Triglops dorothy, a new species of sculpin (Teleostei: Scorpaeniformes: Cottidae) from the southern Sea of Okhotsk

2006·3.9 MB·

by  T W Pietsch

238
—
Abstract Anewspeciesofthecottid Triglops dorothy, a new species of
ognentuhseTbraisgilsoposf2R1eisnpheacridmtenissdceoslclreicbteedd sculpin (Teleostei: Scorpaeniformes: Cottidae)
in Aniva Bay, southern Sakhalin from the southern Sea of Okhotsk
Island,Russia,andoffKitami,onthe
northerncoastofHokkaido,Japan,at
depthsof73-117 m. Ofthe ten spe- TheodoreW.Pietsch
ciesofTriglops nowrecognized,the
sniemwilsaprectioesT,.TpriinggleolpisRdeoirnothhayr,dtis,mwoesltl CSoclhloeogleofofAOqucaetaincaannddFFiisshheerryySScciieenncceess
known from the NorthAtlantic and UniversityofWashington
North Pacific oceans and through- CampusBox355020
outcoastalwatersoftheArctic.The Seattle,Washington98f95-5020
newspeciesdiffersfromT.pingeliin E-mailaddress:twp'g'u.Washingtonedu
a combination ofmorphometric and
meristic characters that includes JamesW.Orr
mostimportantlythenumberofdorso-
lateralscales;thenumberofoblique, ResourceAssessmentandConservationEngineeringDivision
scaleddermalfoldsbelowthelateral AlaskaFisheriesScienceCenter
line; andthe numberofgillrakers. NationalMarineFisheriesService,NOAA
7600SandPointWayNE
Seattle,Washington98115-6349
SculpinsoftheteleostfamilyCottidae 19 nominalspecies and subspecies, of
are nearly ubiquitous in cold-water which nine are currently recognized
benthichabitatsoftheNorthernHemi- as valid (Pietsch, 1993). Members
sphere,comprisingnearly200species ofthe genus are characterized most
in the North Pacific alone (Yabe and strikingly by having a small head, a
Nakabo, 1984; Sheiko and Federov, narrow, elongate body and a slender
2000; Mecklenburgetal., 2002; Love caudal peduncle, a long anal fin con-
et al., 2005), where they are found taining 18-32 rays, pelvic fins with
in almost every benthic habitat from a single spine and three soft rays,
theintertidaltotheuppercontinental branchiostegal membranes united
slope. Many species are preyed upon on the ventral midline but lying free
bylargerfishesandmarinemammals from the isthmus, and scales below
(Browne et al., 2002), and are them- the lateral line modified to form dis-
selves predators primarilyofsmaller creterowsoftinyserratedplatesthat
fishesandcrustaceans(e.g.,Tokranov, lie in close-set, oblique dermal folds.
1998;Hoff,2000;TokranovandOrlov, The species are generallydistributed
2001). Although many members of in rather deep water (primarily be-
the family are also commonly found tween 18 and 600 m, but specimens
in bycatch of commercial fisheries have been taken more or less on the
(Stevenson, 2004; Orlov, 2005), the surface and as deep as 930 m; An-
systematics andlifehistoriesofmost driashev, 1949; Hart, 1973; Fedorov.
species are poorly known (Nelson, 1986) throughout cold-water, conti-
1994;Hoff,2000;Hoff,2006)andnew nental shelfor slope regions ofthe
speciescontinuetobedescribed(e.g., North Pacific, North Atlantic, and
Yabe, 1995; Yabe and Maruyama, Arcticoceans. All appeartofeed pri-
2001;Yabeetal., 2001).Amorecom- marily on small planktonic and ben-
pleteunderstandingofthediversityof thicinvertebrates(Andriashev, 1949;
thefamilyisnecessarytounderstand Fedorov, 1986). Spawningtakesplace
the role ofcottids in the dynamics of fromlate summertowinter; theeggs
North Pacificecosystems. are demersal (Andriashev, 1949; Mu-
MtoantuhescSrciiepnttisfuibcmEidtitteord.28March2005 haTrhdte(c1o8t3t0i)d,giennculsudiTnrgigPlroiposniRsetiinu-s sicTkriagnldopAsblwea,s1f9i6r9s;t pFerdooproosve,d1b9y86J)o.-
M1a6nAuusgcursitptbyaptphreoSvceidenftoirfipcubEldiictaort.ion BanedanSt(e1r88n4i)a,sEJloarnduarnaaGnidlbeErvte(r1m8a96n)n, haasninneglseRsepienchiamredntf(r1o8m30W)e,sbtasGerdeenon-
Fish.Bull.104:238-246(2006). (1898) asjunior synonyms, contains land, but the type species T pingeli
PietschandOrr: Tnglopsdorothy.anewspeciesfromthesouthernSeaofOI<hotsk 239
was not named until 1837, after Reinhardt received theventralmid-line.Symboliccodesforinstitutionsare
two additional specimens, also from West Greenland. thoseprovidedbyLevitonetal. (1985).
During the next 130 years, 18 additional species or Statistical significance was assessed by ANOVA; all
subspecies were described, but little work of a com- differencesatP<0.05wereconsideredsignificant.Toaid
parative nature was done. Schmidt (1930) reviewed in the discrimination ofthe new species andits closest
some North Pacific specimens ofthe group, and later ally, T. pingeli, a standard principal component analy-
(Schmidt, 1950)discussedspecies identifiedin hiswork sis (PCA) was conductedon meristic characters, and a
on the ichthyofauna ofthe Sea ofOkhotsk. Taranets shearedPCAwasconductedonmorphometriccharacters
(1941) placed the genus, together with Sfer?uas, in its forasize-freeanalysis. Theanalyseswereconductedon
own subfamily. The Arctic and North Pacific species the correlation matrix ofraw meristic data and the co-
ofTriglops were studied byJensen (1944), Andriashev variance matrix oflog-transformed raw morphometric
(1949, 1954), and McAllister (1963), and a world-wide data. Differences between species were illustrated by
revision ofthe genus was subsequently published by plottingprincipal component(PC) 1 againstPC2 ofthe
Pietsch (1993). meristic analysis, sheared PC2 against sheared PCS of
Within the material ofTnglops examinedbyPietsch the morphometric analysis, and sheared morphometric
(1993)are 19specimensthatwereoriginallyrecognized PC2 against the standard meristic PCI.
by Charles Henry Gilbert as new to science (see "Dis- Following the PCA, a linear discriminant function
cussion" below). Labeled as such, but left unpublished analysis (DFA) was conducted on morphometric and
byGilbert, Pietsch (1993)cametothe sameconclusion, meristic data to establish the relative significance of
citinga lackofagreement ofthese specimens with the thosecharactersindistinguishingthespecies. Morpho-
diagnosis ofany ofthe nine recognized species ofthe metric datawerestandardizedby dividingbystandard
genus. However, still not totally convinced that the length. Each character was tested to verify assump-
specimens represented a new species, Pietsch (1993) tions ofmultivariate normality and for statistical sig-
again postponed aformal description. LabeledTriglops nificance; those found to be distributed nonnormally
sp. and setaside atthattime,thesespecimensarenow or that were nonsignificant were eliminated from fur-
re-examined in light ofthe discoveryoftwo additional theranalysis. The robustnessoftheDFAwastestedby
specimens, and evidence is provided to formally recog- conducting a leave-one-out cross-validation procedure.
nize the material as a new species. Statistical analyses were performed with S-Plus (vers.
6.1 for Windows, 2002, Insightful, Inc., Seattle, WA)
and SPSS (vers. 11.5.1 for Windows, 2002, SPSS Inc.,
Methods and materials Chicago, IL).
Standard length (SL) is used throughout unless indi-
catedotherwise.Measurementsareexpressedinpercent Systematics
oMfetshtaonddsarudseldenfgotrhtoarkiinngpecrocuennttsofanhdeamdelaesnugrtehm(eHnLt)s. TriglopsdorothyPietschandOrr,n.sp.
follow those described and figured by Miller and Lea Dorothy'ssculpin
w(1i9d7t2h:7is-1m2e)a,suwritehdbtehtewfeoelnlolwatienrgaladmdairtgiionnss:ofinttheerofrrbointtaall Figures 1-4,Tables 1-3
bonesattheirnarrowestwidths;headdepthismeasured Triglopssp.:Pietsch, 1993:379(probablyanundescribed
atmid-orbit;firstinfraorbitalporewidthismeasuredat species, descriptionpostponed).
the greatest width, anterior to posterior; all gill-raker
countsaretakenfromthefirstarch;dorsolateralscale- Holotype
rowcountsincludeallscales(orfusedclustersofscales)
withenlargedplate-likebases;lateral-linescalecounts USNM(U.S.NationalMuseumofNaturalHistory,Wash-
alsoinclude scales that occasionallyextend beyondthe ington, D.C.) 74578, female, 155 mm.Albatross Station
posteriormarginofthehypuralplate. Pectoral-finrays 5013,AnivaBay,46=17'N, 143°09'E,3-mAgassiztrawl,
arecountedfromdorsaltoventral.Countingtheoblique 77 m, substrataunknown, 25 September 1906.
dermalfoldsontheventralhalfofthetrunkpresentsa
specialproblem.Becausemanydermalfolds(thenumber Paratypes
depending on the species), which originate along the
lateral line, terminate irregularly before reaching the Twentyspecimens, 15males(72.5-148.0mm),5females
ventralmidline, countscanbe considerablyhigherand (68.5-143.0 mm). The followingmaterial was collected
more highly variable (than the values reported in the by the U. S. Fish Commission SteamerAlbatross in or
present study) if made along a transect close to the adjacenttoAniva Bay, Sakhalin Island, Russia, with a
lateral line; thus, it is important to count the dermal 3-mAgassiztrawl:CAS-SU22305,3females(68.5-133.0
folds along a transect beginning beneath the pectoral mm),station5007,46"03'N, 142°31'E,77m,greenmud,
fin and continuing along, close to and strictly parallel fine gray sand, 24 September 1906. USNM 74575, 2
with, the base ofthe anal fin (see Pietsch. 1993:337, males(72.5-80.0mm),station5006,46°04'N, 142°29'E,
Fig. 1). Dermal folds ofthe breast are counted along 79 m, green mud, fine gray sand, 24 September 1906;
240 FisheryBulletin 104(2)
Figure 1
Triglopsdorothyn. sp.,holotype.female, 155 mm, USNM74578.
USNM 74576, 3 males (73.0-77.5 mm), 1 female (83.0
mm), station 5008, 46°07'50"N, 142°37'20"E, 73 m,
green mud, fine gray sand, 24 September 1906; USNM
74577, female (143.0 mm), station 5016. 46°44'30"N,
oo 143°45'E, 117m,brownmud,fineblacksand,rock,clay.
25 September 1906; USNM 74579, 8 males (73.5-80.0
5 15 mm), station 5005, 46°04'40"N, 142°27'30"E, 77 m,
green mud, fine gray sand. 24 September 1906. The
O o • *•• fkaoildloo,winJgapmaant.erSieapltweamsbecrol1l9ec6t4e:dHfrUoMmZoff41K5i2t4a,mi1,mHaolke-
5 (132.0mm); HUMZ41525, 1male(148.0 mm).
Diagnosis
10 20 30 40 50 60 70
Headlength A species of Triglops distinguished from all other
describedmembersofthegenusinhavingthefollowing
combinationofcharacterstates:interorbitalwidth8.9-
11.5% HL;caudalfintruncatetoslightlyrounded; dor-
solateralscaleswelldeveloped. 18-31;lateral-linescales
47-50; transverse dermal folds ofbreast 4-10; oblique
dermal folds 67-99; dorsal-fin rays 23-25; pectoral-fin
rays17-19;gillrakers7-12;firstinfraorbitalporelength
3.6-5.0% HL; peritoneum everywhere unpigmented or
atmostwith few, widelyspaced melanophores.
Description
Body relatively compressed, tapering gradually poste-
riorly(Fig. 1); headcompressed, length 28.4-32.3% SL
(Table 1), depth 11.3-13.5% SL; snout pointed, upper
profilelineartoslightlyconvex,length27.4-36.3% HL,
%V-s 15 20 Dorso2l5ateralscales aHbLo;utuepqpuearljtoawleengxtthenodfionrbgit;anlteenrgitohrloyfosrlbiigth2t9l.y0-b3e2y.o0n%d^
lower;firstinfraorbitalporelarge, length3.6-5.0% HL
Figure 2 (aFricgh.,26A-)1;1giolnlrlaokweerrspa7r-t12of(aTracbhl,es2h),or0t-1anodnbulpupnte;rlpoanrgtesotf
gPillnoopttssheodfSorseioagtnhoiyffiOnck.ahnsotpt.cshk(a:or)a(fcAtr)eorwmsidTdtrihifgfloeforpefsnitrpisitantiginenflgriaTor(ir•--) pbeecltoowr)a;l-tfhiinrdrapyel2v0i.c-9f-i2n5r.a0y%1S2L.6(-s1e7x.u7al%lySLd,imloornpgheirc;thsaene
bital (10)poreversushead length; (B)Counts of firstandsecond;lengthofpelvicfinsexuallydimorphic
gillrakersversusdermalfoldsversusdorsolateral (see below); caudal peduncle compressed or roughly
scales. circular in cross section, length 14.7-18.3%f SL, depth
3.1-4.1% SL.Skincoveringelementsofupperandlower
PietschandOrr Tnglopsdorothy,anewspeciesfromthesouthernSeaofOkhotsk 241
242 FisheryBulletin 104(2)
jaws, throat, ventral portion ofcheeks, interopercle, spot usually more distinct than dorsal; males with a
extreme ventral tip ofsubopercle, and extreme dorsal dark spot near distal end ofdorsal and ventral lobes
marginofgillmembranenaked;remainingpartofhead ofcaudal fin, sometimes faint and difficult to discern
anddorsalhalfofbodyeverywherecoveredwithrough, in preserved specimens.
granular scales; approximately 10-16 close-set rows of
tiny granular scales on anterior and dorsal periphery Sexual dimorphism
ofeyeball,rowsusuallyextendingnearlytodorsalmost
margin ofeye. Oblique dermal folds terminating on Females with pectoral-fin rays slightly shorter (22.6-
caudal peduncle well before reaching ventral midline 23.4% SL)thanthoseofmales(20.9-25.0%SL).Females
butsometimescrossingspacebetweenanusandbaseof with pelvic-fin rays slightly shorter (12.6-14.7% SL)
anal fin. Dorsal-fin spines 10-11, anal-fin rays 23-25; thanthoseofmales(12.7-17.7% SL);tissuesurrounding
vertebrae45-47. pelvic-fin spine and first rayexpandedin males.
Color in preservation: dark brown dorsally, light
brown, tan, tocream ventrally, with four dark saddles Statisticalanalyses
alongdorsal length ofbodyextendingventrallyto, and
sometimes slightly below, lateral line, anteriormost Univariate analyses ofmorphometric characters indi-
saddle situated beneath spinous dorsal fin, two be- catedthatonlyinterorbitalwidthandthelengthofthe
neath soft dorsal fin, posteriormost saddle on caudal first infraorbital pore exhibited significant differences
peduncle; most specimens with a small saddle or nar- between T. dorothy and T. pingeli (Table 1). The analy-
row band of pigment crossing soft-dorsal fin at mid- ses ofmeristic characters, on the other hand, revealed
length (homologous tofifth saddleofmany congeners). significant differences in several characters, including
A series ofdark, more-or-less equally spaced spots or fewerdorsolateralscales,lateral-linescales, andpecto-
streaks extending along length of body just ventral ral-fin rays, and a greater numberofdermal folds and
to lateral line, spots nearly always interconnected gill rakers in T. dorothy than in T. pingeli (Table 1).
in males to form a conspicuous dark stripe across Three-wayplotsofdorsolateralscales,dermalfolds,gill
length ofbody, butnotextendingontocaudal-fin base. rakers, and pectoral-fin rays exhibit aclearseparation
Anterior portion oflower lip darkly pigmented, some- between T.pingeli and T. dorothy(Fig. 2B).
times interrupted at symphysis ofjaw; upper lip with InplotsofscoresfromthePCAofmeristiccharacters,
a darkstreak across symphysis and twoorthree simi- two distinct clusters, one for each ofthe two species,
lar, more-or-less equally spaced streaks on each side, were evident (Fig. 3A). The primary axis ofseparation
posteriormost streak contiguous with similar spot or was PCI (30.2%: oftotal variance), which was heavily
streak on maxilla (pigmentation oflips more intense loaded positively on dorsolateral scales and gill rakers
in males); males with a streak ofpigment extending andnegativelyonlateral-linescales;loadingsalongthe
alongdorsal margin ofupperjaw from ventral margin PC2 axis (22.4% oftotal variance) were heavily loaded
oflacrimal to beneath and slightly past eye; a large, positivelyforbreast-scale rows andnegativelyforanal-
diffuse spot posterior to distal tip of maxilla (more finrays anddorsal-finrays(Table3). Incontrast,broad
heavily pigmented in males), sometimes forming an overlap was displayed in the analysis ofmorphometric
oblique streak extending from posterodorsal margin characters, with individuals of T. dorothy clustering
ofmaxillaposteroventrally to preopercle. Dorsomedial among the larger group of T. pingeli from the Sea of
surface of opercle and posterodorsal margin of gill Okhotsk (Fig. 3B). The first principle component (PC)
membrane densely covered with small melanophores, was interpreted as a size component accounting for
pigment extending ventrally along medial surface of 95.5% ofthe total variance. In plots ofthe scores of
branchiostegal rays (pigmentationoften bleached away the sheared second and third shape components (1.5%c
in old, poorly preserved material); lateral surface of and 1.1% oftotal variance, respectively), the great-
distal portionoffourorfive posteriormostbranchioste- est dispersion was along the PC2 axis (Fig. 3B, C),
gal rays, as well as gill membranein between, heavily and loadings along this axis were highest for interor-
peppered with small melanophores. Each spine and bital width, snout length, and caudal-peduncle depth
ray ofdorsal fin (but not membrane in between) with (Table3). Loadings alongthePC3 axiswerehighestfor
two to four, dark, more-or-less equally spaced spots. pectoral-finraylength, ventralcaudal-pedunclelength,
Four to seven narrow, more-or-less equally spaced and orbit length (Table 3).
bands of dark pigment across dorsalmost or nonex- The single linear discriminant function equa-
erted rays ofpectoral fin; a small, dark spot at dorsal tion produced was highly significant (x~=96.5, 6 df,
insertion of pectoral fin often continuous with pig- P<0.0001):
ment extending ventrally to mid-base offin. Anal fin
usually unpigmented, but some specimens with rays D = 0.894(rfs)-0.691(g)-i- 0.530(p)-0.508(c?/')
darkly pigmented or linedwith rows ofmelanophores, -0.284(/o) + 0.098(//),
pigmentation especially intense in males; pelvic fins
usuallyunpigmented, but some males with distal tips where D=thediscriminantscoreofan individual;
of first and second rays lightly pigmented. Base of ds =numberofdorsolateral scales;
caudal finwith dark dorsal andventral spots, ventral g=numberofgill rakers;
PletschandOrr: Tng/opsdorolhy,anewspeciesfromthesouthernSeaofOkhotsk 243
Table3
244 FisheryBulletin 104(2)
Although the USNM catalog ledger has
"cotype"handwrittennexttoeachcatalog
number(all 19CAS-SUandUSNMspeci-
mens arelabeled as such) and Smithson-
ian Institution accession file no. 54484,
dated 6 September 1912, lists the mate-
rialas"co-typesdescribedbyDr.Gilbert"
(PalmerM, a description was never pub-
lished.
WhyGilbert,oncloserinspection,chose
not to formally recognize T. dorothy is
probably because ofits extreme similar-
ityto T.pingeli. Triglopsdorothyiseasily
confused with the latter species, but can
bedistinguished satisfactorilyon the ba-
sisofonly three meristic characters that
overlap slightly between the two species
in the Sea ofOkhotsk (Tables 1-2): the
number ofoblique, scaled dermal folds
belowthelateralline(67-99inT. dorothy
versus 50-87 in T. pingeli), the number
ofdorsolateral scales (18-31 in T. doro-
thy versus 28-37 in T. pingeli). and the
numberofgill rakers (7-12 in T. dorothy
versus 5-9 in T. pingeli). A significant
differenceinthelength ofthe firstinfra-
orbital pore (Fig. 2A) reflects differences
in the overall shape ofthe snout ofthe
two species. In T. dorothy. the snout is
slightly shorter and slightly more ven-
trallyoriented than in T. pingeli.
Triglopsdorothyisknownonlyfromthe
shallow continental shelfofthe southern
SeaofOkhotskbetween Sakhalin Island
and Hokkaido, an area that is thought
tohaveserved as arefugiaforotherspe-
cies duringthe Quaternaryglacial maximum (see Noll
et al., 2001; Brykov et al., 2003). During this period,
glaciers extended to just north ofTerpeniya Bay on
Sakhalin Island and to central Iturup Island in the
southern Kurils, leavingthe southern and central part
ofthe Sea ofOkhotsk ice-free. This region is thought
to have been relatively isolated during this time from
thePacificOcean andtheSeaofJapanbecauseoflower
sea levels that connected the southern coastlines from
Sakhalin to Hokkaido and the southern Kuril Islands
(Kryvolutskaya, 1973; Pietschetal., 2001; 2003). These
land connections may have served to split and isolate
populationsofbenthicmarinefishes, such as T. pingeli.
providing the opportunity for allopatric speciation of
T. dorothy.
PietschandOrr Tnglopsdorothy.anewspeciesfromthesouthernSeaofOkhotsk 245
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loroiggiicnhesofkatihae3f:a4m2i7l-y447Co.tti[dOane,thteracnlsalsastiefidcaftrioonmatnhde T5r4i5g9l4op(sfpeimnagleel,r.18H0UmMmZ),54H5U93MZ(fe5m4a6le7,41(6m4almem,)1,58HUmmM)Z,
RussianbyN.J.WilimovskyandE. Lanz,Instituteof HUMZ 55266(female, 184mm), HUMZ 55267 (female,
Tokra1n9pmFoI9lisv8uisl,.ishasAener.d(uSisCMem.oos.tm,tceiJoU.dnnaftIieercv!aihetbtiruuhnstryiieottolshy.neoo3nfPfo8a.t:cBh6ir.e57fi,i7tcb1-i9is6-woh75la99ot.C.g1eoyrlsuomfobfiThany,ortiVhsaecnrucnsouKvauenrroi,-l mc5(1hm65maa50)lt5,ekm2,am,H)(1U,f55e94Mm°HaZ0mUl0me'M,)5NZ5,,01715H50685U82°m(M60mf80Ze)'m,E(a5f,le5Hem80,aU08l19Mem6,,Z7(m15a7m5l6mJ5e)u0,m,n5me51)H5.10(U9fW7eMme6mm;Zsa)tlH,e5,KU5Ea0Ma1m6s7Z-21t
Tokranov,A.M.,andA.M.Orlov. Kamchatka, 51°25N, 158°04'E, 135-140 m, 21 May
2001. SomebiologicalfeaturesofPsychrolutidaeinthe 1976; HUMZ 55154 (female, 140 mm), Penjinskii Bay.
PacificwatersoffsoutheasternKamchatkaandthenorth- 59''59'N, 160"00'E, 112-113 m, 6 June 1976; HUMZ
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compositionandfeeding. J. Ichthyol. 41:605-614. 157°52'E, 242-245 m, 22 May 1976; HUMZ 56466
YYaabbee,,19MM(O9..S5k,c.hoSor.tpAAsaykeann,Seieawfano.drsmpKeeJ.csai:Ape.msCaJoo.otkftIaics.dhcatuehl)ypo,iln.f,r4Zo2em:s1t7ti-hc2ee0l.suosuotchhwoetsetnesrins m(OHf,kUehm7oMatlJZseuk,n,5e8152016759°174m60;'m(Nm)H,a,lU1eG4,Mi3Zz^1h21i254g'7aEm9,,8m43)61,1(0°f0Hem1m,U'aNlM2,e2,Z1S15e628p00°t81em75mm'5bE),e.(rf9Se14me9a-a7l96eo5;,f
20m0i1s.fesIcsemlitihniu.sfprieotmscthhiesspo.utnhoevr.naKnudrilaArracrheipspeelcaigeos,(SSciogr-- S11e9ptmemm)b.erSe1a97o6f;OHkUhoMtsZk,605750°230('mNa,le1,391°2030'mEm,)1.17Sema,o7f
paeniformes: CottidaeI. Ichthyol. Res. 48:65-70. Okhotsk, 57"38'N, 141^00'E, 121 m, 14 October 1976;
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(Scorpaeniformes: Cottidae), with thedescriptionofa 140mm),HUMZ60810(female, 126mm),HUMZ60811
newspeciesfromnorthernJapanandtheRussianFar (female, 124 mm). Sea ofOkhotsk, 54°54'N, 142°57'E,
East. Ichthyol. Res.48:51-63. 110 m, 6 September 1976; HUMZ 60919 (female, 88
Yabe,19MA8,r4c.ahnidpFeaTl.maiNgloayk(aHCb.ootM.taisduade.a,KI.nATmhaeokfais,heCs.Aorfatghae,JTa.pUayneensoe, mHmU)M,ZGiz6h0i9g4a1,(6f1e°m0a0l'eN,,112558°m41m')E,,H88U-M90Zm6,0795J2un(efe1m9a7l6e;,
JT.apYaonsh.ino,eds.),p.323-330. TokaiUniv.Press,Tokyo, S12e5ptmemm)b,erSe19a76o;fOHkUhoMtZsk,615056°300('mNa,le1,421°2002'mEm,),12H1UmM,Z6
61068 (female, 128 mm). Sea of Okhotsk, 55°09'N,
139°38'E, 124 m, 7 September 1976.