PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF'V^SH'INGTON 113(4):1064-1078. 2000. Brachycalanus flemingeri and B. brodskyi, two new (Crustacea: : Phaennidae) from benthopelagic waters of the tropical Pacific

Frank D. Ferrari and E. L. Markhaseva (PDF) Department of Invertebrate Zoology, National Museum of Natural History; Smithsonian Institution; Washington, DC. 20560-0534, U.S.A.; (ELM) Russian Academy of Sciences, Zoological Institute; Universitetskaya nab. 1, 199034, St. Petersburg, Russia

Abstract.—Brachycalanus flemingeri, new species, has three setae on coxal endite of maxilla 1, a bifurcate worm-like sensory seta on the exopod of max- illa 2, a distal, semicircular protrusion posteriorly on the basis of swimming leg 1, and lacks a medial seta on coxa of switnming leg 4. Brachycalanus brodskyi, new species, is differentiated by its large size, 4.0 mm, and a semi- circular row of small denticles distally and posteriorly on the basis of swim- ming leg 1. Brachycalanus ordinarius (Grice, 1972) is redescribed and sepa- rated from the remaining species by the exopod of maxilla 2 with a distally constricted worm-like seta with a small lobe near the point of constriction. All three species have nine sensory setae on the distal basal lobe plus exopod of maxilla 2; this number of sensory setae has been reported for one other phaen- nid and several scolecitrichids. Suggested shared derived character states for the species oi Brachycalanus Farran, 1905 are epicuticular extensions of the female genital complex and the following two abdominal somites, and knife-like aesthetascs on female antenna 1.

The Brachycalanus Farran, 1905 m (Grice 1972). No adult males of a species includes four species: B. atlanticus (Wol- of Brachycalanus have been collected. The fenden, 1904), B. bjornbergae Campaner, two new Brachycalanus species described 1978; B. ordinarius (Grice, 1972) and B. here are the first records of the genus from rothlisbergi Othman & Greenwood, 1988 the eastern tropical Pacific Ocean. Brachy- although Othman & Greenwood (1988) calanus ordinarius is redescribed, based a suggested that specimens of B. atlanticus paratype. reported by Farran (1905) do not belong to the same species as the specimens of Wol- Materials and Methods fenden (1904). Different species have been reported from the North Atlantic Ocean The specimens from Hawaii collected on (Wolfenden 1904, Farran 1905, Grice 6 July 1997 may have been in a 1972), the South Atlantic Ocean (Campaner net for up to 12 hours prior to sample fix- 1978) and the Gulf of Carpentaria in the ation with 4% formaldehyde; few internal western tropical Pacific Ocean (Othman & tissues remain in these specimens. The Greenwood 1988); only B. atlanticus has specimen collected from Volcano 7 during been reported more than once, by Wolfen- November 1988 was fixed at its depth of den (1904) and Farran (1905). Specimens capture with gluteraldehyde; most internal of Brachycalanus usually are collected tissues remain. Ferrari & Markhaseva from near-bottom localities at depths from (1996, 2000), give further details about col- 72-100 m (Campaner 1978) to 992-1000 lections from Volcano 7 and Hawaii. In the

VOLUME 113, NUMBER 4 1065 laboratory, specimens from both localities toward the distal end of an appendage are were preserved in 0.5% propylene phen- distally polarized; an array arranged in a oxytol/4.5% propylene glycol/95.0% deion- semicircle are radially polarized. Specimens ized freshwater. During examination, spec- remain deposited in the National Museum imens were cleared in steps through 50% of Natural History, Smithsonian Institution, lactic acid/50% deionized freshwater to Washington, D.C. (USNM). 100% lactic acid, stained by adding a so- lution of chlorazol black E dissolved in Brachycalanus flemingeri new species 70% ethanol/30% deionized freshwater, and Figs. 1-3 examined with bright-field and with differ- ential interference optics. Drawings were Specimens.—\\o\oXyY>e^ (USNM 307711) a made with a camera lucida. Dissected and dissected 2.41 nmi female; prosome 2.02 undissected specimens are preserved in mm, urosome 0.44. Paratypes: (USNM 70% ethanol/30% deionized freshwater. 307712) 4 mature females (2.01; 2.04, two Cephalic appendages are abbreviated Al specimens; 2.37 mm) and 3 males CV (2.01; = antenna 1; A2 = antenna 2; Mn = man- 2.07; 2.07 mm) and 1 CIV female (1.65 dible; Mxl = maxilla 1; Mx2 = maxilla 2. mm). Type locality: eastern tropical Pacific Thoracic somites are Thl-7. Appendages Ocean, 19°43'27.01"N, 156°04'35.46"W, off on thoracic somites are Mxp = maxilliped Kona, Island of Hawaii, 6 Jul 1997. (thoracopod 1); PI-5 = swimming legs 1- CVI Female.—(Fig. lA) Cephalon and 5 (thoracopods 2-6). The caudal ramus is Thl fused; Th2-Th4 separate; Th5 and Th6 CR. Measurements are lengths; the length separated but not well articulated. Laterally, of the whole is measured from the posterior comers of prosome triangular; anterior edge of the cephalothorax to the dorsally, acutely triangular and almost posterior edge of the caudal ramus, and reaching mid-length of genital complex does not include the overlap onto the gen- (Fig. IG). Th4 with 2 sensilla laterally; ven- ital complex of the posterior edge of the sixth thoracic somite. Designations of ap- tral longer; Th5 with 1 short dorsal sensilla pendage segments follow Ferrari (1995), laterally. Genital complex and following 2 Ferrari & Dahms (1998) and Ferrari & abdominal somites (Fig. ID, E, H) with Markhaseva (2000). The coxa of the max- small, scale-like spinules. Genital complex illiped of copepods has one lobe with a symmetrical. group of setae (Ferrari & Ambler 1992, Rostrum (Fig. IB, C): subdivided plate Martinez Arbizu 1997); the remaining three with 2 filaments. groups of setae on the calanoid syncoxa be- Al: reaching Th3 and of 24 articulating long to the praecoxa. Articulating arma- segments with groups of setae: 3?, 7?, 3?, ment elements of appendages are termed 2?, 3?, 2?, 3?, 5?, 1, 1, 3?, 1, 3?, 1, 2?, 1, setae regardless of their morphology or de- 2?, 2?, 1, 1, 2?, 2?, 2?, 5-1-1. gree of rigidity. Two setae and one aesthe- A2 (Fig. 2A): coxa with 1 seta and den- tasc on a segment of antenna 1 are desig- ticles. Basis with 2 setae. Ri 2-segmented nated 2-1-1; "?" indicates that a setal ele- with 2 and 14 (6 terminal and 8 subtermi- ment was broken so that its identity on an- nal) setae. Re 7 articulated segments with tenna 1 could not be determined and only 0, 1, 1, 1, 1, 1, and H-3 setae; 2nd segment the scar at the location of attachment of the with several small indentations on medial seta was counted. Setules are epicuticular face and seta minute. extensions of a seta; denticles are epicutic- Mn: coxal gnathobase (Fig. 2C) some- ular extensions of an appendage segment; what elongate with 3 compound and 4 sim- spinules are epicuticular extensions of a so- ple teeth and 1 .seta. Basis with 3 setae (Fig. mite. An array of denticles whose tips point 2B). Re indistinctly segmented apparently 1066 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Brachycalanus flemmaeri new species. A, habitus, right lateral; B, anterior part of cephalon, ventral; C, rostrum, anterior-ventral; D, Th5-6 and urosome, right lateral; E, urosome, ventral; F. genital complex, ventral; G, posterior Th4, Th5-6 and genital complex, dorsal; H, urosome dorsal. Scale lines are 0.1 mm. VOLUME 113, NUMBER 4 1067

with 1, 1, 1, 1, 2 terminal setae; Ri 2-seg- P2 (Fig. 3C, D): all segments anteriorly mented with 2 and 9 setae. and posteriorly with complex arrays of Mxl: praecoxa elongate with 9 terminal, large and small denticles; arrays may be 1 anterior and 4 posterior setae (Fig. 2E); distally or radially polarized. Coxa with coxal epipodite with 1 thin, short seta and medial seta. Basis without seta. Re 3-seg- 8 long, thick setae; coxal endite with 3 se- mented, proximal with 1 medial and 1 lat- tae; 1st and 2nd basal endites with 4 and 5 eral seta, middle with 1 medial and 1 lateral setae respectively (Fig. 2D). Ri with 11 se- seta, distal with 4 medial, 1 terminal, 3 lat- tae (3-1-3+5); Re with 10 setae. eral setae. Ri 2-segmented, proximal with 1 Mx2 (Fig. 2G): proximal praecoxal en- medial seta, distal with 2 medial, 2 termi- dite with 5 setae; distal with 3 setae, 1 poor- nal, 1 lateral setae. ly sclerotized. Proximal coxal endite with 1 P3 (Fig. 3E, F): all segments anteriorly short, poorly sclerotized sensory seta and 2 and posteriorly with complex arrays of long well-.sclerotized setae; distal coxal en- large and small denticles; arrays may be dite with 1 thick and 2 thin setae. Proximal distally or radially polarized. Coxa with basal endite with 1 long, thick seta, 1 long, medial seta. Basis without seta. Re 3-seg- thin seta and 2 poorly sclerotized sensory mented, proximal with 1 medial and 1 lat- setae. Distal basal lobe + Re with 9 sensory eral seta, middle \)Pith 1 medial and 1 lateral setae; 7 brush-like setae with short setules seta, distal with 4 medial, 1 terminal, 3 lat- and 2 worm-like setae, 1 of which is bifur- eral setae. Ri 3-segmented, proximal with 1 cate toward its distal end. medial seta, middle with 1 medial seta, dis- Mxp (Fig. 2F): syncoxa with 1 worm- tal with 2 medial, 2 terminal, 1 lateral setae. like seta on proximal lobe; 1 sclerotized and P4 (Fig. 3G-I): all segments except prox- 1 worm-like sensory seta on middle lobe; 1 imal endopodal anteriorly and posteriorly sclerotized and 1 brush-like sensory seta with complex arrays of large and small den- with short setules on distal lobe; coxal lobe ticles; arrays may be distally or radially po- with 3 setae and distal denticles. Basis with larized. Coxa without seta. Basis without 3 setae on unattentuated proximal lobe and seta. Re 3-segmented, proximal with 1 me- 2 setae on distal lobe; denticles proximally. dial and 1 lateral seta, middle with 1 medial Ri segments from proximal to distal with 4, and 1 lateral seta, distal with 4 medial, 1 4, 3, 4 (1 lateral), 4(1 lateral) setae respec- terminal, 3 lateral setae. Ri 3-segmented, tively. proximal with 1 medial seta, middle with 1 PI (Fig. 3A, B): all segments with den- medial seta, distal with 2 medial, 2 termi- ticles, anteriorly and posteriorly. Coxa nal, 1 lateral setae. without seta. Basis with medial seta and P5 (Fig. 3J): all 3 segments posteriorly distal semicircular protrusion on posterior with denticles of various lengths. Coxa and face. Re 3-segmented, proximal with 1 lat- basis without a seta. Re 1-segmented with eral seta, middle with 1 medial and 1 lateral 1 medial, 2 terminal and 1 lateral setae. seta, distal with 3 medial, 1 terminal, 1 lat- CR (Fig. 2E): 4 large, terminal setae, 1 eral setae; lateral seta on proximal segment small medial-dorsal seta, and 1 small lat- reaches to base of lateral seta on middle eral-ventral seta. segment and lateral seta on middle segment Etymology.—The species name honors reaches to base of lateral seta on distal seg- Abraham Fleminger of Scripps Institution ment. Ri a 1-segmented complex with 3 of Oceanography for his contributions to medial and 2 terminal setae. Anterior face the systematics of calanoid copepods. of Von Vaupel Klein's organ with well-de- Remarks.—Autapomorphies for B. fle- veloped triangular protuberence, anterior mingeri include: one bifurcate, worm-like row of long denticles proximally and pos- sensory seta on exopod of maxilla 2; basis terior pore. of swimming leg 1 with semicircular pro- 1068 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Brachycalanus flemin^eri. new species. A, antenna 2, ventral; B, mandibular palp, dorsal; C, man- dibular gnathobase. distal: D, maxilla 1. anterior; E, maxilla I, praecoxal lobe, posterior, di.stal down; F, max- illiped, anterior; G, maxilla 2 anterior. Scale lines are 0.1 mm. VOLUME 113. NUMBER 4 1069

Fig. 3. Brachycalanus flemingeri, new species. A, PI, anterior; B, basis and rami of PI, posterior; C, coxa, basis and endopod of P2, posterior; D, exopod of P2. posterior, medial seta of proximal segment missing; E, coxa, basis, endopod and proximal and middle segment of exopod of P3, posterior; F, distal part of middle segment and distal segment of exopod of P3, posterior; G, middle segment and distal segment of exopod of left P4. posterior; H, basis and proximal segment detached from middle and distal segments of endopod of right P4, posterior; 1, coxa, basis and proximal ramal segments of left P4. posterior, medial seta of proximal exopodal segment missing; J, P5, posterior, coxa detached. Scale lines are 0.1 mm. 1070 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON trusion at the base of medial seta; three se- Zoological Institute of the Russian Acade- tae on coxal endite of maxilla 1 (rather than my of Sciences, for his contributions to the two setae); and swimming leg 4 without a systematics of calanoid copepods. medial coxal seta. Remarks.—Brachycalanus brodskyi new species differs from the other species of the Brachycalanus brodskyi, new species genus by its size 4.0 mm (remaining spe- Figs. 4-7 cies: 1.52-3.40 mm); basis of swimming leg 1 with semicircular row of small den- Specimens.—Ro\o\.y^& (USNM 307714) ticles near origin of the medial seta. a dissected 4.0 mm female; prosome 3.3 Brachycalanus brodskyi shares eight mm, urosome 1.0 mm. Type locality: east- brush-like setae with long filaments and 1 em tropical Pacific Ocean 1—5 m above bot- worm-like seta on distal basal lobe + exo- tom at base of Volcano 7 (13°23'N, pod of maxilla 2 with B. bjornbergae; this 102°27'W), at depths of 2945-3010 m, dive composition apparently is shared also with 2147, Nov 1988. The anterior seta on prae- B. rothlisbergi (Othman & Greenwood coxal endite of Mxl of the single known 1988 fig. 2C). B. brodskyi shares with B. specimen apparently broken; one seta on flemingeri the acute triangular shape of pos- the middle praecoxal lobe of Mxp appar- terior corners of prosome. ently broken. CVI female.—Differs from B. flemingeri Brachycalanus ordinarius (Grice, 1972) as follows: in habitus, dorsally posterior Fig. 8 comers of prosome exceeding the middle length of genital somite (Fig. 4H). Genital Xanthocalanus ordinarius.—Grice, 1972: complex with long denticles posterior-ven- 237, figs. 101-119. trally. Brachycalanus ordinarius.—Campaner, Al (Fig. 5A-C): 3, 6-1-1, 2-1-1, \ + \, 1978: 976-977. Othman & Greenwood, 24-1, 2?, 2-1-1, 4+1, 1, 1, 2+1, 1, 2+1, 1, 1988: 357. 2, 1,2, 1 + 1, 1, 1,2?, 2, 2, 5+1. Many setae and aesthetascs present on this specimen. CVI female, paratype.—VSNM 137176; A2 (Fig. 6A), Mn (Fig. 6B, C), Mxl differs from B. flemingeri as follows: in (Fig. 6D) as illustrated. habitus, posterior corners of Th6 rounded Mx2: proximal and distal coxal endite (Fig. 8A). each with 1 poorly sclerotized sensory seta Al (Fig. 8B-D): 24 articulated segments (Fig. 6E). Distal basal lobe + Re with 9 with setal groups containing: 3, 6+1, 2+1, sensory setae, 8 brush-like with longer se- 2?, 2+1, 2, 2+1, 4+1, 1, 1, 2+1, I, 2+1, tules (Fig. 6F) and 1 worm-like seta. 1, 2, 1, 1 + 1, 1 + 1, 1, 1, 2, 2, 2, 5+1 setae. Mxp: (Fig. 6G, H) as illustrated. A2: without small distal seta on Re2. PI (Fig. 7A, B): basis with a semicircular Mx2: distal praecoxal endite without a row of small denticles posterior and distal poorly sclerotized seta; distal basipod lobe near origin of endopod, and without semi- + Re with 7 brush-like and 2 worm-like circular protrusion. sensory setae (Fig. 8F); 1 worm-like seta P2 (Fig. 7C, D), P3 (Fig. 7E, F) as illus- constricted distally with a small lobe near trated. point of constriction (Fig. 8G), the other a P4 (Fig. 7G, H): coxa with medial set; simple worm-like seta. middle and distal segments of both rami Mxp: worm-like sensory seta of proximal lost. lobe with long setules (Fig. 8H). P5 (Fig. 71) as illustrated. PI: basis without semicircular row of Etymology.—The species name honors small denticles or semicircular protrusion Konstantin Abramovich Brodsky of the near origin of medial seta; lateral seta on VOLUME 113, NUMBER 4 1071

H

Fig. 4. Brachycakmus hrodskyi. new species. A, habitus, left lateral; B, prosomc and genital complex, dorsal; C, rostrum, lateral; D, urosome. dorsal; E, posterior Th5-6 and urosome, left lateral: F. genital complex, left lateral; G, genital complex, ventral; H, genital complex, dorsal. Scale lines are 0.1 mm. 1072 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 5. Brachycalanus hrodskyi, new species. A, antenna 1, aniculating segments 1-6, ventral down; B, antenna 1, articulating segments 7-17, ventral down; C, antenna 1, articulating .segments 18-24, ventral down. Broken line indicates poorly resolved edge of aesthetasc. All scale lines are 0.1 mm. VOLUME 113, NUMBER 4 1073

Fig. 6. Brachycalanus hrodskyi, new species. A, antenna 2. dorsal; B, mandibular palp, ventral: C, mandib- ular gnathobase, distal; D, maxilla 1, anterior; E, maxilla 2, posterior, distal basal lobe + cxopod omitted; F, exopod of maxilla 2 showing 3 terminal aestheta.scs; G, syncoxa and basis of maxilliped, anterior; H, endopod of maxilliped, anterior. All scale lines are 0.1 mm. 1074 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 7. BrachycaUmus hrodskyi, new species. A, PL posterior; B, PL anterior; C, P2, anterior, terminal seta cutoff; D, P2, posterior; E, P3, anterior, terminal seta cut off; F, P3, posterior; G, coxa, basis and proximal ramal segments of P4, posterior; H, coxa, basis and proximal exopodal segment of P4, anterior; I, P5, posterior. All scale lines are 0.1 mm. VOLUME 113, NUMBER 4 1075

Fig. 8. Brachycakmus oiclinurius (Gricc, 1972). A, posterior Th5-6 and genital complex, right lateral; B, antenna I, articulating segments 1-4, ventral toward right; C. antenna I. articulating segments 5-11, ventral toward right; D, antenna 1, articulating segments 12-21, ventral right; E, antenna 1. articulating segments 22- 24, ventral right; R distal basal lobe and exopod of maxilla 2; G, worm-like setae of exopod of maxilla 2; H, syncoxa and basis, with distal lobe omitted, of maxilliped. anterior. All scale lines arc 0.1 mm. 1076 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON middle segment not reaching base of lateral Othman & Greenwood (1988: 355) also seta of distal segment. suggested that Farran's (1905) specimens of Remarks.—Brachycalanus ordinarius B. atlanticus were not conspecific with differs from the remaining species of the those of Wolfenden (1904). Ohtsuka et al. genus in possessing 1 worm-like seta con- (1998) pointed out that Brachycalanus stricted distally with a small lobe near point shares with at least some species of Xan- of constriction on the distal basal lobe + thocalanus a proximal basal endite of max- exopod of maxilla 2. illa 2 with two worm-like sensory setae, and a syncoxa of the maxilliped with one Discussion or two worm-like sensory setae in addition to one brush-like sensory seta. These have The genus Brachycalanus was estab- not been reported for other phaennid gen- lished by Farran (1905: 41) for three spec- era. imens of B. atlanticus which were "... We found very few differences to sepa- washed from sand brought up by a townet rate the three species of Brachycalanus on the trawl . . . (off the west coast of Ire- studied here from the remaining species. land) .... They measured 2.5, 2.0 and 1.52 However, the number of autapomorphies mm respectively." Farran described the and degree of divergence among the species largest specimen, except for P5, "which should become more apparent after careful were only found in the smallest specimen, redescription, particularly of the setation of which while appearing fully developed in maxilla 1 and maxilla 2, and discovery of other respects, still showed immature seg- males of the five species. Among the three mentation of the abdomen". Farran men- species here that we place in Brachycalan- tioned in the description of the species us, there are different combinations of "While the above was in press a description worm-like and brush-like setae among the of this species under the name of Xantho- nine sensory setae of the distal lobe and calanus atlanticus, was published by Dr. exopod of maxilla 2: one worm-like and Wolfenden" (Farran 1905: 42). In establish- eight brush-like setae in B. bjornbergae and ing the genus, Farran mentioned that it is B. brodskyi; two worm-like and seven closely allied to Xanthocalanus Giesbrecht, brush-like in B. ordinarius and B. flemin- 1892 and noted as distinguishing characters geri. However, one worm-like seta each of the shape of rostrum and an extremely short B. ordinarius and B. flemingeri appears to Al. be more specialized; one worm-like seta of Brachycalanus was diagnosed by Cam- B. ordinarius is similar to a fleshy seta paner (1978: 976) based on the then three (Campaner 1978: 968) with a small lobe, known species; he removed B. minutus Gri- and one worm-like setae of B. flemingeri is ce, 1972 and gave the total number and bifurcate. It appears that a simple combi- kind of sensory setae on the ramus of max- nation of the different kinds of sensory se- illa 2 as 1 worm-like and "(?5) 7-8" brush- tae cannot be used to diagnose the genus like setae. In a later diagnosis, Bradford et Brachycalanus. Whether the number of al. (1983) added that the urosome, and dor- sensory setae which are not worm-like are sal and ventral (actually anterior and pos- phylogenetically significant, remains to be terior) surfaces of P1-P5 were covered with determined. At this time we suggest that spines, an important character distinguish- synapomorphies for the species of Brachy- ing the genus from Xanthocalanus. A later calanus are the presence of spinules on the discussion of Brachycalanus by Othman & female genital complex and following two Greenwood (1988) noted the presence of abdominal somites, and knife-like aesthe- knife-shaped aesthetascs on the antenna 1 tascs on female antenna 1. as derived character state for the genus. Species of the genus Brachycalanus pres- VOLUME 113. NUMBER 4 1077 ently are placed in the Phaennidae, one of brush-like seta is open distally so that its five families within the superfamily Clau- dendritic cilia are in direct contact with the socalanoidea that possess sensory setae or aquatic environment, as described by Nish- aesthetascs on the distal basal lobe plus ex- ida & Ohtsuka (1997). The number present opod of maxilla 2, and on the maxilliped on the ancestral clausocalanoidan and the (Nishida & Ohtsuka 1997). Bradford (1973) ancestor of the lineage which includes the and Bradford et al. (1983) suggested that Aetideidae, Phaennidae and Scolecitrichidae, the number and the kinds of these sensory and the number and kinds of transformations setae be used in diagnosing the Phaennidae of these ancestral states must await a careful and the Scolecitrichidae. Detailed descrip- redescriptions of the species of these fami- tions of this part of maxilla 2 were not pro- lies. vided for the type species of Brachycalanus (Wolfenden, 1904) or in the original de- Acknowledgments scription of the genus (Farran 1905). In his diagnosis, Campaner (1978: 976) men- We thank Dr. Karen R Wishner, Univer- tioned one worm-like plus "(5?) 7-8" sity of Rhode Island, for the specimens brush-like sensory setae; Othman & Green- from Volcano 7 and Dr. Edward J. Buskey, wood (1988) described this limb of B. roth- University of Texas, for those from Hawaii! lisbergi with one worm-like and ten brush- E. L. M. thanks the Smithsonian Institution like setae (although only eight brush-like and the American Philosophical Society for setae can be counted in their fig. 2C); fi- travel support to the National Museum of nally, illustrations by Grice (1972: 235, fig. Natural History; Smithsonian Institution; 109b) for B. ordinarius show two worm- Washington, D.C. like plus six brush-like sensory setae. We have observed nine such setae for B. fle- Literature Cited mingeri, B. brodskyi, and a paratype of B. Bradford. J. 1973. Revision of family and some ge- ordinarius. neric definitions in the Phaennidae and Scole- The presence of nine setae on the distal cithricidae (Copepoda: Calanoida).—New Zea- land Journal of Marine and Freshwater Re- basal lobe plus exopod of maxilla 2 of these search 7:133-152. three Brachycalanus, another phaennid, , L. Haakonssen, & J. Jillett. 1983. The marine Xanthocalanus pavlovskii Brodsky, 1955 as fauna of New Zealand: pelagic calanoid cope- well as all five species of the scolecitrichid pods: families Euchaetidae, Phaennidae. Scole- Landrumius (see Park 1983) and the scole- cithricidae, Diaixidae, and Tharybidae.—New Zealand Oceanographic Institute Memoir 90 citrichid Xantharus renatehaassae Schulz, 150 pp. 1998 indicates that the number of such setae Brodsky, K. A. 1955. On the copepod (Calanoida) fau- has not been conserved in these families as na of the Kuril-Kamchatka trench.—Trudy In- they are understood today. Although most stituta Okeanologii Akademii Nauk SSSR 12: descriptions of Clausocalanoida .species with 184-209. [in Russian] Campaner, A. F. 1978. On some new planktobenthic simple, sclerotized setae on the distal basal Aetideidae and Phaennidae (Copepoda, Calan- lobe plus exopod mention eight of these se- oida) from the Brazilian continental shelf 11. tae, a recent redescription of Pseudochirella Phaennidae.—Ciencia e Cultura 30:966-982. obesa Sars, 1920 by Vaupel Klein & Rijer- Farran. G. R 1905. Report on the Copepoda of the kerk (1997) states and figures nine such se- Atlantic slope off Counties Mayo and Gal- way.—Annual Report of the Fisheries Ireland, tae. Ohtsuka et al. (1998) suggested that 1902-1903, part 2, (2):23-52. brush-like setae may be derived from worm- Ferrari, F D. 1995. Six copepodid stages of RUIge- like setae because the dendritic cilia of a wayia klausruetzleri. a new species of copepod worm-like setae are enclosed within a thin (Ridgewayiidae, Calanoida) from the cuticle, homologous to the thick cuticle of barrier reef in Belize, with comments on ap- pendage development.—Proceedings of the Bi- sclerotized setae, while the cuticle of a ological Society of Washington 108:180-200. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1078 Smirnovipina barentsiana comb. nov. (Smirnov, — & J. W. Ambler. 1992. Nauplii and copepod- 1931)._Sarsia 82:313-323. ids of the cyclopoid copepod Dioithona oculata Nishida, S.. & S. Ohtsuka. 1997. Ultrastructure of the (Oithonidae) from a mangrove cay in Belize.— mouthpart sensory setae in mesopelagic cope- Proceedings of the Biological Society of Wash- pods of the family Scolecitrichidae.—Plankton ington 105:275-298. Biology and Ecology 44:81-90. —, & H.-U. Dahms. 1998. Segmental homologies Ohtsuka, S., I. Takeuchi, & A. Tanimura. 1998. Xan- of the maxilliped of some copepods as inferred thocalanus grticilis and Thanbis manna (Co- by comparing setal numbers during copepodid pepoda. Calanoida) rediscovered from the Ant- development.—Journal of Crustacean Biology arctic Ocean with baited traps.—Journal of Nat- 18:298-307. ural History 32:785-804. , & E. L. Markhaseva. 1996. Parkius karen- Othman, B. H. R., & J. G. Greenwood. 1988. Brachy- wishneriie. a new genus and species of calanoid calanm rolhlisherni. a new species of plankto- copepod (Parkiidae. new family) from bentho- bcnthic copepod (Calanoida, Phacnnidae) from pelagic waters of the eastern tropical Pacific the Gulf of Carpentaria, Australia.—Records of Ocean.—Proceedings of the Biological Society the Australian Museum 40:353-358. of Washington 109:264-285. Park, T. 1983. Calanoid copepods of some scolecith- , & . 2000. Griceus buskeyi. a new ge- ricid genera from antarctic and subantarctic wa- ters.—Antarctic Research Series 38:165-213. nus and species of calanoid copepod (Crusta- Sars, G. O. 1920. Calanoides recueillis pedant les cam- cea) from benthopelagic waters off Hawaii.— pagnes de S. A. S. Ic Prince Albert de Monaco Proceedings of the Biological Society of Wash- (Nouvcau supplement).—Bulletin Institut ington 113:78-88. Occanographic (Monaco) 377:1-20. Giesbrecht, W. 1892. Systematik und Faunistik der pe- Schulz. K. 1998. A new species of Xantluinis Andron- lagischen Copepoden des Golfes von Neapel ov. 1981 (Copepoda: Calanoida) from the me- und der angrcnzenden Meeres-abschnitte.— sopelagic zone of the Antarctic Ocean.—Hel- Fauna und Flora des Golfes von Neapel und der golaender Meeresuntersuchungen 52:41-49. angrcnzenden Meeres-abschnitte 19:1-831 + Vaupel Klein, J. C. von, & C. D. M. Rijcrkerk. 1997. 54 pis. A detailed redescription of the Pseiidochirella Grice, G. D. 1972. The existence of a bottom-living obesa female (Copepoda. Calanoida). II. Anten- calanoid copepod fauna in deep-water with de- nae and mouthparts.—Crustaceana 70:394-417. scriptions of five new species.—Crustaceana Wolfenden. R. N. 1904. Notes on the Copepoda of the 23:219-243. North Atlantic Sea and the Faroe Channel.— Martinez Arbizu, R 1997. A new genus of cyclopinid Journal of the Marine Biological Association of copepods (Crustacea), with a redescription of the United Kingdom 7:110-146.