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A New Copepod Species From California, Usa: Hesperodiaptomus Californiensis (Crustacea: Copepoda: Calanoida: Diaptomidae) PDF

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PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 109(1):103-1U. 1996 A new copepod species from California, U.S.A.: Hesperodiaptomus californiensis (Crustacea: Copepoda: Calanoida: Diaptomidae) Megan Scanlin and Janet W. Reid (MS) College Station Box 3505, The College of William and Mary, Williamsburg, Virginia 23186-3505, U.S.A.; (JWR*) Research Associate, Department ofInvertebrate Zoology/MRC-163, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, U.S.A. (*Corresponding author) — Abstract. Hesperodiaptomus californiensis, new species (Copepoda: Cal- anoida: Diaptomidae) was collected from vernal pools in Lassen County, north- em California, U.S.A. It differs from its three most morphologically similar congeners, Hesperodiaptomus schefferi, Hesperodiaptomus victoriaensis and Hesperodiaptomus kiseri in having both leg 5 endopods of the male relatively long, the left lateral protrusion on the genital double somite of the female directed dorsally, and in other details. Co-occurring diaptomids were Hesper- odiaptomus novemdecimus and Leptodiaptomus tyrrelli. A new species ofdiaptomid copepod was females cannot be reliably assigned to a collected in Lassen County, California, as particular species with present knowledge. part of a study of vernal pools (described These possible species complexes are H. by King et al. 1996). The new copepod oc- breweri (Wilson, 1958a), H. eiseni (Lillje- & curred in three pools and at the time ofcol- borg in Guerne Richard, 1889), and H. lection was abundant in every pool. arcticus; as well as H. kiseri (Kincaid, Wilson (1959) listed 16 species of Hes- 1953) and H. victoriaensis (Reed, 1958). perodiaptomus in North America; since The high degree of morphological similar- then no new species has been recognized ity between H. kiseri and H. victoriaensis from this continent. However, the taxon creates some difficulty in establishing the Hesperodiaptomus arcticus (Marsh, 1920) species from California as a new taxon, be- s.l. may be composed of several cryptic cause it shares several characters with both & species. Boileau Hebert (1988) and Bo- these species. ileau (1991) found genetic differences be- Hesperodiaptomus species and other tween different North American popula- diaptomid copepods are characterized clas- tions of H. arcticus, which were accompa- sically by the features of the fifth legs of nied by subtle morphological differences. the males and females, the right antennule On the other hand, Stepanova (in Borutskii of the male and setation of the left anten- et al. 1991) rejected efforts by Streletskaya nule ofthe male and both antennules ofthe H (1983, 1986) to split Siberian arcticus female, and features of the somites, partic- into three taxa: H. koolensis Streletskaya, ularly the shape of the thoracic "wings" 1983, H. judayi Streletskaya, 1986 and H. (expansions of the sixth thoracic somite). kurenkovi Streletskaya, 1986, because of Pending eventual revision ofthe genus Hes- the supposedly minordegree ofmorpholog- perodiaptomus and the possible discovery ical differences. of additional morphological and genetic Some other members of Hesperodiapto- characteristics, we describe the new species mus are so similar morphologically that the using these traditional discriminators. 104 PROCEEDINGS OFTHE BIOLOGICAL SOCIETY OF WASHINGTON The copepod specimens were originally 1.5 km south of intersection with Route fixed in buffered formalin and transferred A21), 2 m tow of wet meadow lake bottom to 70% ethanol for long term storage. Field with plankton net, sample 220, USNM sampling methods were described by King 264062. 10 66 9 9, pool #43 (Long Lake), et al. (1996). Descriptions were made from sample 219, LACM 95-50.2. All collected whole specimens in glycerin and/or lactic 23 April 1992 by J. L. King, D. Gluesen- acid; dissected specimens were mounted ei- kamp and J. Tritt. Undissected specimens ther in CMC-10 or in commercial polyvinyl preserved in 70% ethanol. lactophenol with a little Chlorazol Black E Co-—occurring species (determined by added. All measurements were made from JWR). Pool #41: Hesperodiaptomus nov- specimens in glycerin. Specimens were de- emdecimus (Wilson, 1953), Leptodiaptomus posited in the collections of the United tyrrelli (Poppe, 1888), Acanthocyclops car- States National Museum ofNatural History, olinianus (Yeatman, 1944), Diacyclops Smithsonian Institution (USNM) and the crassicaudis var. brachycercus (Kiefer, Natural History Museum of Los Angeles 1927), Microcyclops rubellus (Lilljeborg, County, California (LACM). 1901), Bryocamptus washingtonensis Wil- son, 1958b; pool #42: H. novemdecimus, L. Family Diaptomidae G. O. Sars tyrrelli, A. carolinianus, Acanthocyclops Genus Hesperodiaptomus Light, 1938 vernalis (Fischer, 1853) s.l., Diacyclops na- Hesperodiaptomus californiensis, vus (Herrick, 1882); pool #43 (Long Lake): new species H. novemdecimus, L. tyrrelli, A. carolini- Figs. 1-3 anus, B.—washingtonensis. — Male. Length (mm) of holotype 2.21; Type material. Holotype S, USNM of ten paratypes, mean = 2.14, median = 264057; allotype 9, USNM 264058; para- 2.18, range = 1.93-2.32. Pediger 5 (sixth types: 3 SS 3 9 9, each dissected on slide thoracic somite) (Fig. la), lateral "wings" in PVL or in CMC-10, and 464 SS 9 9 + symmetrical, each with two sensilla. Uro- copepodids, USNM 264059; 463 c?c? 9 9 some (Fig. la) symmetrical except urosom- + copepodids, LACM 95-50.1; all from ites 1 and 4 slightly produced laterally and California, Lassen County, pool #41, posteriorly. Urosomites 2-4 and caudal 40°37'N, 121°03'W, immediately adjacent rami with pore-canals, middle dorsal pore- to west side of Route 44, 1.4 km north of canal of urosomite 4 displaced slightly to Bogart Safety Rest Area/Ranger Station, 2 left of dorsal midline. Inner sides of caudal m tow of wet meadow edge with plankton rami with long hairs, rest of ramal surfaces net, sample 215. — with short sparse hairs. Rostral points (Fig. Additional, non-type material. 372 SS lb) short, acute. 9 9 + copepodids, California, Lassen Antennules (Fig. la, c, d) reaching mid- m County, pool #42, 40°32'N, 12rO'W, 30 dle of urosomite 2. Right antennule with west of Route 44, just south of railroad two setae on segment6; segment 8 with one track crossing, 2 m tow ofwet meadow lake seta and one spine; segment 10 with one USNM edge with plankton net, sample 218, seta and one stout spine reaching middle of 264060. 894 SS 9 9, pool #42 bottom, segment 12; segment 11 with one seta and sample 217, LACM 95-50.3. 41 SS 9 9, one stout spine, reaching distal end of seg- USNM pool #41, sample 214, 264061. ment 13; segment 13 with one stout spine 400+ 66 9 9, pool #42 surface, sample reaching past middle of segment 14; seg- LACM 216, 95-50.4. 2S 66 9 9, Califor- ment 14 without spine; segment 15 with nia, Lassen County, pool #43 (Long Lake), small acute process pointing distally; seg- m 40°31'N, 120°59'W, 100 west of Route ment 16 with two setae. Segment 23 (Fig. 44, about 0.36 km south ofpool #41 (about Id), process at distal end straight, reaching VOLUME NUMBER 109, 1 105 50 jam Fig. 1. Hesperodiaptomus californiensis, new species, male: a. Habitus, dorsal; b. Rostral points (indicated by arrows);c. Rightantennule, segments6-17; d. Rightantennule, segments23-25; e. Leg2endopod, segments 1 and 2, posterior. 106 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Hesperodiaptomus californiensis, new species, male: a. Leg 5, posterior; b. Left leg 5 exopod, posterior; c, Left leg 5 exopod and endopod, anterior; d. Left leg 5 endopod, posterior; e. Right leg 5 endopod, anterior. Both scales = 50 jjim. end of last segment of antennule. Setation cess. Inner margin of exopod 2 with two of left antennule as in female. pads, proximal pad haired, proximal hairs Leg 2 endopod segment 2 (Fig. le) with- of this pad thinner and shorter than distal out defined Schmeil's organ, but with small hairs; distal pad covered with rows of teeth transverse ridge, more or less developed on and few tiny hairs near tip of exopod. Pos- different specimens. terior surface ofexopod 2 with longitudinal Left leg 5 (Fig. 2a-d): Coxa, small pro- groove. Endopod reaching midlength ofex- cess tipped with sensillum near outer pos- opod 2, ofone segment, narrowing at distal terodistal margin. Basis with lateral seta. Vs. Tip of endopod narrowing in acute pro- Exopod segment 1 longer than segment 2, cess; also with five small subterminal with haired pad on inner margin, hairs thick spines and group of subterminal hairs. and as long as about halfwidth ofendopod. Right leg (Fig. 2a, e): Coxa with small Exopod 2, tip forming stout, blunt, smooth posterodistal process ending in sensillum. lateral process; also with serrate digital pro- Basis with lateral seta, shallow longitudinal VOLUME NUMBER 109, 1 107 Fig. 3. Hesperodiaptomuscalifomiensis, new species, female: a. Habitus, dorsal; b. Rostralpoints (indicated by arrows); c, Antennule, segments 1-9; d, Leg 5, anterior; e. Tip ofleg 5 endopod, posterior. groove on posterior surface, and rounded, margin; lateral spine at distal % of exopod unornamented protrusion on middle of in- 2, V2 thickness of endopod and Vi length of ner margin; length of protrusion equal to exopod 2, straight, finely denticulate. Ter- width of endopod, its width equal to V2 minal claw tapering gradually from en- width of endopod. Outer margin of exopod larged base, twice length of exopods 1 and 1 with large, distally directed conical pro- 2 combined, with row of teeth extending cess on distolateral comer. Exopod 2 almost from tip of claw almost to base. Right en- twice length ofexopod 1, with shallow lon- dopod without suture on posterior surface, gitudinal groove along distal V2 of inner suture visible on anterior surface where en- 108 PROCEEDINGS OFTHE BIOLOGICAL SOCIETY OF WASfflNGTON dopod narrows slightly at midlength; en- one or both margins, longer and stouter dopod tapering to blunt point; with 3-5 than outer, naked spiniform seta. spines near outer distal margin, and subter- Type locality.—Pool #41, 40°37'N, minal grou—p of small hairs. 121°03'W, Las—sen County, Cahfornia. Female. Length (mm) of allotype 2.39; Etymology. The species name is given of ten paratypes, mean = 2.20, median = after the State of California, in which the 2.22, range = 1.95-2.33. Prosome (Fig. 3a) type locality is located.— symmetrical, thoracic wings symmetrical, Habitat desc—ription. The three pools each with two sensilla locatedon lateral and (41, 42 and 43 Long Lake) are relatively posterior margins. Genital double somite large for vernal pools (400 X 400 m, 300 extended laterally in asymmetrical conical X 300 m and 800 X 200 m respectively at projections each tipped with sensillum, left time of sampling), shallow (maximum projection directeddorsally, right projection depths 0.15-0.6 m), clear, and covered 70- laterally. Caudal rami haired in pattern sim- 90% with aquatic grasses. The electrical ilar to that of male. Rostral points (Fig. 3b) conductivity was low (30, 60 and 10 low, rounded. IjlMHO), water temperature 16, 16 and 23 Antennule (Fig. 3a, c) reaching mid- C, pH 7.7, 7.7 and 8.2, alkalinity 20, 28 and length of genital double somite. Segments 8 ppm, and total dissolved solids 10, 30 and 1-8 or9 each with few tiny hairs irregularly ppm respectively. — Discussion and comparisons. Hespero- scattered on posterior surface. Seta on seg- ment 1 long, reaching midlength ofsegment diaptomus californiensis differs from H. 4. Appendages per segment as follows (Ro- schejferi (Wilson, 1953) in the male leg 5: man numerals = segment, Arabic numerals the conical process on the coxa of the right = number of setae, a = aesthetasc, sp = leg is larger; the inner protrusion on the ba- sis of the right leg of H. californiensis is spine): 1(1+a), II(3+a), 111(1+a), IV(1), V(l+a), VI(1), VII(l+a), VIII(l-Hsp), rounded, not quadrate as in H. schejferi; IX(2+a), X(l), XI(2), Xn(l+sp+a), XIII(l), and the endopods ofboth left and right legs are longer in H. californiensis. In leg 5 of XIV(l+a), XV(1), XVI(l+a), XVII(l), the female, H. californiensis has the endo- XVIII(l), XIX(l+a), XX(1), XXI(l), pod extended in a terminal point. The gen- XXn(2), XXIII(2), XXIV(5+a). ital double somite in H. schefferi has very Leg 2 (not figured) of all females ex- slight lateral projections. amined without trace of Schmeil's organ or Hesperodiaptomus victoriaensis is mor- transverse ridge on endopod 2. phologically closest to H. californiensis. Leg 5 (Fig. 3d, e): Coxa with posterior The male of H. californiensis differs from lateral protrusion ending in sensillum. Basis H. victoriaensis in having the right leg 5 with lateral seta reaching Vz oflength ofex- with longer endopod, and in having the in- opod 1. Endopod two-segmented, distal ner process on the basis larger and placed segment twice length of proximal segment. at about midlength rather than at the inner Endopod 2 with two setae at tip, tip pro- proximal corner. In the female of H. victo- truding in acute point between these setae, riaensis, the lateral projections of the gen- and 3-5 subterminal teeth surrounding tip ital double somite are slightly asymmetri- of endopod. Exopod 2 a little longer than cal, the left projection being directed pos- exopod 1, claw with inner margin toothed. terolaterally rather than dorsally. Exopod 2 with small articulated spine lat- Both H. schefferi and H. victoriaensis eral to outer margin of exopod 3, reaching possess an acute process on the distal mar- from slightly beyond middle to end of ex- gin of segment 16 of the right antennule of opod 3. Exopod 3 distinct from exopod 2. the male. Such a process is lacking in H. Inner spiniform seta of exopod 3 serrate on californiensis. VOLUME NUMBER 109, 1 109 The differences between H. kiseri and H. cies are found in eastern Canada and the californiensis are primarily in leg 5 of the U.S.A., and the ranges of three species ex- male: in H. californiensis there is no prox- tend into Siberia (Borutskii et al. 1991). imal protrusion on the inner margin of the The species that are morphologically clos- right basis, and only one protrusion on the est to H. californiensis occur well north of middle ofthe inner margin. There is no dis- its range. Hesperodiaptomus kiseri is found tally placed spiniform projection on the in- in the State ofWashington and in Saskatch- ner margin ofthe right exopod 1. The small ewan (Wilson 1959). Hesperodiaptomus protrusion on the posterior surface of the schefferi is found in the Pribilof Islands, right exopod 2, present in H. kiseri, is lack- Alaska, and the northern Rocky Mountain ing in H. californiensis. The claw is States (Wilson 1959). Hesperodiaptomus smoothly tapered in H. californiensis, with- victoriaensis has been collected in the out angles as in H. kiseri. The lateral pro- mountains of southwestern Alberta (Ander- jections of the genital double somite of the son 1967, 1971), near Churchill, Manitoba & female are asymmetrical in H. californien- (Hebert 1985; Boileau Hebert 1988), and sis, symmetrical in H. kiseri. The thoracic on Victoria Island, N.W.T. (Reed 1958). wings are not expanded laterally in H. cal- Two other species of diaptomids, Lepto- iforniensis. diaptomus tyrrelli and Hesperodiaptomus In H. californiensis, the left lateral pro- novemdecimus, occurred in large numbers jection on the female double somite is dor- together with H. californiensis in all three sally directed. The second (middle) uro- pools sampled. Many instances of co-oc- somite is relatively long. On leg 5 of the curring diaptomid species have been re- male, the endopodites are relatively longer ported and have been much discussed in the than in other similar species. published literature, including reviews by In the most recent diagnosis ofthe genus Cole (1961) and Hutchinson (1967). Of 34 Hesperodiaptomus, Borutskii et al. (1992) examples of co-occurrence listed by Cole stated that the left antennule of the male is (1961), 10 involved congeneric species. like that of the female. However, H. hirsu- Hutchinson (1967) observed that co-occur- tus (Wilson, 1953) always, and //. schefferi ring diaptomid species tend to be of differ- sometimes, display sexual dimorphism in ent sizes and suggested that these size dif- setation on some segments (Wilson 1953). ferences may indicate non-overlapping The observed setation pattern ofH. califor- feeding niches, and that differences in feed- niensis was constant, similar in both sexes, ing would likely be found to be related to and the most common pattern found in the structural differences. genus (cf. Wilson 1953). Several co-occurrences of species of Species of the genus Hesperodiaptomus Hesperodiaptomus have been reported. are supposed to lack Schmeil's organ (Wil- Wilson (1953) reported H. wardi (Wilson, son 1953). Schmeil's organ (Schmeil 1896) 1953) together with H. novemdecimus. An- is a small protuberance of unknown func- derson (1971) collected H. shoshone (S. A. tion, which in diaptomids if present is lo- Forbes, 1893) with H. arcticus, and H. cated on the posterior surface of leg 2 en- shoshone with H. victoriaensis. Hammer & dopod 2. Ifthe small transverseridge in this Sawchyn (1968) found H. arcticus in the position which was noted in some of the same pond as H. kiseri, although the inci- males of H. californiensis examined, does dences were separated by a five-day inter- correspond to Schmeil's organ, this is the val. These authors noted distinct size dif- first reported occurrence of this structure in ferences between the two species, but pos- the genus. tulated mutual exclusion between them. An Most species ofHesperodiaptomus occur especially interesting example was given by in northwestern North America. A few spe- Anderson (1967), who found H. shoshone —— 110 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON with H. victoriaensisand L. tyrrellitogether , & R D. N. Hebert. 1988. Genetic differenti- in a pond in Alberta. There, H. shoshone ation of freshwater pond copepods at Arctic individuals averaged 1.6 times longer than Borutskisii,teEs..—V.H,ydLr.oAb.ioSltoegpiaano1v6a7,/1&68M:.39S3.-4K0o0s.. 1991. H. victoriaensis and preyed actively upon RevisionoftheCalanoidaofFreshwatersofthe the much smaller L. tyrrelli. Anderson USSR. Zoological Institute, USSR Academy of (1967) described the enlarged clawlike se- Sciences, SaintPetersburg,503 pp. (inRussian; tae ofthe maxilliped ofH. shoshone, which abstract in English). were apparently an adaptation for this pred- Cole, G. A. 1961. Some calanoid copepods from Ar- atory activity. The corresponding setae of izona with notes on—congeneric occurrences of Diaptomus species. Limnology and Oceanog- H. victoriaensis were found to be unmodi- raphy 6(4):432-442. fied, and this species apparently did notpre- Fischer, S. 1853. Beitrage zur Kenntnis der in der date upon the smallerdiaptomid. In the case Umgegend von St. Peters—burg sich findenden of the Lassen County Hesperodiaptomus Cyklopiden; (Fortsetzung). Bulletin de la So- pair, the setae of the mouthparts and max- ciete Imperiale des Naturalistes de Moscou 26: illiped ofboth species are unmodified. Hes- Forbes,7S4.-A1.0018+93p.laAtesprIeI,liImIiI.naryreporton the aquatic perodiaptomus novemdecimus averaged invertebrate fauna of the Yellowstone National larger than H. californiensis in all three Park, Wyo—ming, and of the Flathead region of ponds, and the two could be reliably sorted Montana. Bulletin of the United States Fish because of this size difference. In accor- Commissionfor 1891:207-258 + platesXXXVH- dance with Hutchinson's (1967) hypothesis, XLn. & we suggest that differences between these Guerne, J. de, J. Richard—. 1889. Revision des Cal- anides d'eau douce. Memoires de la Societe species in feeding niches are likely to exist, Zoologique de France 2:53-181 + plates I-IV. although we have made no direct observa- Hammer, U. T, & W. W. Sawchyn. 1968. Seasonal tions. successionandcongenericassociationsofDiap- tomus—spp. (Copepoda) in some Saskatchewan Acknowledgments ponds. Limnology and Oceanography 13(3): 476-484. This article was prepared during the se- Hebert, P. D. N. 1985. The ecology of th—e dominant nior author's participation in the Mentor- copepod species at alow arctic site. Canadian ship Program ofthe Thomas Jefferson High Journal ofZoology 63:1138-1147. School for Science and Technology, Alex- Herrick, C. L. 1882. Cyclopida—e of Minnesota with andria, Virginia. We thank the collector, Ja- notes on other copepods. Report of the Geo- logical and Natural History Survey of Minne- mie L. King, University of California, Da- sota 10:221-223 + plates I-VII. vis for providing habitat information and a Hutchinson, G. E. 1967. A treatise on limnology. II. draftofthe articleby King et al. Jamie King John Wiley & Sons, Inc., New York, 1115 pp. and two anonymous reviewers made valu- Kiefer, F. 1927. Freile—bende Siisswasser-Copepoden able suggestions on an earlier draft. aus Nordamerika. Zoologischer Anzeiger 72: 262-268. Kincaid,T. 1953. Acontributiontothetaxonomyand Literature Cited distribution ofthe American fresh water calan- Anderson, R. S. 1967. Diaptomid copepods fromtwo oid Crustacea. The Calliostoma Company, Se- mofouZnotoaliongypo4n5d:s10i4n3A-l1b0e4r7t.a.—Canadian Journal King, Ja.ttLl.e,, WMa.shAi.ngStiomno,vi7c3h,pp&. +R.plCa.teBsru1s-c5a.. 1996. . 1971. Crustaceanplanktonof146 alpineand Endemism, species richness, and ecology of subalpinelakesandpondsinwesternCanada. crustacean as—semblages in northern California JournaloftheFisheriesResearchBoardofCan- vernal pools. Hydrobiologia (in press). ada 28:311-321. Light,S.F. 1938. NewsubgeneraandspeciesofDiap- Boileau, M. G. 1991. A genetic determination of tomid copepods from—the inland waters ofCal- crypticspecies (Copepoda: Calanoida)andtheir ifornia and Nevada. University of California postglacial biogeography in North America. PubliWcations in Zoology 43:67-78. Zoological Journal ofthe Linnean Society 102: Lilljeborg, 1901. Synopsisspecierumhueu—squein 375-396. Suecia observatorum generis Cyclopis. Kon- —— VOLUME 109, NUMBER 1 111 glige Svenska Vetenskaps-Akademiens Han- Calanoida). Pp. 64—99 in Biogeography of the dlingar 35:3-118 + Plates I-VI. Beringian sector of the Subarctic: Proceedings Marsh, C. D. 1920. The fresh waterCopepod—a ofthe of the Tenth All-Union Symposium, Vladivos- Canadian Arctic Expedition 1913-18. Report tok, 1986 (in Russian) (not seen; cited in Bo- of the Canadian Arctic Expedition 1913-18, rutskii et al. 1991). 7(Crustacea), Part J:3-25. Wilson, M. S. 1953. New and inadequately known Poppe, S. A. 1888. Diagnoses de deux espec—es nou- Nort—h American species of the genus Diapto- velles du genre Diaptomus Westwood. Bulle- mus. Smithsonian Miscellaneous Collections 122:1-30. t1i5n9.de la SocieteZoologiquede France 13:158- . 1958a. New records and species ofcalanoid Reed, E. B. 1958. Two new species of Diaptomus copepods from Saskatchewan and Louisiana. Canadian Journal ofZoology 36:489^97. Cforpoempoadrcat)i.c—anCdanasdubiaarnctJiocurCnaanlaodfaZ(oCoalloagnyoid3a6:, p.ods1.9548.b.DiaNgonrotshesAmoefrniecawnspheacripeasctoifcofirdeshc-owpae-- 663-670. Schmeil, O. 1896. DeutschlandsfreilebendeSiisswas- ter Canthocamp—tidae and Cletodidae (genus Huntemannia). Proceedings of the Biological ser-Copepoden. Ill Teil: Centropagidae. Erwin Society ofWashington 71:43^8. StreletsNkaagyeal,eE.VerYl.ag1,98St3u.ttgTahret,"1e43wepnp.f group of the 7.381-95799.4 FirneeW-.livTi.ngEdCmoopnedpsoodna,: Ceda.l.anWoaidrad.aPnp.d genus Hesperodiaptomus (Copepoda, Calanoi- Whipple's fresh-waterbiology. Second edition. da) and a new sp—ecies H. koolensis from the John Wiley & Sons, Inc., New York, 1248 pp. ChukotPeninsula. ZoologicheskiiZhurnal62: Yeatman, H. C. 1944. American cyclopoid copepods 1474-1480 (in Russian). of the viridis-vernalis group (including a de- . 1986. Toward a revision of the Beringian scription of Cyclops carolinianus n. sp.). freshwater calanoid crustaceans (Copepoda, American Midland Naturalist 32:1-90.

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