Caprellids (Crustacea: Amphipoda: Caprellidae) from ... · Org. Divers. Evol. 5, Electr. Suppl. 10 (2005) Díaz & al.: Caprellids (Crustacea: Amphipoda) from Venezuela 2 and Fig

Org. Divers. Evol. 5, Electr. Suppl. 10 (2005)

Org. Divers. Evol. 5, Electr. Suppl. 10: 1 - 25 (2005)© Gesellschaft für Biologische SystematikURL: http://www.senckenberg.de/odes/05-10.htmURN: urn:nbn:de:0028-odes0510-4

Caprellids (Crustacea: Amphipoda: Caprellidae) from shallow waters of the Caribbean coast of VenezuelaYusbelly J. Díaza, José M. Guerra-Garcíab, *, Alberto Martína

a Universidad Simón Bolívar, Departamento Estudios Ambientales e INTECMAR, Apartado 89000, Caracas 1086-A, Venezuela

b Laboratorio de Biología Marina, Departamento Fisiología y Zoología, Facultad de Biología, Universidad de Sevil-la, Avda Reina Mercedes 6, 41012 Sevilla, Spain

* Corresponding author, e-mail: [email protected]

Received 30 April 2004 • Accepted 2 November 2004

Abstract

The caprellidean fauna of Venezuela is investigated. Twenty-seven stations in shallow waters of the states Falcón, Carabobo, Aragua, An-zoátegui, Sucre and Nueva Esparta were sampled, and the caprellids were sorted and identified. A total of 3984 specimens were identified; twelve species in eight genera are reported. Two species, Caprella penantis Leach, 1814 and Paracaprella digitimanus Quitete, 1971, are new records for Venezuelan waters. An identification key and illustrations are provided for all caprellid species known from Venezuela, as well as comments on their distribution and ecology.

Keywords: Crustacea; Amphipoda; Caprellidae; New records; Venezuela; Identification key

IntroductionThe caprellidean amphipods of Venezuela have been scarcely studied. McCain (1968) and McCain & Stein-berg (1970) reported the presence of six species: Ca-prella equilibra Say, 1818, Deutella incerta (Mayer, 1903), Hemiproto wigleyi McCain, 1968, Mayerella redunca McCain, 1968, Paracaprella pusilla Mayer, 1890, and Phtisica marina Slabber, 1769. Stoner & Lewis (1985), conducted an ecological study in the Ar-chipelago Los Roques and recorded the species Pseu-daeginella biscaynensis (McCain, 1968) and Hemiae-gina minuta Mayer, 1890. Villarroel & Graciani (1997) published the first record of Caprella danilevskii Czer-niavskii, 1868 for Venezuela. Recently, Guerra-García (2003a), as a part of the revision of the genus Deutella, described Deutella margaritae Guerra-García, 2003 based on material from Isla Margarita. Consequently, to our knowledge, only ten species have been reported from Venezuelan waters.

The lack of studies dealing with the caprellids from Venezuela is also applicable to other areas of Central and South America. McCain & Steinberg (1970) poin-ted out that the coasts of these areas are virtually unstu-

died; undoubtedly, many new records and species will be reported from there. Recent attempts to improve the knowledge of caprellids along these coasts have been carried out in Chile (Guerra-García 2001; Guerra-Gar-cía & Thiel 2001; Thiel et al. 2003), Brazil (Wakabara et al. 1991; Wakabara & Serejo 1998), and the Gulf of Mexico (Ortíz et al. 2002; Escobar-Briones & Win-field 2003). In addition, the Laboratory of Peracaridean Crustaceans from Simón Bolívar University, Venezu-ela, has been developing a sampling programme, focu-sing on collecting peracaridean crustaceans, especially amphipods, from the shallow waters of Venezuela. The caprellids from these collections have been studied, and the results are included in the present paper. Two new species records for Venezuelan waters are repor-ted, illustrations are provided for all twelve caprellid species known from Venezuela, and an identification key based on these figures is presented.

Material and methodsSamples were collected from a total of 27 stations along the Venezuelan coast from 1980-2003 (Table 1


Díaz & al.: Caprellids (Crustacea: Amphipoda) from Venezuela 2

and Fig. 1). All sites were sampled from the intertidal to 3m depth using snorkeling, SCUBA, van Veen grabs or trawling, except for station 3 where sampling was carried out at 15 m using a van Veen grab only. Samp-les of potentially suitable substrates for caprellids were collected (mainly algae, seagrasses, hydroids, bryozo-ans, mussels and oysters fouling, and sediments). All the samples were fixed in 70% ethanol, and the speci-mens were sorted using a stereo microscope.

Although the phylogeny and higher classification of the caprellids is still under debate (e.g. Laubitz 1993; Takeuchi 1993), we follow the recent classification of Myers & Lowry (2003) in the present paper, and our study has focused on members of the family Caprel-lidae.

The specimens examined for this study are deposi-ted in the following institutions:

MNCN = Museo Nacional de Ciencias Naturales, Madrid, Spain.

USB = Colección del Laboratorio de Crustáceos Pe-racáridos de la Universidad Simón Bolívar, Caracas, Venezuela.

USNM = National Museum of Natural History, Smithsonian Institution, Washington D.C., USA.

Abbreviated lists of synonymies are included for each species. A more extensive synonymy is found in McCain & Steinberg (1970). A lateral view of a gene-ralized caprellid (Fig. 2) demonstrates the morphologi-cal terms used in the key. In the illustrations for each species (Figs. 3-16), the respective most important cha-racteristics are indicated by arrows.

Station Locality Coordinates State

1 Playa Sur 10º55’13” N, 68º16’22” W Falcón (FA) 2 Cayo Sombrero 10º52’74” N, 68º12’54” W Falcón (FA) 3 Boca Grande 10º50’55” N, 68º13’29” W Falcón (FA) 4 Boca Seca 10º51’05” N, 68º14’07” W Falcón (FA) 5 Playa Caimán 10º61’10” N, 68º14’09” W Falcón (FA) 6 Tumba Cuatro 10º50’72” N, 68º15’57” W Falcón (FA) 7 Las Luisas 10º51’47” N, 68º17’69” W Falcón (FA) 8 Caño León 10º51’19” N, 68º19’50” W Falcón (FA) 9 Caño Capuchinos 10º49’40” N, 68º18’13” W Falcón (FA) 10 Planta Centro 10º30’00” N, 68º09’20” W Carabobo (CA) 11 Isla Larga 10º29’16” N, 67º57’00” W Carabobo (CA) 12 Veluz estación 2 10º30’16” N, 67º43’15” W Aragua (AR) 13 Veluz estación 4 10º30’21” N, 67º41’46” W Aragua (AR) 14 Veluz estación 5 10º30’24” N, 67º41’23” W Aragua (AR) 15 Petrozuata DII PDSE A-R1 11º03’01” N, 64º55’50” W Anzoátegui (AN) 16 MMCJ 14-00 B-08/r4 10º06’32” N, 64º47’19” W Anzoátegui (AN) 17 Chacopata 10º21’28” N, 64º20’49” W Sucre (SU) 18 Playa Medina 10º42’52” N, 63º00’45” W Sucre (SU) 19 Cipara 10º44’59” N, 62º41’57” W Sucre (SU) 20 Guayacán 10º39’08” N, 63º49’44” W Sucre (SU) 21 Isla Caribe 10º41’24” N, 63º51’07” W Sucre (SU) 22 Laguna de Punta de Piedras 10º53’55” N, 63º55’30” W Nueva Esparta (NE) 23 Puente de la Laguna de La

Restinga 10º58’44” N, 64º10’00” W Nueva Esparta (NE)

24 Ostrero La Salle 10º59’05” N, 64º09’50” W Nueva Esparta (NE) 25 Laguna de La Restinga 11º01’89” N, 63º49’33” W Nueva Esparta (NE) 26 Playa Guacuco 11º03’38” N, 63º48’77” W Nueva Esparta (NE) 27 Playa El Agua 11º08’61” N, 63º48’77” W Nueva Esparta (NE)

Table 1: List of stations sampled for the present study. See also Fig. 1.



List of speciesFamily Caprellidae Leach, 1814

Subfamily Caprellinae Leach, 1814Caprella danilevskii Czerniavskii, 1868Caprella equilibra Say, 1818Caprella penantis Leach, 1814Deutella incerta (Mayer, 1903)Deutella margaritae Guerra-García, 2003Hemiaegina minuta Mayer, 1890Mayerella redunca McCain, 1968Paracaprella digitimanus Quitete, 1971Paracaprella pusilla Mayer, 1890Pseudaeginella biscaynensis (McCain, 1968)

Subfamily Phtisicinae Vassilenko, 1968Hemiproto wigleyi McCain, 1968Phtisica marina Slabber, 1769

Key to the species of Caprellidae from VenezuelaThis key is meant as an easy field guide to be used without dissections. For illustration of morphological terms see Fig. 2.1. Gills on pereonites 2-4. Pereopods 3 and 4 six-arti-

culate .................................................................... 2– Gills on pereonites 3 and 4. Pereopods 3 and 4 redu-

ced (one- or two-articulate) or absent .................. 32. Abdomen with two pairs of appendages in males,

one pair in females ................................................. ................................... Hemiproto wigleyi (Fig. 15)– Abdomen with three pairs of appendages in males,

two pairs in females ................................................ ....................................... Phtisica marina (Fig. 16)3. Pereopod 5 reduced to three minute articles ...........

Mayerella redunca (Figs. 10, 11)– Pereopod 5 six-articulate ..................................... 44. Pereopods 3 and 4 absent .................................... 5– Pereopods 3 and 4 present, though reduced to one

or two articles ...................................................... 75. Head with a well-developed rostrum ....... Caprella

penantis (Fig. 5)– Head without a well-developed rostrum .............. 66. Pereonite 2 with a ventral projection between gna-

thopods 2. Propodus of pereopods 5-7 with grasping spines .......................... Caprella equilibra (Fig. 4)

– Pereonite 2 without ventral projection between gan-thopods 2. Propodus of pereopods 5-7 without gra-sping spines ............. Caprella danilevskii (Fig. 3)

7. Pereopods 3 and 4 one-articulate ......................... 8– Pereopods 3 and 4 two-articulate ........................ 98. Head smooth. Pereonite 2 with a ventral projection.

Abdomen with distinct appendages visible in lateral view .......................... Hemiaegina minuta (Fig. 9)

– Head and pereonite 1 with dorsal projections. Per-eonite 2 without ventral projection. Abdomen with shorter appendages ................................................ ....................Pseudaeginella biscaynensis (Fig. 14)

9. Head with one or two dorsal projections . Deutella incerta (spiny form) (Fig. 7)

– Head smooth, without dorsal projections .......... 1010. Antenna 1 longer than combined length of head

plus pereonites 1 and 2 ...................................... 11– Antenna 1 shorter than combined length of head

plus pereonites 1 and 2 ...................................... 1211. Anterolateral projections on pereonites 3 and 4 well

developed ................................................................ .................................. Deutella margaritae (Fig. 8)– Anterolateral projections on pereonites 3 and 4 not

well developed ....................................................... ................ Deutella incerta (smooth form) (Fig. 6)12. Propodus of gnathopod 2 with a ventral projection

medially .................................................................. ........................ Paracaprella digitimanus (Fig. 12)– Propodus of gnathopod 2 without ventral projec-

tion ......................... Paracaprella pusilla (Fig. 13)

Data on individual speciesSubfamily Caprellinae Leach, 1814

Caprella danilevskii Czerniavskii, 1868(Fig. 3)Caprella Danilevskii Czerniavskii, 1868: 92, pl. 6, figs. 21-34. – Mayer (1890: 58, pl. 5, fig. 44; pl. 7, figs. 12-13)

Caprella Danilewskii Czerniavskii. – Chevreux and Fage (1925: 454, fig. 432).

Caprella danilevskii Czerniavskii. – McCain (1968: 22, figs. 10-11); McCain and Steinberg (1970: 16); Ca-vedini (1982: 499); Krapp-Schickel (1993: 779, fig. 531); Camp (1998: 132); Guerra-García and Takeuchi (2002: 683-684, fig. 6).

Caprella inermis Haswell, 1879: 319-351.Material examined (76 specimens). Station 18: 2

females, 3 juveniles (USB-SU0187), 03/15/2001; 2 males, 1 female (USB-SU0263), 03/15/2001. Station 26: 2 males, 1 female, 1 juvenile (USB-NE0042), 08/13/2000. Station 27: 16 males, 48 females (USB-NE0048), 1 male, 1 female (both used for lateral-view figures) (MNCN 20.04/5894), 08/15/2000.

Remarks. Although Caprella danilevskii is widely distributed around the world (McCain and Steinberg 1970; Guerra-García and Takeuchi 2004), the first re-cord for Venezuela is recent (Villarroel and Graziani 1997). The Venezuelan specimens are in agreement with the Mediterranean material described by Krapp-Schickel (1993) and the Atlantic specimens figured by McCain (1968). The distinctive male abdomen, the



absence of grasping spines on the pereopods, and the short dactylus of male gnathopod 2 distinguishes this species from other species of Caprella. The life history of C. danilevskii is one of the most well known in the Caprellidae; Takeuchi and Hirano (1991) studied the growth and reproduction of this species based on Japa-nese specimens.

Habitat. This species has been found living on al-gae, seagrasses, sponges, tunicates and bryozoans (Mc-Cain 1968). In the present study, Caprella danilevskii was found clinging on the algae Sargassum sp., Acan-tophora sp., Gracilaria sp., Ulva lactuca Linnaeus, 1753, and Hypnea muscifornis J.V. Lamouroux, 1813.

Caprella equilibra Say, 1818(Fig. 4)

Caprella equilibra Say, 1818: 391-392. – McCain (1968: 25, figs. 12-13); McCain and Steinberg (1970: 19); Cavedini (1982: 500); Krapp-Schickel (1993: 782-783, fig. 533); Camp (1998: 132); Guerra-García and Thiel (2001: 878-879, fig. 6).

Caprella aequilibra Say. – Mayer (1882: 45, pl. 1, fig. 7; pl. 2, fig. 1-11; pl. 4, figs. 20-25; pl. 5, figs. 16-18); Chevreux and Fage (1925: 455, fig. 433).

Material examined (390 specimens). Station 17: 27 juveniles (USB-SU0544), 05/09/2003. Station 19: 1 female, 2 juveniles (USB-SU0208), 03/18/2003; 15 males, 14 females, 21 juveniles (USB-SU0254), 03/18/2003. Station 20: 1 male, 4 females (USB-SU0347), 1 male, 1 female (both used for lateral-view figures) (MNCN 20.04/5895), 05/08/2003; 148 speci-mens (USB-SU0348), 05/08/2003; 100 specimens (USB-SU0349), 05/08/2003; 11 males, 15 females, 30 juveniles (USB-SU0555), 05/06/2003; 1 male (USB-SU0656), 05/06/2003.

Remarks. Caprella equilibra, similarly to C. dani-levskii, is widely distributed around the world (McCain and Steinberg 1970; Krapp-Schickel 1993). The Vene-zuelan specimens have a distinctive ventral projection between gnathopods 2 that is characteristic for this species. The presence of C. equilibra in Venezuela was reported by McCain (1968), based on specimens from Puerto Cabello, Carabobo state. All the material inclu-ded in the present study is from Sucre state.

Habitat. Caprella equilibra has been found on seagrasses, algae, sponges, hydroids, bryozoans, co-lonial ascidians and alcyonarians, from the intertidal to 3000m depth (Krapp-Schickel 1993). The species is also found on artificial devices such as aquaculture nets, water duct pipes, power plants, as well as on floa-ting substrates, e.g. buoys and drifting algae (Takeuchi and Sawamoto 1998; Thiel et al. 2003). Caprella equi-libra feeds principally by filtering, frequently using grooming behaviour (Guerra-García et al. 2002). In the present study, this species was found on mussels, tur-

key wings (Arca zebra (Swainson, 1833)), on Thalas-sia testudinum Banks ex König, 1805, and algae.

Caprella penantis Leach, 1814(Fig. 5)Caprella Penantis Leach, 1814: 404.

Caprella acutifrons Mayer, 1882: 48. – Mayer (1890: 50, pl. 2, figs. 36-37, 39-41, pl. 4, figs. 52-53, 55, 57-61, 65-69; including the ‘forms’ neglecta, tabi-da, gibbosa, carolinensis, lusitanica, virginia); Mayer (1903: 79, pl. 3, figs. 4-28; pl. 7, figs. 62-65).

Caprella penantis Leach. – McCain (1968: 33, figs. 15-16); McCain and Steinberg (1970: 33); Cavedini (1982: 508); Krapp-Schickel (1993: 791-793); Camp (1998: 132); Guerra-García and Takeuchi (2002: 692-693, fig. 12).

Material examined (1 specimen). Station 19: 1 male (used for lateral-view figure) (USB-SU0281), 03/13/2001.

Remarks. The specimen represents the first record of Caprella penantis from Venezuela. The species has been recorded under several species- or subspecies na-mes from temperate regions worldwide (McCain and Steinberg 1970). Further morphological and molecular studies are required to evaluate the respective status of these nominal species or subspecies (McCain 1968; Laubitz 1972; Takeuchi and Hirano 1995).

Habitat. This species has been found on red and brown algae, seagrasses such as Posidonia, on hydro-ids, alcyonarians, zoantharians, bryozoans, sponges, and attached to echinoderms (Arbacia) and decapods (Libinia) (McCain 1968; Krapp-Schickel 1993). The specimen found during the present study was collected among mussels, Perna perna (Linnaeus, 1758) and P. viridis (Linnaeus, 1758).

Deutella incerta (Mayer, 1903)(Figs. 6,7)

Luconacia incerta Mayer, 1903: 49-50, pl. 2, figs. 11-14; pl.6, figs. 73-75; pl. 9, figs. 21, 40, 57. – McCain (1968: 53-54, 68-72, figs. 33-35); McCain and Stein-berg (1970: 53).

Protellopsis stebbingii Kunkel, 1910: 111-113, fig. 43.

Deutella incerta (Mayer). – Steinberg & Dougherty, 1957: 281, 285-286. – Gable and Lazo-Wasem (1987: 635-636, fig. 4); Camp (1998: 132); Guerra-García (2003a: 1062-1065, fig. 3).

Material examined (113 specimens). Station 1: 1 female (USB-FA0632), 05/08/2000; 1 male, 1 fema-le, 1 juvenile (USB-FA0787), 08/02/2000; 3 males, 2 females, 1 juvenile (USB-FA0965), 08/02/2000; 1 juvenile (USB-FA1024), 08/02/2000; 1 female (USB-FA1464), 07/16/2001. Station 4: 1 male, 1 fe-male (USB-FA0187), 12/1993. Station 6: 1 male, 4



females, 3 juveniles (USB-FA0544), 02/08/2000; 1 female (USB-FA0630), 05/2000; 1 male, 1 female, 2 juveniles (USB-FA0701), 08/02/2000; 2 females, 3 ju-veniles (USB-FA0784), 08/02/2000; 1 juvenile (USB-FA0785), 08/02/2000; 1 juvenile (USB-FA0786), 08/02/2000; 1 juvenile (USB-FA0788), 08/02/2000; 1 juvenile (USB-FA0999), 02/12/2001; 1 male (USB-FA1010), 11/05/2000; 1 juvenile (USB-FA1011); 1 fe-male (USB-FA1017), 11/05/2000. Station 7: 1 female (USB-FA0192), 12/1993; 1 juvenile (USB-FA1480), 05/08/2000. Station 8: 1 female (USB-FA0305), 10/02/1999; 1 female (USB-FA0607), 05/08/2000; 5 males, 1 female (USB-FA0634), 1 male, 1 female (used for lateral-view figure, Fig. 6) (MNCN 20.04/5896), 05/08/2000; 2 females (USB-FA0950), 08/02/2000; 1 juvenile (USB-FA0976), 02/12/2001; 1 juvenile (USB-FA1007), 11/05/2000; 4 juveniles (USB-FA1008), 11/05/2000; 1 female (USB-FA1452), 07/16/2001. Sta-tion 9: 7 males, 6 females, 4 juveniles (USB-FA0633), 05/08/2000; 1 male, 2 females (USB-FA0708), 08/02/2000; 1 female (USB-FA0940), 08/02/2000; 4 males, 2 females (USB-FA1014), 11/05/2000; 1 ju-venile (USB-FA1015), 11/05/2000; 2 females (USB-FA1019), 11/2000; 1 male (USB-FA1020), 11/05/2000; 1 female (USB-FA1239), 07/16/2001; 5 males, 2 fema-les, 6 juveniles (USB-FA1479), 06/1993. Station 12: 1 female (USB-AR0247), 07/25/1980. Station 13: 1 female (USB-AR0302), 08/1981. Station 15: 1 male, 2 females (USB-AN0059), 01/1999. Station 16: 1 female (USB-AN0094), 03/24/2000. Type material (Fig. 7): 1 male, 1 female (paralectotypes) (USNM 026001) coll-ected from 29º28’N, 87º56’W, Albatros, station 2389, 49 meters.

Remarks. Deutella incerta is widely distributed in the temperate and tropical areas of the western North Atlantic (McCain 1968) and seems to be one of the most common caprellids in Bermuda (Gable and Lazo-Wasem 1987). McCain (1968) had already reported the presence of this species in Venezuelan waters. A detailed discussion regarding the status of D. incerta is included in Guerra-García (2003a). McCain (1968) pointed out that this species varies considerably in the degree of body spination; larger individuals bear dor-sal projections, but the latter are absent in the smaller specimens. All the specimens examined for the present study lack dorsal projections, and their morphology is in agreement with the small specimens figured by McCain (1968), but differs from the type specimens, which are provided with distinct projections (Fig. 7).

Habitat. This species has been collected on man-grove roots, Sargassum sp., Thalassia sp., sponges, hydroids, alcyonarians and ascidians, and occasional-ly it has been taken in plankton tows (Guerra-García

2003a). The material studied here was collected from sandy bottoms and from Thalassia testudinum.

Deutella margaritae Guerra-García, 2003(Fig. 8)

Deutella margaritae Guerra-García, 2003a: 1065-1070, figs. 4-7.

Material examined (966 specimens). Station 22: 1 male (USB-NE0007), 04/08/1998. Station 23: 34 males, 15 females, 53 juveniles (USB-NE0192), 12/18/2003; 14 males, 8 females, 97 juveniles (USB-NE0193), 12/18/2003; 25 males, 30 females, 64 ju-veniles (USB-NE0194), 12/18/2003; 230 specimens (USB-NE0195), 12/18/2003; 156 specimens (USB-NE0202), 12/18/2003. Station 24: 13 males, 11 fema-les, 7 juveniles (USB-NE0174), 12/18/2003. Station 25: 1 female (USB-NE0008), 07/05/1998; 25 males, 9 females, 30 juveniles (USB-NE0157), 11/05/2003; 23 males, 27 females, 4 juveniles (USB-NE0180), 11/05/2003; 30 males, 35 females, 14 juveniles (USB-NE0187), 11/05/2003. Type material (Fig. 8): Holo-type male (USNM 1000206), allotype female (USNM 1000207) and paratypes (3 males, 7 females, USNM 1000208), 01/11/1964 from Puente de la Restinga, Isla Margarita, Venezuela.

Remarks. This is the first record of Deutella mar-garitae after the original description (Guerra-García 2003a), based on material from Isla Margarita, Vene-zuela. The specimens have been assigned to D. mar-garitae mainly on the basis of the similar morphology of pereopods 3 and 4, mouthparts and abdomen, and the presence of distinctive anterolateral projections on pereonites 2-4. However, in the examined specimens pereopod 5 is inserted at midlength of the pereonite 5 (similar to Deutella incerta) and not on the posterior end as in the type specimens. We have found a con-siderable degree of variation in Deutella species from Venezuela, thus do not discount the possibility that a complex of several more species of Deutella may exist there. Further morphological and, especially, molecular studies would be necessary to clarify the systematics of Deutella in Venezuelan waters. At this time, we maintain the two species already recorded from Vene-zuela, and prefer not to erect any new species without clear evidence of discrete characteristics.

Habitat. The type material was found in boulders near Rhizophora sp. The material examined in the present study was collected from oysters and hydro-ids, Halocordyle disticha (Goldfuss, 1820), on roots of Rhizophora mangle L.

Hemiaegina minuta Mayer, 1890(Fig. 9)



Hemiaegina minuta Mayer, 1890: 40, pl. 1, figs. 25–27; pl. 3, figs. 32–35;pl. 5, figs. 52–53; pl. 6, figs. 13, 33–34, pl. 7, fig. 4. – McCain (1968: 61–64, figs. 29–30); McCain and Steinberg (1970: 51); Gable and Lazo-Wasem (1987: 637); Müller (1990: 836); Serejo (1997: 631, fig. 1); Camp (1998: 132); Guerra-García (2003b: 6-7, fig. 3; 2003c: 105-106, fig.10).

Hemiaegina quadripunctata Sundara Raj, 1927: 126–127, pl. 18.

Hemiaegina costai Quitete, 1972: 165–168, pls. 1–2.

Material examined (3 specimens). Station 20: 1 juvenile (USB-SU0722), 1 male, 1 female (both used for lateral-view figures) (MNCN 20.04/5897), 05/06/2003.

Remarks. Hemiaegina minuta is widely distributed in tropical and temperate waters of the world’s oceans (Müller 1990). The species was reported from Vene-zuela by Stoner and Lewis (1985) – from Gran Roque Island, Archipiélago Los Roques (11º55’N, 66º40’ W) – during an ecological study. The material studied from Sucre state is scarce and in poor condition (e.g. male lacks gnathopods 2).

Habitat. Hemiaegina minuta has been collected from Sargassum sp. and taken in plankton tows (Mc-Cain and Steinberg 1970). Müller (1990) reported H. minuta as preferring more or less exposed reef locati-ons. The species has been collected also from Thalas-sia testudinium (Stoner and Lewis 1985). Guerra-Gar-cía (2003b) found it associated with Turbinaria ornata (Turner) J. Agardh on the coast of Mauritius, and Guer-ra-García (2003c) reported it from Papua New Guinea, living on Dictyota sp., Halimeda sp., Gracilaria sp., Galaxaura sp., and Amansia glomerata (Agardh) Nor-ris, 1979. Hemiaegina minuta also has been found as-sociated with many different substrata in Queensland: green algae such as Halimeda sp., brown and red algae, sponges, tunicates, seagrasses such as Posidonia sp., dead corals encrusted with algal turf, and under small boulders (Guerra-García, personal observations). The specimens studied in the present work were found on turkey wings (Arca zebra).

Mayerella redunca McCain, 1968(Figs. 10-11)

Mayerella redunca McCain, 1968: 75-78, figs. 37-38, 50. – McCain and Steinberg (1970: 53); Guerra-García (2003d: 189, 191, 194, figs. 8, 11).

Material examined (5 specimens). Station 5: 1 female (USB-FA0216), 03/29/2000. Station 10: 1 fe-male (used for lateral-view figure, Fig. 10) (MNCN 20.04/5898), 12/1981; 1 juvenile (USB-CA0445), 01/1982. Type material (Fig. 11): Holotype male (USNM 120176), allotype female (USNM 120177)

collected from North of Coche Island, Nueva Esparta state, Venezuela, 10º50’N, 63º54’W, 04/15/1939.

Remarks. The specimens reported here represent the first record of the species from Venezuela since the original description by McCain (1968). As far as is known, Mayerella redunca is endemic to Venezuela. A revision of the genus Mayerella can be found in Guer-ra-García (2003d).

Habitat. McCain (1968) reported that specimens were collected between 35m and 60m depth, but he did not mention any substrates. The female from Falcon state was collected from sandy bottoms, the remaining two specimens in plankton samples.

Paracaprella digitimanus Quitete, 1971(Fig. 12)

Paracaprella digitimanus Quitete, 1971: 161, figs. 1-3. – Wakabara and Serejo (1998: 585).

Material examined (2 specimens). Station 20: 1 male (MNCN 20.04/5899), 1 juvenile (USB-SU0721), 03/13/2001.

Remarks. The original description of Paracaprella digitimanus was based on material from Brazil (Quite-te 1971); the present finding is the first for Venezuela. Although the genus Paracaprella is still awaiting revi-sion, P. digitimanus can be easily distinguished from its congeners by a striking projection on the propodus of gnathopod 2.

Habitat. Quitete (1971) collected Paracaprella di-gitimanus from hydroids. The two specimens reported here were found on mussels (Perna perna and P. viri-dis).

Paracaprella pusilla Mayer, 1890(Fig. 13)

Paracaprella pusilla Mayer, 1890: 41, pl. 1, figs. 28-30; pl. 3; figs. 45-47; pl. 5, figs. 48-49; pl. 6, fig. 10; 1903: 67, pl. 2, figs. 36-37; pl. 7, fig. 52. – Stein-berg and Dougherty (1957: 283-284, figs. 16, 19, 24, 30); McCain (1968: 82-86, figs. 41-42); Wakabara et al. (1991: 73); Camp (1998: 132); Guerra-García and Thiel (2001: 880, fig. 8).

Caprella nigra Reid, 1951: 283-284, 289, fig. 58.Material examined (2324 specimens). Station 2: 1

female (USB-FA0564), 08/30/2000. Station 10: 1 juve-nile (USB-CA0054), 01/1982; 1 male (USB-CA0055), 04/1982; 1 juvenile (USB-CA0065), 04/1982; 1 fema-le (USB-CA0071), 02/1982; 1 female (USB-CA0099), 01/1986; 1 juvenile (USB-CA0132), 05/1982; 1 fema-le (used for lateral-view figure) (MNCN 20.04/5900), 04/1982; 1 male (USB-CA0143), 04/1982; 1 juvenile (USB-CA0169); 1 juvenile (USB-CA0212); 1 male (USB-CA0237); 2 juveniles (USB-CA0300), 07/1982; 2 males, 1 female (USB-CA0305), 10/1982; 3 males (USB-CA0312), 11/1982; 4 males, 4 females, 1 juvenile



(USB-CA0316), 08/1982; 3 juveniles (USB-CA0321), 12/1982; 1 male, 2 juveniles (USB-CA0334), 1 male (MNCN 20.04/5901), 11/1982; 3 males (USB-CA0465), 03/1981. Station 11: 6 males, 13 females, 8 juveniles (USB-CA0431), 1 male, 1 female (MNCN 20.04/5309), 08/29/2000. Station 14: 1 female (USB-AR0013), 07/24/1980. Station 20: 22 males, 14 females, 86 juve-niles (USB-SU0346), 1 male (used for lateral-view fi-gure), 1 female (MNCN 20.04/5902), 05/08/2003; 210 specimens (USB-SU0350), 05/08/2003; 213 specimens (USB-SU0351), 05/08/2003; 201 specimens (USB-SU0352), 05/08/2003; 169 specimens (USB-SU0353), 05/08/2003; 193 specimens (USB-SU0354), 08/05/2003; 215 specimens (USB-SU0355), 05/08/2003; 255 speci-mens (USB-SU0356), 05/08/2003; 209 specimens (USB-SU0357), 05/08/2003; 147 specimens (USB-SU0358), 05/08/2003; 194 specimens (USB-SU0366), 05/08/2003. Station 21: 2 females (USB-SU0575), 05/07/2003. Station 23: 46 males, 49 females, 21 ju-veniles (USB-NE0210), 12/18/2003. Station 24: 2 ma-les (USB-NE0213), 12/18/2003. Station 25: 1 female (USB-NE0214), 11/05/2003; 2 males, 2 females (USB-NE0215), 11/05/2003.

Remarks. Mayer (1890) described Paracaprella pusilla based on material collected from Brazil. Later, the species was reported from Venezuela by McCain (1968). Paracaprella pusilla is similar to Paracaprella tenuis Mayer, 1903; however, males of the former can be distinguished by the large, sharp-pointed projection on the anteroventral margin of pereonite 2, the proxi-mal knob on the basis of gnathopod 2, and by the pres-ence of setae on the dactylus of gnathopod 2 (McCain 1968). Large males of P. pusilla are very similar to lar-ge males of P. barnardi McCain, 1967 in that both bear a small anterodorsal tubercle on pereonite 2; however, the tubercle is not as well developed in the former as in the latter species, and the ventrolateral projection on the anterior margin of pereonite 2 is much larger in P. pusilla (see McCain 1967).

Habitat. Paracaprella pusilla has been collected from mangrove roots, seagrasses, hydroids and ascidi-ans (McCain 1968). The specimens collected here were found on gravel bottoms, ropes, mussels, oysters, sa-bellariid worm rock (Phragmatopoma lapidosa (Kin-berg, 1867)), and on hydroids (e.g. Halocordyle disti-cha) associated with mangrove roots.

Pseudaeginella biscaynensis (McCain, 1968)(Fig. 14)

Fallotritella biscaynensis McCain, 1968: 58-61, figs. 27-28, 53. – McCain and Steinberg (1970: 51); Gable and Lazo-Wasem (1987: 637-638).

Pseudaeginella biscaynensis (Mc Cain). – Laubitz (1995: 88); Camp (1998: 132).

Remarks. Laubitz (1995) considered Fallotritella as synonymous with Pseudaeginella. Pseudaeginella bis-caynensis is common in Caribbean waters (McCain and Steinberg 1970). In spite of the considerable number of stations sampled along the Venezuelan coast during the present work, we have not found P. biscaynensis. Ho-wever, the species has been reported from Venezuela by Stoner and Lewis (1985), based on material from Gran Roque Island, Archipiélago Los Roques (11º55’ N, 66º40’ W), collected during an ecological study. The figures included here are taken from McCain (1968) and based on the type material from Florida.

Habitat. The species is associated with red algae (McCain 1968), the green alga Avrainvillea sp. and the seagrass Thalassia sp. (Gable and Lazo-Wasem 1987; Stoner and Lewis 1985), with corals, Cymodocea sp. and Syringodium sp. (Guerra-García 2002), and with the seaweed Galaxaura sp. (Guerra-García 2003c).

Subfamily Phtisicinae Vassilenko, 1968

Hemiproto wigleyi McCain, 1968(Fig. 15)

Hemiproto wigleyi McCain, 1968: 65-68, figs. 31, 32(c-e), 50. – McCain and Steinberg (1970: 51); Camp (1998: 132).

Material examined (13 specimens). Station 6: 1 female (USB-FA1012), 11/05/2000; 1 female (USB-FA1016), 11/05/2000; 1 female (USB-FA1018), 11/05/2000; 1 male (USB-FA1123), 08/02/2000; 1 ju-venile (USB-FA1481), 02/12/2000. Station 7: 1 male, 1 female (USB-FA0609), 1 female (used for lateral-view figure) (MNCN 20.04/5903), 05/08/2000; 1 male (used for lateral-view figure) (MNCN 20.04/5904), 05/08/2000. Station 8: 1 female (USB-FA1009), 11/05/2000; 1 female (USB-FA1025), 11/05/2000. Sta-tion 9: 1 male (USB-FA0012), 06/1993. Station 21: 1 female (MNCN 20.04/5905), 05/07/2003.

Remarks. Hemiproto is, so far, a monotypic genus and includes the single species H. wigleyi, described by McCain (1968). This species is known from Coche Island, Nueva Esparta state, Venezuela, collected at 35-60m depth, and from Fort Lauderdale, Florida, at 32 m. Watling (1997) reported H. wigleyi from the San-ta Maria Basin region, California. We have assigned the specimens collected from Falcon state to H. wigley mainly on the basis of the combination of the following characteristics: molar absent; mandibular palp three-articulate; pereopods 3 and 4 six-articulate; pereopod 5 five-articulate; abdomen with two pairs of uni-arti-culate appendages in males (one of them reduced), and only one pair in females.

Habitat. The ecology of this species is unknown. McCain (1968) only reported that specimens were col-



lected between 35m and 60 m depth, but he did not mention any substrates. The specimens reported here were collected from sandy bottoms and living on Tha-lassia testudinum and Halimeda opuntia J.V. Lamou-roux, 1816.

Phtisica marina Slabber, 1769(Fig. 16)

Phtisica marina Slabber, 1769: 77, pl. 10. – Chev-reux and Fage (1925: 434, fig. 422); McCain (1968: 91, figs. 46-47); McCain and Steinberg (1970: 65); Camp (1998: 132); Guerra-García and Takeuchi (2002: 705).

Proto brunneovittata Haller, 1879: 231. – Haller (1880: 339, pl. 22, figs. 19-22).

Proto pedata Haller, 1879: 230. – Haller (1880: 398).

Proto ventricosa Mayer, 1882: 22, pl. 1, fig. 1; pl. 3, figs. 16-29; pl. 4, figs. 12-13; pl.5, figs. 1-5. – Mayer (1890: 12, pl. 3, figs. 4-6; pl. 5, figs. 3-6; pl. 6, fig. 1; pl. 7, fig. 1); Mayer (1903: 20, pl. 6, fig. 23).

Material examined (1 specimen). Station 3: 1 im-mature female (used for lateral-view figure) (USB-FA0465), 05/25/2000.

Remarks. Phtisica marina is widely distributed in the Atlantic and Pacific Oceans, and in the Mediterra-nean Sea (McCain 1968; Krapp-Schickel 1993). Mc-Cain (1968) reported the species from Cubagua and Margarita Islands, Nueva Esparta state, Venezuela. Phtisica marina can be differentiated from Hemiproto wigleyi mainly by the abdomen (see also the remarks under H. wigleyi).

Habitat. This species has been reported from green and brown algae, seagrasses, sponges, hydroids, bryo-zoans, echinoderms (Asteroidea), and to a depth of 660 m (McCain 1968; Krapp-Schickel 1993). The speci-men reported here was collected from sandy bottoms at 15m depth. A predatory mode of life is suggested for this species (Guerra-García et al. 2002).

AcknowledgementsWe are very grateful to J. Gutiérrez (“Fundación La Salle de Ciencias Naturales”, Venezuela), J. Bolaños (“Universidad de Oriente”, Venezuela), and W. Fera-gotto (“Benthos Buceo Profesional”, Venezuela) for providing some of the samples. Thanks are also due to the “Agenda Parque Nacional Morrocoy”-FONACIT, Venezuela (project number 96001837-V), and to the University of Sevilla, Spain (“III Plan Propio de Apoyo a Programas Internacionales”) for financial support.

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72º 60º

12º

2º

ATLANTICOCEAN

CARIBBEAN SEA

COLOMBIA

BRAZIL

VENEZUELA

N

1-9

10

11

12-1416

1523-25

26-2722

18-1920-21

17

Los Roques

Fig. 1: Map of Venezuela, showing the sampling localities. See also Table 1.



Fig. 2: Lateral view of a generalized caprellid (modified from McCain 1968).



Fig. 3: Caprella danilevksii Czerniavskii, 1868. Lateral view. A. Male. B. Female (MNCN 20.04/5894). Scale bar: 1 mm.



Fig. 4: Caprella equilibra Say, 1818. Lateral view. A. Male. B. Female (MNCN 20.04/5895). Scale bar: 1 mm.



Fig. 5: Caprella penantis Leach, 1814. Male, lateral view (USB-SU0281). Scale bar: 1 mm.



Fig. 6: Deutella incerta (Mayer, 1903). Lateral view. A. Male. B. Female (MNCN 20.04/ 5896). Scale bar: 1 mm.



Fig. 7: Deutella incerta (Mayer, 1903). Paralectotypes (USNM 026001). A. Male, lateral view. B. Female, lateral view. C. Male gnathopod 2. D. Female gnathopod 2 (refigured from Guerra-García 2003a). Scale bars: 1 mm.



Fig. 8: Deutella margaritae Guerra-García, 2003. Lateral view. A, Holotype male (USNM 1000206). B. Allotype fema-le (USNM 1000207) (refigured from Guerra-García 2003a). Scale bar: 1 mm.



Fig. 9: Hemiaegina minuta Mayer, 1890. Lateral view. A. Male. B. Female (MNCN 20.04/5897). Scale bar: 1 mm.



Fig. 10: Mayerella redunca McCain, 1968. Female (MNCN 20.04/5898), lateral view. Scale bar: 1 mm.



Fig. 11: Mayerella redunca McCain, 1968. A, C. Holotype male (USNM 120176). B, D. Allotype female (USNM 120177). A. Male, lateral view. B. Female, lateral view. C. Male gnathopod 2. D. Female gnathopod 2 (refigured from Guerra-García 2003d). Scale bars: 1 mm.



Fig. 12: Paracaprella digitimanus Quitete, 1971. A. Male (MNCN 20.04/5899), lateral view. B. Juvenile (USB-SU0721), lateral view; C, male gnathopod 2. Scale bars: A, B: 1 mm; C: 0.5 mm.



Fig. 13: Paracaprella pusilla Mayer, 1890. A. Male (MNCN 20.04/5902), lateral view. B. Female (MNCN 20.04/5900), lateral view. C. Male gnathopod 2. Scale bars: A, B: 1 mm; C: 0.5 mm.



Fig. 14: Pseudaeginella biscaynensis (McCain, 1968). Lateral view. A. Holotype male (USNM 120179). B. Allotype female (USNM 120180) (refigured from McCain 1968). Scale bar: 1 mm.



Fig. 15: Hemiproto wigleyi McCain, 1968. A. Male (MNCN 20.04/5904), lateral view. B. Female (MNCN 20.04/5903), lateral view. C. Male abdomen, ventral view. D. Female abdomen, ventral view. Scale bars: A, B: 1 mm; C, D: 0.05 mm.



Fig. 16: Phtisica marina Slabber, 1769. Premature female (USB-FA0465). A. Lateral view. B. Abdomen, ventral view. Scales bar: A: 1 mm; B: 0.1 mm.

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Caprellids (Crustacea: Amphipoda: Caprellidae) from ... · Org. Divers. Evol. 5, Electr. Suppl. 10 (2005) Díaz & al.: Caprellids (Crustacea: Amphipoda) from Venezuela 2 and Fig