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Home , Caprella equilibra, Caprella penantis, Caprellidae, Coche Island

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

Caprellids (Crustacea: : ) from shallow waters of the coast of

Yusbelly 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 were sampled, and the caprellids were sorted and identified. A total of 3984 specimens were identified; twelve in eight genera are reported. Two species, 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

Introduction The caprellidean amphipods of Venezuela have been died; undoubtedly, many new records and species will scarcely studied. McCain (1968) and McCain & Stein- be reported from there. Recent attempts to improve the berg (1970) reported the presence of six species: Ca- knowledge of caprellids along these coasts have been prella equilibra Say, 1818, Deutella incerta (Mayer, carried out in Chile (Guerra-García 2001; Guerra-Gar- 1903), Hemiproto wigleyi McCain, 1968, Mayerella cía & Thiel 2001; Thiel et al. 2003), Brazil (Wakabara redunca McCain, 1968, Paracaprella pusilla Mayer, et al. 1991; Wakabara & Serejo 1998), and the Gulf 1890, and Phtisica marina Slabber, 1769. Stoner & of Mexico (Ortíz et al. 2002; Escobar-Briones & Win- Lewis (1985), conducted an ecological study in the Ar- field 2003). In addition, the Laboratory of Peracaridean chipelago Los Roques and recorded the species Pseu- from Simón Bolívar University, Venezu- daeginella biscaynensis (McCain, 1968) and Hemiae- ela, has been developing a sampling programme, focu- gina minuta Mayer, 1890. Villarroel & Graciani (1997) sing on collecting peracaridean crustaceans, especially published the first record of Caprella danilevskii Czer- amphipods, from the shallow waters of Venezuela. The niavskii, 1868 for Venezuela. Recently, Guerra-García caprellids from these collections have been studied, (2003a), as a part of the revision of the genus Deutella, and the results are included in the present paper. Two described Deutella margaritae Guerra-García, 2003 new species records for Venezuelan waters are repor- based on material from Isla Margarita. Consequently, ted, illustrations are provided for all twelve caprellid to our knowledge, only ten species have been reported species known from Venezuela, and an identification from Venezuelan waters. key based on these figures is presented. The lack of studies dealing with the caprellids from Venezuela is also applicable to other areas of Central Material and methods and South America. McCain & Steinberg (1970) poin- Samples were collected from a total of 27 stations ted out that the coasts of these areas are virtually unstu- along the Venezuelan coast from 1980-2003 (Table 1

Org. Divers. Evol. 5, Electr. Suppl. 10 (2005) Díaz & al.: Caprellids (Crustacea: Amphipoda) from Venezuela 2 and Fig. 1). All sites were sampled from the intertidal MNCN = Museo Nacional de Ciencias Naturales, to 3m depth using snorkeling, SCUBA, van Veen grabs Madrid, Spain. or trawling, except for station 3 where sampling was USB = Colección del Laboratorio de Crustáceos Pe- carried out at 15 m using a van Veen grab only. Samp- racáridos de la Universidad Simón Bolívar, Caracas, les of potentially suitable substrates for caprellids were Venezuela. collected (mainly algae, seagrasses, hydroids, bryozo- USNM = National Museum of Natural History, ans, mussels and oysters fouling, and sediments). All Smithsonian Institution, Washington D.C., USA. the samples were fixed in 70% ethanol, and the speci- Abbreviated lists of synonymies are included for mens were sorted using a stereo microscope. each species. A more extensive synonymy is found in Although the phylogeny and higher classification of McCain & Steinberg (1970). A lateral view of a gene- the caprellids is still under debate (e.g. Laubitz 1993; ralized caprellid (Fig. 2) demonstrates the morphologi- Takeuchi 1993), we follow the recent classification of cal terms used in the key. In the illustrations for each Myers & Lowry (2003) in the present paper, and our species (Figs. 3-16), the respective most important cha- study has focused on members of the family Caprel- racteristics are indicated by arrows. lidae. The specimens examined for this study are deposi- ted in the following institutions:

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

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

Org. Divers. Evol. 5, Electr. Suppl. 10 (2005) Díaz & al.: Caprellids (Crustacea: Amphipoda) from Venezuela 3

List of species – Head and pereonite 1 with dorsal projections. Per- eonite 2 without ventral projection. Abdomen with Family Caprellidae Leach, 1814 shorter ...... Pseudaeginella biscaynensis (Fig. 14) Subfamily Caprellinae Leach, 1814 9. Head with one or two dorsal projections . Deutella Caprella danilevskii Czerniavskii, 1868 incerta (spiny form) (Fig. 7) Say, 1818 – Head smooth, without dorsal projections ...... 10 Leach, 1814 10. 1 longer than combined length of head Deutella incerta (Mayer, 1903) plus pereonites 1 and 2 ...... 11 Deutella margaritae Guerra-García, 2003 – Antenna 1 shorter than combined length of head Hemiaegina minuta Mayer, 1890 plus pereonites 1 and 2 ...... 12 Mayerella redunca McCain, 1968 11. Anterolateral projections on pereonites 3 and 4 well Paracaprella digitimanus Quitete, 1971 developed ...... Paracaprella pusilla Mayer, 1890 ...... Deutella margaritae (Fig. 8) Pseudaeginella biscaynensis (McCain, 1968) – Anterolateral projections on pereonites 3 and 4 not well developed ...... Subfamily Phtisicinae Vassilenko, 1968 ...... Deutella incerta (smooth form) (Fig. 6) Hemiproto wigleyi McCain, 1968 12. Propodus of gnathopod 2 with a ventral projection Phtisica marina Slabber, 1769 medially ...... Paracaprella digitimanus (Fig. 12) Key to the species of Caprellidae from Venezuela – Propodus of gnathopod 2 without ventral projec- This key is meant as an easy field guide to be used tion ...... Paracaprella pusilla (Fig. 13) without dissections. For illustration of morphological terms see Fig. 2. Data on individual species 1. Gills on pereonites 2-4. Pereopods 3 and 4 six-arti- Subfamily Caprellinae Leach, 1814 culate ...... 2 – Gills on pereonites 3 and 4. Pereopods 3 and 4 redu- Caprella danilevskii Czerniavskii, 1868 ced (one- or two-articulate) or absent ...... 3 (Fig. 3) 2. Abdomen with two pairs of appendages in males, Caprella Danilevskii Czerniavskii, 1868: 92, pl. 6, figs. one pair in females ...... 21-34. – Mayer (1890: 58, pl. 5, fig. 44; pl. 7, figs. 12- ...... Hemiproto wigleyi (Fig. 15) 13) – Abdomen with three pairs of appendages in males, Caprella Danilewskii Czerniavskii. – Chevreux and two pairs in females ...... Fage (1925: 454, fig. 432)...... Phtisica marina (Fig. 16) Caprella danilevskii Czerniavskii. – McCain (1968: 3. Pereopod 5 reduced to three minute articles ...... 22, figs. 10-11); McCain and Steinberg (1970: 16); Ca- Mayerella redunca (Figs. 10, 11) vedini (1982: 499); Krapp-Schickel (1993: 779, fig. – Pereopod 5 six-articulate ...... 4 531); Camp (1998: 132); Guerra-García and Takeuchi 4. Pereopods 3 and 4 absent ...... 5 (2002: 683-684, fig. 6). – Pereopods 3 and 4 present, though reduced to one Caprella inermis Haswell, 1879: 319-351. or two articles ...... 7 Material examined (76 specimens). Station 18: 2 5. Head with a well-developed rostrum ...... Caprella females, 3 juveniles (USB-SU0187), 03/15/2001; 2 penantis (Fig. 5) males, 1 female (USB-SU0263), 03/15/2001. Station – Head without a well-developed rostrum ...... 6 26: 2 males, 1 female, 1 juvenile (USB-NE0042), 6. Pereonite 2 with a ventral projection between gna- 08/13/2000. Station 27: 16 males, 48 females (USB- thopods 2. Propodus of pereopods 5-7 with grasping NE0048), 1 male, 1 female (both used for lateral-view spines ...... Caprella equilibra (Fig. 4) figures) (MNCN 20.04/5894), 08/15/2000. – Pereonite 2 without ventral projection between gan- Remarks. Although Caprella danilevskii is widely thopods 2. Propodus of pereopods 5-7 without gra- distributed around the world (McCain and Steinberg sping spines ...... Caprella danilevskii (Fig. 3) 1970; Guerra-García and Takeuchi 2004), the first re- 7. Pereopods 3 and 4 one-articulate ...... 8 cord for Venezuela is recent (Villarroel and Graziani – Pereopods 3 and 4 two-articulate ...... 9 1997). The Venezuelan specimens are in agreement 8. Head smooth. Pereonite 2 with a ventral projection. with the Mediterranean material described by Krapp- Abdomen with distinct appendages visible in lateral Schickel (1993) and the Atlantic specimens figured view ...... Hemiaegina minuta (Fig. 9) by McCain (1968). The distinctive male abdomen, the

Org. Divers. Evol. 5, Electr. Suppl. 10 (2005) Díaz & al.: Caprellids (Crustacea: Amphipoda) from Venezuela 4 absence of grasping spines on the pereopods, and the key wings (Arca zebra (Swainson, 1833)), on Thalas- short dactylus of male gnathopod 2 distinguishes this sia testudinum Banks ex König, 1805, and algae. species from other species of Caprella. The life history of C. danilevskii is one of the most well known in the Caprella penantis Leach, 1814 Caprellidae; Takeuchi and Hirano (1991) studied the (Fig. 5) growth and reproduction of this species based on Japa- Caprella Penantis Leach, 1814: 404. nese specimens. Caprella acutifrons Mayer, 1882: 48. – Mayer Habitat. This species has been found living on al- (1890: 50, pl. 2, figs. 36-37, 39-41, pl. 4, figs. 52-53, gae, seagrasses, , and bryozoans (Mc- 55, 57-61, 65-69; including the ‘forms’ neglecta, tabi- Cain 1968). In the present study, Caprella danilevskii da, gibbosa, carolinensis, lusitanica, virginia); Mayer was found clinging on the algae sp., Acan- (1903: 79, pl. 3, figs. 4-28; pl. 7, figs. 62-65). tophora sp., Gracilaria sp., Ulva lactuca Linnaeus, Caprella penantis Leach. – McCain (1968: 33, figs. 1753, and Hypnea muscifornis J.V. Lamouroux, 1813. 15-16); McCain and Steinberg (1970: 33); Cavedini (1982: 508); Krapp-Schickel (1993: 791-793); Camp Caprella equilibra Say, 1818 (1998: 132); Guerra-García and Takeuchi (2002: 692- (Fig. 4) 693, fig. 12). Caprella equilibra Say, 1818: 391-392. – McCain Material examined (1 specimen). Station 19: 1 (1968: 25, figs. 12-13); McCain and Steinberg (1970: male (used for lateral-view figure) (USB-SU0281), 19); Cavedini (1982: 500); Krapp-Schickel (1993: 782- 03/13/2001. 783, fig. 533); Camp (1998: 132); Guerra-García and Remarks. The specimen represents the first record Thiel (2001: 878-879, fig. 6). of Caprella penantis from Venezuela. The species has Caprella aequilibra Say. – Mayer (1882: 45, pl. 1, been recorded under several species- or subspecies na- fig. 7; pl. 2, fig. 1-11; pl. 4, figs. 20-25; pl. 5, figs. 16- mes from temperate regions worldwide (McCain and 18); Chevreux and Fage (1925: 455, fig. 433). Steinberg 1970). Further morphological and molecular Material examined (390 specimens). Station 17: studies are required to evaluate the respective status of 27 juveniles (USB-SU0544), 05/09/2003. Station 19: these nominal species or subspecies (McCain 1968; 1 female, 2 juveniles (USB-SU0208), 03/18/2003; Laubitz 1972; Takeuchi and Hirano 1995). 15 males, 14 females, 21 juveniles (USB-SU0254), Habitat. This species has been found on red and 03/18/2003. Station 20: 1 male, 4 females (USB- brown algae, seagrasses such as Posidonia, on hydro- SU0347), 1 male, 1 female (both used for lateral-view ids, alcyonarians, zoantharians, bryozoans, sponges, figures) (MNCN 20.04/5895), 05/08/2003; 148 speci- and attached to echinoderms (Arbacia) and decapods mens (USB-SU0348), 05/08/2003; 100 specimens (Libinia) (McCain 1968; Krapp-Schickel 1993). The (USB-SU0349), 05/08/2003; 11 males, 15 females, 30 specimen found during the present study was collected juveniles (USB-SU0555), 05/06/2003; 1 male (USB- among mussels, Perna perna (Linnaeus, 1758) and P. SU0656), 05/06/2003. viridis (Linnaeus, 1758). Remarks. Caprella equilibra, similarly to C. dani- levskii, is widely distributed around the world (McCain Deutella incerta (Mayer, 1903) and Steinberg 1970; Krapp-Schickel 1993). The Vene- (Figs. 6,7) zuelan specimens have a distinctive ventral projection Luconacia incerta Mayer, 1903: 49-50, pl. 2, figs. between gnathopods 2 that is characteristic for this 11-14; pl.6, figs. 73-75; pl. 9, figs. 21, 40, 57. – McCain species. The presence of C. equilibra in Venezuela was (1968: 53-54, 68-72, figs. 33-35); McCain and Stein- reported by McCain (1968), based on specimens from berg (1970: 53). Puerto Cabello, Carabobo state. All the material inclu- Protellopsis stebbingii Kunkel, 1910: 111-113, fig. ded in the present study is from Sucre state. 43. Habitat. Caprella equilibra has been found on Deutella incerta (Mayer). – Steinberg & Dougherty, seagrasses, algae, sponges, hydroids, bryozoans, co- 1957: 281, 285-286. – Gable and Lazo-Wasem (1987: lonial ascidians and alcyonarians, from the intertidal 635-636, fig. 4); Camp (1998: 132); Guerra-García to 3000m depth (Krapp-Schickel 1993). The species (2003a: 1062-1065, fig. 3). is also found on artificial devices such as aquaculture Material examined (113 specimens). Station 1: 1 nets, water duct pipes, power plants, as well as on floa- female (USB-FA0632), 05/08/2000; 1 male, 1 fema- ting substrates, e.g. buoys and drifting algae (Takeuchi le, 1 juvenile (USB-FA0787), 08/02/2000; 3 males, and Sawamoto 1998; Thiel et al. 2003). Caprella equi- 2 females, 1 juvenile (USB-FA0965), 08/02/2000; libra feeds principally by filtering, frequently using 1 juvenile (USB-FA1024), 08/02/2000; 1 female grooming behaviour (Guerra-García et al. 2002). In the (USB-FA1464), 07/16/2001. Station 4: 1 male, 1 fe- present study, this species was found on mussels, tur- male (USB-FA0187), 12/1993. Station 6: 1 male, 4

Org. Divers. Evol. 5, Electr. Suppl. 10 (2005) Díaz & al.: Caprellids (Crustacea: Amphipoda) from Venezuela 5 females, 3 juveniles (USB-FA0544), 02/08/2000; 1 2003a). The material studied here was collected from female (USB-FA0630), 05/2000; 1 male, 1 female, 2 sandy bottoms and from . juveniles (USB-FA0701), 08/02/2000; 2 females, 3 ju- veniles (USB-FA0784), 08/02/2000; 1 juvenile (USB- Deutella margaritae Guerra-García, 2003 FA0785), 08/02/2000; 1 juvenile (USB-FA0786), (Fig. 8) 08/02/2000; 1 juvenile (USB-FA0788), 08/02/2000; Deutella margaritae Guerra-García, 2003a: 1065- 1 juvenile (USB-FA0999), 02/12/2001; 1 male (USB- 1070, figs. 4-7. FA1010), 11/05/2000; 1 juvenile (USB-FA1011); 1 fe- Material examined (966 specimens). Station male (USB-FA1017), 11/05/2000. Station 7: 1 female 22: 1 male (USB-NE0007), 04/08/1998. Station 23: (USB-FA0192), 12/1993; 1 juvenile (USB-FA1480), 34 males, 15 females, 53 juveniles (USB-NE0192), 05/08/2000. Station 8: 1 female (USB-FA0305), 12/18/2003; 14 males, 8 females, 97 juveniles (USB- 10/02/1999; 1 female (USB-FA0607), 05/08/2000; 5 NE0193), 12/18/2003; 25 males, 30 females, 64 ju- males, 1 female (USB-FA0634), 1 male, 1 female (used veniles (USB-NE0194), 12/18/2003; 230 specimens for lateral-view figure, Fig. 6) (MNCN 20.04/5896), (USB-NE0195), 12/18/2003; 156 specimens (USB- 05/08/2000; 2 females (USB-FA0950), 08/02/2000; 1 NE0202), 12/18/2003. Station 24: 13 males, 11 fema- juvenile (USB-FA0976), 02/12/2001; 1 juvenile (USB- les, 7 juveniles (USB-NE0174), 12/18/2003. Station FA1007), 11/05/2000; 4 juveniles (USB-FA1008), 25: 1 female (USB-NE0008), 07/05/1998; 25 males, 11/05/2000; 1 female (USB-FA1452), 07/16/2001. Sta- 9 females, 30 juveniles (USB-NE0157), 11/05/2003; tion 9: 7 males, 6 females, 4 juveniles (USB-FA0633), 23 males, 27 females, 4 juveniles (USB-NE0180), 05/08/2000; 1 male, 2 females (USB-FA0708), 11/05/2003; 30 males, 35 females, 14 juveniles (USB- 08/02/2000; 1 female (USB-FA0940), 08/02/2000; 4 NE0187), 11/05/2003. material (Fig. 8): Holo- males, 2 females (USB-FA1014), 11/05/2000; 1 ju- type male (USNM 1000206), allotype female (USNM venile (USB-FA1015), 11/05/2000; 2 females (USB- 1000207) and paratypes (3 males, 7 females, USNM FA1019), 11/2000; 1 male (USB-FA1020), 11/05/2000; 1000208), 01/11/1964 from Puente de la Restinga, Isla 1 female (USB-FA1239), 07/16/2001; 5 males, 2 fema- Margarita, Venezuela. les, 6 juveniles (USB-FA1479), 06/1993. Station 12: Remarks. This is the first record of Deutella mar- 1 female (USB-AR0247), 07/25/1980. Station 13: 1 garitae after the original description (Guerra-García female (USB-AR0302), 08/1981. Station 15: 1 male, 2 2003a), based on material from Isla Margarita, Vene- females (USB-AN0059), 01/1999. Station 16: 1 female zuela. The specimens have been assigned to D. mar- (USB-AN0094), 03/24/2000. Type material (Fig. 7): 1 garitae mainly on the basis of the similar morphology male, 1 female (paralectotypes) (USNM 026001) coll- of pereopods 3 and 4, mouthparts and abdomen, and ected from 29º28’N, 87º56’W, Albatros, station 2389, the presence of distinctive anterolateral projections on 49 meters. pereonites 2-4. However, in the examined specimens Remarks. Deutella incerta is widely distributed in pereopod 5 is inserted at midlength of the pereonite 5 the temperate and tropical areas of the western North (similar to Deutella incerta) and not on the posterior Atlantic (McCain 1968) and seems to be one of the end as in the type specimens. We have found a con- most common caprellids in Bermuda (Gable and Lazo- siderable degree of variation in Deutella species from Wasem 1987). McCain (1968) had already reported Venezuela, thus do not discount the possibility that a the presence of this species in Venezuelan waters. A complex of several more species of Deutella may exist detailed discussion regarding the status of D. incerta there. Further morphological and, especially, molecular is included in Guerra-García (2003a). McCain (1968) studies would be necessary to clarify the systematics pointed out that this species varies considerably in the of Deutella in Venezuelan waters. At this time, we degree of body spination; larger individuals bear dor- maintain the two species already recorded from Vene- sal projections, but the latter are absent in the smaller zuela, and prefer not to erect any new species without specimens. All the specimens examined for the present clear evidence of discrete characteristics. study lack dorsal projections, and their morphology Habitat. The type material was found in boulders is in agreement with the small specimens figured by near Rhizophora sp. The material examined in the McCain (1968), but differs from the type specimens, present study was collected from oysters and hydro- which are provided with distinct projections (Fig. 7). ids, Halocordyle disticha (Goldfuss, 1820), on roots of Habitat. This species has been collected on man- Rhizophora mangle L. grove roots, Sargassum sp., Thalassia sp., sponges, hydroids, alcyonarians and ascidians, and occasional- Hemiaegina minuta Mayer, 1890 ly it has been taken in tows (Guerra-García (Fig. 9)

Org. Divers. Evol. 5, Electr. Suppl. 10 (2005) Díaz & al.: Caprellids (Crustacea: Amphipoda) from Venezuela 6

Hemiaegina minuta Mayer, 1890: 40, pl. 1, figs. collected from North of , Nueva Esparta 25–27; pl. 3, figs. 32–35;pl. 5, figs. 52–53; pl. 6, figs. state, Venezuela, 10º50’N, 63º54’W, 04/15/1939. 13, 33–34, pl. 7, fig. 4. – McCain (1968: 61–64, figs. Remarks. The specimens reported here represent 29–30); McCain and Steinberg (1970: 51); Gable and the first record of the species from Venezuela since Lazo-Wasem (1987: 637); Müller (1990: 836); Serejo the original description by McCain (1968). As far as is (1997: 631, fig. 1); Camp (1998: 132); Guerra-García known, Mayerella redunca is endemic to Venezuela. A (2003b: 6-7, fig. 3; 2003c: 105-106, fig.10). revision of the genus Mayerella can be found in Guer- Hemiaegina quadripunctata Sundara Raj, 1927: ra-García (2003d). 126–127, pl. 18. Habitat. McCain (1968) reported that specimens Hemiaegina costai Quitete, 1972: 165–168, pls. were collected between 35m and 60m depth, but he did 1–2. not mention any substrates. The female from Falcon Material examined (3 specimens). Station 20: state was collected from sandy bottoms, the remaining 1 juvenile (USB-SU0722), 1 male, 1 female (both two specimens in plankton samples. used for lateral-view figures) (MNCN 20.04/5897), Paracaprella digitimanus Quitete, 1971 05/06/2003. (Fig. 12) Remarks. Hemiaegina minuta is widely distributed Paracaprella digitimanus Quitete, 1971: 161, figs. in tropical and temperate waters of the world’s oceans 1-3. – Wakabara and Serejo (1998: 585). (Müller 1990). The species was reported from Vene- Material examined (2 specimens). Station 20: 1 zuela by Stoner and Lewis (1985) – from Gran Roque male (MNCN 20.04/5899), 1 juvenile (USB-SU0721), Island, Archipiélago Los Roques (11º55’N, 66º40’ W) 03/13/2001. – during an ecological study. The material studied from Remarks. The original description of Paracaprella Sucre state is scarce and in poor condition (e.g. male digitimanus was based on material from Brazil (Quite- lacks gnathopods 2). te 1971); the present finding is the first for Venezuela. Habitat. Hemiaegina minuta has been collected Although the genus Paracaprella is still awaiting revi- from Sargassum sp. and taken in plankton tows (Mc- sion, P. digitimanus can be easily distinguished from Cain and Steinberg 1970). Müller (1990) reported H. its congeners by a striking projection on the propodus minuta as preferring more or less exposed reef locati- of gnathopod 2. ons. The species has been collected also from Thalas- Habitat. Quitete (1971) collected Paracaprella di- sia testudinium (Stoner and Lewis 1985). Guerra-Gar- gitimanus from hydroids. The two specimens reported cía (2003b) found it associated with Turbinaria ornata here were found on mussels (Perna perna and P. viri- (Turner) J. Agardh on the coast of Mauritius, and Guer- dis). ra-García (2003c) reported it from Papua New Guinea, living on Dictyota sp., Halimeda sp., Gracilaria sp., Paracaprella pusilla Mayer, 1890 Galaxaura sp., and Amansia glomerata (Agardh) Nor- (Fig. 13) ris, 1979. Hemiaegina minuta also has been found as- Paracaprella pusilla Mayer, 1890: 41, pl. 1, figs. sociated with many different substrata in Queensland: 28-30; pl. 3; figs. 45-47; pl. 5, figs. 48-49; pl. 6, fig. green algae such as Halimeda sp., brown and , 10; 1903: 67, pl. 2, figs. 36-37; pl. 7, fig. 52. – Stein- sponges, tunicates, seagrasses such as Posidonia sp., berg and Dougherty (1957: 283-284, figs. 16, 19, 24, dead corals encrusted with algal turf, and under small 30); McCain (1968: 82-86, figs. 41-42); Wakabara et boulders (Guerra-García, personal observations). The al. (1991: 73); Camp (1998: 132); Guerra-García and specimens studied in the present work were found on Thiel (2001: 880, fig. 8). turkey wings (Arca zebra). Caprella nigra Reid, 1951: 283-284, 289, fig. 58. Material examined (2324 specimens). Station 2: 1 Mayerella redunca McCain, 1968 female (USB-FA0564), 08/30/2000. Station 10: 1 juve- (Figs. 10-11) nile (USB-CA0054), 01/1982; 1 male (USB-CA0055), Mayerella redunca McCain, 1968: 75-78, figs. 37- 04/1982; 1 juvenile (USB-CA0065), 04/1982; 1 fema- 38, 50. – McCain and Steinberg (1970: 53); Guerra- le (USB-CA0071), 02/1982; 1 female (USB-CA0099), García (2003d: 189, 191, 194, figs. 8, 11). 01/1986; 1 juvenile (USB-CA0132), 05/1982; 1 fema- Material examined (5 specimens). Station 5: 1 le (used for lateral-view figure) (MNCN 20.04/5900), female (USB-FA0216), 03/29/2000. Station 10: 1 fe- 04/1982; 1 male (USB-CA0143), 04/1982; 1 juvenile male (used for lateral-view figure, Fig. 10) (MNCN (USB-CA0169); 1 juvenile (USB-CA0212); 1 male 20.04/5898), 12/1981; 1 juvenile (USB-CA0445), (USB-CA0237); 2 juveniles (USB-CA0300), 07/1982; 01/1982. Type material (Fig. 11): Holotype male 2 males, 1 female (USB-CA0305), 10/1982; 3 males (USNM 120176), allotype female (USNM 120177) (USB-CA0312), 11/1982; 4 males, 4 females, 1 juvenile

Org. Divers. Evol. 5, Electr. Suppl. 10 (2005) Díaz & al.: Caprellids (Crustacea: Amphipoda) from Venezuela 7

(USB-CA0316), 08/1982; 3 juveniles (USB-CA0321), Remarks. Laubitz (1995) considered Fallotritella as 12/1982; 1 male, 2 juveniles (USB-CA0334), 1 male synonymous with Pseudaeginella. Pseudaeginella bis- (MNCN 20.04/5901), 11/1982; 3 males (USB-CA0465), caynensis is common in Caribbean waters (McCain and 03/1981. Station 11: 6 males, 13 females, 8 juveniles Steinberg 1970). In spite of the considerable number of (USB-CA0431), 1 male, 1 female (MNCN 20.04/5309), stations sampled along the Venezuelan coast during the 08/29/2000. Station 14: 1 female (USB-AR0013), present work, we have not found P. biscaynensis. Ho- 07/24/1980. Station 20: 22 males, 14 females, 86 juve- wever, the species has been reported from Venezuela niles (USB-SU0346), 1 male (used for lateral-view fi- by Stoner and Lewis (1985), based on material from gure), 1 female (MNCN 20.04/5902), 05/08/2003; 210 Gran Roque Island, Archipiélago Los Roques (11º55’ specimens (USB-SU0350), 05/08/2003; 213 specimens N, 66º40’ W), collected during an ecological study. The (USB-SU0351), 05/08/2003; 201 specimens (USB- figures included here are taken from McCain (1968) SU0352), 05/08/2003; 169 specimens (USB-SU0353), and based on the type material from Florida. 05/08/2003; 193 specimens (USB-SU0354), 08/05/2003; Habitat. The species is associated with red algae 215 specimens (USB-SU0355), 05/08/2003; 255 speci- (McCain 1968), the green alga Avrainvillea sp. and the mens (USB-SU0356), 05/08/2003; 209 specimens seagrass Thalassia sp. (Gable and Lazo-Wasem 1987; (USB-SU0357), 05/08/2003; 147 specimens (USB- Stoner and Lewis 1985), with corals, Cymodocea sp. SU0358), 05/08/2003; 194 specimens (USB-SU0366), and Syringodium sp. (Guerra-García 2002), and with 05/08/2003. Station 21: 2 females (USB-SU0575), the seaweed Galaxaura sp. (Guerra-García 2003c). 05/07/2003. Station 23: 46 males, 49 females, 21 ju- veniles (USB-NE0210), 12/18/2003. Station 24: 2 ma- Subfamily Phtisicinae Vassilenko, 1968 les (USB-NE0213), 12/18/2003. Station 25: 1 female (USB-NE0214), 11/05/2003; 2 males, 2 females (USB- Hemiproto wigleyi McCain, 1968 NE0215), 11/05/2003. (Fig. 15) Remarks. Mayer (1890) described Paracaprella Hemiproto wigleyi McCain, 1968: 65-68, figs. 31, pusilla based on material collected from Brazil. Later, 32(c-e), 50. – McCain and Steinberg (1970: 51); Camp the species was reported from Venezuela by McCain (1998: 132). (1968). Paracaprella pusilla is similar to Paracaprella Material examined (13 specimens). Station 6: 1 tenuis Mayer, 1903; however, males of the former can female (USB-FA1012), 11/05/2000; 1 female (USB- be distinguished by the large, sharp-pointed projection FA1016), 11/05/2000; 1 female (USB-FA1018), on the anteroventral margin of pereonite 2, the proxi- 11/05/2000; 1 male (USB-FA1123), 08/02/2000; 1 ju- mal knob on the basis of gnathopod 2, and by the pres- venile (USB-FA1481), 02/12/2000. Station 7: 1 male, ence of setae on the dactylus of gnathopod 2 (McCain 1 female (USB-FA0609), 1 female (used for lateral- 1968). Large males of P. pusilla are very similar to lar- view figure) (MNCN 20.04/5903), 05/08/2000; 1 male ge males of P. barnardi McCain, 1967 in that both bear (used for lateral-view figure) (MNCN 20.04/5904), a small anterodorsal tubercle on pereonite 2; however, 05/08/2000. Station 8: 1 female (USB-FA1009), the tubercle is not as well developed in the former as 11/05/2000; 1 female (USB-FA1025), 11/05/2000. Sta- in the latter species, and the ventrolateral projection on tion 9: 1 male (USB-FA0012), 06/1993. Station 21: 1 the anterior margin of pereonite 2 is much larger in P. female (MNCN 20.04/5905), 05/07/2003. pusilla (see McCain 1967). Remarks. Hemiproto is, so far, a monotypic genus Habitat. Paracaprella pusilla has been collected and includes the single species H. wigleyi, described from mangrove roots, seagrasses, hydroids and ascidi- by McCain (1968). This species is known from Coche ans (McCain 1968). The specimens collected here were Island, Nueva Esparta state, Venezuela, collected at 35- found on gravel bottoms, ropes, mussels, oysters, sa- 60m depth, and from Fort Lauderdale, Florida, at 32 bellariid worm rock (Phragmatopoma lapidosa (Kin- m. Watling (1997) reported H. wigleyi from the San- berg, 1867)), and on hydroids (e.g. Halocordyle disti- ta Maria Basin region, California. We have assigned cha) associated with mangrove roots. the specimens collected from Falcon state to H. wigley mainly on the basis of the combination of the following Pseudaeginella biscaynensis (McCain, 1968) characteristics: molar absent; mandibular palp three- (Fig. 14) articulate; pereopods 3 and 4 six-articulate; pereopod Fallotritella biscaynensis McCain, 1968: 58-61, 5 five-articulate; abdomen with two pairs of uni-arti- figs. 27-28, 53. – McCain and Steinberg (1970: 51); culate appendages in males (one of them reduced), and Gable and Lazo-Wasem (1987: 637-638). only one pair in females. Pseudaeginella biscaynensis (Mc Cain). – Laubitz Habitat. The ecology of this species is unknown. (1995: 88); Camp (1998: 132). McCain (1968) only reported that specimens were col-

Org. Divers. Evol. 5, Electr. Suppl. 10 (2005) Díaz & al.: Caprellids (Crustacea: Amphipoda) from Venezuela 8 lected between 35m and 60 m depth, but he did not Shallow-water Marine Invertebrates of Florida. Florida mention any substrates. The specimens reported here Mar: Res: Inst: Techn: Rep: TR-3, 123–189. were collected from sandy bottoms and living on Tha- Cavedini, P., 1982. Contributo alla conoscenza dei Caprelli- lassia testudinum and Halimeda opuntia J.V. Lamou- di del Mediterraneo (Crustacea, Amphipoda). Boll. Mus. Civ. Stor. Nat. Verona 8, 493–531. roux, 1816. Chevreux, E., Fage, L., 1925. Amphipodes. Faune de Phtisica marina Slabber, 1769 France 9, 1–488. Czerniavskii, V., 1868. Materialia ad zoographiam Ponticam (Fig. 16) comparatam. Trav. Soc. Natural. St. Pétersbg. 1, 19–136. Phtisica marina Slabber, 1769: 77, pl. 10. – Chev- Escobar-Briones, E., Winfield, I., 2003. Checklist of the reux and Fage (1925: 434, fig. 422); McCain (1968: 91, benthic Gammaridea and Caprellidea (Crustacea: Pera- figs. 46-47); McCain and Steinberg (1970: 65); Camp carida: Amphipoda) from the Gulf of Mexico continental (1998: 132); Guerra-García and Takeuchi (2002: 705). shelf and slope. Belg. J. Zool. 133, 37–44. Proto brunneovittata Haller, 1879: 231. – Haller Gable, M.F., Lazo-Wasem, E.A., 1987. The Caprellids (Am- (1880: 339, pl. 22, figs. 19-22). phipoda: Caprellida) of Bermuda: a survey of specimens collected from 1876-1987, including cave inhabitants, Proto pedata Haller, 1879: 230. – Haller (1880: and the description of Deutella aspiducha, new species. 398). Proc. Biol. Soc. Wash. 100, 629–639. Proto ventricosa Mayer, 1882: 22, pl. 1, fig. 1; pl. 3, Guerra-García, J.M., 2001. Redescription of Caprellina figs. 16-29; pl. 4, figs. 12-13; pl.5, figs. 1-5. – Mayer longicollis (Nicolet, 1849) (Amphipoda, Caprellidea, (1890: 12, pl. 3, figs. 4-6; pl. 5, figs. 3-6; pl. 6, fig. 1; pl. Phtiscidae) from Chile, with notes on ontogenetic 7, fig. 1); Mayer (1903: 20, pl. 6, fig. 23). development and clinging behaviour. Crustaceana 74, Material examined (1 specimen). Station 3: 1 im- 1291–1303. mature female (used for lateral-view figure) (USB- Guerra-García, J.M., 2002. Redescription of five rare FA0465), 05/25/2000. caprellids (Crustacea: Amphipoda: Caprellidea) collected Remarks. Phtisica marina is widely distributed in from Tanzanian coasts. Helgol. Mar. Res. 55, 221–231. Guerra-García, J.M., 2003a. Revision of the genus Deutella the Atlantic and Pacific Oceans, and in the Mediterra- (Crustacea: Amphipoda: Caprellidea) with description nean Sea (McCain 1968; Krapp-Schickel 1993). Mc- of a new species, redescription of Deutella venenosa Cain (1968) reported the species from and Mayer, 1890 and a key to the species of Deutella. J. Nat. Margarita Islands, Nueva Esparta state, Venezuela. Hist. 37, 1059–1984. Phtisica marina can be differentiated from Hemiproto Guerra-García, J.M., 2003b. The Caprellidea (Crustacea: wigleyi mainly by the abdomen (see also the remarks Amphipoda) from Mauritius Island, Western Indian under H. wigleyi). Ocean. Zootaxa 232, 1–24. Habitat. This species has been reported from green Guerra-García, J.M., 2003c. Caprellids (Crustacea: Amphi- and brown algae, seagrasses, sponges, hydroids, bryo- poda) from Papua New Guinea, with the description of a new species. Helgol. Mar. Res. 57, 100–109. zoans, echinoderms (Asteroidea), and to a depth of 660 Guerra-García, J.M., 2003d. Redescription of Mayerella m (McCain 1968; Krapp-Schickel 1993). The speci- magellanica McCain & Gray, 1971 (Crustacea: Am- men reported here was collected from sandy bottoms phipoda: Caprellidea) from Chile with a revision of the at 15m depth. A predatory mode of life is suggested for genus Mayerella. Hydrobiologia 490, 187–195. this species (Guerra-García et al. 2002). Guerra-García, J.M., Corzo, J., García-Gómez, J.C., 2002. Clinging behaviour of the Caprellidea (Amphipoda) from Acknowledgements the Strait of Gibraltar. Crustaceana 75, 41–50 Guerra-García, J.M., Takeuchi, I., 2002. The Caprellidea We are very grateful to J. Gutiérrez (“Fundación La (Crustacea: Amphipoda) from Ceuta, North Africa, with Salle de Ciencias Naturales”, Venezuela), J. Bolaños the description of three species of Caprella, a key to the (“Universidad de Oriente”, Venezuela), and W. Fera- species of Caprella, and biogeographical discussion. J. gotto (“Benthos Buceo Profesional”, Venezuela) for Nat. Hist. 36, 675–713. providing some of the samples. Thanks are also due to Guerra-García, J.M., Takeuchi, I., 2004. The Caprellidea the “Agenda Parque Nacional Morrocoy”-FONACIT, (Crustacea: Amphipoda) from Tasmania. J. Nat. Hist. 38, Venezuela (project number 96001837-V), and to the 967–1044. University of Sevilla, Spain (“III Plan Propio de Apoyo Guerra-García, J.M., Thiel, M., 2001. La fauna de capréli- a Programas Internacionales”) for financial support. dos (Crustacea: Amphipoda: Caprellidea) de la costa de Coquimbo, centro-norte de Chile, con una clave taxonó- mica para la identificación de las especies. Rev. Chil. References Hist. Nat. 74, 873–883. Camp, D.K., 1998. Checklist of shallow-water marine Haller, G., 1879. Vorläufige Notizen über die Systematik malacostracan Crustacea of Florida, in: Camp, D.K., der im Mittelmeer vorkommenden Caprelliden. Zool. Lyons, W.G., Perkins, T.H. (Ed.), Checklists of Selected Anz. 2, 230–233.

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Haller, G., 1880. Miscellanea arthropodologica. I. Beschrei- Reid, D.M., 1951. Report on the Amphipoda (Gammaridea bung zweier neuer Caprellen. Z. Ges. Naturwiss. 6, and Caprellidea) of the coast of tropical West Africa. 742–749. Atlantide Rep. 2, 189–291. Haswell, W.A., 1879. On some additional new genera and Say, T., 1818. An account of the Crustacea of the Uni- species of amphipod crustaceans. Proc. Linn. Soc. N. S. ted States (Caprellidae). J. Acad. Nat. Sci. Philad. 1, Wales 4, 319–351. 390–392. Krapp-Schickel, T., 1993. Suborder Caprellidea, in: Ruffo, Serejo, C., 1997. Hemiaegina costai Quitete, 1972 a syno- S. (Ed.), The Amphipoda of the Mediterranean. Mém. nym of Hemiaegina minuta Mayer, 1890 (Amphipoda: Inst. Oceanogr., Monaco, pp. 773–809. Caprellidae). Crustaceana 70, 630–632. Kunkel, B.W., 1910. The Amphipoda of Bermuda. Trans. Slabber, M., 1769. Natuurkundige verlustigingen, behel- Connecticut Acad. Arts Sci. 16, 1–116. zende microscopise waarneemingen van in- en uitlandse Laubitz, D.R., 1972. The Caprellidea (Crustacea, Amphi- water- en land-dieren. J. Bosch, Haarlem. poda) of Atlantic and Arctic Canada. Natl. Mus. Canada Steinberg, J.E., Dougherty, E.C., 1957. The skeleton Publ. Biol. Oceanogr. 4, 1–82. (Crustacea: Caprellidae) of the Gulf of Mexico. Laubitz, D.R., 1993. Caprellidea (Crustacea: Amphipoda) Tulane Stud. Zool. 5, 267–288. towards a new synthesis. J. Nat. Hist. 27, 965–976. Stoner, A.W., Lewis, F.G., 1985. The influence of quantita- Laubitz, D.R., 1995. Caprellidea (Crustacea: Amphipoda) tive aspects of habitat complexity in tropical sea-grass from the southern and western Indian ocean. Mésogée meadows. J. Exp. Mar. Biol. Ecol. 94, 19–40. 54, 81–100. Sundara Raj, B., 1927. Suborder Caprellidea (Laemodipo- Leach, W.E., 1814. Crustaceology. Vol 7, in: Brewster, da). The littoral fauna of Krusadai Island in the Gulf of D. (Ed.), Edinburgh Encyclopaedia. Caprellidae, pp. Mannaer. Bull. Madras Gov. Mus., N. Ser., Nat. Hist. 383–384. Sect. 1, 125–128. Mayer, P., 1882. Die Caprelliden des Golfes von Neapel Takeuchi, I., 1993. Is the Caprellidea a monophyletic und der angrenzenden Meeres-Abschnitte. Fauna Flora group? J. Nat. Hist. 27, 947–964. Golf. Neapel 6, 1–201. Takeuchi, I., Hirano, R., 1991. Growth and reproduction of Mayer, P., 1890. Die Caprelliden des Golfes von Neapel Caprella danilevskii (Crustacea: Amphipoda) reared in und der angrenzenden Meeres-Abschnitte. Fauna und the laboratory. Mar. Biol. 110, 391–397. Flora des Golfes von Neapel 17, 1–55. Takeuchi, I., Hirano, R., 1995. Clinging behaviour of the Mayer, P., 1903. Die Caprellidae der Siboga-Expedition. epifaunal caprellids (Amphipoda) inhabiting the Sargas- Siboga Exped. 34, 1–160. sum zone on the Pacific Coast of Japan, with its evolutio- McCain, J.C., 1967. Paracaprella barnardi, a new species nary implications. J. Crust. Biol. 15, 481–492. of caprellid (Crustacea; Amphipoda) from the west coast Takeuchi, I., Sawamoto, S., 1998. Distribution of caprellid of Panamá. Proc. Biol. Soc. Wash. 80, 219–222. amphipods (Crustacea) in the western North Pacific McCain, J.C., 1968. The Caprellidea (Crustacea: Amphipo- based on the CSK International Zooplankton collection. da) of the western north Atlantic. U.S. Natl. Mus. Bull. Plankton Biol. Ecol. 45, 225–230. 278, 1–116. Thiel, M., Guerra-García, J.M., Lancellotti, D.A., Vás- McCain, J.C., Steinberg, J.E., 1970. Amphipoda-I, Caprelli- quez, N., 2003. The distribution of littoral caprellids dea-I, in: Gruner, H.E., Holthuis, L. B. (Eds.), Crustaceo- (Crustacea: Amphipoda: Caprellidea) along the Pacific rum Catalogus 2, pp. 1–78. coast of continental Chile. Rev. Chil. Hist. Nat. 76, Müller, H.G., 1990. New species and records of coral reef 297–312. inhabiting Caprellidae from Bora Bora and Moorea, So- Villarroel, E.J., Graziani, C.A., 1997. Primer registro de ciety Islands (Crustacea: Amphipoda). Rev. Suisse Zool. Caprella danilevskii Czerniavskii (Amphipoda: Caprelli- 97, 827–842. dae) en Venezuela. Saber, Univ. Oriente 9, 102–103. Myers, A.A., Lowry, J.K., 2003. A phylogeny and a new Wakabara, Y., Serejo, C.S., 1998. – Peracari- classification of the Corophiidea Leach, 1814 (Amphipo- da. Amphipoda. Gammaridea and Caprellidae, in: Young, da). J. Crust. Biol. 23, 443–485. P.S. (Ed.), Catalogue of Crustacea of Brazil. Museu Ortíz, M., Alvárez, F., Winfield, I., 2002. Caprellid am- Nacional, Rio de Janeiro, Ser. Livros 6, pp. 561–594. phipods (Amphipoda: Caprellidae) illustrated key for Wakabara, Y., Tararam, A.S., Valerio-Berardo, M.T., Dule- the genera and species from the Gulf of Mexico and ba, W., Pereira F.P., 1991. Gammaridean and caprellide- the . Universidad Nacional Autónoma de an fauna from Brazil. Hydrobiologia 223, 69–77. México. Facultad de Estudios Superiores Iztacala. Watling, L., 1997. The Suborder Caprellidea, in: Blake, Quitete, J.M.P.A., 1971. Paracaprella digitimanus, nova es- J.A., Watling, L., Scott P.V. (Eds.), Taxonomic Atlas of pécie de Caprellidae da costa brasileira (Crustacea: Am- the Benthic Fauna of the Santa Maria Basin and the Wes- phipoda). Atas Soc. Biol. Rio de Janeiro 14, 161–164. tern Santa Barbara Channel. Santa Barbara, California, Quitete, J.M.P.A., 1972. Hemiaegina costai, nova espécie pp. 223-240. de Caprellidae da costa brasileira (Crustacea: Amphipo- da) Atas Soc. Biol. Rio de Janeiro 15, 165–168.

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72º 60º CARIBBEAN SEA 12º Los 23-25 Roques 15 N 11 22 26-27 1-9 12-14 ATLANTIC 10 18-19 OCEAN 16 20-21 17 VENEZUELA

COLOMBIA

2º BRAZIL

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

Org. Divers. Evol. 5, Electr. Suppl. 10 (2005) Díaz & al.: Caprellids (Crustacea: Amphipoda) from Venezuela 11

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

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

Org. Divers. Evol. 5, Electr. Suppl. 10 (2005) Díaz & al.: Caprellids (Crustacea: Amphipoda) from Venezuela 13

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

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

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

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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.

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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.

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

Org. Divers. Evol. 5, Electr. Suppl. 10 (2005) Díaz & al.: Caprellids (Crustacea: Amphipoda) from Venezuela 19

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

Org. Divers. Evol. 5, Electr. Suppl. 10 (2005) Díaz & al.: Caprellids (Crustacea: Amphipoda) from Venezuela 20

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.

Org. Divers. Evol. 5, Electr. Suppl. 10 (2005) Díaz & al.: Caprellids (Crustacea: Amphipoda) from Venezuela 21

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.

Org. Divers. Evol. 5, Electr. Suppl. 10 (2005) Díaz & al.: Caprellids (Crustacea: Amphipoda) from Venezuela 22

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.

Org. Divers. Evol. 5, Electr. Suppl. 10 (2005) Díaz & al.: Caprellids (Crustacea: Amphipoda) from Venezuela 23

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.

Org. Divers. Evol. 5, Electr. Suppl. 10 (2005) Díaz & al.: Caprellids (Crustacea: Amphipoda) from Venezuela 24

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.

Org. Divers. Evol. 5, Electr. Suppl. 10 (2005) Díaz & al.: Caprellids (Crustacea: Amphipoda) from Venezuela 25

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.

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