Invertebrate Zoology, 2007. 4(1): 1-14 ? INVERTEBRATE ZOOLOGY, 2007 Redescription of Tegastes nanus Sars, 1904 (Copepoda: Harpacticoida: Tegastidae) from Spitsbergen in the Arctic Ocean Frank D. Ferrari\ Pavel V. Rybnikov^, Hans-U. Dahms^ ' IZ/MSC; MRC-534, National Museum of Natural History, Smithsonian Institution, 4210 Silver Hill Rd., Suitland, MD 20746, U.S.A. ^ P.P. Shirshov Institute of Oceanology RAS, Moscow, Russia ^ National Taiwan Ocean University, Institute of Marine Biology, 2 Pei-Ning Road, KEEIUNG, Taiwan, 202 ABSTRACT: Tegastes nanus Sars, 1904 (Copepoda, Harpacticoida, Tegastidae) is reported from Ny Alesund, Spitsbergen (Svalbard) in the Arctic Ocean. The morphology of adult females and males is redescribed. Specimens from Ny Alesund are similar to Norwegian specimens collected by Sars from Eggesbones and Alesund, as well as to published descriptions of specimens of this species by Chislenko (1967) from the White Sea. Specimens from Ny Alesund, Eggesbones and Alesund differ from published description of the species collected in the Gulf of Maine (McAlice & Coffin, 1990) by the shape of the extension of the genital somite plus 6th thoracic somite of females, the spermatophore reservoir of males, and several limbs. This is the first detailed description of the mouthparts of T. nanus and the first report of a harpacticoid copepod from Spitsbergen. KEY WORDS: systematics, morphology, phytal, Harpacticoida, Copepoda, Crustacea. riepeonMcaHiie Tegastes nanus Sars, 1904 (Copepoda: IHarpacticoida: Tegastidae) c apxunejiara LUnMM6epreH B CeeepHOM JleAOBMTOM OKeane O.fl. ct>eppapM\ n.B. PBIGHMKOB^, FaHC-y. flaMC^ ' IZ/MSC; MRC-534, National Museum of Natural History, Smithsonian Institution, 4210 Silver Hill Rd., Suitland, MD 20746, U.S.A. ^ Hncmumym OKeanonozuu UM. TI.II. Uluptuoea PAH, MocKea, Poccun ^ National Taiwan Ocean University, Institute of Marine Biology, 2 Pei-Ning Road, KEEIUNG, Taiwan, 202 PE3IOME: Tegastes nanus Sars, 1904 (Copepoda, Harpacticoida, Tegastidae) oxMeneH B CcBepHOM JleaoBHTOM oKcane y apxHoenara UlnHuGepreH (paiioH Ny Alesund). HepeonHcana Mop(|)oiiorHH Bspocntix caMOK H caMuoB 3Toro BH^ia. 3K3eMnjiHpti H3 lUnHuGepreHa CXO^KH C 3K3eMniiapaMH, coGpanntiMH CapcoM y 6eperoB HopBernx (Eggesbones H Alesund), H cooTBeTCTByioT onncaHHK) axoro BH^ia H3 Benoro Mopa (^HcneHKo, 1967). 3K3eMnjiHpi.i, coGpaHHBie y 6eperoB lUnHuGepreHa H HopBernn, oxuHHaioTCH OT onHcanHx /laHHoro BH/ja H3 3aiiHBa Man (McAlice & Coffin, 1990) (|>opMOH Bbipocxa reHHxaiiBHoro coMHxa caMKH, cpocmeroca c 6-M xopaKanBHtiM COMHXOM, (JjopMofi BLipocxa reHHxantHoro coMHxa caivma, necymero 2 F.D. Ferrari, P.V. Rybnikov, H.-U. Dahms cnepMaTo4)op, H neKOTopbiMH KoneHHocTHMH. B ^laHHoii pa6oTe BnepBtie jinsi T. nanus aaexcH onHcaHHe Mop(|>oiiorHH poTOBtix KOHeHHocxeH, H coo6maeTCH o ero naxo/iKe y apxHnenara IIInHLi,6epreHa. KJIIOHEBLIE CJIOBA: cHcxeMaxHKa, Mop(J)OJiorHH, (jjHxajii., Harpacticoida, Copepoda, Crustacea. Introduction The harpacticoid copepod genus Tegastes Norman, 1903 is one of six genera in Tegas- tidae. There are 38 nominal species of Tegastes, all distinctive in their laterally compression and ventrally protruding cephalosome. Species of Tegastes are encountered infrequently and rel- atively little is known about their biology. Dur- ing a survey of the marine fauna of Spitsbergen, an island archipelago (75?-85?N; 10?-30?E) in the Arctic Ocean, three specimens of Tegastes nanus Sars, 1904 were washed from red algae collected in the harbor of Ny Alesund. Here we redescribe females and males of T. nanus from Ny Alesund, Spitsbergen, and compare these to Sars's Norwegian specimens from Eggesbones and Alesund, Norway. Materials and methods Samples of the red alga Ceramium sp. from the rocky shore of Ny Alesund, Spitsbergen (78?33'N, 12?E) were taken during July 1998. The algal thalli were washed and the wash water passed through a 42|am sieve. Specimens re- tained in the sieve were then fixed in 4% form- aldehyde solution and subsequently transferred into glycerin for slide preparation. Specimens of Tegastes nanus were observed with a Leitz Diaplan microscope using phase contrast at 1250x magnification for preliminary observa- tions and for the line drawings. Systematic ac- counts were compared with Bodin (1997) and amended. One female (# 31032) and one male (#31031) are deposited in the collection of the Senckenberg Museum, Frankfurt/a.M. Germa- ny. The specimens studied by Sars' cannot be located because he did not designate type spec- imens of this species or label a vial as new species. However, vial F20359 with one female and two males from Eggesbones, Norway (62?19'N, 05?39'E), and vial F20366 with one female and one male from Alesund (62?19'N, 5?30'E) belonging to Sars and now in the Crus- tacea Collection of the Zoological Museum at the University of Oslo were kindly sent by Ase Wilhelmsen, Principal Engineer of the Crusta- cean Collection to one of us (FDF) for study. These specimens were cleared through 50% lactic acid/ 50% de-ionized freshwater to 100% lactic acid, stained by adding a solution of chlorazol black E dissolved in 70% ethanol/ 30% de-ionized freshwater, and examined with bright-field and with differential interference optics. SpecimensofT. nanMS studied byChislen- ko could not be located. Specimens of T. nanus are curved ventrally along the anterior-posterior axis so that they are difficult to measure in either dorsal or lateral view. Total length was measured laterally, in sections along the anterior-posterior curve, and includes caudal rami and rostrum. In general, descriptive terms follow (Ferra- ri, 1995). Interpretations of homologies of the first maxilla follow Boxshall (1985), of the second maxilla follow Ferrari & Ivanenko (2005), of the maxilliped follow Ferrari & Dahms (1998); the protopod of these three limbs has a coxa with one setiferous endite. Descriptive part Tegastes nanus Sars, 1904 Female. Colour pale yellowish brown, dark- ening to chestnut along sutures between somites. Total body length 0.30 mm. Body laterally compressed; dorsal margin strongly curved an- teriorly and posteriorly (Fig. 1). Cephalosome Redescription of Tegastes nanus Sars, 1904 Fig. 1. Tegastes nanus Sars, 1904. Female. A ? right lateral habitus; B ? left lateral urosome without leg 5. Scale bars: A ? 100 \\.ra; B ? 50 (im. PHC. 1. Tegastes nanus Sars, 1904. CaMKa. A ? BHeniHHH BHA cnpaea; B ? ypocoMa 6e3 Horn 5-H HOFH, BHA cueBa. MacmTa6Hbie 0Tpe3KH: A ? 100 MKM; B - 50 MKM. F.D. Ferrari, P.V. Rybnikov, H.-U. Dahms Fig. 2. Tegastes nanus Sars, 1904. Female. Al ? first antenna; A2 ? second antenna; Md ? mandible; Mxl ? first maxilla (praecoxal endite separate); Mx2 ? second maxilla. Scale bar 50 ^m. PHC. 2. Tegastes nanus Sars, 1904. CaMKa. Al ? aHTeHHa nepBaa; A2 ? aHTenna BTopaa. Md ? MaHflH6yjia. Mxl ? MaKCHHna nepeaa (3HaHT npeKOKCti OTflenen); Mx2 ? MaKCHJina BTopaa. MacniTaSHtiii oxpeaoK 50 MKM. Redescription of Tegastes nanus Sars, 1904 Fig. 3. Tegastes nanus Sars, 1904. Female. PI ? swimming leg 1; P2 ? swimming leg 2. Scale bar 50 \aa. PHC. 3. Tegastes nanus Sars, 1904. CaMKa. PI ? HuaBaTeJibHaa Hora 1; P2 ? njiaBaxeJibHaa Hora 2. MacmxaSHbiH 0Tpe30K 50 MKM. F.D. Ferrari, P.V. Rybnikov, H.-U. Dahms strongly produced ventrally on both sides to a narrowly rounded tip. Anterodorsal face of ceph- alosome covered by dense tiny chitin platelets. Cephalosome [including 1st and 2nd thoracic somites] separated from metasome by a dis- tinctly sclerotized strip (Fig. 1 A). Urosome (Fig. IB): rugose; genital somite fused to 6th thoracic somite and anterior abdominal somite; these segments quadrate posteroventrally and extend- ing as well-pronounced comers (Fig. IB); both posterior comers rounded. Last three abdomi- nal somites small and articulating. Caudal rami small, about twice as long as wide; each ramus one thick dorsal seta on a lobe, plus one thin seta medial and one thin seta lateral to the thick seta, and 4 terminal setae along a curved, lateral edge (Fig. IB). First antenna (Fig. 2 Al): 8-segmented, aes- thetascs on the 4th segment (exceeding slightly the tip of distal segment) and on the 8th segment (same length but more slender). Setal armature 1, 7, 7,2+1 aesthetasc, 2,3,3,10+1 aesthetasc. Second antenna (Fig. 2 A2). Coxa short, engaging an irregular pedestal proximally; ba- sis without ornamentation. Exopod small, 2- segmented; proximal segment with 1 medial seta, distal with 1 medial and 2 distal setae. Proximal endopodal segment with 1 medial seta and anterior scale-like attenuation; distal seg- ment with longitudinal row of denticles laterally and with 2 medial, 1 lateral and 4 terminal setae. Mandible (Fig. 2 Mn). Gnathobase with 2 groups of tooth-like attenuations, 3 teeth in proximal group, 3 teeth distally, plus a proximal seta. Basis with 2 medial setae. Exopod a simple knob-like segment with 1 terminal seta. Endo- pod 2 segmented; proximal segment with 2 medial setae; distal with 3 terminal setae. First maxilla (Fig. 2 Mxl). Praecoxal endite with 7 short, thick setae and 1 longer, thinner seta distally. Coxal exite with 1 seta. Basis fused distally to endopod; proximal basal endite with 1 seta, distal basal endite unattenuated, with 2 setae. Endopod apparently 1-segmented, elon- gate, with 4 terminal setae. Second maxilla (Fig. 2 Mx2). Syncoxa with transverse row of spinules proximolaterally; praecoxal endite with 3 setae; coxal endite with 2 setae. Basis elongate with transverse row of spinules proximolaterally; proximal basal en- dite with 2 setae; distal basal endite with 3 setae, middle seta curved and thick. Ramus a very small unarticulated segment with 3 setae. Maxilliped (Fig. 5 Mxp). A subchela; syn- coxa with 1 mediodistal seta with setules, prox- imomedial and proximolateral set of denticles plus another set of lateral denticles at mid- length. Basis with 1 mediodistal seta directed proximally and reaching about a 4"" the length of palmar edge; palmar edge with plate-like seta medial row of larger denticles and row of slen- der ones anteriorly. Proximal endopodal seg- ment with 1 long seta; distal endopodal segment attenuate, appressed to medial edge of basis, with 2 slender setae towards its base. Swimmingleg 1 (Fig. 3 PI). Coxa small with lateral longitudinal row of denticles proximally. Basis elongate with denticles along medial and lateral margin; distally with 1 medial seta and 1 lateral seta. Rami 1 -segmented. Endopod wider and slightly shorter than exopod, with denticles along lateral edge and with 1 proximomedial seta, 1 distomedial seta and 4 terminal setae. Exopod with medial denticles proximally and with 1 proximal and 1 distal seta laterally, and 3 terminal setae, middle thickest and longest. Swimming leg 2 (Fig. 3 P2). Coxa with a set of denticles on pronounced distolateral comer. Basis with denticles along medial margin and 1 lateral seta. Rami 3-segmented. Proximal seg- ment of exopod with 1 lateral seta and 1 medial seta, and lateral denticles; middle segment with 1 lateral seta and 1 medial seta, and lateral denticles; distal segment with 1 lateral, 3 termi- nal and 2 medial setae, and lateral denticles. Proximal segment of endopod with 1 medial seta and lateral denticles; middle segment with 2 medial setae and lateral denticles; distal seg- ment with 1 lateral, 2 terminal and 2 medial setae, and lateral denticles. Swimming leg 3 (Fig. 4 P3). Coxa with denticles on pronounced distolateral corner. Basis with denticles on distolateral corner and 1 lateral seta. Rami 3-segmented. Proximal segment of exopod with 1 lateral seta and 1 medial seta, and lateral denticles; middle seg- Redescription of Tegastes nanus Sars, 1904 Fig. 4. Tegastes nanus Sars, 1904. Female. P3 ? swimming leg 3; P4 ? swimming leg 4. Scale bar 50 \aa. PHC. 4. Tegastes nanus Sars, 1904. CaMKa. P3 ? HuaBaTeJibHaa Hora 3; P4 ? njiaBaxeJibHaa Hora 4. MacniTaSHbifl 0Tpe30K 50 MKM. ment with 1 lateral seta and 1 medial seta, and lateral denticles; distal with 2 lateral, 2 termi- nal and 2 medial setae, and lateral denticles. Proximal segment of endopod with 1 medial seta, and lateral denticles; middle segment with 2 medial setae, and lateral denticles; distal segment with 2 medial, 2 terminal and 1 lateral setae. Swimming leg 4 (Fig. 4 P4). Coxa without denticles. Basis with medial denticles, and with 1 lateral seta. Rami 3-segmented. Proxi- mal segment of exopod with 1 lateral seta and 1 medial seta, and lateral denticles; middle segment with 1 lateral seta and 1 medial seta, and lateral denticles; distal segment with later- al denticles and 2 lateral, 2 terminal and 3 medial setae; middle medial seta long and thick. Proximal segment of endopod with 1 medial seta and lateral denticles; middle seg- ment with 2 medial setae and lateral denticles; F.D. Ferrari, P.V. Rybnikov, H.-U. Dahms Fig. 5. Tegastes nanus Sars, 1904. Female. P5 ? leg 5; Mxp ? maxilliped. Scale bar 50 |xm. PHC. 5. Tegastes nanus Sars, 1904. CaMKa. P5 ? Hora 5; Mxp ? MaKCHJiJiHnefl. MacmxaSHbiH 0Tpe30K 50 MKM. distal segment with 2 medial, 2 terminal and 1 lateral setae. Leg 5 (Fig. 5). Baseoendopod well-devel- oped, with denticles on proximomedial comer and 3 medial setae without setules, 1 apical pinnate seta, 1 small, lateral seta, and 1 proxi- molateral seta; surface relief an irregular pattern of thickened cuticle. Exopod2-segmented, elon- gate, 2/3 length of endopod; proximal segment with 1 lateral seta, distal segment with 3 lateral setae and 2 terminal setae. Leg 6 not found. Male. Length 0.29 mm. Similar to female in size, color, shape and surface relief of somites (Fig. 6A). Comers of cephalosome somewhat less narrowly produced ventrally. Sixth and 7th thoracic somites (Fig. 6B) protrading more ven- trally as spermatophore reservoir (Sars 1904: 68) with a slight bump between 2 pronounced extensions; spermatophore large, ovoid. Each abdominal somite compressed within adjacent anterior somite. Terminal setae of caudal rami shorter and broader than female. First antenna (Fig. 7 Al). A 9-segmented haplocer with geniculations between 4th and 5th, and 8th and 9th segments. Seta formula: 1, 6 (2+2+2), 3+1 aesthetasc, 11+2 aesthetascs, 1, 1, 1, 1, 8+1 aesthetasc. Redescription of Tegastes nanus Sars, 1904 Fig. 6. Tegastes nanus Sars, 1904. Male. A ? left lateral habitus; B ? left lateral urosome. Scale bars: A ? 100 |.im; B ? 50 nm. PHC. 6. Tegastes nanus Sars, 1904. Caivieii. A ? BHeniHHH BHfl cjiesa; B ? ypocoMa, BHA cueBa. MacniTaSHbie 0Tpe3KH: A ? 100 MKM; B ? 50 MKM. Second antenna, mandible, maxilla 1, max- illa 2, maxilliped similar to female. Swimming legs \-A similar to female with the following exceptions: distal endopodal seg- ment of P2 and P4 (Figs. 8 P2, 9 P4) with 3 medial setae; middle medial seta on distal seg- ment of P4 long and thick (Fig. 9 P4). P5 (Fig. 6B). Exopod long, 2-segmented; proximal segment with 1 lateral seta, distal segment with 2 lateral and 2 terminal setae. Discussion Tegastid harpacticoids have been collected with few exceptions from surfaces of benthic invertebrates and macroalgae (Ivanenko & De- fay, 2005). Tegastes nanus Sars, 1904 is a temperate to boreal harpacticoid copepod orig- inally described from western Norway (Sars, 1904). Records from the eastern Atlantic Ocean include the coasts of eastern Ireland (Roe, 195 8) 10 F.D. Ferrari, P.V. Rybnikov, H.-U. Dahms Fig. 7. Tegastes nanus Sars, 1904. Male. Al ? first antenna. Scale bar 50 |.im. PHC. 7. Tegastes nanus Sars, 1904. Caivieit. Al ? aHTeHHa nepsaa. MacniTaSHtiH 0Tpe30K 50 MKM. and western Ireland (Farran, 1913), and the western Baltic Sea (Kunz, 1935). The species also has been recorded from the White Sea of the Arctic Ocean (Chislenko, 1967), the western North Atlantic Ocean from James Bay, an ex- tension of Hudson Bay (Willey, 1923), from the Gulf of St. Lawrence (Tremblay, 1944), and from the Damariscotta River, Gulf of Maine (McAlice & Coffin, 1990). Reports of this spe- cies from the subtropics, including the Suez Canal (records summarized by Lang, 1948) are considered doubtful, following the opinion of Todaro et al. (1996). The phytal substrate known for T. nanus includes two genera of red algae, Delesseria (see Kunz, 1935) and Ptilota (see McAlice & Coffin, 1990), and two genera of brown algae, Laminaria (see Chislenko, 1967, McAlice & Coffin, 1990) and Fucus (see Chislenko, 1967). Sars' (1904) original description was very brief; the figures include only the lateral view of the female P5, the male urosome and maxil- liped. The five specimens we received in two lots from the University of Oslo were very Redescription of Tegastes nanus Sars, 1904 11 Fig. 8. Tegastes nanus Sars, 1904. Male. PI ? swimming leg 1; P2 ? swimming leg 2. Scale bar 50 \aa. PHC. 8. Tegastes nanus Sars, 1904. Caivieii. PI ? Hora 1; P2 ? Hora 2; Mxp ? MaKCHJinnnea. MacmTaSHbiii 0Tpe30K 50 MKM. brittle, presumably from long exposure to the dehydrating effects of the ethanol preservative. The 0.45 mm female (remaining male is 0.38 mm) from Alesund (F 20366) and the 0.42 mm male from Eggesbones (remaining male is 0.40 mm and female is 0.49 mm) (F 20357) were dissected and are morphologically similar to specimens from Ny Alesund although both sex- es are larger. Since Sars' (1904) original description of the species, r. nanus has been redescribed twice. More illustrations (lateral habitus, Al, P5 of both sexes, PI and second maxilla) were pro- vided by Chislenko (1967) from White Sea specimens. The most recent redescription by McAlice and Coffm (1990) included a descrip- tion and illustration of a male from a phytal habitat of the Gulf of Maine. This description is much more complete, but it lacks necessary detail for a complete comparative analysis. Unfortunately, the material upon which these authors based their descriptions is not available. Females from Alesund and Ny Alesund agree with Sars' (1904) and Chislenko's (1967) brief 12 F.D. Ferrari, P.V. Rybnikov, H.-U. Dahms Fig. 9. Tegastes nanus Sars, 1904. Male. P3 ? swimming leg 3; P4 ? swimming leg 4. Scale bar 50 \aa. PHC. 9. Tegastes nanus Sars, 1904. Caivieii. P3 ? HuaBaTeJibHaa Hora 3; P4 ? njiaBaxeJibHaa Hora 4. MacmxaSHbiH 0Tpe30K 50 MKM. descriptions. Chislenko observed that females of T. nanus possessed small bristles on the lateral surface of the baseoendopod of P5; Sars' (1904) drawings show the same, although he did not mention this in the text. The female and male from Ny Alesund ex- press sexual dimorphism in the shape of the cephalosome, thoracic somites 6 and 7, and the anterior abdominal somite, as well as antenna 1, leg 5, and swimming legs 2 and 4. The penulti- mate segment of the 8-segmented female anten- na 1 corresponds to the antepenultimate and penultimate segment of the 9-segmented male antenna 1; an arthrodial membrane does not form on the female's limb. The female from Alesund and male from Eggesbones expressed a dimorphism in swimming leg 3 similar to those of swimming legs 2 and 4 in specimens from Ny Alesund. Illustrations and setal formula of the male described by McAlice and Coffin (1990) differ from our specimens. The antenna 1 of the male from the Gulf of Maine has only one aesthetasc on the S'*" segment. The male from Ny Alesund and the male from Eggesbones have 1, 2 and 1 aesthetascs on S"^"*, 4"" and 8"" segments respec- tively. McAlice and Coffin (1990) note that in contrast to Chislenko's figures, the 5* segment Redescription of Tegastes nanus Sars, 1904 13 of Al of their specimens are much longer. The specimens studied here have a 5"" segment length similar to the ones given by Chislenko. Males from Ny Alesund and from Eggesbones have a seta on the P' segment of the exopod of antenna 2 (not present on the male from the Gulf of Maine) and 4 terminal setae on the distal en- dopodal segment of antenna 2 (3 setae on the male from the Gulf of Maine). The Gulf of Maine male also expresses several differences in swimming legs compared to the males from Ny Alesund and from Eggesbones: there is no medial seta on the proximal exopodal segment of swimming legs l-A (present in the males from Ny Alesund and from Eggesbones); there are 2 medial setae on the distal endopodal segment of swimming legs 2 and 4 (3 males from Ny Ale- sund and from Eggesbones). It should be noted the differences in medial setae on the distal endopodal segment of swimming legs 2 and 4 also result in gender dimorphism in specimens from Ny Alesund; it is possible that this state is simply polymorphic in this species. The other differences are more substantial. Equally im- portant are differences in the posteroventral extension of the genital somite fused to the 6th thoracic somite and anterior abdominal somite of the female with less pronounced tips in the Gulf of Maine specimens, and spermatophore reservoir of the male without a bump between the tips of the Gulf of Maine specimens. Spec- imens of r. nanus from Ny Alesund are conspe- cific with those from Alesund and Eggesbones, as described by Sars (1904) and the White Sea. Tegastes nanus from the Gulf of Maine (McAl- ice & Coffin 1990) probably is not conspecific with those from Ny Alesund, Alesund, Egges- bones and the White Sea. Acknowledgements We thank Mr. J. Diirbaum for collecting the tegastid specimens during a summer campaign at Ny Alesund on Spitsbergen in July 1998. Late P.R. acknowledged a grant from the Deutscher Akademischer Austauschdienst (DAAD - A/ 99/09723) during his stay at H.U.D's laboratory at the AG Zoosystematik und Morphologic. 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