Notes on some alpheid shrimps (Decapoda: Caridea) of Thalassia testudinum meadows, from the Central–Southern Mexican Caribbean

Notas sobre algunos alfeidos (Decapoda: Caridea) de praderas de Thalassia testudinum, del Caribe Centro–Sur Mexicano

Ramiro Román–Contreras and Mario Martínez–Mayén

Universidad Nacional Autónoma de México (UNAM), Instituto de Ciencias del Mar y Limnología, Laboratorio de Carcinoparasitología, Apartado Postal 70–305, México D. F. 04510, México, E–mail: rokon@cmarl.unam.mx.

Recibido: 03 de febrero 2010
Aceptado: 22 de noviembre 2010

ABSTRACT

This study provides an account of some shrimps of the family Alpheidae collected on Thalassia testudinum meadows in Bahía de la Ascensión, Bahía del Espíritu Santo and Mahahual reef lagoon, Quintana Roo, on the Caribbean coast of Mexico. A total of 657 specimens of alpheid shrimps were collected; they represent two genera and seven species. Alpheus was the most diverse genus with five species. An updated geographic distribution is provided for all the seven species recorded; four of them represent new records for Bahía del Espíritu Santo and two for the Mahahual reef lagoon.

Key words: Alpheidae, Decapoda, Caridea, Mexican Caribbean.

RESUMEN

Se presenta el listado de algunas especies de camarones de la familia Alpheidae recolectados en praderas de Thalassia testudinum en la Bahía de la Ascensión, la Bahía del Espíritu Santo y la laguna arrecifal de Mahahual, Quintana Roo, en la costa del Caribe mexicano. Se capturaron un total de 657 especímenes de camarones alfeidos que representan a dos géneros y siete especies. Alpheus fue el género más diverso, con cinco especies. Se proporciona la distribución geográfica actualizada de todas las especies registradas. Cuatro de ellas representan nuevos registros para la Bahía del Espíritu Santo y dos para la laguna arrecifal de Mahahual.

INTRODUCTION

Shrimps of the family Alpheidae, commonly known as snapping shrimps, are an abundant and ecologically diverse group of decapod crustaceans in most marine tropical and subtropical waters (Anker et al., 2006a), but are also distributed in cool–temperate (e.g., Anker & Komai, 2004), freshwater (De Grave et al. 2008), and brackish water habitats, such as mangroves and estuaries (Christoffersen, 1984). Alpheids are distributed from the intertidal down to about a 1000 m in depth (see Anker et al., 2006a), but reach their greatest diversity in waters shallower than 50 m, particularly on coral reefs and adjacent habitats, e.g., sea–grass beds or sand flats with coral rubble. Many species are symbiotic with other organisms, such as sponges, anemones, cnidarians, mollusks, echinoderms, annelids, fishes, and other crustaceans (Kim & Abele, 1988; Anker et al., 2006a).

Several studies on the caridean fauna including the snapping shrimps have been carried out in the Caribbean region, most of them focused on their taxonomic and zoogeographic aspects (e.g., Armstrong, 1949; Chace, 1972; Carvacho, 1979, 1982; Duffy, 1996; Martínez–Iglesias et al., 1996, 1997; Vargas & Cortés, 1999; Anker, 2007; Anker et al., 2006b; Ríos & Duffy, 2007; Macdonald et al., 2009); however, studies for the Mexican Caribbean are rather scarce, with Chace (1972) being the main reference presenting extensive taxonomic information on this group for the tropical western Atlantic, including the Yucatán Peninsula. Other important contributions are by Markham and Donath–Hernández (1990) and Markham et al. (1990), who provided a synopsis of the shallow–water crustaceans of Quintana Roo, including several new records of alpheids. Briones–Fourzán and Lozano–Álvarez (2002) reported some data about the alpheids from Cozumel. The deep water caridean fauna was studied by Escobar–Briones and Villalobos–Hiriart (2003), who mentioned three new records of snapping shrimps from the coast of Quintana Roo.

During a study on the biology and ecology of bopyrid parasites of the caridean fauna based on recent field work in the Bahía de la Ascensión, Bahía del Espíritu Santo and the Mahahual reef lagoon, alpheid shrimps were collected by trawling on turtle–grass (Thalassia testudinum Banks ex König) meadows. In the present paper those shrimps are listed and some notes about them are provided.

MATERIAL AND METHODS

Field work was conducted at five stations in the Bahía de la Ascensión (19°30’–19°50’ N, 87°25’–87°50’ W), five in Bahía del Espíritu Santo (19°12’–19°25’ N, 87°41’–88°47’ W), and one in Mahahual reef lagoon (18°42.88’ N, 87°42.44’ W) along the central–southern coast of Quintana Roo, Mexico (Fig. 1), during July 1997, May and November 2001, May 2002, and January 2003. All specimens were collected with a Colman–Seagrove sledge net with a mesh size of 800 µm in meadows of turtle–grass (T. testudinum) at a depth range of 0.4–1.0 m. Immediately after sampling, all the specimens were fixed in 10% formaldehyde and then transferred to 70% ethanol. The bottom was sand–mud with fragments of shells at Bahía de la Ascensión and Bahía del Espíritu Santo and sand in the Mahahual reef lagoon. Additional metadata (currents, temperature, salinity, sediment type, and vegetation) of these study areas can be found in Merino Ibarra (1986), Jordán–Dahlgren et al. (1994), Castellanos Osorio and Suárez–Morales (1997) and Suárez–Morales & RiveraArriaga (1998).

The species treated here were arranged alphabetically; synonymies, material examined, geographic ranges, ecological notes, as well as new local records in the Mexican Caribbean are provided for each species. All specimens are deposited in the collection at the authors laboratory, in the Universidad Nacional Autónoma de México (UNAM).

RESULTS

A total of 657 specimens from two genera and seven species were collected during the surveyed period. The genus Alpheus Fabricius, 1798 was the most diverse, with five species. Four species represent new records for the Bahía del Espíritu Santo, while two species are new records for Mahahual reef lagoon.

Order Decapoda Latreille, 1802

Infraorder Caridea Dana, 1852

Superfamily Alpheoidea Rafinesque, 1815

Family Alpheidae Rafinesque, 1815

Alpheus cf. armillatus H. Milne–Edwards, 1837

Alpheus cf. armillatus. Almeida et al., 2007: 11.

Crangon armillatus. Schmitt, 1935: 142.

Material examined: Bahía del Espíritu Santo, 2 specimens, November 2001.

Distribution: North Carolina (Williams, 1984) to nearby Port Aransas, Texas (Felder & Chaney, 1979); from Isla Lobos, Veracruz (Hernández Aguilera et al., 1996) and Isla Pérez, Arrecife Alacrán, Yucatán, Gulf of Mexico (Martínez–Guzmán & Hernández–Aguilera, 1993), Isla Cozumel, Quintana Roo (Chace, 1972; Briones–Fourzán & Lozano–Álvarez, 2002), south to Santa Catarina, Brazil (Christoffersen, 1998); West Indies from Bermuda (Markham & McDermott, 1980) to Tobago (Chace, 1972).

Ecological notes: Under rocks and shells between interstices of coral (Chace, 1972); in sponges (Markham et al., 1990), and turtle–grass (McClure, 2005). Shallow–water to 14 m (Felder & Chaney, 1979), on calcareous bottom covered with T. testudinum, depth 0.5 m (present study).

Local new records: Bahía del Espíritu Santo, Quintana Roo.

Remarks: The material examined fits well with Chace’s (1972) key, and the figures and keys presented by Martínez–Iglesias et al. (1997) and McClure (2005) for the species. Alpheus armillatus is a large species complex that includes many cryptic species (Mathews & Anker, 2009). Those authors, based on molecular analyses of this species complex from the Caribbean Sea, Gulf of Mexico, Florida, Brazil, and the tropical eastern Pacific, reported the presence of six clades broadly overlapped in their geographic distribution, including 11 lineages in the western Atlantic and eight in the eastern Pacific, for a total number of at least 19 putative species contained in the A. armillatus species complex.

Alpheus bouvieri A. Milne–Edwards, 1878

Alpheus Edwarsii . Dana, 1852: 542 [not Audouin, 1826].

Not A. bouvieri. Christoffersen, 1979: 303 (part., material from eastern Pacific; see Anker et al., 2009); Kim & Abele, 1988: 58, fig. 24 (see Williams et al., 2001 and Anker et al., 2009).

Material examined: Bahía de la Ascensión, one specimen, May 2002.

Distribution: Eastern and southern Florida (Abele & Kim, 1986); Gulf of Mexico: Veracruz and Campeche (Hernández Aguilera et al., 1996), Punta Xamach, Bahía de la Ascensión, Quintana Roo, Mexico (Markham et al., 1990); Bermuda (Markham & McDermott, 1980) and Cuba (Martínez–Iglesias et al., 1996) to Trinidad (Christoffersen, 1979); south to Torres, Rio Grande do Sul, Brazil (Christoffersen, 1979); eastern Atlantic from Cape Verde Islands and Senegal to Ilha de Sao Tomé and Congo (Christoffersen, 1979); south Atlantic: Ascension Island (Manning & Chace, 1990).

Ecological notes: Among rocks, dead coral, sometimes in sponges (Christoffersen, 1979), on zoanthids and coralline algae (Anker et al., 2009). Shallow subtidal to about 10 m depth (Anker et al., 2009), in beds of T. testudinum on sand, depth 0.6 m (present study).

Remarks: The specimen examined agrees in general with Chace’s (1972) key, and the description and figures by Christoffersen (1979) and Anker et al. (2009) for A. bouvieri. This species was reported previously from the eastern Pacific and the Atlantic by Kim and Abele (1988). Anker et al. (2009) reviewed the Alpheus bouvieri species complex and restricted A. bouvieri to the Atlantic, while the eastern Pacific populations were assigned to Alpheus javieri Anker, Hurt and Knowlton, 2009. This species is genetically distinct and with little differences in morphology and color pattern respect to A. bouvieri, such as a slightly narrower and deeper dorsal groove on the palm of the major chela, and the slightly shorter first carpal segment of the second pereiopod, as well as tan or brown–colored spines on the propodus of third and fourth pereiopods, whereas in A. bouvieri such spines do not bear traces of color (Anker et al., 2009).

A recently described species that could be confused with A. bouvieri in the western Atlantic is A. agilis Anker, Hurt & Knowlton, 2009, which is distributed only in Brazil and West Africa, distinguished from A. bouvieri by the presence of a spine on ischium of the third and fourth pereiopods (see Anker et al., 2009).

Alpheus cf. floridanus Kingsley, 1878

Alpheus floridanus. Chace, 1972: 65, figs. 17–20 (part.); Herbst et al., 1978: 991; Christoffersen, 1979: 311, figs. 6–8; Markham & McDermott, 1980: 1270; Rodríguez, 1980: 144, fig. 41a–d; Martínez Iglesias et al., 1997: 423, fig. 11; Christoffersen, 1998: 358; Escobar Briones & Villalobos–Hiriart, 2003: 108; McClure, 2005: 139 (part., not fig. 13).

Alpheus cf. floridanus. Almeida et al., 2007: 12.

Material examined: Bahía de la Ascensión, 9 specimens, May 2002; 1 specimen, January 2003. Bahía del Espíritu Santo, 1 specimen, May 2001. Mahahual reef lagoon, 19 specimens, July 1997.

Distribution: Western Atlantic from southeast of Cape Hatteras (35°01.5’N, 75°30.8’W), North Carolina (Herbst et al., 1978); Bermuda (Markham & McDermott, 1980); Dry Tortugas, Florida throughout the Gulf of Mexico (Christoffersen, 1979, 1998); West Indies from Bahamas to Curaçao and Bonaire (Christoffersen, 1979); Mexico: Bahía de la Ascensión, Quintana Roo (Chace, 1972) and north of Mahahual (Escobar–Briones & Villalobos–Hiriart, 2003); south to Rio Grande do Sul, Brazil (Christoffersen, 1998). Actually the records of A. floridanus from the eastern Atlantic and eastern Pacific refer to closely related species (Anker, 2001; Williams et al., 2001; Almeida et al., 2007).

Ecological notes: Bottoms with calcareous algae, among hydrozoans and bryozoans (Christoffersen, 1979); in soft muddy sediments in mangroves, estuaries and coastal lagoons (Christoffersen, 1979, Martínez–Iglesias et al., 1997). Shallow–water to 240 m (Escobar–Briones & Villalobos–Hiriart, 2003), on sand and mud bottoms with Thalassia, 0.5–1 m depth (present study).

Local new records: Bahía del Espíritu Santo, Quintana Roo.

Remarks: Alpheus floridanus form also a species complex, with four species in the western and eastern Atlantic and eastern Pacific (see Anker, 2001; Almeida et al., 2007). On the basis of a phylogenetic analysis, Bracken et al. (2007) mentioned the existence of at least five different species within this complex when populations of the eastern Pacific, western Atlantic and Gulf of Mexico were compared.

Alpheus cf. packardii Kingsley, 1880

Alpheus packardii. De Grave et al., 2006: 1421–1422.

Crangon packardii. Schmitt, 1935: 144.

Alpheus normanni Kingsley 1878 . Chace, 1972: 68; Christoffersen, 1979: 322; Williams, 1984: 97, fig. 66; Bauer, 1985: 152; Martínez–Guzmán & Hernández–Aguilera, 1993: 615; Martínez–Iglesias et al., 1997: 424, fig. 13; Christoffersen, 1998: 359; Briones–Fourzán & Lozano–Álvarez, 2002: 200; McClure, 2005: 145, fig. 17, pl. 3, fig. A.

Distribution: Listed as A. normanni from Bermuda; around Cape Charles, Virginia (Williams, 1984) to Key West Florida and Texas (Christoffersen, 1979); throughout the Gulf of Mexico to Isla Pérez, Arrecife Alacrán, Yucatán (Martínez–Guzmán & Hernández–Aguilera, 1993); Isla Cozumel and Bahía de la Ascensión, Quintana Roo (Chace, 1972; Briones–Fourzán & Lozano–Álvarez, 2002); to São Paulo, Brazil (Christoffersen, 1998); West Indies from Bahamas (Christoffersen, 1979) to Tobago (Chace, 1972).

Ecological notes: Among coral reefs (Chace, 1972), on sand, mud, broken shells, algae and sponges (Christoffersen, 1979) and meadows of T. testudinum (Bauer, 1985). Subtidal to 73 m (Chace, 1972), collected on sand bottom covered with T. testudinum, depth 0.5–1 m (present study).

Local new records: Bahía del Espíritu Santo and Mahahual reef lagoon, Quintana Roo.

Remarks: Alpheus packardii form also is a species complex, with at least three probable cryptic species in the western Atlantic, and two in the eastern Pacific (Almeida et al., 2007). Chace (1937) considered the eastern Pacific species A. normanni and the western Atlantic A. packardii as synonyms. Kim and Abele (1988) compared specimens of A. normanni from the eastern Pacific with material from Florida and find that the male specimens from the eastern Pacific presented the minor chela of the first pereiopods more elongated than the minor chela of male specimens from Florida; therefore, the eastern Pacific form was considered by those authors as a different species from the Atlantic form and, according to Christoffersen (1998) the western Atlantic specimens previously named as A. normanni must be attributed to A. packardii. The minor chela of 50 males examined in the present study ranges from 3.64 to 4.38 times as long as broad, compared to 5.8 reported by Kim & Abele (1988) for A. normanni from the eastern Pacific, so, we concluded that the specimens examined belong to the A. packardii species complex.

Alpheus viridari (Armstrong, 1949)

Crangon viridari. Armstrong, 1949: 8–12, fig. 2.

Alpheus viridari. Chace, 1972: 73; Carvacho, 1979: 455; Markham & McDermott, 1980: 1270; Martínez–Iglesias et al., 1997: 424, fig. 17; Alvarez–León, 2003: 265; McClure, 2005: 155, fig. 23; Hernández–Ávila et al., 2007: 37.

Material examined: Bahía de la Ascensión, 1 specimen, May 2002. Mahahual reef lagoon, 4 specimens, July 1997.

Distribution: Western Atlantic: Bermuda (Markham & McDermott, 1980); Florida Keys to Trinidad and Curaçao (Chace, 1972); Isla Cozumel and Bahía de la Ascensión, Quintana Roo, Mexico (Chace, 1972); South Lagoon of Chengue, Tayrona, Colombia (Alvarez–León, 2003); Isla Cubagua, Venezuela (Hernández–Ávila et al., 2007).

Local new records: Mahahual reef lagoon, Quintana Roo.

Synalpheus apioceros Coutière, 1909

Synalpheus apioceros. Coutière, 1909: 27, fig. 9; Chace, 1972: 86; Carvacho, 1979: 463; Felder & Chaney, 1979: 25; Rodríguez, 1980: 155; Abele & Kim, 1986: 19, 226–227, figs. i–k; Christoffersen, 1998: 362; Markham et al., 1990: 421; Wicksten, 2005: 32.

Synalpheus cf. apioceros. Almeida et al., 2007: 14.

Material examined: Bahía del Espíritu Santo, one specimen, May 2001.

Distribution: Southern Florida (Chace, 1972); Stetson and West Flower Garden Bank (Wicksten, 2005); Seven and One–half Fathom Reef, Texas (Felder & Chaney, 1979); Isla Cozumel and Cayo Culebras, Bahía de la Ascensión, Quintana Roo, Mexico (Chace, 1972); the Antilles, from Anegada Island to Tobago (Chace,1972); to Santa Catarina, Brazil (Christoffersen, 1998).

Ecological notes: Rubble fields near sea grass beds and mangroves (Carvacho, 1979); eroded rocks, mollusk shells, cavities of sponges and tubes of polychaetes in 10–12 m (Felder & Chaney, 1979), sand and rocky substrate with T. testudinum, depth less than 1 m (present study).

Local new record: Bahía del Espíritu Santo, Quintana Roo.

Remarks: The morphology of the specimen examined agrees well with the description and figures provided by Coutière (1909), and Abele & Kim´s (1986) key and illustrations to S. apioceros. Almeida et al. (2007) stated that S. apioceros is a species complex with at least two species in the western Atlantic.

Synalpheus fritzmülleri. Coutière, 1909: 35–37, fig. 18.

Synalpheus fritzmuelleri. Chace, 1972: 92; Christoffersen, 1979: 341; Felder & Chaney, 1979: 25; Markham & McDermott, 1980: 1270; Williams, 1984: 102, fig. 70; Manning & Chace, 1990: 22; Markham et al., 1990: 422; Hernández Aguilera et al., 1996: 36; Martínez–Iglesias et al., 1996: 35; Christoffersen, 1998: 362; McClure, 2005: 175, fig. 38; Wicksten, 2005: 32; Hermoso–Salazar & Solís–Weiss, 2010 : 65–68.

Synalpheus cf. fritzmuelleri. Almeida et al., 2007: 14.

Material examined: Bahía de la Ascensión, one specimen, January 2003.

Distribution: Bermuda (Markham & McDermott, 1980); off Cape Hatteras, North Carolina; west coast of Florida (Christoffersen, 1979); East Flower Garden Bank (Wicksten, 2005) and Seven and One–Half Fathom Reef, Texas (Felder & Chaney, 1979) to Isla Pérez, Alacrán Reef, Yucatán, Mexico (Hernández Aguilera et al., 1996); Puerto Morelos and Punta Xamach (Markham et al., 1990), Isla Cozumel, Bahía de la Ascensión and Bahía del Espíritu Santo, Quintana Roo, Mexico (Chace, 1972); Cuba (Martínez–Iglesias et al., 1996) to Tobago (Chace, 1972); south to Santa Catarina, Brazil (Christoffersen, 1998); also in South Atlantic: Saint Helena (Christoffersen, 1979) and Ascension Island (Manning & Chace, 1990).

Ecological notes: Sea grass flats with corals, Porites Link, 1807, eroded coral, among roots of mangroves (Chace, 1972) and sponges (Markham et al., 1990); intertidal to 75 m (Christoffersen, 1979), among T. testudinum and sand bottom, depth 0.5 m (present study).

Remarks: Previous records of S. fritzmuelleri from Tres Marías Archipelago (Isla María Madre) in the eastern Pacific actually belong to Synalpheus nobilii Coutière, 1909; therefore, according to Hermoso–Salazar and Solís–Weiss (2010), A. fritzmuelleri must be removed of the list from the eastern Pacific alpheids.

DISCUSSION

The species recorded belong to the genera Alpheus and Synalpheus, both highly diverse and known to contain cryptic species that are difficult to distinguish based on morphologic characters (see Knowlton, 1986 for a review of cryptic species; Knowlton, 1993; Mathews & Anker, 2009). Hence, it is required to consider other data such as colour patterns (Knowlton, 1986; Knowlton & Mills, 1992; Anker, 2001) and the ecology and genetics of the species (Knowlton, 1993; Williams et al., 2001; Anker et al., 2009; Mathews & Anker, 2009). According to Anker (2001), with the incorporation of those data to the taxonomy, many species of alpheids formerly considered as widely distributed, have been recognized as species complexes.

Of the species here treated, A. cf. armillatus, A. bouvieri, A. cf. floridanus, A. cf. packardii, and S. apioceros are currently recognized as part of species complexes (Almeida et al., 2007). Only the A. bouvieri species complex has been solved (Anker et al., 2009); the taxonomic review of the other mentioned complexes is also urgent (see Anker et al., 2007). On the other hand, S. fritzmuelleri is considered as a possible species complex (Almeida et al., 2007), whereas A. viridari may be a sister clade of the A. armillatus complex (Mathews & Anker, 2009).

The occurrence of snapping shrimps in our samples is explained by the proximity to reefs in the studied areas, since many associated reef–species of crustaceans utilize the surrounding seagrass to live (Heck, 1977). The capture of species associated with other organisms such as S. fritzmuelleri (reported from sponges and among ascidians and gorgonians, see Heck, 1977; Christoffersen, 1979; Markham et al., 1990) may be attributed to the collecting activities that detached these shrimps from their hosts.

The species recorded herein are restricted to tropical and subtropical waters (Christoffersen, 1982; Manning & Chace, 1990; Markham et al., 1990) and according to previous geographic records, all those species are distributed in the western Atlantic, although A. bouvieri and S. fritzmuelleri also occur in the eastern Atlantic. At present, a detailed analysis of the distributional patterns in marine zoogeographic provinces of the species collected is not possible, as most belong to species complexes (Anker, 2001; Almeida et al., 2007), and further changes are expected in their nomenclatural status and geographical distribution (Anker et al., 2007). Therefore, a review of the alpheid cryptic fauna from the Mexican coasts using the multidisciplinary approaches mentioned is recommended in order to determine the taxonomic status of the species herein reported.

ACKNOWLEDGMENTS

Our gratitude to A. Reda Deara (Estación El Carmen–ICMyL) for his support in the field; to J. Romero Rodríguez, M. A. Carballido Carranza, A. Sánchez Quiñones and V. Vega González for processing partially the biological material, to the Dirección General de Vida Silvestre de la SEMARNAP and to the staff of the Reserva de la Biosfera de Sian Ka´an for providing facilities during the field work. Thanks to three anonymous referees who helped to improve the first draft of the manuscript.

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