Clavellotis sebastidis sp. nov. (Copepoda, Lernaeopodidae) parasitic on Sebastes oculatus Valenciennes, 1833 from Argentina

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Acta Parasitologica, 2005, 50(1), 74 79; ISSN 1230-2821 Copyright 2005 W. Stefañski Institute of Parasitology, PAS Clavellotis sebastidis sp. nov. (Copepoda, Lernaeopodidae) parasitic on Sebastes oculatus Valenciennes, 1833 from Argentina Stefański Raúl Castro 1* and M. Teresa González 2 1 Departamento Acuicultura, Facultad de Recursos del Mar, Universidad de Antofagasta, Casilla 170, Antofagasta; 2 Instituto de Ecología y Evolución, Facultad de Ciencias, Universidad Austral de Chile, Casilla 567, Valdivia; Chile Abstract A new species of Clavellotis Castro et Baeza, 1984 (Copepoda, Lernaeopodidae) parasitic on Patagonian redfish, Sebastes oculatus Valenciennes, 1833 from southern coast of Argentina is described and illustrated. The new species of Clavellotis differs from its congeners by the length of second maxilla, shape of the female trunk, length of genital process, and a combination of characters. The male also shows differences in the second maxilla, the maxilliped and in the shapes of the trunk and genital process. This is the first record of Clavellotis from the southern Atlantic Ocean and also the first record on a Sebastes species. Key words Copepoda, Lernaeopodidae, Clavellotis, new species, fish, Sebastes, Argentina Introduction Skóra Among the copepods that parasitize marine fishes, the family Lernaeopodidae includes 45 genera (Benz et al. 2000), someon teleost and others on elasmobranch fishes. All of them are characterized by their mode of attachment by means of a bulla at the end of the second maxilla (with the exception of Dendrapta Kabata, 1964, Schistobrachia Kabata, 1964 and Brianella Wilson, 1915). The bulla is inserted into the host tissue, facilitating a strong and permanent attachment allowing the copepod to have a range of action, depending upon the length of the second maxilla, in order to obtain its food. There is no record in the literature for lernaeopodids that are parasitic on Sebastes in Argentinean marine waters. The only lernaeopodids reported from Argentinean waters are Lernaeopoda bivia Leigh-Sharpe, 1930 on Halaelurus bivius [valid name Schroederichthys bivius (Müller et Henle, 1838)], from Puerto Deseado (Kabata 1986), and Lernaeopoda mustelicola Leigh-Sharpe, 1919 on Mustelus schmitti Springer, 1939 (cf. Brian 1944 after Etchegoin and Ivanov 1999). The Sebastes species are parasitized in other latitudes by some lernaeopodids, such as Neobrachiella robusta (Wilson, 1912) (cf. Dojiri 1981, Kabata 1987). The study of Sebastes oculatus collected in Argentine waters has permitted the detection of lernaeopodid specimens from the Clavella group (Kabata 1979) attached to their branchial arches (gill rakers). For the lernaeopodids, the male type gives an important clue that permits the definition of the taxonomic position of some specimens (Wilson 1915; Kabata 1979, 2004; Castro and Baeza 1984). In this case, the male characteristics permit us to identify the present specimens as belonging to the genus Clavellotis Castro et Baeza, 1984. This genus includes eight species (Kabata 1990) plus the most recent addition of C. briani (cf. Benmansour et al. 2001). Characteristics of the female and male specimens are described, and their taxonomic identity is determined by comparison with all Clavellotis species, leading to the proposal of a new species, C. sebastidis. Materials and methods Drawings were made with the aid of a camera lucida attached to a stereomicroscope or compound microscope. Measure- * Corresponding author: rcastro@uantof.cl

Clavellotis sebastidis sp. nov. on S. oculatus from Argentina 75 Śląski 1 2 4 5 3 Figs 1 4. Female Clavellotis sebastidis sp. nov.: 1. Female non-ovigerous, entire lateral view, with male attached; B bulla, M male, SM second maxilla, arrow indicating the genital process. 2. Female ovigerous, entire lateral; arrow indicating the genital process. 3. Female trunk dorsal view; arrow indicating the genital process. 4. First antenna; W whip, 1, 3, 4, 5, 6 numbers of setae

76 Stanisła Zdzisław Raúl Castro and M. Teresa González ments were made in micrometres, with a reticulated eye piece attached to the microscope. The terminology adopted follows that of Kabata (1979). Results Clavellotis sebastidis sp. nov. (Figs 1 14) Female: Cephalothorax cylindrical, longer than trunk; latter suboval with developed genital process. Some specimens with short anal process protruding from genital process. Second maxilla long, about as long as trunk or slightly shorter than cephalothorax, fused at apex, bearing distally short bulla, with rounded swelling on each side. Measurements (based on 20 specimens): Cephalothorax length 1702.6 (1281 2205), width 337 (256 462); second maxilla length 1174.8 (974 1461), width 387 (231 641); trunk length 1255 (949 1590), width 811.8; genital process length 212.6 (103 282), width 174 (103 256); egg sac length (n = 8), 947.5 (641 1103), diameter 362 (259 462). Appendages: First antenna (Fig. 4) apparently three-segmented, distal armature with 5 visible elements (setae: 1, 3, 4, 5 and 6). Seta 6 longer than others. Second antenna (Fig. 5): Exopod long, axis in line with sympod, bulbous, armed with spinules on outer surface. Endopod apparently two-segmented, distal segment armed with three spines, two of them of equal size, third smaller. Mandible (Fig. 6) with three secondary teeth, dental formula P1S1, P1S1, P1S1, B4. First maxilla (Fig. 7) biramous; exopod with two digitiform papillae, each bearing apical long seta; short seta at base of dorsal papilla; exopod lateral, short, with two subequal apical setae. No other armature detected. Second maxilla (Figs 1 and 2) as long as trunk and/or cephalothorax, slender, with rounded lateral swellings at base, fused at apex. Bulla short, typical for Clavellotis. Maxilliped (Fig. 8): Robust corpus, myxa bearing single seta. Subchela cylindrical, with spiniform process on lateral surface, distally with basal seta and sparse denticulation on distal part of inner margin; distal claw slightly curved. Male (Fig. 9) typical of Clavellotis Castro et Baeza, 1984. Body sack-like; trunk abbreviated, much smaller than cephalothorax, genital process prominent. First antenna (Fig. 10A) three-segmented, basal segment with long whip. Distal segment armature of 5 elements, seta 6 longer than others. Second antenna (Fig. 10B): Exopod bulbous and long, covered on distal outer surface with small spiniform processes and short spine; endopod much longer than exopod, with two segments, distal segment with outer distal margin armed with two spines, and a pad of short spinules ventrally, also pad of short spinules at base of first segment. Mandible (Fig. 11) with three secondary teeth, dental formula P1S1, P1S1, P1S1, B4. First maxilla (Fig. 12): Endopod with two papillae, each surmounted by long seta; short seta at base of dorsal papilla. Exopod ventrolateral; short papilla with two setae of about equal size. Second maxilla (Fig. 13) globose, strong, slightly curved claw closing against prominent inner part of corpus. Maxilliped (Fig. 14) subrectangular, with long, slender base, bearing two rows of spiniform processes, each row separated from other, but appearing fused in dorsal view. Locality: Golfo San Jorge (43 S), Argentina. Host: Patagonian redfish, Sebastes oculatus. Habitat: Branchial arches (gill rakers). Material deposited in the British Museum of Natural History: holotype no. 2004.556; paratypes nos. 2004.557 558, allotype no. 2004.559. Etymology: The specific name sebastidis refers to the generic name of the host. Discussion The males of the specimens recorded in this study show morphology typical of the Clavella group. Consequently, these specimens can be assigned to Clavellotis Castro et Baeza, 1984, based principally on the males, which permit clear discrimination among all those genera within the Clavella group, and fit well within the amended diagnosis of Kabata (1990). To date, the genus Clavellotis included nine species: C. dilatata (Krøyer, 1863) type species, and 7 species that were transferred from the genus Clavellopsis Wilson, 1915 by Kabata (1990): C. bilobata (Pillai, 1962), C. branchiostegi (Yamaguti, 1939), C. characis (Richiardi, 1880), C. fallax (Heller, 1856), C. pagri (Krøyer, 1863), C. sargi (Kurz, 1877), C. strumosa (Brian, 1906), and a recently added C. briani Benmansour, Ben-Hassine, Diebakate et Raibaut, 2001, plus a tentative, unnamed member of this genus (Roubal 1981), whose female was never fully described and more information is required for a definitive decision on the status of this species (Kabata 1990). Among all species of Clavellotis, the new species differs from a group that shares a well-developed aliform lateral projection from the cephalothorax base (C. bilobata, C. briani, C. branchiostegi, C. dilatata, C. fallax and C. strumosa). The remaining species are more closely related to the new species, due to the presence of a subcircular lateral projection (C. characis, C. pagri and C. sargi). The new species can be differentiated from C. sargi because, in the latter, the female bears a very long genital process, and the trunk is subcircular. The male shows differences in the trunk and genital process, in the second maxilla and in the maxilliped shape. C. pagri also differs in the female trunk shape and in the length of the second maxilla, which is longer in the new species. Other differences are found in the male second maxilla, maxilliped, and in the shapes of the trunk and genital process. The present specimens, parasitic on S. oculatus, can be distinguished from C. characis by the length of the female genital process, which is longer in C. characis, by the trunk

Clavellotis sebastidis sp. nov. on S. oculatus from Argentina 77 Roborzyński rosbśźćv fjad kadsććżć shape, and especially by the length of the second maxilla, which is shorter in C. characis. The male shows other differences in the length of the trunk and the genital process, as well as in the shapes of the second maxilla and maxilliped. Consequently, the presently described Clavellotis specimens cannot be considered conspecific with any of the previously mentioned species. The second maxilla is longer than in those species. Clavellotis shows typically distinctive ap- Figs 5 8. Female Clavellotis sebastidis sp. nov., appendages: 5. Second antenna; EN endopod, E exopod. 6. Mandible. 7. First maxilla; E exopod, EN endopod. 8. Maxilliped; B basal seta, C corpus, CL claw, D denticles, SH shaft

78 Raúl Castro and M. Teresa González pearances in the trunk and genital process. This, in addition to differences in the maxilliped and second maxilla, preclude its inclusion within the previously described species of Clavellotis. The authors suggest the creation of a new taxon for the specimens that parasitize S. oculatus; the name proposed is Clavellotis sebastidis. This is the first record of Clavellotis from the southern Atlantic, and the first on Sebastes species. Figs 9 14. Male Clavellotis sebastidis sp. nov.: 9. Male entire, lateral view; BC buccal cone, GP genital process, SM second maxilla, M maxilliped. 10. First antenna and second antenna; A first antenna, B second antenna, E exopod, EN endopod, W whip, 6 seta six. 11. Mandible. 12. First maxilla. 13. Second maxilla. 14. Maxilliped

Clavellotis sebastidis sp. nov. on S. oculatus from Argentina 79 Acknowledgements. The collection of the parasites of S. oculatus from the Argentine coast was possible with economic support given to M.T. González by the D.I.D. of the Universidad Austral de Chile (project D-2003-09). References Benmansour B., Ben-Hassine O.K., Diebakate D., Raibaut A. 2001. Sur deux species des copepodes Lernaeopodidae (Siphonostomatoida) parasite du marbré Lithognathus mormyrus (Linnaeus, 1758) (Pisces, Sparidae). Zoosystema, 23, 695 703. Benz G.W., Kabata Z., Bullard S.A. 2000. Margolisius abditus n. gen., n. sp. (Copepoda: Lernaeopodidae) from gill lamellae of a remora (Remora remora) collected in the Gulf of California. Journal of Parasitology, 86, 241 244. Castro R., Baeza H. 1984. Clavellotis, new genus (Copepoda, Lernaeopodidae), and re-description of Clavellotis dilatata (Krøyer, 1863). Journal of Crustacean Biology, 4, 688 694. Dojiri M. 1981. Copepods of the families Lernaeopodidae and Naobranchiidae, parasitic on fishes from the Southern California inshore waters. Journal of Crustacean Biology, 1, 251 264. Etchegoin J., Ivanov U.A. 1999. Parasitic copepods of narrownose smooth-hound shark Mustelus schmitti (Chondrichthyes: Triakidae) from Argentina. Folia Parasitologica, 46, 149 153. Kabata Z. 1979. Parasitic Copepoda of British fishes. The Ray Society, London. Kabata Z. 1986. Redescriptions of and comments on four little-known Lernaeopodidae (Crustacea: Copepoda). Canadian Journal of Zoology, 64, 1852 1859. Kabata Z. 1987. The developmental stages of Neobrachiella robusta (Wilson, 1912), a parasitic copepod of Sebastes (Teleostei: Scorpaeniformes). Canadian Journal of Zoology, 65, 1331 1336. Kabata Z. 1990. Revision of the genus Clavellopsis Wilson, 1915 (Copepoda: Lernaeopodidae). Canadian Journal of Zoology, 68, 2564 2566. Kabata Z. 2004. Some comments on the genus Lernaeopodina Wilson, 1915 (Copepoda: Siphonostomatoida: Lernaeopodidae). Systematic Parasitology, 57, 15 17. Roubal F.R. 1981. The taxonomy and its site specificity of the metazoan ectoparasites on the black bream Acanthopagrus australis (Gunther), in Northern South Wales. Australian Journal of Zoology, Suppl., Ser. N. 84. Wilson C.B. 1915. North American parasitic copepods belonging to Lernaeopodidae, with a revision of the entire family. Proceedings of the United States National Museum, 47, 565 729. (Accepted October 13, 2004)