The Copepod Genus Herrmannella (Poecilostomatoida) Associated with Marine Bivalve Mollusks at Kodiak Island, Alaska 1

The Copepod Genus Herrmannella (poecilostomatoida) Associated with Marine Bivalve Mollusks at Kodiak Island, Alaska 1

ARTHUR G. HUMEs 2

ABSTRACT: The sabelliphilid copepods Herrmannella kodiakensis Humes, n. sp., and H. saxidomi (lllg, 1949) are reported from the marine bivalve Sax idomus giganteus (Deshayes). The new species may be differentiated from its congeners by the shape of the genital double-somite in the female (with "shoulders"). Copepodids of both species were found in S. giganteus and Pro tothaca staminea (Conrad). These are the first records of Herrmannella in Alaska.

THE GENUS Herrmannella currently contains longicaudata Avdeev, 1975, and H. longi 22 species living in the mantle cavity of chaeta Avdeev, 1975; Korea: H. soleni Kim marine intertidal or shallow-water Bivalvia. & Ho, 1991, H. hoonsooi Kim, 1992, and H. Several species occur in Europe: H. pecteni exigua Kim, 1993. (Sowinski, 1884), H. rostrata Canu, 1891, H. In this paper Herrmannella kodiakensis haploceras (Bocquet & Stock, 1959), H. bar Humes, n. sp., and H. saxidomi Illg, 1949 neae (pelseneer, 1929), and H. duggani Holmes are reported from the venerid bivalve Sax & Minchin, 1991; one in West Africa: H. in idomus giganteus (Deshayes) at Kodiak jlatipes (Humes & Cressey, 1958); two in the Island, southern Alaska. Copepodids of both West Indies: H. caribaea Humes, 1970 and species of Herrmannella are recorded from S. H. dissidens Humes, 1970; and two in Chile: giganteus and the venerid Protothaca stami H. mesodesmatis Humes, 1967 and H. proto nea (Conrad). These are the first records of thacae Humes, 1967. The European species Herrmannella from Alaska. Herrmannella parva Norman & T. Scott, The copepods were cleared in lactic acid 1905, and H. valida G. O. Sars, 1918, are not and the dissections prepared using the method known to be associated with a host (see of Humes and Gooding (1964). In the figure Marine Biological Association 1957, Holmes captions, the letter after the explanation of and Gotto 1992). each figure refers to the scale at which it was Thirteen species of Herrmannella are as drawn. sociated with various shallow-water or inter tidal bivalves in the Pacific Ocean north ofthe equator, as follows. California and Wash Order POECILOSTOMATOIDA Thorell, 1858 ington: H. columbiae (Thompson, 1897) (see Family SABELLIPHILIDAE Gurney, 1927 Illg 1949), H. panopeae Illg, 1949 (see also Stout, 1949), H. saxidomi Illg, 1949, H. Herrmannella kodiakensis Humes, n. sp. tivelae Illg, 1949, H. perplexus Illg, 1949, Figures la-i, 2a-l, 3a-i and H. bullata Humes & Stock, 1973; Peter the Great Bay, Sea of Japan: H. dentata TYPE MATERIAL. 20 n, 7 c3c3 from 13 Avdeev, 1987, H. hiatellai Avdeev, 1975, H. Saxidomus giganteus (Deshayes), intertidal, mouth ofLarsen Bay, Kodiak Island, Alaska, 57° 31.3' N, 153° 58.7' W, 1 September 1993, IThis work was supported by a grant from the Na Thomas K. Duncan collector. Holotype 5j2 tional Science Foundation (BSR 88-21979). Manuscript accepted 21 October 1994. (USNM 268329), allotype c3 (USNM 268330), 2Boston University Marine Program, Marine Bio and 21 paratypes (16 n, 5 c3c3) (USNM logical Laboratory, Woods Hole, Massachusetts 02543. 268331) deposited in the National Museum 289 A E E o ~ h

FIGURE 1. Herrmannella kodiakensis, n. sp. Female. a, dorsal (scale A); b, lateral (A); c, urosome, dorsal (B); d, somite bearing leg 5 and genital double~somite, ventral (B); e, somite bearing leg 5 and genital double-somite, lateral (B);f, genital area, dorsal (D); g, anal somite and caudal ramus, dorsal (C); h, rostrum, ventral (C); i, rostrum, lateral

(D). AI> antennule; A2, antenna.

'waya- Herrmannella Associated with Marine Bivalves-HUMEs 291 of Natural History, Smithsonian Institution, and 35 J1.m at greatest width. Claw 50 J1.m. Washington, D.C. Seta near base of claw slightly jointed. Lab rum (Figure 2e) with 2 widely diverging FEMALE: Body (Figure la,b) moderately lobes. Mandible (Figure 2d), paragnath, and elongate, prosome flattened, with rostral area maxillule (Figure 2e) similar to those of con projecting in dorsal view. Length (not includ geners. Maxilla (Figure 2f) with lash bearing ing setae on caudal rami) 1.30 (1.13-1.43) large, stout spines grading distally to small, mm and greatest width 0.41 (0.37-0.44) mm, slender spines. Maxilliped (Figure 2g) slen based on 10 specimens in lactic acid. Greatest der, third segment with 1 minute spinule. dorsoventral thickness 0.29 mm. Somite Ventral area between maxillipeds and first bearing leg I separated from cephalosome pair of legs (Figure 2h) projecting slightly in by dorsal transverse suture, and with epi lateral view (Figure Ib). mera not expanded. Somites bearing legs 2 Legs 1-4 (Figures 2i,k,l, 3a) segmented 4 with epimera expanded and rounded. Ratio and armed as in congeners. Exopod of leg 1 of length to width of prosome 1.95: 1. Ratio with outer spines having prominent lateral of length of prosome to that of urosome spinules along proximal sides (Figure 2j). 1.15: 1. Somite bearing leg 5 (Figure Ie) 104 Inner coxal seta on leg 4 33 J1.m long. Leg 5 by 148 J1.m. Genital double-somite (Figure (Figure 3b) with unomamented free segment Id) 156 J1.m long, 180 J1.m wide in laterally 86 by 31 J1.m, width taken at proximal inner expanded anterior two-thirds, with rounded expansion. Ratio 2.97: 1. Two unequal ter anterior "shoulders" and indented poster minal setae, 35 J1.m and 78 J1.m. Dorsal seta iorly, posterior third 104 J1.m wide with paral short, 20 J1.m. All setae smooth. Leg 6 prob lel sides. In lateral view (Figure Ie) this ably represented by 2 setae on genital area somite slightly expanded dorsally at mid (Figure If). region. Genital areas located dorsolaterally Color of living specimens slightly opaque in expanded anterior two-thirds. Each area gray to hyaline, eye red. (Figure If) with 2 small setae 13 J1.m. Three postgenital somites, from anterior to pos MALE: Body (Figure 3e,d) resembling that terior, 70 by 88, 60 by 83, and 93 by 75 J1.m. of female. Length (excluding setae on caudal Caudal ramus (Figure Ig) elongate, 152 rami) 1.15 (1.07-1.29) mm and greatest width by 26 J1.m (greatest width), ratio of length to 0.30 (0.26-0.32) mm, based on 6 specimens width 5.85 : 1. Outer lateral seta, located at in lactic acid. Greatest dorsoventral thick midlength, 42 J1.m; dorsal seta minute, 8 J1.m; ness 0.23 mm. Ratio of length to width of outermost terminal seta 47 J1.m; innermost prosome 2.08 : 1. Ratio of length of prosome terminal seta 50 J1.m; and 2 median terminal to that of urosome 1.18: 1. Somite bearing setae, 121 J1.m (outer) and 242 J1.m (inner), leg 5 (Figure 3e) 44 by 83 J1.m. Genital somite both slightly swollen beyond articulation. subspherical in dorsal view, 138 by 130 J1.m. All setae smooth. Body surface smooth, with Four postgenital somites, from anterior to out visible sensilla. Egg sac not seen. posterior, 57 by 78, 57 by 65, 57 by 59, and Rostrum (Figure Ih) projecting slightly 68 by 58 J1.m. Caudal ramus resembling that and rounded posteriorly (Figure 1i). Anten of female. Body surface as in female. nule (Figure 2a) 286 J1.m long, its 7 segments Rostrum like that of female. Antennule armed with 4, 13, 6, 3, 4 + 1 aesthete, 2 + 1 similar to that of female but 3 aesthetes aesthete, and 7 + 1 aesthete. Lengths of seg added (at points indicated by dots in Figure ments (measured along their posterior non 2a). Antenna like that of female. Labrum, setiferous margins): 18 (55 J1.m along anterior mandible, paragnath, maxillule, and maxilla margin), 78, 26, 36, 36, 27, and 21 J1.m, re as in female. Maxilliped (Figure 3f) slender, spectively. All setae smooth. Antenna (Fig elongate, 4-segmented, proximal part of claw ure 2b) 4-segmented, 190 J1.m long. Armature assumed to represent fourth segment. First 1, 1, 3, and 1+3. Second segment with distal segment unarmed. Second segment with 2 inner comer projecting. Fourth segment 47 inner setae, 1 slender, 1 stout, and numerous J1.m along inner side, 29 J1.m along outer side, small spines. Small third segment unarmed. c ~

FIGURE 2. Herrmannella kodiakensis, n. sp. Female. a, antennule, dorsal (scale D); b, antenna, anterior (inner) (0); c, labrum, with positions of paragnaths indicated by broken lines, ventral (0); d, mandible, anterior (E); e, max illule, anterior (E);f, maxilla, anterior (F); g, maxilliped, antero-inner (F); h, area between maxillipedg (MXPD) and first pair oflegs (PI), ventral (C); i, leg 1 and intercoxal plate, anterior (C);j, spines on exopod ofleg I, anterior (F); k, leg 2 and intercoxal plate, anterior (C); I, leg 3 and intercoxal plate, anterior (C).

• FIGURE 3. Herrmannella kodiakensis, n. sp. Female. a, leg 4 and intercoxal plate, anterior (scale C); b, leg 5, dorsal (F). Male. c, dorsal (scale A); d, lateral (A); e, urosome, dorsal (D); f, maxilliped, inner (F); g, endopod of leg 2, anterior (D); h, genital somite showing leg 6, ventral (C); i, spermatophores, attached to female, ventral (D). 294 PACIFIC SCIENCE, Volume 49, July 1995

Claw 150 Jlm, with 2 very unequal proximal LITERATURE CITED setae. Ventral area between maxillipeds and first pair of legs as in female. AVDEEV, G. Y. 1975. Three new species of Legs 1, 3, and 4 similar to those offemale. commensal copepods of the genus Herr Exopod of leg 2 like that of female, but en mannella Canu (Cyclopoida, Sabelliphili dopod showing slight sexual dimorphism in dae) from Bivalvia in Peter the Great Bay. spiniform processes on third segment (Figure Izv. Tikhookean. Nauchno-Issled. Inst. 3g). Leg 5 (Figure 3e) with small rectangular Rybn. Khoz. Okeanogr. 98: 222-232 (in free segment 13 by 4 Jlm. Leg 6 (Figure 3h) Russian). posteroventral flap on genital somite bearing ---. 1987. Two new copepod species 2 setae, 21 Jlm and 34 Jlm. (Sabelliphilidae, Poecilostomatoida) from Spermatophore (Figure 3i) oval, ca. 70 by bivalvian molluscs in the Peter the Great 39 Jlm not including neck. Bay of the Japan Sea. Zool. Zh. 66: 608 Color of living specimens as in female. 613 (in Russian). BOCQUET, C., and J. H. STOCK. 1959. Cope ElYMOLOGY: The specific name kodiakensis podes parasites d'invertebres des cotes refers to the island where the copepods were de France. X. Sur les especes de Paran found. thessius (Cyclopoida, Lichomolgidae) du REMARKS: One feature of the new species groupe des Herrmannella, associes aux serves to distinguish it from all congeners: pelecypodes. Proc. K. Ned. Akad. Wet. the shape of the genital double-somite in the Ser. C BioI. Med. Sci. 62:238-249. female, with anterior "shoulders." In addi CANU, E. 1891. Les copepodes marins du tion, the free segment of leg 5 in the female, Boulonnais. Y. Les semi-parasites. Bull. with a distinct proximal inner expansion, dif Sci. Fr. Belg. 23 :467-487. fers from all congeners except Herrmannella HOLMES, J. M. C., and R. Y. Gorro. 1992. A longicaudata Avdeev, 1975. The female of list of the Poecilostomatoida (Crustacea: that Asian species, however, has an elongate Copepoda) of Ireland. Bull. Jr. Biogeogr. genital double-somite with subparallel sides. Soc. 15: 1-32. HOLMES, J. M. C., and D. MINCHIN. 1991. A Herrmannella saxidomi rug, 1949 new species of Herrmannella (Copepoda, Poecilostomatoida, Sabelliphilidae) asso MATERIAL EXAMINED: 4 ~~, 1 d' from 13 ciated with the oyster Ostrea edulis L. Saxidomus giganteus (Deshayes), intertidal, Crustaceana 60: 258-269. mouth of Larsen Bay, Kodiak Island, 0 0 HUMES, A. G. 1967. Cyclopoid copepods of Alaska, 57 31.3' N, 153 58.7' W, 1 Septem the genus Paranthessius associated with ber 1993, Thomas K. Duncan collector. marine pelecypods in Chile. Proc. U.S. REMARKS: This species is known from Natl. Mus. 124(3628): 1-43. Saxidomus nuttallii Conrad in California --. 1970. Cyclopoid copepods of the (rug 1949). genus Paranthessius associated with ma Several hundred copepodids of H sax rine pelecypods in the West Indies. Bull. idomi were recovered from the 13 specimens Mar. Sci. 20: 605-625. of Saxidomus giganteus and a much smaller HUMES, A. G., and R. F. CRESSEY. 1958. number from six Protothaca staminea (Con Copepod parasites of mollusks in West rad) at Kodiak Island. Africa. Bull. Inst. Fr. Afr. Noire Ser. A Sci. Nat. 20: 921-942. HUMES, A. G., and R. U. GOODING. 1964. A ACKNOWLEDGMENTS method for studying the external anat omy of copepods. Crustaceana 6: 238 I thank Thomas K. Duncan for collecting 240. the bivalve hosts and transporting them to HUMES, A. G., and J. H. STOCK. 1973. A re Woods Hole. vision of the family Lichomolgidae Koss-

dZSi Herrmannella Associated with Marine Bivalves-HuMES 295

mann, 1877, cyclopoid copepods mainly PELSENEER, P. 1929. Copepodes parasites de associated with marine invertebrates. mollusques. Ann. Soc. R. Zoo!. Belg., 1928, Smithson. Contrib. Zoo!' 127: i-v, 1-368. 59: 33-49. ILLG, P. L. 1949. A review of the copepod SARS, G. O. 1918. An account of the Crusta genus Paranthessius. Proc. U.S. Nat!. Mus. cea of Norway with short descriptions 99: 391-428. and figures of all the species. Vo!. VI, Co KIM, I.-H. 1992. Herrmannella hoonsooi, a pepoda, Cyclopoida, parts XIII, XIV, new species of Copepoda (Poecilostoma Lichomolgidae (concluded), Oncaeidae, toida, Sabelliphilidae) associated with a Corycaeidae, Ergasilidae, Clausiidae, Eu bivalve from Korea. Korean J. Zoo!., nicicolidae, Supplement. Pages 173-225. special issue 3: 77-84. Bergen, Norway. ---. 1993. Two new species of poecilo SOWINSKY, W. 1884. K faune rakoobraznikh stomatoid Copepoda associated with razor Cernago Morya. I. 0 njekotorykh para clams (Bivalvia, Solenidae) in the Yellow sitnykh formakh iz gruppy Copepoda. Sea. Korean J. Zoo!' 9: 191-202. Mem. Kiev Soc. Nat. Hist. 7:225-262. KIM, I.-H., and J.-S. Ho. 1991. Copepod STOUT, W. E. 1970. Some associates of Tresus parasites of commercial bivalves from nuttallii (Conrad, 1837) (pelecypoda: Korea. 1. Two new poecilostomatoid spe Mactridae). Veliger 13: 67-70. cies from Solen grandis Dunker in the THOMPSON, I. C. 1897. Appendix: Note on Yellow Sea. Korean J. Syst. Zoo!. 7: 1-12. dredging and tow-netting in Puget Sound, MARINE BIOLOGICAL ASSOCIATION OF THE Pacific Coast. In W. A. Herdman, I. C. UNITED KINGDOM. 1957. Plymouth marine Thompson, and A. Scott. On the plankton fauna, 3 ed. Plymouth, England. collected continuously during two tra NORMAN, A. M., and T. SCOTT. 1905. Crus verses ofthe North Atlantic in the summer tacea Copepodanew to science from Devon of 1897; with an appendix on dredging in and Cornwall. Ann. Mag. Nat. Hist. (7) Puget Sound. Proc. Trans. Liverpool Bio!. 15: 284-300. Soc. 12: 35-90.