo- logical Club 19: 79-92. Chace FA (1972) The shrimps of the Smithsonian—Bredin Caribbean Expeditions with summary of the West Indian shallow-water spe- cies (Crustacea: Decapoda: Natantia). Smithsonian Contributions to Zoology 98: 1-179. Christoffersen ML (1979) Campagne de la Calypso au large des cétes Atlantiques de l’Amerique du Sud (1961-1962) I. Decapod Crus- tacea: Alpheoida. Annales de |’ Institut Océanographique, Monaco 55: 297-377. Christoffersen ML (1998) Malacostraca—Eucarida, Caridea, Crangonoi- dea and Alpheoidea (except Glyphocrangonidae and Crangonidae). In: Young PS (Ed) Catalogue of Crustacea of Brazil. Museu Nacio- nal, Rio de Janeiro, 351-372. Clarke A (1987) Temperature, latitude and reproductive effort. Marine Ecology Progress Series 38: 89-99. Coen LD, Heck KL, Abele LG (1981) Experiments on competition and predation among shrimps of seagrass meadows. Ecology 62 (6):1484—1493. http://doi.org/10.2307/1941505 Contreras F, Castafieda O (2004) Las Lagunas Costeras y Estuarios del Golfo de México: Hacia el establecimiento de indices ecologicos. In: Caso M, Pisanty I, Ezcurra E (Eds) Diagnoéstico Ambiental del Golfo de México, Ciudad de México, 373-415 Dawes CJ, Phillips RC, Morrison G (2004) Seagrass communities of the Gulf Coast of Florida: status and ecology. Florida Fish and Wildlife Conservation Commission, Fish and Wildlife Research Institute and the Tampa Bay Estuary Program, St Petersburg, Florida, 78 pp. De Grave S, Ashelby CW (2013) A re-appraisal of the systematic status of selected genera in Palaemoninae (Crustacea: Decapoda: Palae- monidae). Zootaxa 3734 (3): 331-344. http://doi.org/10.11646/ zootaxa.3734.3.3. De Grave S, Fransen CHJM (2011) Carideorum catalogus: the Recent species of the dendrobranchiate, stenopodidean, procarididean and caridean shrimps (Crustacea: Decapoda). Zoologische Mededelin- gen 85 (9): 195-589. Diaz H (1980) The mole crab Emerita talpoida Say: a case of changing life history pattern. Ecological Monographs 50: 437-456. http:// doi.org/10.2307/1942652 Dominguez JC, Sanchez AJ, Florido R, Barba E (2003) Distribucion de Macrocrustaceos en Laguna Mecoacan, al sur del Golfo de México. Hidrobioldgica 13 (2): 127-136. d’Udekem d’Acoz C (1997) Redescription of Hippolyte obliquimanus Dana, 1852, and comparisson with Hippolyte williamsi Schmitt, 1924 (Decapoda: Caridea). Crustaceana 70: 469-479. http://doi. org/10.1163/156854097X00050 Escobar EG (1984) Comunidades de macro-invertebrados bentonicos en la Laguna de Términos, Campeche: composicion y estructura. Master’s thesis, Universidad Nacional Autonoma de México, Ciu- dad de México, 193 pp. Escobar-Briones E (2004) Estado del conocimiento de las comunidades bénticas en el Golfo de México. In: Caso M, Pisanty I, Ezcurra Check List 14 (2) E (Eds) Diagnoéstico Ambiental del Golfo de México. Primera Edicion, Ciudad de México, 201-246. Estavillo GC, Aguayo CE (1985) Ambientes Sedimentarios Recien- tes en la Laguna Madre, NE de Mexico. Boletin de la Sociedad Geoldgica Mexicana 46 (1-2): 29-63. Ferreira RS, Ramos-Vieira RR, D’Incao F (2010) The marine and estua- rine shrimps of the Palaemoninae (Crustacea: Decapoda: Caridea) from Brazil. Zootaxa 2606: 1—24. http://doi.org/10.15468/trz6dp Giraldes BW, Freire AS (2015) Extending the southern range of four shrimps (Crustacea: Decapoda: Stenopodidae, Hippolytidae and Alpheidae) in southwestern Atlantic (27°S) and confirming of Mediterranean Stenopus spinosus Risso, 1827 in Brazil. Zootaxa 3972 (3): 419-431. http://doi.org/10.11646/zootaxa.3972.3.7 Glancy TP, Frazer TK, Cichra CE, Lindberg WJ (2003) Comparative patterns of occupancy by decapod crustaceans in seagrass, oyster and marsh-edge habitats in a northeast Gulf of Mexico estuary. Estuaries 26 (5): 1291-1301. http://doi.org/10.1007/BF0280363 1 Hermoso-Salazar M, Martinez-Guzman LA (1991) Estudio taxondémico de ocho familas de camarones (Crustacea: Decapoda) en cinco arrecifes del Golfo de México. Thesis, Universidad Nacional Autonoma de México, Ciudad de México, 87 pp. Hermoso-Salazar M, Arvizu-Coyotzi K (2015) Crustaceos del Sistema Arrecifal Veracruzano. In: Granados-Barba A, Ortiz-Lozano LD, Salas-Monreal D, Gonzalez-Gandara C (Eds) Aportes al cono- cimiento del Sistema Arrecifal Veracruzano: hacia el corredor arre- cifal del suroeste del Golfo de México. Universidad Autonoma de Campeche, Campeche, 47—72. Hernandez JL, Toral-Almazan RE, Ruiz-Nufio JA (1996) Especies cata- logadas de crustaceos estomatodpodos y decapodos para el Golfo de México, Rio Bravo, Tamaulipas a Progreso, Yucatan. Comision Nacional para el Conocimiento y Uso de la Biodiversidad / Direc- cion General de Oceanografia Naval, Secretaria de Marina, Ciudad de México, 132 pp. Hernandez JL, Ruiz-Nufio A, Toral-Almazan RE, Arenas-Fuentes V (2005) Camarones, Langostas y Cangrejos de la Costa Este de México. Comision Nacional para el Conocimiento y Uso de la Biodiversidad (Conabio), Ciudad de México, 350 pp. Hildebrand HH (1958) Estudios bioldgicos preliminares sobre la Laguna Madre de Tamaulipas. Ciencia 17 (1—9):151—173. Hildebrand HH (1969) Laguna Madre, Tamaulipas: observations on its hydrography and fisheries. In: Ayala—Castafiares A (Ed) Lagunas Costeras, un Simposio. Memorias del Simposio Internacional de Lagunas Costeras (origen, dinamica y productividad). UNAM- UNESCO, Ciudad México, 679-686. Holthuis LB (1950) The Palaemonidae collected by the Siboga and Snellius Expeditions with Remarks on other Species. I. Subfamily Palaemoninae. The Decapoda of the Siboga Expedition. Part X. E.J. Brill, Leiden, 268 pp. Holthuis LB (1952) A general revision of the Palaemonidae (Crus- tacea, Decapoda, Natantia) of the Americas II. The subfamilies Palaemonidae. Allan Hancock Foundation, Occasional Papers 12: 1-396. Hooks TA, Heck KL, Livingston RL (1976) An inshore marine inverte- brate community: structure and habitat associations in the northe- asthern Gulf of Mexico. Bulletin of Marine Science 26 (1): 99-109. Howard RK (1984) The trophic ecology of caridean shrimps in an eelgrass community. Aquatic Botany 18: 155-174. https://doi. org/10.1016/0304-3770(84)90085-8 Kennish MJ, Paerl HW (2010) Coastal lagoons: critical habitats of environmental change. In: Kennish MJ, Paerl HW (Eds) Coastal Lagoons: Critical Habitats of Environmental Change. United States of America. CRC Press, Boca Raton, Florida, 1-16. Kim W, Abele L (1988) The snapping shrimp genus A/pheus from the eastern Pacific (Decapoda: Caridea: Alpheidae). Smithsonian Con- tributions to Zoology 454: 1-119. King PS, Sheridan P (2006) Nekton of new seagrass habitats colonizing a subsided salt marsh in Galveston Bay, Texas. Estuaries 29 (2): 286-296. http://doi.org/10.1007/BF02781997 Herrera-Barquin et al. | Caridean shrimps from Laguna Madre, Tamaulipas 493 Kingsley JS (1878) Notes on the North American Caridea in the Museum of the Peabody Academy of Science at Salem, Mass. Proceedings of the Academy of Natural Sciences of Philadelphia 30: 89-98. Kingsley JS (1880) On a collection of Crustacea from Virginia, North Carolina, and Florida, with a revision of the genera of Crangonidae and Palaemonidae. Proceedings of the Academy of Natural Sci- ences of Philadelphia 31 (3): 383-427. Kneib RT (1988) Testing for indirect effects of predation in an inter- tidal soft-bottom community. Ecology 69: 1795-1805. http://doi. org/10.2307/1941158 Knowlton RE, Vargo CK (2004) The larval morphology of Palaemon floridanus Chace, 1942 (Decapoda, Palaemonidae) compared with other species of Palaemon and Palaemonetes. Crustaceana 77: 683-715. http://doi.org/10.1163/1568540041958563 Knowlton N, Weigt LA (1998) New dates and new rates for divergence across the Isthmus of Panama. Proceedings of the Royal Society of London B: Biological Science 265: 2257-2263. http://doi. org/10.1098/rspb. 1998.0568 Knowlton N, Weigt LA, Solorzano LA, Mills DK, Bermingham E (1993) Divergence in proteins, mitochondrial DNA, and reproduc- tive compatibility across the Isthmus of Panama. Science 260: 1629-1632. http://doi.org/10.1126/science.8503007 Leya-Tristan A (2005) Ecologia y estructura poblacional de los stocks reproductora-recluta, del Camarén Café Farantepenaeus aztecus IVES, 1891, en la Laguna Madre y Plataforma Continental de Tamaulipas, México. PhD thesis, Autonomous University of Nuevo Leon, San Nicolas de los Garza, 185 p. Leija-Tristan A, Contreras-Arquieta A, Garcia-Garza ME, Contreras-Bal- deras AJ, Lozano-Vilano ML, Contreras-Balderas S, Garcia-Ramirez ME, Ortiz-Rosales J, Segoviano-Salinas F, Jiménez-Guzman F, Lazcano-Villarreal D, de Leon-Gonzalez A, Martinez-Lozano S, Rodriguez-Almaraz GA, Guzman-Lucio MA, Gonzalez de la Rosa MC, Garcia-Salas JA, Guajardo-Martinez G, Gonzalez-Rojas JI, Guzman-Velasco A (2000) Taxonomic, Biological and Biogeo- graphic aspects of selected Biota of the Laguna Madre, Tamaulipas, Mexico. In: Munawar M, Lawrence SG, Munawar IF, Malley DF (Eds) Acuatic Ecosystems of México: Status & Scope. Backhuys Publishers, Leiden, 399-435. Leite FPP, Turra A (2003) Temporal variation in Sargassum biomass, Hypnea epiphytism and associated fauna. Brazilian Archives of Biology and Technology 46 (4): 665-671. http://doi.org/10.1590/ $1516-89132003000400021 Lemaitre E, Alvarez-Leon R (1992) Crustaceos decapodos del Pacifico Colombiano: lista de especies y consideraciones zoogeograficas. Anales del Instituto de Investigaciones Marinas Punta Betin 21: 33-76. Llanso RJ, Bell SS,Vose FE (1998) Food habits of red drum and spot- ted seatrout in a restored mangrove impoundment. Estuaries 21: 294-306. https://doi.org/10.2307/1352476 Lopez de la Rosa I, Garcia-Raso JE, Rodriguez A (2002) Evolution of a decapod community (Crustacea) of a shallow bottom with seaweeds from southern Europe. Journal of the Marine Bio- logical Association of the United Kingdom 82: 85-95. https://dol. org/10.1017/S0025315402005209 Mantelatto FL, Martinelli JM, Garcia RB (1999) Fecundity of Hip- polyte obliquimanus Dana, 1852 (Decapoda, Caridea, Hippolyti- dae) from the Ubatuba region, Brazil. In: FR Schram, von Vaupel Klein JC (Eds) Crustaceans and the Biodiversity Crisis. Procced- ings of the Fourth International Crustacean Congress, Amsterdam, 1: 691-700. Markham JC, McDermott JJ (1980) A tabulation of the Crustacea: Decapoda of Bermuda. Proceedings of the Biological Society of Washington 93: 1266-1276. Markham JC, Donath-Hernandez FE, Villalobos-Hiriart JL, Diaz-Bar- riga AC (1990) Notes on the shallow-water marine crustacean of the Caribbean coast of Quintana Roo, México. Anales del Instituto de Biologia, Universidad Nacional Autonoma de México, Serie Zoologia 61 (3): 405-446. Martinez-Iglesias JC, Carvacho A, Rios R (1996) Catalogo de los cari- deos marinos (Crustacea, Decapoda, Caridea) de las aguas someras de Cuba. Avicennia 4/5: 27-40. Mathews LM, Schubart CD, Neigel JE, Felder DL (2002) Genetic, eco- logical and behavioural divergence between two sibling snapping shrimp species (Crustacea: Decapoda: A/pheus). Molecular Ecology 11: 1427-1437. http://doi.org/10.1046/j.1365-294X.2002.01539.x McClure M (2005) Snapping shrimps. In: Hernandez JL, Ruiz-Nufio A, Toral-Almazan RE, Arenas-Fuentes V (Eds) Camarones, Langostas y Cangrejos de la Costa Este de México, Vol. I. Comision Nacional para el Conocimiento y Uso de la Biodiversidad (Conabio), Ciudad de México, 350 pp. McClure MR, Greenbaum IF (1994) Biochemical variation in Alpheus (Decapoda, Caridea, Alpheidae) from the coast of Texas: Evidence for cryptic species. Southwestern Naturalist 39: 63-66. http://doi. org/10.2307/3672194 Pachelle PPG, Anker A, Mendes CB, Bezerra LEA (2016) Decapod crustaceans from the state of Ceara, northeastern Brazil: an updated checklist of marine and estuarine species, with 23 new records. Zoo- taxa 4131 (1): 001-063. http://doi.org/10.11646/zootaxa.4131.1.1 Pereira G (1997) A cladistic analysis of the freshwater shrimps of the family Palaemonidae (Crustacea, Decapoda, Caridea). Acta Bioldgica Venezuelica 17:1—69. Pralon BGN, Negreiros-Fransozo ML (2006) Population biology of Palaemon (Paleander) northropi Rankin, 1898 (Crustacea, Decapoda, Palaemonidae) in a tropical South American estuary. Acta Limnologica Brasiliensia 18 (1): 77-87. Rendon-von Osten J, Garcia-Guzman J (1995) Evaluacion del impacto ambiental de las actividades humanas en la Laguna Madre, Tamau- lipas. In: Botello AV, Rojas-Galaviz JL, Benitez JA, Zarate-Lomeli D (Eds) Golfo de México, Contaminacion e Impacto Ambiental: Diagnostico y Tendencias. Universidad Autonoma de Campeche, Campeche, 521-540. Rodriguez-Almaraz GA, Leija-Tristan A, Mendoza-Alfaro R (2000) Records of the Caridean Shrimps (Crustacea: Decapoda) from the Coasts of the Mexican Pacific Ocean, Gulf of Mexico and Mexican Caribbean. Bulletin of Marine Science 67 (2): 857-867. Roman-Contreras R (1988) Caracteristicas ecologicas de los crustaceos decapodos de Laguna de Términos. In: Yafiez-Arancibia A, Day JW Jr (Eds) Ecologia de los ecosistemas costeros en el sur del Golfo de México: La region de la Laguna de Términos. Instituto de Ciencias del Mar y Limnologia, Universidad Nacional Autonoma de México. Coastal Ecology Institute, Louisiana State University. Editorial Universitaria, Ciudad de México, 305-322. Roman-Contreras R, Martinez-Mayén M (2010) Palaemonidae (Crusta- cea: Decapoda: Caridea) from the shallow waters of Quintana Roo, Mexican Caribbean coast. Revista Mexicana de Biodiversidad 81 (1): 43-51. Roman-Contreras R, Romero-Rodriguez J (2005) Incidence of infestation by Bopyrina abbreviata Richardson, 1904 (Isopoda: Bopyridae) on Hippolyte zostericola (Smith, 1873) (Decapoda: Hippolytidae) in Laguna de Términos, Gulf of Mexico. Nauplius 13: 83-88. Sanchez AJ, Raz-Guzman A, Barba E (1996) Habitat value of sea- grasses for decapods in tropical coastal lagoons of the southwestern Gulf of Mexico: an overview. In: Kuo J, Phillips RC, Walker DI, Kirkman K (Eds) Seagrass Biology: Proceedings of an Interna- tional Workshop, Rottnest Island, Western Australia. University of Western Australia, Nedlands, 233-240. Santana-Moreno LD, Rojas-Gonzalez RI, Wakida-Kusunoki AT (2016) First record of Ambidexter symmetricus (Decapoda: Caridea: Pro- cessidae) from the Yucatan Peninsula. Revista Mexicana de Biodi- versidad 87: 1138-1140. https://doi.org/10.1016/j.rmb.2016.07.008 Santos PS, Soledade GO, Almeida AO (2012) Decapod crustaceans on dead coral from reef areas on the coast of Bahia, Brazil. Nauplius 20: 145-169. http://doi.org/10.1590/S0104-649720 12000200007 Sheridan P (2004) Comparison of restored and natural seagrass beds 494 near Corpus Christi, Texas. Estuaries 27 (5): 781-792. http://doi. org/10.1007/BF029 12040 Sheridan PF, Minello TJ (2003) Nekton use of different habitat types in seagrass beds of lower Laguna Madre, Texas. Bulletin of Marine Science 72: 37-61. Smith SI (1873) Crustacea. In: Verrill AE (Ed) Report upon the inver- tebrate animals of Vineyard Sound and the adjacent waters, with an account of the physical characters of the region. United States Commission of Fish and Fisheries 1: 545-580. Soledade OG, Almeida AO (2013) Snapping shrimps of the genus Alpheus Fabricius, 1798 from Brazil (Caridea: Alpheidae): updated list and key for identification. Nauplius 21 (1): 89-122. http://doi. org/10.1590/S0104-64972013000100010 Souza JAF, Schwamborn R, Barreto AV, Farias ID, Fernandes LMG, Coelho PA (2011) Marine and estuarine shrimps (Dendrobranchi- ata, Stenopodidea, and Caridea), of Pernambuco state (Brazil) and northeastern Brazilian oceanic islands. Atlantica 33: 33-63. http:// doi.org/ 10.5088/atl. 2011.33.1.33 Strenth NE, Chace FA (1995) New records of natant decapods (Crus- tacea: Palaemonidae) from the south Texas coast. Texas Journal of Science 47: 315-31. Terossi M, Mantelatto FL (2010) Sexual ratio, reproductive period and seasonal variation of the gonochoric shrimp Hippolyte obliqui- manus (Caridea: Hippolytidae). Marine Biology Research 6 (2): 213-219. http://doi.org/10.1080/1745 1000903078630 Terossi M, Wehrtmann IS, Mantelatto F (2010) Interpopulation Com- parison of reproduction of the Atlantic shrimp Hippolyte obliqui- manus (Caridea:Hippolytidae). Journal of Crustacean Biology 30 (4): 571-579. http://doi.org/10.1651/09-3233.1 Tunnell J, Judd FW (2002) The Laguna Madre of Texas and Tamauli- pas. Texas A&M University Press, College Station, Texas, 335 pp. Vera-Caripe J, Lira C, Hernandez-Flores X, Montoya H (2012) Algunas especies del género A/pheus Fabricius, 1798 (Crustacea Decapoda) de la Isla de Coche, con una adicion a la carcinofauna venezolana. Boletin del Instituto Oceanografico de Venezuela 52 (2): 11-18 p. Verrill A (1922) Decapod Crustacea of Bermuda. Part I]. Macrura. Transactions of the Connecticut Academy of Arts and Sciences 26: 1-179. http://doi.org/10.5962/bh1.title.23648 Check List 14 (2) Wicksten MK (1989) Hippolyte zostericola (Crustacea: Decapoda) in the Eastern Pacific. Proceedings of the Biological Society of Wash- ington 102: 644-645. Wicksten M (2005a) Hippolytid shrimps. In: Hernandez JL, Ruiz-Nufio A, Toral- Almazan RE, Arenas-Fuentes V (2005) Camarones, Lan- gostas y Cangrejos de la Costa Este de México, Vol. I. Comision Nacional para el Conocimiento y Uso de la Biodiversidad (Cona- bio), Ciudad de México, 350 pp. Wicksten M (2005b) Palaemonid shrimps. In: Hernandez JL, Ruiz-Nufio A, Toral- Almazan RE, Arenas-Fuentes V (2005) Camarones, Lan- gostas y Cangrejos de la Costa Este de México, Vol. I. Comision Nacional para el Conocimiento y Uso de la Biodiversidad (Cona- bio), Ciudad de México, 350 pp. Williams A (1965) Marine decapod crustaceans of the Carolinas. Fish- ery Bulletin 65 (1): 1-298. Williams AB (1974) Two new axiids (Decapoda: Thalassinidea: Calo- caris) from North Carolina and the Straits of Florida. Proceedings of the Biological Society of Washington 87 (39): 451-464. Williams AB (1984) Shrimps, lobsters, and crabs of the Atlantic coast of the eastern United States, Maine to Florida. 1-566 pp. Williams ST, Knowlton N, Weigt LA, Jara JA (2001) Evidence for three major clades within the snapping shrimp genus A/pheus inferred from nuclear and mitochondrial gene sequence data. Molecular Phylogenetics and Evolution 20: 375-389. http://do1.org/10.1006/ mpev.2001.0976 Woodward IO, White RWG (1981) Effects of temperature and food on the fecundity and egg development rates of Boeckella symmetrica (Copepoda: Calanoida). Australian Journal of Marine and Fresh- water Research 32: 997-1001. https://doi.org/10.1071/MF9810997 Zarur A (1962) Algunas consideraciones geobiologicas de la Laguna de Términos, Campeche. Revista de la Sociedad Mexicana de Historia Natural 23: 51—70. Zupo V, Nelson WG (1999) Factors influencing the association pat- terns of Hippolyte zostericola and Palaemonetes intermedius (Decapoda: Natantia) with seagrasses of the Indian River Lagoon, Florida. Marine Biology 134: 181-190. http://doi.org/10.1007/ s002270050536