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Miscel.lania Zoologica 23.2 (2000) 69 Redescription of hirsuta Mayer, 1890 (Crustacea, , Caprellidea) from the Strait of

J. M. Guerra-García, J. E. Sánchez-Moyano & J. C. García-Gómez

Guerra-García, J. M., Sánchez-Moyano, J. E. & García-Gómez, J. C., 2000. Redescription of Caprella hirsuta Mayer, 1890 (Crustacea, Amphipoda, Caprellidea) from the . Misc. Zool., 23.2: 69-78.

Redescription of Caprella hirsuta Mayec 1890 (Crustacea, Amphipoda, Caprellidea) from the Strait of Gibraltar.- Caprella hirsuta Mayer, 1890 is redescribed based on specimens collected from the Strait of Gibraltar (Southern -Northern ) during a study of the amphipod fauna from these coasts. Careful examination of these caprellids revealed differences with the previous descriptions, mainly the structure of gnathopod 2, pereopods and abdomen.

Key words: Crustacea, Amphipoda, Caprellidea, Caprella hirsuta, Strait of Gibraltar, Redescription.

(Rebut: 8 1 01; Acceptació condicional: 11 VI 01; Acc. definitiva: 3 VI1 01)

J. M. Guerra-García ('j, J. E. Sánchez-Moyano, J. C. García-Gómez, Lab. de Biología Marina, Dept. de Fisiología y Biología , Fac. de Biología, Univ de Sevilla, Apdo. 1095, E41080, Sevilla, Esparia (Spain).

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This study was supported by a grant "Programa de Formación de Profesorado Universitario y Personal Investigador AP 98 2861 7065" from the Spanish Ministry of Education, Culture and Sport and by the "Asamblea de ".

O 2000 Museu de Zoologia 7O Guerra-García et al. lntroduction a stereo-microscope; permanent mounts were made in polyvinyl-lactophenol. All the Caprella is the largest of the suborder figures were drawn with the aid of a camera Caprellidea (Crustacea, Amphipoda). It is lucida. widely distributed from the temperate to boreal regions occurring primarily on seaweeds, seagrasses and hydroids (TAKEUCHI, Results 1993). A total of 109 species of Caprella were Ca~rellahirsuta Maver. 1890 ífias. 2-51 described around the world before 1970 (MCCAIN& STEINBERG, 1970) and since then, Caprella hirsuta MAYER,1890, p. 77, pl. 2, fig. 19, more than 30 new species have been pl. 4, figs. 26-29; MONTEROSSO,1915, pp. 12-13; described (e.g. ARIMOTO,1970; Lnuei~z,1970; CHEVREUX& FAGE, 1925, p. 449, figs. 429; RANCUREL, LAUBIT~& LEWBEL,1974; VASSILENKO,1974; 1949, p. 168; RUFFO& WIESER,1952, p. 22; COSTA, ARIMOTO,1977, 1978, 1979a, 1979b, 1980, 1960, pp. 99-1 00; GOTTLIEB,1960, p. 164; MCCAIN 1982; KRAPP-SCHICKEL& RUFFO, 1986; TAKEUCHI, & STEINBERG, 1970, p. 23; CAVEDINI,1982, p. 503; 1986; VASSILENKO,1992; TAKEUCHI,1993; PLATVOET KRAPP-SCHICKEL,1993, pp. 785-786, fig. 535. et al., 1995; SANCHEZ-MOYANOet al., 1995b; MARTIN& PETTIT,1998; KRAPP-SCHICKEL& VADER, Material examined 1998; MORI, 1999). Two mature males (3.5 and 3.76 mm in body In this paper Caprella hirsuta Mayer, 1890 length), four mature females (2.1 6-3.16 mm), is described on the basis of material collected three premature females (2.06-2.23 mm), one from the Strait of Gibraltar between 1990 juvenile (1.66 mm), VI1 90 on the seaweed and 1998. This species, considered as Halopteris scoparia (L.) Sauvageau., 4 m in Mediterranean endemic since the original depth, San Garcia, Bahía de description, has also been recently reported (36"IO'N. 5'25'W), coll. J. E. Sánchez-Moyano; from the Atlantic African coast (BELLAN- six mature males (3.16-3.7 mm), 12 mature SANTINI & RUFFO, 1998). females (1.66-3.26 mm), five premature Since the morphological study of Caprella females (1.8-2.7 mm), seven juveniles (0.86- hirsuta specimens from the Strait of 1.7 mm), VI1 92, clinging to the seaweed Gibraltar, several characteristics which were Jania rubens (L.) Lamouroux, 3 m in depth, not included in previous descriptions and Torreguadiaro, Cádiz (36'20fN, 5"15'W), coll. several differences with the previous J. E. Sánchez-Moyano; two mature males descriptions have been observed. (3.4-3.7 mm), April 1995, clinging to Jania Specimens of C. hirsuta from the Strait of rubens (L.) Lamouroux, 2 m in depth, Gibraltar have been deposited in the Museu Sarchal, Ceuta (35"53'N, 5"18'W), coll. J. E. de Zoologia de , Spain (MZB 2001- Sánchez-Moyano; one mature male (3.6 mm), 0321) and the Museo Nacional de Ciencias August 1998, clinging to Halopteris scoparia Naturales de , Spain (MNCN 20.041 (L.) Sauvageau, 5 m in depth, Rompecala, 4650 and 20.0414651) Ceuta (35"54'N, 5"17'W), coll. J. M. Guerra- García; one mature male (3.73 mm) Vlll 98, clinging to Cladostephus spongiosus (Hudson) Material and methods C. Agardh, 10 m in depth, El Desnarigado, Ceuta (35"53'N, 5'17'W) coll. J. M. Guerra- The specimens of Caprella hirsuta were García. collected by scuba diving in shallow waters (2-10 m depth) in the infralittoral zone of Redescription Torreguadiaro and San García, Cádiz (Southern Male "a" (MNCN 20.0414650) from El Sarchal, Spain) and El Sarchal, El Desnarigado and Ceuta (35"53'N, 5"18'W). Rompecala, Ceuta (Northern Africa) (fig. 1). Body length. 3.7 mm. The samples were fixed using formalin Lateral view. Head rounded, without 4% in seawater solution and placed in ethyl rostrum or projections. Body smooth. alcohol 70%. Gills. Oval, length ca. twice width. Several specimens were dissected under Antennae. Antenna 1 approx. 112 body Misce1,lania Zooloaica 23.2 (2000) 7 1

t Bahía

Algeciras

San Garcla Atlantic Mediterranean Ocean Sea

Ceuta El Desnarigado

length. Peduncular article 1 a little more rounded, with dense setulae close to margin. robust than the others. Flagellum with nine Mandibular process strong, bordered by articles. Antenna 2, shorter than peduncle rectangular teeth. lncisor and lacinia mobilis of antenna 1, carrying many short simple both 5-toothed. Maxillae 1 outer lobe with setae. Swimming setae absent. Article 2 of seven forked, 2-pronged spines distally. flagellum very short. Distal article of palp with four strong spines Mouthparts. Upper lip semicircular, on end and a row of five setae medially. syrnmetrically bilobed, pubescent apically. Maxilla 2 outer lobe with two rows of simple lnner and outer lobes of the lower lip setae and one plumose setae on dista1 end. 7 2 Guerra-García et al.

propodus, with a distal round projection. lschium rectangular, as long as wide. Merus slend, length ca. twice width. Carpus short. Propodus as broad as long, with very long, dense setae dorsally. Palm with median acute tooth and deep incision. Proximally, one long tooth followed by 2 smaller teeth. Pereopods. Palm of propodus with five grasping spines following the formula 1, 2, 2 (from proximal to distal end). Dactylus with 1 plumose setae. Lateral penes, as long as wide. Abdomen with a pair of 2-articulate appendages. Proximal article of each appendage carrying several setulae and about 113 distal article in length. Distal article slender, length about 1.5 times width. Distal end serrated, with fine short setae. Dorsal lobe with a pair of plumose setae.

Female "b" (MNCN 20.0414651) from Torreguadiaro, Cádiz (36"20'N, Sol 5'W) Body length. 3.26 mm. Gnathopod 2 inserted in anterior half of pereonite 2. Propodus twice as long as broad without long setae on dorsal surface. Median acute tooth and deep incision lacking in Fig. 2. Caprella hirsuta Mayer, 1890. palm. Two pairs of iong proximal teeth. Lateral view: a. a"; b. 9. Basis without dista1 round projection. Capre/la hirsuta Mayer, 7890. vis- Anterior brood lamellae setose al1 round, ta lateral: a. a"; b. 9. posterior pair not setose. Abdomen without the 2-articled appendages.

Discussion lnner lobe with one plumose setae and a In the present paper the previous descriptions row of simple setae. lnner plate of maxilliped of Caprella hirsuta (MAYER,1890; CHEVREUX& rectangular, as big as outer plate, carrying FAGE, 1925; KRAPP-SCHICKEL,1993) are completed. two teeth and several plumose setae. Outer Furthermore, several differences with these plate rounded, provided with one tooth descriptions were found: and two simple setae distally. Palp 4- 1. Male specimens of C. hirsuta from the articulate. Article 1 short, article 2 almost as Strait of Gibraltar have a round distal wide as long, covered with numerous long projection in the basis of gnathopod 2. setae on medial margin, article 3 slender, 2. There are three proximal teeth on the article 4 with rows of setulae on grasping propodus palm of gnathopod 2 in males margin. and two pairs in females from the Strait of Gnathopods. Gnathopod 1 basis longer Gibraltar. In previous descriptions of C. than ischium to carpus combined. Propodus hirsuta from Portovenere, ltaly (see figures 4 times longer than carpus. Length of in KRAPP-SCHICKEL,1993) only two teeth are propodus ca. 1.5 times width. Palm with a reported. pair of proximal grasping spines. Gnathopod 3. Specimens from the Strait of Gibraltar 2 inserted at the middle of pereonite 2. have five grasping spines on the propodus Basis about 314 of pereonite 2, as long as palm of pereopods. In previous descriptions Miscel.lania Zoologica 23.2 (2000) 73

Fig. 3. Caprella hirsuta Mayer, 1890, a-d a*, e 9: a. Antenna 1; b. Antenna 2; c. Gnathopod 1; d. Gnathopod 2; e. Gnathopod 2. Caprella hirsuta Mayer, 1890, a-d a*, e 9: a. Antena 1; b. Antena 2; c. Gnatópodo 1; d. Gnatópodo 2; e. Gnatópodo 2. 74 Guerra-García et al. Miscel.lania Zoologica 23.2 (2000) 75

Fig. 5. Caprella hirsuta Mayer, 1890, 13:a. Upper lip; b. Lower lip; c. Maxilliped; d. Right rnandible; e. Left mandible; f. Maxillae 1; g. Maxillae 2. Caprella hirsuta Mayer, 1890, a*: a. Labio superior; b. Labio inferior; c. Maxilipedo; d. Mandíbula derecha; e. Mandíbula izquierda; f. Maxila 7; g. Maxila 2. Guerra-García et al. only three grasping spines are described Ecology (one medial and two dista1 accessory spines). 4. Male specimens from the Strait of Caprella hirsuta had previously been found Gibraltar present a striking abdomen. The on Balanus (CHEVREUX& FAGE, 1925), hydroids dista1 article of the 2-articulate appendages and Asteroidea between O and 30 m (KRAPP- has a clearly serrated margin with fine short SCHICKEL, 1993). In the present study we setae. This serration, not previously reported collected C. hirsuta from several species of in the genus Caprella, has recently also been algae with very different morphological observed in other species of caprellids from characteristics, ¡.e. Jania rubens, Halopteris Ceuta, Caprella tuberculata Bate and scoparia and Cladostephus spongiosus. This Westwood and C. santosrosai Sánchez- caprellid does not therefore seem very Moyano, Jiménez-Martín and García-Gómez specific in choosing its substrate. Never- (pers. obs.). theless, at al1 times we found C. hirsuta on substrates with much detritus. Specimens Biogeographical distribution from the Strait of Gibraltar were usually found covered with sediment. This sediment Caprella hirsuta has been considered as might be of use in evading predators, as Mediterranean endemic (KRAPP-SCHICKEL,1993) described for another caprellid species since its original description. Nevertheless, (SANCHEZ-MOYANOet al., 1995a). using information from BITAR (1987) and MENIOUI(1 988), BELLAN-SANTINI& RUFFO (1 998) recently also reported the presence of this Resumen species on the Atlantic African coast (from Cape Spartel to Cape Blanc. This represents Redescripción de Caprella hirsuta Mayer, the most occidental site for C. hirsuta (fig. 1). 1890 (Crustacea, Amphipoda, Caprellidea) On the other hand, the most oriental place del Estrecho de Gibraltar in the where C. hirsuta has been found is Israel. GOTTLIEB(1960) Se redescribe Caprella hirsuta a partir de reported the presence of C. hirsuta in ejemplares recolectados en el Estrecho de Caesarea, where this species was found during Gibraltar (sur de España-norte de África) a study of the benthonic Amphipoda of the (fig. 1) durante un estudio de la fauna de Mediterranean coast of Israel. The most cited anfípodos de esta zona. Se incluyen ilustra- localities for C. hirsuta are on the Medi- ciones completas de la especie (figs. 2-5). La terranean coast of and . In France observación detallada de los ejemplares re- the species has been found in Marseille veló algunas diferencias con las descripcio- (CHEVREUX& FAGE, 1925; BELLAN-SANTINI,1969, nes previas, principalmente en la estructura 1971). Port-Vendres and Villafranche-sur-Mer del gnatópodo 2, los pereiópodos y el abdo- (CHEVREUX& FAGE, 1925). In Italy, the presence men. of C. hirsuta has been reported in Capo Caccia (CAVEDINI,1982). Catania (MONTEROSSO,1915), Napoli (MAYER,1890). Portovenere and Acknowledgements Sampieri (RUFFO& WIESER,1952). On the Mediterranean coasts of C. We wish to express our thanks to Compañía hirsuta has been found in , del Mar and Club Calypso for field assistance. and in La Galite, (CHEVREUX& FAGE, 1925). ln Spain, JIMENO (1993) found the species in Blanes and BALLESTEROSet al. (1987) reported References its presence in the . This study shows that C. hirsuta is also present in the ARIMOTO,l., 1970. A new caprellid Amphipod Strait of Gibraltar. In Algeciras Bay its Caprella nagaoi n. sp. from Akkeshi Bay, presence is restricted to San García. In Ceuta Hokkaido. Bull. Japan. Soc. Sci. Fish., 36(7): we only found C. hirsuta in the Mediterranean 676-677. area Rompecala-El Desnarigado-El Sarchal - 1977. A new caprellid (Amphipoda, (fig. 1). Caprellidea) Caprella (Rostricephala) Miscel.lania Zooloqica 23.2 (2000)

generosa n. sp., collected in Tateyama dei Caprellidi del Mediterraneo (Crustacea: Bay, in Japan. Ann. Rep. Stud. Bunkyo Amphipoda). Boll. Mus. Civ. St. Nat. Univ. Women's College, 21: 12-14. Verona, 8: 493-53 1. - 1978. A new caprellid Caprella (Rostricephala) CHEVREAUX,E. & FAGE, L., 1925. Amphipodes. equina n. sp. (Amphipoda, Caprellidea) from Faune de France, 9: 1-488. Kunimi-ho, O'ita Prefecture, in Japan. Bull. COSTA,S., 1960. Premier apercu sur la Biogeogr. Soc. Japan, 33(3): 33-35. répartition des Caprelles dans la région - 1979a. A new species Caprella (Caprella) de Villefranche-sur-Mer. Rec. trav. Sta. liliata (Amphipoda, Caprellidea) from mar d'Endoume, 33: 99-101. Uchiura Bay, Fukuri Prefecture, Japan~Ann. GOTTLIEB,E., 1960. The benthonic Amphipoda Rep. Stud. Bunkyo Univ. Women's College, of the Mediterranean coast of Israel. l. 23: 1-2. Notes on the geographical distribution. - 1979b. A new caprellid Caprella (Rostrhicephala) Bull. Res. council Israel, zoology, 9(2-3): dissona n. sp. (Amphipoda, Caprellidea) from 157-1 66. Toyoma, Fukushima Prefecture, Japan. Proc. JIMENO, A., 1993. Contribución al estudio de Japan. Soc. Syst. Zool., 16: 33-34. los Anfípodos de las costas mediterráneas - 1980. A new species Caprella (Caprella) catalanas. Estudio faunísitico, ecológico, corallina n. sp. (Amphipoda, Caprellidea) biológico y biogeográfico. Tesis doctoral, from Ogasawara, Japan. Ann. Rep. Stud. Universidad de Barcelona. Bunkyo Univ. Women's College, 24: 55-56. KRAPP-SCHICKEL,T., 1993. Suborder Caprellidea. - 1982. Three species of caprellid amphipods In: The Amphipoda of the Mediterranean from Ogasawara Islands, with a descrip- (S. Ruffo, Ed.). Mém. I'lnst. Ocean. tion of a new species. Proc. Japan. Soc. Monaco, 13(3): 773-809. Syst. Zool., 22: 21-23. KRAPP-SCHICKEL,T. & RUFFO, S., 1986. A new BALLESTEROS,M., CASTELL~, J., CALLES,M. & SARDA, Caprella of the Mediterranean Sea ., 1987. Invertebrados alguícolas marinos (Crustacea: Amphipoda). Boll. Mus. Civ. de las Islas Pitiusas. Consell Insular d'Eivissa St. Nat Verona, 13: 437-441. i Formentera. Conselleria dlEcologia i Medi KRAPP-SCHICKEL,T. & VADER, W., 1998. What is, Ambient. and what is not, Caprella acanthifera BELLAN-SANTINI,D., 1969. Contribution a Leach, 1814 (Amphipoda, Caprellidea)? I'étude des peuplements infralittoraux sur Part 1: the acanthifera-group. J. Nat. Hist., substrat rocheux (étude qualitative et 32: 949-967. quantitative de la frange supérieure). LAUBITZ,D. R., 1970. Studies on the Caprellidea Recueil des travaux de la Station marine (Crustacea: Amphipoda) of the American dlEndoume, 63: 1-294. North Pacific. Publications in Biological - 1971. Étude des Crustacés amphipodes de Oceanography, 1. Natural Museum of la biocénose des Algues photophiles dans Natural Sciences, Otawa, Canada. la région provencale. Rapports etprocés- LAUBIR,D. R. & LEWBEL,G. S., 1974. A new verbaux des réuninos. Commission species of caprellid (Crustacea: Amphipoda) internationale pour I'exploration scien- associated with gorgonian octocorals. Can. tifique de la mer Méditerranée, 20(3): J. ZOO~.,52(5): 549-551. 221-223. MARTIN,J. W. & PETTIT,G., 1998. Caprella BELLAN-SANTINI,D. & RUFFO, S., 1998. Faunistics bathytatos new species (Crustacea, and Zoogeography in The Amphipoda of Amphipoda, Caprellidea), from the the Mediterranean, S. Ruffo (Ed.). Mém. mouthparts of the crab Macroregonia I'lnst. Ocean. Monaco, 13(4): 895-91 1. macrochira Sakai (Brachyura, Majidae) in BITAR,C., 1987. Étude de peuplements the vicinity of deep-sea hydrotermal vents benthiques littoraux des c6tes atlantiques off British Columbia. Bull. Mar. Sci., 63(1): et méditerranéennes du Maroc. lmpact de 189-1 98. la pollution. Comparaisons biogéo- MAYER,P., 1890. Die Caprelliden des Golfes graphiques. These pour I'obtention du von Neapel und der angrenzenden grade de Docteur es Sciences de I'Univer- Meeres-Abschnitte. Fauna Flora Golf. sité dfAix-Marseille 11. Neapel, 17: 1-55. CAVEDINI,P., 1982. Contibuto alla conoscenza MCCAIN,J. C. & STEINBERG, J. E., 1970. Guerra-García et al.

Amphipoda-1, Caprellidea-l. Crustaceorum sistematiche ed ecologiche su alcuni Catalogus, 2: 1-78. Anfipodi delle xoste mediterranee MENIOUI,M., 1988. Contribution a la italiane. Mem. Mus. Civ. St. Nat. Verona, connaissance des peuplements infra- 3: 11-30. littoraux superficiels des c6tes atlanto- SANCHEZ-MOYANO,J. E., CARBALLO, J. L. & ESTACIO, méditerranéennes du Maroc (Étude F. J., 1995a. Pedoculina garciagomezi faunistique, écologique et biogéo- (Amphipoda: Caprellidea), new species graphique, These pour obtenir le grade from Bahía de Algeciras (Southern Spain). de Doctorat d'État es-Sciences Biolo- Crustaceana, 68: 41 8-427. giques, Université Mohammed V, . SANCHEZ-MOYANO,J. E., JIMÉNEZ-MART~N,J. A. & MONTEROSSO,B., 191 5. del golfo GARC~A-G~MEZ,J. C., 1995b. Caprella di Catania. Atti della Accademia Gioenia santosrosai n. sp. (Amphipoda: Capre- di scienze naturali in Catania, 8: 1-16. Ilidea) from the Strait of Gibraltar MORI,A., 1999. Caprella kuroshio, a new (Southern Spain). Ophelia, 43(3): 197-204. species (Crustacea: Amphipoda: Caprelli- TAKEUCHI,l., 1986. Caprella kominatoensis n. dea), with a redescription of Caprella sp. (Amphipoda: Caprellidea) from circur Mayer, 1903, and an evaluation of Amatsu-Kominato, Chiba, Japan. Bull. the genus Metacaprella Mayer, 1903. Proc. Nat. Sci. Mus. Tokyo, Ser. A (Zool.), 12: Biol. Soc. Wash., 112(4): 722-738. 67-73 PLANOET,D., DEBRUYNE, R. H. & GMELIGMEYLING, - 1993. Caprella arimotoi, a new species A. W., 1995. Description of a new (Crustacea: Amphipoda: Caprellidea) from Caprella-species from the Netherlands: the Seto lnland Sea, Japan. Proc. Biol. Caprella macho nov. spec. (Crustacea, Soc. Wash., 106: 11 5-121. Amphipoda, Caprellidea). Bull. Zool. VASSILENKO,S. V., 1974. Caprellids of the sea Museum, 5(1): 1-4. of USSR and adjacent waters. Opredelpo RANCUREL,P. ,1949. Notes sur les Amphipodes Faune USSR, 107: 1-287. marins de la région de Marseille. Bull. - 1992. A new species of the genus Caprella Mus. Hist. Nat, Marseille, 9(3): 165-172. from the sublittoral of the Komandor RUFFO,S. & WIESER,W., 1952. Untersuchungen Islands, North West Pacific (Crustacea: über die Algenbewohnende Mikrofauna Amphipoda: Caprellidae).Zoosystematica mariner Hartboden. II. Observazioni Rossica, 1: 13-17.