PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 112(1):181-188. 1999. Collocherides brychius, a new species (Copepoda: Siphonostomatoida: Asterocheridae) from a deep-water hydrothermal site in the northeastern Pacific Arthur G. Humes Boston University Marine Program, Marine Biological Laboratory, Woods Hole, Massachusetts 02543, U.S.A. — m Abstract. Collocherides brychius is described from a depth of 2253 at a hydrothermal site on the Juan de Fuca Ridge in the northeastern Pacific. The species is close to C astroboae Stock 1971, but may be distinguished from that species by its relatively small size (length of female 0.57 mm) and by the inner terminal seta on the caudal ramus of the female being approximately as long as the caudal ramus, instead of more than 3 times its length. Although its three congeners are associated with ophiuroid echinoderms in shallow water, the new species was recovered free in microfaunal samples in the deep sea. The copepod genus Collocherides Stock phrioids, cyclopoids, poecilostomatoids, 1971 (Siphonostomatoida: Asterocheridae), and harpacticoids. In this paper a new spe- contains three species. Collocherides astro- cies of the siphonostomatoid genus Col- boae Stock 1971, lives in the stomach of locherides is described from a deep-water the basket star Astroboa nuda (Lyman) at hydrothermal site in the northwestern Pa- Eilat in the Gulf ofAqaba and at the Dahlak cific. With the addition of this new species, Archipelago (depth about 50 cm) in the Red a new erebonasterid poecilostomatoid re- Sea (Stock 1971). It has also been recov- ported by Martinez Arbizu (1999), and a ered from the stomach of Astroboa alba- new aegisthid harpacticoid described by & trossi Doderlein in Indonesia (precise lo- Conroy-Dalton Huys (1999), the number cality unknown but probably Java Sea) of copepods known from deep-sea hydro- (Stock 1971). Large numbers of C. astro- thermal vents and cold seeps rises to 70. m boae were found on A. nuda in 18 at Nosy Be, northwestern Madagascar (Humes Materials and Methods 1973). Collocherides singularis Humes 1986, lives with Astroboa nuda (depth 5 m) The copepods were collected at a low at Poelau Gomumu, Moluccas, 01°50'00"S, temperature vent, where the highest reading 127°30'54"E (Humes 1986). Collocherides was 52° Celcius. The vent (Marker M) was bleptus Humes 1993, occurs intertidally on situated on new lava flows, and is believed the ophiuroid Macrophiothrix sp. at Nosy to be relatively young. (Vents at older ma- Be, northwestern Madagascar (Humes ture lavas are believed to support more het- 1993). erogeneous faunas, with more species, than & Sixty-seven species of copepods have young vents (Milligan Tunnicliffe been recorded in deep water from hydro- 1994).) The diving submersible Alvin used thermal vents and cold seeps in the world's its arm or "claw" to gather tube worms & oceans (Humes Segonzac 1998). These (Vestimentifera) and associated fauna and species consist for the most part of siphon- deposit them in a biobox mounted on a bas- ostomatoids, with fewer calanoids, miso- ket. The sample was preserved for later 182 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON sorting, during which copepods were recov- postgenital somites from anterior to poste- ered (Tsurumi, pers. comm.). rior 36 X 53, 31 X 40, and 23 X 36 fjim. The copepods, which had been preserved Genital double-somite and 3 postgenital so- in ethanol, were cleared and dissections mites with pairs of small posterolateral made in lactic acid, using the wooden slide spinelike processes or spinules (Fig. Id, e), method described by Humes & Gooding those on anal somite prominent. (1964). All figures were drawn with the aid Caudal ramus (Fig. le) moderately elon- of a camera lucida. The letter after the ex- gate, unomamented, length including ter- planation of each figure refers to the scale minal pointed process 44 iJim, length with- at which it was drawn. out process 35 ixm, width 16 iJim. Ratio of length (without process) to width 2.19:1, ra- Siphonostomatoida Thorell, 1859 tio including process 2.75:1. With 6 smooth Asterocheridae Giesbrecht, 1899 setae, 1 long and 5 short, longest seta 52 Collocherides Stock, 1971 |jLm long, much longer than other setae and Collocherides brychius, new species slightly swollen proximally. Long seta Figs. 1-3 slightly longer than ramus, ratio 1.18:1. — Body surface with few sensilla on uro- Type material. 9 ? $, 3 S S, in 2253 some, otherwise unomamented. m, Juan de Fuca Ridge, Segment Cleft, Egg sac (Fig. Id), seen on only 1 female, North Field, Vent Marker M, northeastern large, oval, 179 X 99 jxm. Pacific, 44°58.97'N, 130°12.35'W, 28 Aug Rostral area (Fig. If) triangular, not pro- 1990. Holotype 9 (USNM 243645), allo- truding. Antennule (Fig. 2a) 20-segmented, type S (USNM 243646), and 6 paratypes 187 |jLm long, with aesthetasc 60 jJim long (5 $ ?, 1 (?) (USNM 243647) deposited in on segment 18. Formula for armature: 1, 2, the National Museum of Natural History, 2, 2, 2, 2, 2, 2, 6, 2, 2, 2, 2, 2, 2, 2, 2, 2+ Smithsonian Institution, Washington, D.C. 1 aesthetasc, 2, and 9. All setae smooth. Remaining specimens (dissected) in the col- Antenna (Fig. 2b) 99 ixm long, including lection of the author. terminal spine 22 |xm. First segment (coxa) — Female. Body slender (Fig. la), not short and unarmed. Second segment (basis) flexed between prosome and urosome (Fig. with 1 minute seta (exopod). Third segment lb). Average length (not including setae on long and unarmed. Short fourth segment mm caudal rami) 0.57 (0.55-0.58 mm) and bearing 2 short setae and 1 long terminal average width 0.18 nam (0.17-0.19 nam), spine, its truncated tip with few extremely based on 9 specimens. Dorsoventral thick- minute setules. ness (at level of slight protuberance be- Oral cone (Fig. If) short, oval in ventral tween maxillipeds and first pair oflegs) 127 view, prominent in lateral view (Fig. lb). |xm. Epimera of metasomal somites round- Mandible (Fig. 2c) with slender 2-jointed ed (Fig. lb). Ratio of length to width of palp 60 |jLm long. Gnathobase (Fig. 2d) 50 prosome 2.03:1. Ratio of length ofprosome jjim long., with several blunt terminal teeth. to that of urosome 1.31:1. Maxillule (Fig. 2e) with 1 seta on slender Urosome with 5 somites. Somite bearing outer lobe and 4 setae on stout inner lobe. leg 5 (Fig. Ic) 47 X 83 jjim. Genital double- Maxilla (Fig. 2f) 2-segmented, first segment somite in dorsal view 81 ixm long and 81 unarmed, second segment bearing recurved |jLm wide at widest part. In lateral view (Fig. claw with truncated minutely spinulose tip. Id), this double-somite with dorsal and ven- Maxilliped (Fig. 2g) 5-segmented. First tral sides slightly rounded. Genital areas lo- segment with inner distal seta. Second seg- cated laterally in anterior half of double- ment elongate (52 jjim long) with 1 seta on somite (Fig. Ic, d). Each genital area with inner margin. Third and fourth segments small seta and minute spine (Fig. Id). Three short with 1 seta. Fifth segment elongate VOLUME 112, NUMBER 1 183 Fig. 1. Collocherides brychius, new species. Female, a, body, dorsal (scale A); b, body, lateral (A); c, urosome, dorsal (B); d, urosome, lateral (B); e, anal somite and caudal ramus, ventral (C); f, cephalosome, ventral (B). 184 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Collocherides brychius, new species. Female, a, antennule, ventral (scale C); b, antenna, dorsal (C); c, mandible, ventral (C); d, gnathobase ofmandible, flat view (D); e, maxillule, anterior (C); f, maxilla, posterior (C); g, maxilliped, posterior (C); h, leg 1 and intercoxal plate (E); i, leg 2 and intercoxal plate, posterior (E); j, leg 3 and intercoxal plate, anterior (E); k, leg 4 and intercoxal plate, anterior (E); 1, leg 5, ventral (C). VOLUME 112, NUMBER 1 185 Fig. 3. Collocherides brychius, new species. Male, a, body, dorsal (scale A); b, urosome, dorsal (B); c, antennule, ventral (C); d. leg 5, ventral (C); e, somites 1-3 of urosome, showing legs 5 and 6, ventral (E). 1 186 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON with 1 distal seta. Claw (26 jjiin long) with 37 X 15 |jLm including terminal process. In- truncated tip bearing minute spinules. ner terminal seta 143 jxm long, relatively Ventral area between maxillipeds and much longer than in female, 3.86 times lon- first pair of legs (Fig. If) slightly protuber- ger than ramus. ant (Fig. lb). Rostral area like that of female. Anten- Legs 1-4 (Fig. 2h-k) biramous with 3- nule (Fig. 3c) 18-segmented, geniculate. segmented rami. Formula for spines (Ro- Aesthetasc on segment 17 59 jxm long. Ar- man numerals) and setae (Arabic numerals) mature: 1, 2, 2, 2, 2, 2, 2, 2, 6, 2, 2, 2, 2, as follows: 2, 2, 1, 1 + aesthetasc, and 9. Antenna as in female. coxa0-0 basis 1-0 expI-1; l-l; 11,2,2 Oral cone, mandible, maxillule, maxilla, enpO-1; 0-2; 1.2,3 maxilliped, and legs 1-4 like those of fe- P2 coxa0-0 basis 1-0 eexxppII--11;; I-l; 111,1,4 male. enpO-1; 0-2; 1,2,3 Leg 5 (Fig. 3d) with second segment 20 P3 coxa0-0 basis 1-0 expI-1; I-l; 111,1,3 X 12 fxm, ratio 1.67:1, bearing 3 slender enpO-1; 0-2; 1,1,3 outer setae and 2 broad, hyaline, inner se- tae, 33 |xm, with blunt tips. P4 coxa0-0 basis 1-'-0 expI-1; I-l; 111,1,3 Leg 6 (Fig. 3e) usual posteroventral flap enp0- 0-2, 1,1,2 on genital somite bearing 2 setae. Inner coxal seta absent in all 4 legs. Out- Color of living specimens unknown. — er margins of exopod segments of all 4 legs Etymology. The specific name bry- with very small spinules. chius, from Greek brychios, meaning from Leg 5 (Fig. 21) 2-segmented. First seg- the depths of the sea, refers to the depth at ment with outer seta and produced medially which specimens of the new species was as broad triangular flap, pointed at tip, and found. — having inner and outer setules. Second seg- Remarks. The female of Collocherides ment 31 X 14 jjim, ratio 2.2:1, bearing 4 brychius may be differentiated from its setae and ornamented along inner margin three congeners as follows: with few setules. From C. astroboae Stock 1971, in which Leg 6 represented by seta and small spi- the long terminal seta on the caudal ramus niform process on genital area (Fig. Id) is approximately 3.6 times longer than the mm Color—of living specimens unknown. ramus, and the body length is 0.62 Male. Body (Fig. 3a) elongate, slender, (measurements based on an average of five and, as in female, not flexed. Length (not specimens from Astroboa nuda from the including setae on caudal rami) 0.50 mm Dahlak Archipelago (Zoologisch Museum (0.50-0.51 mm) and greatest width 0.15 Amsterdam, cat. no. 101.090). mm (0.15-0.16 mm), based on 3 speci- From C. singularis Humes, 1986, in mens. Greatest dorsoventral thickness at which the caudal ramus lacks a terminal level of small ventral protuberance 96 ixm. process, the long terminal seta on the cau- Ratio of length to width of prosome 1.98: dal ramus is approximately 1.8 times longer 1 Ratio of length ofprosome to that ofuro- than the ramus, the second segment of the . some 1.36:1. endopod of legs 1-3 has one inner seta, and Urosome (Fig. 3b) with 6 somites. So- the average body length is 0.64 mm. mite bearing leg 5 32 X 70 |jim. Genital From C bleptus Humes, 1993, in which somite 60 X 83 ixm, with rounded lateral the ratio of the innermost terminal seta on margins. Four postgenital somites from an- the caudal ramus to the length of the ramus terior to posterior 34 X 49, 32 X 39, 26 X itself is approximately 2.51:1, the genital 32, and 21 X 31 fxm. double-somite is 120 X 80 jjim, distinctly Caudal ramus similar to that of female. longer than wide, the average body length —— VOLUME NUMBER 112, 1 187 is 0.77 mm, and the ventral surfaces of the the Mid-Atlantic Ridge by Tyler et al. genital and postgenital somites have nu- (1995). However, no ophiuroids are yet merous small scalelike spines. known from Juan de Fuca vents, where the C C In brychius sexual dimorphism occurs brychius was found (Tunnicliffe, pers. in the length of the inner terminal seta on comm.). the caudal ramus, which in the female is 42 |jLm, but in the male is 143 ixm and less Acknowledgments swollen proximally than in the female. The four species of Collocherides may I thank Dr. Verena Tunnicliffe and Maia Tsurumi, Department of Biology, Univer- be further distinguished by the following sity of Victoria, British Columbia, for send- key. ing the specimens of C. brychius to me for study. I also thank Dirk Platvoet, Zoolo- Key to females of the genus Collocherides gisch Museum, Amsterdam, for arranging the loan of Stock's specimens of Astroboa 1. Genital double-somite distinctly longer than wide, ratio 1.4:1; body length 0.77 nuda from Dahlak Archipelago. mm (0.75-0.78 mm) C. bleptus Genital double-somite quadrate or near- Literature Cited ly so, ratio approximately 1:1; body length not exceeding 0.67 mm 2 Conroy-Dalton, S., & R. Huys. 1999. A new genus of Aegisthidae (Copepoda: Harpacticoida) from 2. Long inner terminal seta on caudal ra- hydrothermal vents on the Galapagos Rift. mus 3.6 times longer than ramus C Journal of Crustacean Biology (in press). astroboae Hessler, R. R., & W. M. Smithey, Jr. 1983. The distri- Longest seta on caudal ramus 1.18 times bution and community structure of megafauma (or less) longer than ramus 3 at the Galapagos Rift hydrothermal vents. In mm 3. Body length 0.57 (0.55-0.58 mm); Hydrothermal processes and seafloor spreading legs 1-3 with second segment of endo- centers. R A. Rona, K. Bostrom, L. Laubier, & pod having 2 inner setae C brychius K. L. Smith, Jr., eds., NATO conference series Body length 0.64 mm (0.63-0.65 mm); 4:735-770. Plenum Press, New York. legs 1-3 with second segment of endo- Humes, A. G. 1973. Cyclopoid copepods associated pod with 1 inner seta C. singularis with—the ophiuroid Astroboa nuda in Madagas- car. Beaufortia 21:25-35. The ecological and host relationships . 1986. Two new species of Copepoda associ- ated with the basket slarAstroboa nuda (Ophiu- within the genus Collocherides are poorly — C roidea) in the Moluccas. Zoologica Scripta 15: known. Collocherides astroboae, singu- 323-332. laris, and C. bleptus are associated with . 1993. Collocherides bleptus n. sp. (Copepoda: shallow-water ophiuroids, while C. bry- Siphonostomatoida) associat—ed with an intertid- chius lives in deep—water hydrothermal al ophiuroid in Madagascar. Bijdragen tot de Dierkunde 63:121-127. sites and was recovered free in meiofaunal & R. U. Gooding. 1964. A method for study- samples. However, much more information , — ing the external anatomy of copepods. Crus- is needed to understand these relationships. taceana 6:238-240. The possibility that there may have been , & M. Segonzac. 1998. Copepoda from deep- ophiuroids at the deep-sea vent site cannot sea hydrothermal sites and cold seeps: descrip- tion ofa new species ofAphotopontius from the be excluded. Ophiuroids, known from abys- — East Pacific Rise and general distribution. Ca- sal depths (e.g., Lauermann and Kaufman hiers de Biologic Marine 39:51-62. 1998), have been reported from deep-sea Lauermann, L. M. L., & R. S. Kaufman. 1998. Deep- hydrothermal sites (Hessler and Smithey sea epibenthic echinoderms and a temporally 1983; Grassle 1986; Tunnicliffe 1991, varying food supply: results f—rom a one year & time series in the N. E. Pacific. Deep-Sea Re- 1998; Segonzac 1992; Sibuet Olu 1998). search 1145:817-843. A new genus and species of ophiuroid, Martinez Arbizu, P. 1999. New Erebonasteridae (Co- Ophioctenella acies, was described from pepoda) from Vilkitzky Strait in the Arctic and U^ — PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON II from a Pacific hydrothermal vent site (northern Stock, J. H. 1971. Collocherides astroboae n. gen., n. — Fiji Basin). Journal ofCrustacean Biology 19: sp., a siphonostome cyclopoid—copepod living 93-105. in the stomach of basket stars. Bijdragen tot Milligan, B. N., & V. Tunnicliffe. 1994. 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Biogeography, biodiver- from hydrothermal mounds along the Mid-At- — sity and fluid dependence ofdeep-sea cold-seep lantic Ridge. Journal ofthe Marine Biological communities at active and passive margins. Association of the United Eangdom 75:977- Deep-Sea Research II 45:517-567. 986.