A New Species Of Kolpotocheirodon (Teleostei : Characidae : Cheirodontinae : Compsurini) From Bahia, Northeastern Brazil, With A New Diagnosis Of The Genus

2004·5.4 MB·English

by  L R Malabarba

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PROCEEDINGSOFTHEBIOLOGICALSOCIETYOFWASHINGTON
117(3):317-329.2004.
A newspeciesofKolpotocheirodon (Teleostei: Characidae:
Cheirodontinae: Compsurini) fromBahia,northeasternBrazil,witha
newdiagnosisofthegenus
LuizR.Malabarba,FlavioC.T.Lima,andStanleyH.Weitzman*
(LRM)MuseudeCienciaseTecnologia,PUCRS,Av.Ipiranga6681,90619-900,PortoAlegre,
RS,Brazil,andDepto.Zoologia,IB,UniversidadeFederaldoRioGrandedoSul,
Av.BentoGon§aIves9500,91501-970,PortoAlegre,RS,Brazil,e-mail:[email protected];
(FCTL)MuseudeZoologiadaUSRCaixaPostal42594,04299-970SaoPaulo,SRBrazil,
e-mail:[email protected];
(SHW)DivisionofFishes,WG12,DepartmentofSystematicBiology,MRC-0159,
SmithsonianInstitution,PO.Box37012,WashingtonD.C.20013-7012,U.S.A,
e-mail:[email protected]
—
Abstract. Kolpotocheirodonfigueiredoi,anewspeciesofthecharacidsub-
familyCheirodontinae,tribeCompsurini,isdescribedfromtheupperrioPa-
raguagubasin,Bahia,Brazil.Anewdiagnosisforthegenusisproposed,based
mostlyonscanningelectronmicroscope(SEM)analysesofthecaudalorgan
ofthenewspeciesandthatofthesinglepreviouslyknownspecies,Kolpoto-
cheirodontheloura.Thegenusisdiagnosednowinpartbythepresenceofa
previouslyundescribed,sexuallydimorphicandapparentlyglandular,structure
found in the lower caudal-fin lobe ofmales. The basal relative position of
KolpotocheirodonwithintheCompsurini,inwhichallspeciesareinseminating,
isfurthersupportedbythepresenceofaquasperminbothspeciesratherthan
the apomorphic elongate spermnuclei presentintheremaining membersof
thetribe.
Resumo. Kolpotocheirodonfigueiredoie descrito para aporgao superior
dabaciadorioParaguagu,Bahia,Brasil.Propoem-seumanovadiagnosepara
ogenero,baseadaprincipalmentenaanalisedeMicroscopiaEletronicadeVar-
redura do orgao caudal da novaespecie e de Kolpotocheirodon theloura, a
linicaespecieconhecidaanteriormente.Ogeneroediagnosticadopelapresenga
deumaestruturaaparentementeglandularepreviamentenaodescritadolobo
ventral da nadadeira caudal dos machos. A posigao relativamente basal de
KolpotocheirodonemCompsurini, umatribo depeixes cominseminagaode
Cheirodontinae,ecorroboradapelapresen9adeespermatozoidesaproximada-
menteesfericos (aquasperm) nas duas especies, ao inves dapresen9adees-
permatozoidesdenucleoalongado,comoobservadonosdemaismembrosda
tribo.
ThegenusKolpotocheirodonwasrecent- abarbaetal., 1998)andwasdiagnosedpri-
ly described by Malabarba & Weitzman marilybythepresenceofauniquespecial-
(2000),fromasinglespecies,K. theloura, izedcaudalorganattheproximalregionof
from the uppermost tributaries ofthe rio the ventral caudal-fin lobe ofmales. This
SaoFranciscoandrioParanacentralBrazil, organ consists of hypertrophied elongate
ThegenusisamemberofthetribeComp- dermalflapsattachedalongthefinraysand
surini,subfamilyCheirodontinae(seeMai- aseriesofrelativelyflattabsandpapillae
PROCEEDINGSOFTHEBIOLOGICALSOCIETYOFWASHINGTON
attachedalongtheexposedborderofthose Institutionalabbreviationsareaslistedin
flaps. These structures were unknown in Levitonetal.(1985).Characterpolarityfor
otherinseminatingorexternallyfertilizing thediagnosesofthetwoKolpotocheirodon
speciesofcharacids. speciesandarevisedanalysisofKolpoto-
AtthetimetheresearchofMalabarba& cheirodon monophyly is here established
Weitzman (2000) was conducted, F. C. T. byuseofparsimonythroughare-analysis
Limaandcolleagueswerecollectinginthe ofthecheirodontinecladeCompsurinithat
rioPratinha,atributaryoftherioParagua- was first diagnosed by Malabarba et al.
§u, Iraquara, Bahia, Brazil andtherethey (1998).Thisnewanalysisalsoincludesspe-
discoveredanewcheirodontinespeciesthat ciesofthegeneraSaccoderma, Compsura,
hasacaudalorgansimilartothatpresentin Macropsobrycon, andthespeciesAcinoch-
K.theloura.Thisnewcompsurinspeciesis eirodon melanogramma ("identified" as
herein described and ecological data and "NewGenusandSpeciesB"inMalabarba
fieldobservationsfromthetypelocalityare etal,1998),andKolpotocheirodontheloura
presented. (then"identified"as"NewGenusandSpe-
Datafromthedescriptionofthenewspe- ciesA").
cies, examination of a new collection of
better-preserved specimens ofK. theloura KolpotocheirodonMalabarba&Weitzman
thanoriginallyavailable,andscanningelec-
tronmicroscopy(SEM)observationsofthe KolpotocheirodonMalabarba&Weitzman,
caudal-finstructuresofthesetwospeciesal- 2000:270 (type species: Kolpotocheiro-
lowareanalysisofthecharactersdiagnos- don theloura Malabarba & Weitzman,
ingKolpotocheirodonandredescriptionof 2000:271 bymonotypyandoriginaldes-
theautapomorphiesthatdistinguishitstype ignation).
species.
Commentspreliminarytothediagnosis.—
MethodsandMaterials TnohseedgeinnuMsalKaoblaprobtaocehteailr.o(d1o9n98)wa(sasdNieagw-
The systematic methods for making GenusandSpeciesA)andinMalabarba&
countsandmeasurementsforallspecimens Weitzman (2000)bythepresenceofthree
studiedherearethesameasthosedescribed apomorphic features thatoccurinitstype
andusedbyMalabarba&Weitzman(1999) species. These characters, as describedby
andarenotre-describedhere.However,un- MalabarbaandWeitzman(2000),areaspe-
liketheconventionforfinrayswhereinthe cialized part ofa caudal organ located at
countfortheraysfortheholotypeisgiven theproximal regionoftheventralcaudal-
firstfollowedby therangeandmean sep- finlobeofmaturemalesandconsistofhy-
aratelyfortheunbranchedandthebranched pertrophiedelongatedermal flaps attached
rays,thecountsofjawteethdonotreport alongthefinraystogetherwithaseriesof
asinglevaluefollowedbyanindicationof relatively flat tabs and papillae attached
variation. Instead, only the range of the alongtheexposedborderoftheseflaps(=
counts, for example, maxilla with 2 or 3 character36inMalabarba1998);hookson
teeth, is provided. This is because we are theanal-finraysofmaturemalesdistributed
confidentonlyincountstakenfromcleared along the most posterior unbranched and
and stained specimens. SEM photographs fiveanteriorbranchedanal-finrays(=char-
were taken from specimens fixed in for- acter 30 in Malabarba, 1998); and the
malinandpreservedin70%ethanol.Before twelfth and thirteenth caudal-fin rays are
metalizationwithgold,thefinswerepassed dorsally concave along their basal halves
through 99% ethanol, then acetone, and and have ventrally expanded segments(=
treatedwithacriticalpointdryer. character34,state2inMalabarba 1998).
VOLUME117,NUMBER3
Fig. I. SEMofcaudalorganinKolpotocheimelonfigiieiredoi,male(MZUSP55219,25.5mmSL),from
rioPratinha,Iraquara,Bahia,Brazil.(A)lowercaudal-finlobe;(B)detailedimageshowingthesmoothborder
oftheflapattachedalongthebasalportionofthenineteenthcaudal-fin;(C)and(D)detailedimagesofthe
pineappleorgansoftheventrallobeofthecaudalfin.
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Diagnosis. By using SEMthespecial- function, possibly pheromone in nature.
izedcaudal-finorgandescribedinthepre- Thispineappleorganhasnotbeenfoundin
vious diagnosis of Kolpotocheirodon is othercheirodontinesorothercharacids,and
now found to be more complex than for- itspresencesupportsahypothesisofclose
merlyknown. Anew caudal organ, previ- relationshipbetweenthetwoKolpotochei-
ouslyundescribed,correspondstoasecond- rodonspecies.
arysexuallydimorphicorganfoundexclu- Both Kolpotocheirodon species have a
sivelyintheventrallobeofthecaudalfin conspicuous small black spot at the mid-
ofmalesofbothKolpotocheirodonspecies. lengthofthefirstbranchedanal-finrayof
This "pineapple-like" organ is easily rec- males(Figs.5,7and8),absentinfemales
ognized by its peculiar shape, somewhat (Fig. 6). Suchaspotisabsentinallother
coneshapedorpapilla-like,butcompletely knowncheirodontines.Itishereconsidered
coveredby smallerpapillae-likebodiesor derivedandasynapomorphyforthegenus.
knobs(seeFigs. 1,2).Thesearedistributed Males of Kolpotocheirodon figueiredoi
amongthelargepapillaeofthecaudalfin andK. thelourahavetheventralbodysur-
ofmales ofK. theloura (see Fig. 3), but face in the area covering the pelvic bone
form the entire caudal-fin organ in K.fi- withadarkbrownmark,nearlyintheshape
giieiredoi(seeFig. 1).Thisorganisfound ofan isosceles triangle. This pigment ap-
only in adult males ofboth species, sug- pearstoexternallydelineateanareacorre-
gesting that it may have a reproductive spondingtothemusclesinsertedonthepel-
PROCEEDINGSOFTHEBIOLOGICALSOCIETYOFWASHINGTON
Fig 2 DetailedSEMimagesotthepineappleorgansfoundbetweenthepapillaeoftheventrallobeofthe
caudal-hnrayotmalesinKolpotocheiiodontheloura MNRJ18081,SL262mm,fromlagoaPertaPe,noSao
Fianciscodrainage,Palmital,MinasGcrais,Brazil (A)bai =50|j,m,(B)bar=20|xm
viebone(Fig. 8). Suchaspotisabsentin eachgenus.Note:Menezesetal.(inpress)
all other cheirodontines, and constitutes a and Weitzman et al. (in press) have de-
synapomorphyforthetwospecies. scribedanddiscussedglandularsofttissue
Malabarba&Weitzman(2000)described intheanalfinsofsexuallyactivemalechar-
thepresenceofwell-developedhooksalong acidsofmanykindsincludingglandulocau-
the last unbranched and five anterior dines, and some non-glandulocaudines.
branchedanal-finraysofmalesasderived, This tissue is most often associated with
and diagnostic for Kolpotocheirodon (= anal-fin hooks, but in one species aglan-
character30inMalabarba 1998).Thenew dularorganwasfound.
specimensavailableofK. theloura, MNRJ
18081,havethelastunbranchedandfiveto Kolpotocheirodontheloura
seven anterior branched anal-fin rays of Malabarba&Weitzman
malesbearinghooks(5branchedraysin7 Fig.7
tbihmmsfsaitairnhnnhnropplooooaeegedeoaoudrl6ccinnnkooe-licigfpctdsnfamomoisshihvenrteaneepea(sndtntrdedr5puisilhrscieanato)e,asicbtcrni.bgboamirgoofiinr6eeeatmlnaoaoosMnrinlpnfnnsatit-csfclnnyii;flbuhsstoyhiueereshtnu2emas8uiddeh3nderibniadsfrinseirnof,afasnn,iarfeynnpgiypar.SsndesorbeoKowhcensn.fAfihcobtiveoelimonoicinaffvtetfnkoctoeishn6haurshgdmplsositoueea-n,oulseaKmfrclhyplgioiitmsesareahnlhtmisaodevcnphexeedireadonscsoatmratb)ohscyieneo7e.nsprohftnmca.teiwoehosahraicbTrioua,ieniWiieihknnvninnedoeesgde-6-3sgs,r MKaotclcMPhWrMm2S(eaaeocoap7eaLNrlualoxo3ilimduakidRt-t2aiadatsrlo44zJtallbia)lc,a.l-moi;aahl-f3ennarr,fei1-xasfynibnii822gha,nlga0r7e7itm.a—8.8oep.ilgeh1,s4d2o2n&oto,)ob0e-haom;en)0d3fk)m,P1i:0Ws;g2e)3:.)ef(tr75,iA;2phitg22lae.7a7t,l7sBrl12rz78apoP7-ats,mfmeu(z6(2yips,rpai,pgp8e.alr.eenfr1c,elisifgivm()oiM1.Mima(1;g(aci.Sxe01d(-lni96an2efhala95eso7siolb9xsnc5la(aF)lra,(oahrirGi,rmvtnosyaepbiefaytonrtinnaleppkcagit-dael.esiiorfdnu))ssnaii&,4d.s;-,;nln
among compsurins are anal-fin hooks re- Diagnosis. Kolpotocheirodon theloura
strictedtotheanterioranal-finrays,inthe isdiagnosedfromthenewKolpotocheiro-
lastunbranchedandfouranteriorbranched donspeciesandothercharacidfishesbythe
rays. By parsimony both conditions are followingautapomoiphies:Asdescribedin
consideredderived and autapomorphicfor Malabarba&Weitzman(2000),K.theloura
VOLUME117,NUMBER3
Fig.3. SEMimagesofcaudalorganinKolpotocheirodontheloiira,male,MNRJ18081,SL26.2mm,from
lagoaPertaPe,rioSaoFranciscodrainage,Palmital,MinasGerais,Brazil.(A)lowercaudal-finlobe,bar=600
oIJfimre;l(aBti)vedleytafillaetdtiabmsagaeloonfgthitesfelxappoastetdachboerddearl,onbgarthe=ba1s0a0lipxomr;ti(oCn)ofdetthaeilneidneitmeaegnethocfautdhael-fflianpsraayttwaicthhedatsoertihees
eighteenththroughthirteenthorfourteenthfin-rayswithasingleseriesofpapillaealongitsexposedborder,bar
=200|jLm.
hashypertrophiedelongatedermalflapsat- or fourteenth fin-rays are relatively short,
tachedalongthefinraysoftheventralcau- narrowandbearpapillaeinasingleseries
dal-fin lobe of males (= character 36 in along its exposed border (Fig. 3A, C).
Malabarba 1998). The flap attachedalong These modified flaps ofthe caudal organ
thebasalportionofthenineteenthcaudal- arenotexclusivetomales inK. theloura,
fin ray has a series ofrelatively flat tabs beingalsofoundinfemales,althoughless
alongitsexposedborder(Fig.3).Theflaps developed(Fig.4).Modifiedflapsarealso
attachedtotheeighteenththroughthirteenth observed inthe dorsal fin ofmales ofK.
PROCEEDINGSOFTHEBIOLOGICALSOCIETYOFWASHINGTON
Fig.4. DetailedSEMimagesoftheflapsbearingpapillaealongthebasalportionoftheventrallobeofthe
caudal-finrayinKolpotocheirodontheloura,female,MNRJ18081,SL26.5mm,fromlagoaPertaPe,rioSao
Franciscodrainage,Palmital,MinasGerais,Brazil.(A)bar=500\x,m;(B)bar=200|xm.
theloura(Fig.9).Thesemodifiedflapsare stripe between pseudotympanum and area
independent of the sexually dimorphic ventraltodorsalfin(Fig.7).Suchbarsare
pineapple-likeorgansdescribedasasyna- absentinthenewKolpotocheirodonspecies
pomorphy for Kolpotocheirodon and are and in othercompsurins and representan
absentinK.figueiredoi.Themodifiedflaps autapomorphyforK. theloura.
constituteanautapomorphyofK.theloura.
AsdescribedinMalabarba&Weitzman Kolpotocheirodonfigueiredoi
(2000),thetwelfthandthirteenthcaudal-fin newspecies
rays of K. theloura are curved, dorsally Figs.5,6
concavealongtheirbasalhalves,andwith
v3e4n,trsatlaltye2exipnanMdaeldabasrebgam,en1t9s98()=. Tchhiasracfteea-r mmHoSlLo,tyBpreaz.i-lM,LB\aJhSiVa,7I0r0a3q7u,ara1,mrailoeP,ra3t0i.n5-
turewasnotobservedinK.figueiredoiand ha, Fazenda Pratinha (12°21'13"S;
is considered autapomorphic for the type 41°32'57"W), 17-21Dec 1998;RGerhard,
species,K. theloura. FC.TLima,—FDiDarioandL.S.Rocha.
Kolpotocheirodonthelourahas3-5very Paratypes. All specimens collected
small vertical bars crossing lateral body with the holotype: MCP 22345, 3 males.
Fig.5. Kolpotocheirodonfigueiredoi.i Vspecies,holotype.
ha,Iraquara,Bahia,Brazil.
VOLUME117,NUMBER3
Fig.6. Kolpotocheirodonfigueiredc ipecies,paratype,female,MZUSP55219,SL30.0mm;rioPra-
tinha,Iraquara,Bahia,Brazil.
25.1-30.5 mm SL, 2 females, 24.0-24.8 followingcharactershavealternativestates
mm SL. MZUSP'55219, (26) 14 males, betweenK.figueiredoiandK.theloura,but
24.2-28.2 mm SL, 8 females, 24.0-31.0 thesealsooccuralternativelyamongother
m26m.9SmL;m(S1LmaAlleiza2r8i.n2rmedmsSaLndanAdlc1iafnemballuee cmoomsptspuarrisnimsopneicoiuess.lyNaecvceerpttheedleesisthetrheaysaaur-e
msStaLaliesneec2dt6i.so2pneemcdimfmoerSnhsLisctaloneldaorge1yd;fe1wmimatalhleet2r2y65p..s45inmm;mm1 mtaoprWophmheoirrcepahfsoircmKa.lfoetrsheolKfo.uKr.fai.fgiugeuierierdeodioiohraveapnoo-
SLsectioned—forTEMstudy). hooks onthe caudal-finrays, whilemales
Diagnosis. Kolpotocheirodon figueire- ofK. thelourahavethetwelfthtothefour-
doi lacks all autapomorphies described teenthorfifteenthprincipalcaudal-finrays
aboveforK.theloura,buthasnounambig- bearing4-6retrorsehooksoneachsidein
uousautapomorphiesforitsdiagnosis.The a row along their dorsal divisions (Mala-
^sk-
Fig.7. Kolpotocheirodontheloura,male,MNRJ18081,SL25.0n aPertaPe,rioPalmital,Minas
Gerais,Brazil.
PROCEEDINGSOFTHEBIOLOGICALSOCIETYOFWASHINGTON
^^K^^Si^^^^
Fig.8. Kolpotocheirodon.figueiredoimale,MCP23455,SL30.5mm.(A)Ventralbodysurfaceinthearea
coveringthepelvicboneshowingadarkbrownmark,nearlyisoscelestriangleshape,apparentlyexternally
delineatingtheareacorrespondingtothemusclesinsertedinthepelvicbone.(B)and(C)Leftlateralviewof
thedorsal(B)andanalfins(C),showingthedarkspotsofthosefins.
barba&Weitzman,2000:fig.4).Thepres- (Malabarbaetal., 1998)indicatedthepres-
enceofhooksonthecaudal finisknown ence ofcaudal-finhooks as a synapomor-
forseveralcompsurins,includingAcinoch- phyforthecompsurincheirodontines,and
eirodonmelanogramma (hooksoncaudal- itsabsenceasecondaryreversalinsomeof
finrays 13-14,rarelyonray 15),Saccod- itsspecies.Theinclusionofanewspecies
ermahastata(hooksoncaudal-finraysIS- bearingnohooks inthemostbasalgenus
IS), "Odontostilbe'' dialeptura (hooks on ofthetribeallowseithertherecognitionof
caudal-finrays 12-16), andMacropsobry- thepresence ofhooks as asynapomorphy
conuruguayanae(hooksoncaudal-finrays forthetribeCompsuriniwithareversalin
12-14, plus several spinelets along the K.figueiredoi, orthe recognition ofinde-
proximalhalfofcaudal-finrays 14to 18). pendent acquisitions ofhooks in K. thel-
However, hooks are absent in Compsura ouraandinthecladeincludingtheremain-
heterura, Compsura gorgonae, and "OJ- ingcompsurins.Thefirsthypothesisispre-
ontostilbe''mitoptera.Malabarba&Weitz- fened,sinceitbetterconformswiththepu-
man(1999,2000)pointedoutthatalthough tativehomologyofcaudal-finhooksamong
thesehooksarepresentontheventrallobe compsurins(dePinna 1991).
ofthecaudal fin inall these species,they MalesofK.figueiredoihaveaconspic-
donotalloccuronthesamecaudal-finrays uoussmallblackspotinthesofttissuebe-
inallspeciesandareofdifferentshapes.A tween midlength offirst and second, and
previous analysis ofcharacterdistribution second and third branched dorsal-fin rays
VOLUME117.NUMBER3
Fig.9. DetailedSEMimagesoftheflapsbearingpapillaeakir rdorsal-finrayinKolpotochei-
rodontheloura,male,MNRJ18081,SL26.2mm,fromlagoaPei 1Pe,rioSaoFranciscodrainage,Palmital,
MinasGerais,Brazil.(A)bar=500(Jim;and(B)bar= 100(jim.
(Figs. 5, 8). This is absent (Fig. 7) in K. ofthedorsalblackspotasasynapomorphy
theloura (= character 65 in Malabarba forthetribeCompsuriniwithareversalin
1998).Amongcompsurins,asimilarspotis K. theloura, ortherecognitionofindepen-
observedinspeciesofCompsura,Macrop- dentacquisitionsofthedorsalblackspotin
sobryconandAcinocheirodon,butisabsent K.figueiredoiandinthecladeincludingre-
in species ofSaccoderma, This spot was mainingcompsurins.Thefirsthypothesisis
previouslyproposedasasynapomorphyfor preferredbecauseitbetterconformstothe
acladeincludingthelastfourgeneracited putativehomologyofthedorsalblackspot
above. Again,the inclusionofanew spe- amongcompsurins.
cies in the most basal genus ofthe tribe A furthercharacterdistinguishingK.fi-
allowsboththerecognitionofthepresence gueiredoi is its caudal-peduncle/caudal-fin
Table1.—MorphometriesofKolpotocheirodonfigueiredoi,newspecies.Standardlengthisexpressedinmm;
measurementsthroughheadlengtharepercentagesofstandardlength;thelastfourentriesarepercentagesof
headlength.Rangeincludestheholotype,MZUSP70037,andparatypesMCP22345,MZUSP55219.
Males Females
"isr „ Low High X SD ^ Low High SD
Standardlength(mm) 30.5 13 25.5 30.5 27.0 10 23.8 31.0 26.4
Snouttoanal-finorigin 59.7 13 57.3 63.0 60.0 1.30 10 61.6 65.8 63.7 1.35
Snouttodorsal-finorigin 50.5 13 45.7 52.9 49.5 1.80 10 48.1 52.4 49.7 1.35
Snouttopelvic-finorigin 42.0 13 42.0 46.7 44.5 1.51 10 45.3 48.3 47.3 1.17
Dorsal-finbaselength 13.1 13 12.1 15.6 13.6 1.20 10 11.9 14.3 13.2 0.72
Anal-finbaselength 25.9 13 24.5 29.0 26.8 1.27 10 23.5 26.8 25.4 1.08
Caudalpedunclelength 15.1 13 12.5 15.5 14.1 0.90 10 11.1 15.4 13.6 1.25
Caudalpeduncledepth 13.8 13 13.6 16.1 14.8 0.76 10 11.3 13.8 12.5 0.83
Depthatdorsal-finorigin 31.1 13 31.1 35.7 33.5 1.13 10 33.6 39.4 35.4 1.86
Dorsal-finheight 29.8 12 26.1 29.8 28.0 1.36 9 25.2 27.5 26.4 0.83
Pelvic-finlength 19.0 13 16.7 19.3 18.2 0.71 10 13.4 16.3 14.6 0.79
Pectoral-finlength 19.3 13 17.5 20.1 18.8 0.77 8 15.7 19.2 17.4 1.40
Bonyheadlength 23.3 13 23.3 25.2 24.6 0.59 10 23.0 25.0 24.1 0.72
Snoutlength 21.1 13 20.3 23.8 22.0 1.10 10 19.4 23.3 21.8 1.35
Upperjawlength 31.0 13 25.4 31.0 27.7 1.47 10 22.7 30.2 27.9 2.13
Horizontaleyediameter 38.0 13 33.3 38.5 36.2 1.39 10 34.8 39.7 37.3 1.68
Leastinterorbitalwidth 29.6 13 26.1 31.1 28.9 1.31 10 25.4 29.5 28.1 1.32
PROCEEDINGSOFTHEBIOLOGICALSOCIETYOFWASHINGTON
spot (Figs. 5, 6) thatis moreorlessrect- 7orfewercusps.Teethposteriortosecond
angularinshapeandhorizontallyelongate. toothasymmetricalwithmostlateralcusps
Itneverreachesthedorsalandventralbor- situatedtowards toothbase andmostme-
dersofthecaudalpeduncle.Thesamespot dialcuspmoredistallylocated.Cuspssmall
is vertically elongate, lozenge-shaped and and regular and approximately equal in
reachesthedorsalandventralbordersofthe size.Cuttingedgeslightlyarchedtoalmost
caudalpeduncl—einK. theloura(Fig.7). straight.
Description. Morphometric data sum- Dorsal-finrays,ii,9,n=22(ii,8inone
marized in Table 1. Body elongate and specimen). First unbranched ray less than
compressed,greatestdepthatdorsal-finor- half-lengthofsecond.Dorsal-finoriginap-
igin.Predorsalprofileslightlyconvex.Pro- proximatelyatmidlengthofbody.Adipose-
file ofbody along dorsal-fin base poster- fin origin nearly at vertical throughinser-
oventrally inclined, nearly straight from tionofposteriormostanal-finray.
base ofposteriordorsal-finraytoadipose Anal-finrays,iii, 18,(iii-ivX=3.5,16-
fin.Ventralbodyprofileconvexfromtipof 19, X = 17.5, n = 22). Anal-fin origin
lowerjawtopelvic-finorigin.Musclescov- slightlyposteriortoverticalpassingthrough
ering pelvic bone strongly prominent in baseofposteriormostdorsal-finrayinfe-
ventral body profile, especially in males. malesandataverticalpassingthroughbase
Areabetween pelvic- and anal-fin origins oftwolastdorsal-finraysinmales.Distal
slightly concave in females and notably borderofanalfinconcaveinfemales,with
concave in males, with apairofconcavi- anterior5-6branchedraysverylong,form-
ties, separatedfromeachotherbyasmall ing prominentanteriorlobe. Distalborder
mediankeel visible only whenpelvic fins ofanalfinofmalesconvexintheanterior
movedoutofway.Theseconcavitieslodge lobe, decreasing in length gradually and
pelvicfinswhenlaterretracted.Ventralpro- forming aposteriorconcaveborder. Anal-
filealonganal-finbaseslightlyconcavein finraysofmaleswithslender,elongatere-
females. In males same profile, slightly trorsehooksonlongestunbranchedray,and
convexinregionofanteriorlobeandslight- anteriorfirst5or6branchedrays(scattered
ly convex along remaining posterior fin hookspresentinbranchedrays7and8in
portion.Dorsalandventralprofilesofcau- one specimen). Hooks insertedatpostero-
dal peduncle nearly straight in females. lateralborderoffinrays,bentoverlateral
Dorsal and especially ventral surfaces of surface offin ray and anteriorly directed.
caudalpeduncleofmalesconvex,withan Hooks located on posterior ray branches,
internaltranslucentcavity,coveredbycau- less numerous on anterior ray branches.
dalpedunclescales. One, rarely two, bilateral pair of bony
Head small. Snout shorterthan eye di- hooksperraysegment.
ameter.Mouthterminal.Maxillashort,po- Pectoral-finrays,i,9(i,8-9,X= 8.6,n
sitionedatanangleofapproximately45de- = 22). Distal ends of longest rays not
greesrelativetolongbody axis. Posterior reachingpelvic-finorigininfemales;reach-
tipofmaxillareachingverticalthatpasses ingornotinmales.Pelvic-finrays,i,7(1,7
throughanteriorborderofeye. inallspecimens,n = 22).Pelvic-finorigin
Premaxilla with 4 teeth, each having 9 anteriortoverticalpassingthroughdorsal-
small evenly spacedcusps allaboutequal fin origin. Distal tip ofpelvic fin passing
insize. Cuttingedgearched. Maxillawith anal-finorigininmales,butnotinfemales.
2or3teethsimilarinformtothoseofpre- Male pelvic fins bearing elongate ventro-
maxilla, with 7-9 cusps. Cutting edge medialretrorsehooksalongbranchedrays
slightly archedto almoststraight. Dentary 2to8.
bone with 4 or 5 large teeth each with 7 Principalcaudal-finrays10/10(10/9,but
cusps;followedby2or3smallerteethwith 10/10and9/9,inonespecimeneach,n =