Biometric Analysis of Nautilus pompilius from the Philippines and the Fiji Islands

Full text

From:NAUrLUS
Editedbyヽ颯BruceSallndersandNeilH.Land■nt
(Plellumhbb価 電c∝pOntiOn,1987)
Chapter7
BiometricAnalysisofNα utilus
pompiliusfromthePhilippinesandthe
Fililslal■ ds
KAZUSHIGETANABEandJUNZOTSUKAHARA
llntroduction
2 ヽ4aterialsandMethods
21Fieldヽ lethods
22LaboratowMe■ OdS
3Results
31AverageRelativeCrowth
320ntogeneticChangeofShellForm
33NepionicShells
340onadDevelopmentandSexualDimorphism
4Conclusions
105
106
106
106
108
108
110
110
111
112
1.Introduction
SinceMoseley(1838)firstanalyzedthegeometryoftheNα utilusshell,itslnor―
phO10gicalfeaturesandallometricgrowthhavebeenstudiedbymanyauthors
(ThOmpson,1917:HuXley,1932,Hirano,1977:HiranoandObata,1979;Hirano
etal.,1980)MostpreviousM′ ork、vasbasedonsmallcollectionsofindividual
specimens,thesexesanddetailedlocalityrecordof、vhichM′ ereunknoM′ nln
1981and1983,aresearchprOjectontheecologyofNpompiliuswascarriedout
inthePhilippinesandtheFililslandsasajointventureofJapanese,Philippine,
andFijianscientists(undertheleadershipofProfSHayasakaofKagoshima
University)Duringthisproject,weexaminedthebiometricpropertiesofsoftand
hardtissuesincapturedanimals(Hayasakaet α
l,1982;Tanabeet αl,1983,1985,
Tsukahara,1985)ThisChaptersummarizesourworkonNpompilius,withspe‐
cialreferencetosexualdimorphismexpressedinallometricrelationshipsand
thevariationintheshellandsoftpartsinthetM′ opopulationsfromル videlysep‐
aratedareas
KAZUSIⅡ GETANABE ●
Ceologicallnstitute,FacultyofScience,UniversityofTokyo,Tokyol13,
JapanJUNZOTSUKAHARA ●
DepammentOfBiology,FacultyofScience,KagoshimaUniver‐
sity,Kagoshima890,Japan

106
2.Materillζ よIaMetlodし ,
ChapteF7
ThischapterisbasedmainlyonthreecollectionsofNpompiliusfromthe
PhilippinesandtheFijilslandsOnecollectionconsistsof52specimensthat
werecapturedintheTa■ onStait(depthOf120‑3101n)abOut2‑3knlo∬ Bindoy
Village,NegrosOriental,thePhilippines,in1981byHayasakaandothers(forthe
actualcatchrecords,seeFig3andTable10inHayasakaet α
l,1982)Theother
露
Lピ￨=l路 鶴
理
嵩
‰肝
1:生∬
器
鷲覇
Iぶき
r版lf∬IF:1黒11:
ml腑￨卜 景
電鳳子
器11辞 ユ
WT蹴詣ゴi響:思鷺L器
details,seeTanabeet αl.,1985)ヽ 〔eaSurementsbyTanabeet α
I(1983)onthe
shellsof31specimensofNα utilusfrOmtheSuvaarea(collectedbyHayasakaand
Shinomiyain1981fromalmostthesamelocationsasthoseofthe101specimens
mentionedabove)M′ erealsousedinthisstudyThePhilippinesandtheFiji
lslandsrepresentthenorthMresternandsoutheasternmarginalareasoftherange
ofNpοmpilius(SeeFiglinChapter3),therefOre,thespecimensfromthesetwo
sitesareespeciallysuitableforanalysisofgeographicVariation
2.1.FieldMethods
Allthespecimensstudied、 verecapturedusingbaitedtrapS,M′ hichwere
spedttydedgnedbriemskbyPhh:IILTl認 ■
灘}識:l鑓1鶴盟■
eachanimal、 vaslabeled,、 veighed,Ine
器載
3T,t離 態
群:ξ∫
繋
e轟潔Wfκ∬
寵樵∫盤lIIttl堪∬
霊を
releasedintheirhabitatafterlaboratoryobservationM√ asdeterminedbythepres―
胤∫
満
Ⅷ
∫献
棚艦
]IL恥:鶴 計鵠
崖
li賜糧
蠍
鍛
鍔
二
誓
saundersandSpinosa,1978)ヽ lⅨimumshelldiameterandaperturalwhorl
breadthandheightweremeasuredoneveryspecimenwithaslidecaliper〔 ac
curacy=005mm)ExceptforonespecimenflomthePacificHalborarea,alll市 e
animalswereweighedusingadialscale(aCCuracy± lg〕 Thesemeasurements
wereusedtoanalyzeaveragerelativegrOン ,thandsexualdimorphismlnaddition
tothesebasiccharacters,freshgonadsfron■ somespecimenscOllectedinboththe
PhilippinesandtheFililslandsM′ ere、veighedtodeterminetherelationshipbe―
t、veensexuallnaturityandshellsize
2.2.LaboratoryMethodS
ヽ4easurementsfromallthespecimenscapturedwereanalyzedtoinvestigate
theallometricrelationshipsamongcharactersineachsampleandthevariation
intotal、veightandshellsizeRelativegro、 vthandontogeneticchangesinshell
formwereexaminedintheshellsdf28animalsfromthePhilippinesandtheF"i
lslandsTheseshellswerecutandpohshedalongthemedianplane,andthe

BiolnetricAnalysisofNα u[iluspompi]ius 107
FigurelBasicmorphology,orientation,andmeasurementsof■ eshellofNpompiliusinmedian
(1け〕
anddorsalvental(■ gllt)sectiOnsseetlletextfordiscussionReprintedwlthpermissionf■ om
Tanabeetai(1985)
nepionicshellsize、 ν
aslneasuredEach half''shell、 vasalsocutalongthelon―
gitudinalaxisofthececumonthemediansectiontoproduceadorsal― ventral
section(Figl)Shelldiameter(D),half‐ lengthOfthe、vhOrlbreadth(B/2),、 vhOrl
height(H),diStancefromcoilingaxistoumbilicalshoulder(C),andinnerwhorl
height(F)weremeasuredonthehalveddorsal― ventralsectionsatintervalsof
180° ヽleasurementsofmediananddorsal― ventralsections、veremadev′ itha
profileprOjector(NikonV‑12),attaChedtoadigitalmicrometer〔 accuracy± lμ
m)
Onthebasisofthesemeasurements,theshell― fOrmratiosproposedbyRaup(1966)
andChamberlain〔 1976),suchas、 vhOrlexpansionrate[(R′ Th1〕 2,n:V′ horinum‐
ber,sincen>2πl,diStanceoftheM′ hOrlstothecoilingaxis〔 C/R),relatiVewhorl
thickness(BH),andflankposition(F′ 1)),ヽVerecalculatedforeachspecimenat
severalgro、 vthstages.ThefolloM′ ingabbreviationswereusedinthebiometric
descriptions:
=Samplesize
=Arithmeticmean
=Coefficientofvariation
=Standarddeviation
=Observedrange
Ｒ
Ｎ・ ＸＶ ｓＯ
nepiOnic
constriction

108 Chapter7
Ox
α
Oα
=Standarderrorofthesamplemean
=Slopeofallometry(grOWthratio)
=StandarderrorofthegroM√ thratio
=Correlationcoefficient
=Probabilitylevel
202122
LOG(SHELLOInHETER〕
202122
LOG(SHELLDInHETER〕
ｒＰ
3.Results
3.1.AverageRela●veGrowth
Weappliedbivariateanalysis(Imbrie,1956)tothiSSpeciestoexpressthe
allometricrelationshipbetlt7eenapairoflnorphologicalcharactersTherelation‐
shipbet、 veen、νhorlbreadth〔 B)andShelldiameter(D)andbetWeenwhorlheight
(H)andShelldiameter(D)forfemaleandmalespecimenstromtheSuvaareais
representedondoublelogarithmicplots(Fig2)Thes10pes(growthratios)ofthe
reducedm● oraxesofBvsDinmalesandfemalesbothindicateslightlynegative
a1lomet理 ァ
,、vhilethoseofHvsDindicateisometryAccordingtoKermackand
Haldane(1950),thedifferenceintheslopesofthetworeducedmajoraxesis
evaluatedbycalculatingthestatisticz:
Z=(α .― α
2)/V′ σ
る
1+σ晨
2
、vhere α
landα
2(α■
>α2)arethes10pesoftheaxesand σ
α
,andσ
α
2arethe
standarderrorsofthetwoslopesUsingthismethod,v′ ecalculatedzvalues
betweenthetworeducedmaloraxesofthemaleandfemalesamplesfromthe
SuvaareaThezvaluesbetweenfemalesandmalesintheSuvasample〔 027for
logBvslogD:056forlogHvslogD)aremuChSmallerthanthecriticalvalue
196atP=005:hence,theslopesinbothsexesfromthesameareadonotshow
21
︵エトロＣ口にめ コと０工
︐一 一０コ
︵Ｆ工０¨ロエ コに０󰿯󰿯︶ ００Ｊ
Figure20oublelogarithmicscatterplotsWithreducedmaioraXesshowingthea1lometricrelationship
bet、veen、 vhorlbreadthandshelldianleter(lel)andbetween、 νhorlheightandshelldiameter(right)
inmaleandfemalespecimensofNpompi]iusfromtheSuvaareaoftheFttilslands
′
・0,74 ■■0111 ■ 0946呟・
O079

・Phippines(Taヽ onAreaN・ 52)
Lo9B‐ 1つ 4LogD― o398
｀
｀
Fil(SuvaAreaN・ 101)
LogB・ 0785Lo90+0167
■ 088%手 0037
2122
LOG(SHELLOinHEIER〕
BiometricAnalysisofNα utiluspOmpilius
Figure4Doublelogarithmicscatterplots
withreducedmaloraxesshowingtheal
lometricrelationshipbetweentissue
weightandshelldiameterinthesamples
(maleSandfemaleJOfNpompiliusfrom
theFijilslandsandthePhilippines
‐ 0805,%‐ 0398
109
＾■卜０こ口∝ω コ∝０エコ︶ ＯＣコ
一トエ０︻ロエ コビ０工󰿯︺ ００コFl,(SuVaAreaN‐ ヽ01)
Lo9H・ 1022Lo9D― o231
・ o969,τ
・0025
17:、 20212223
LOC(SHELL010HETER)
Fittre3.Double10garlhllicscatteFp10tswithreducedmaloraxesshowingthea1lometicrelationship
betweenwhorlbreadthandshelldiameter(left)andbe■ Veenwhorlheightandshelldiameter(right〕
inthesamples(malesandfemales〕 ofNpompiliusfromtheFijilslandsandthePhilippines
anystatisticallysignificantdilferenceSimilarly,malesandfemalesfromtheSuva
areashownosignificantdifferenceintheslopesoftheirreducednlaloraxes
bet、 veenpairsoflogB,logH,and10gDTherefore,thelineartrendsofthereduced
maloraxesbetweenpairsofmorphologicalcharactersinthesamplesfromthe
SuvaandPacificHarborareasofFijiareessentiallyequivalent.
Ontheotherhand,cOmparisonsbetv′ eentheFijianandPhilippinecollec‐
tionsrevealwidedifferences(Figs3and4).Asignificantdifferenceoccursin
thes10pesofthereducedma,oraxesbetweenthetwOsamplesforwhorlbreadth
,PHI[IPPINES(TANONAR[AN・ 34)‐
‐ 0863 咋0297.
︵ 卜 ■ ０ ︼ 口
︐ ＵＤ ０ 〇 一 Ｆ一 〇 ０コ
202122
L00(SHELLDInMElER,
・Phlppines(TanonAreaN・ 33)
201、

1■ 0Chapter7
EVEOAMFTERH000LENCTH
0。
0 ぉ
8。 。
。
αШ卜ｕΣく﹃ｏ 回＞ｕ
TANoNSAMPLE
FI」 ISAMPLE
。MALE
oFEMALE
▲MALE
△
FEMALE
い
８０ ∞
Figure5Dollblescatterplotssho、 villgthea110metricrelatiollshipbetweeneyediameterand
lel■ghinthespecimensofNpompiliusfromtheTa● On,PhilippinesandSuva,Fttiareas
andwhorlheightvsshelldiameter(Z=3.46forlogBvs10gD;z=1623for
logHvs10gD;z>2.58=Zp‑0。 .,wherePく 001iSsignificant)Inaddition,
differencesoccurintherelativegowthpatternofeyediametervshoodlength
(Fig5)
3.2.OntogeneticChangeofShellForm
Ontogeneticchangesinshell‐ formparameters、 vereexalninedin21speci―
mens〔15malesand6females)frOmtheSuvaarea(Fig6)FlankpOsitionand
thedistanceofthe、 νhOrlstothecOilingaxisremainnearlyconstant,whereas
relativev′ hOrlthicknessand、 vhorlexpansionratedecrease、 vithgrOwth.Inthe
earlystagesofdevelopment,malesandfemalesshO、 vnosignificantdilferencein
theseshell― formparameters,butinthematureOralmostmaturestages(55■ ),
maleshavelargervaluesofrelativeM′ hOrlthicknessandMrhOrlexpansionrate
thandofemalesTheselargervaluesindicatethatmaturemalesusuallyhave
larger、vhorlradiiandwhorlbreadthsthallmaturefemales
3.3.NepionicShells
Themeannepionicshellsizeandits95%collfidenceintervalin3Fijian
specimens(2750± 204mm,s=244mm,V=886,andOR=244‑326
mm)areshghtlylargerthanthosein6specimensfromthePhilippines(25.72±

111
Figure6Twoseriesofdoublescaterplotsoftheshell‐ formparametersofRaup{1966)inselected
specimensofNpornpiliusfromtheSuvaacaofFttishOWingtheontogeneticchangesinshellform
betweenmalesandfemalesReprintedwithpenllisslonfromTanabeeta:(1985〕
104mm:s=o99mm,V=3.84,OR=247‑271mm)AStudent'st― test,
however,doesnotshowastatisticallysignificantdifference(P<005)inthe
meannepionicshellsizebet、 veenthet,vosamplesFromtheoxygenisotopic
compositionofNα utilusshells,Cochranetal(1981)andBE.Taylorandや Vard
(1983)concludedthatthenepionicconstrictionformsathatchinglfthisOpinion
iscorrect,theforegoingdataprobablyreflectthesimilarityinthesizeofthe
postembryonicyoung.
3.4.GonadDevelopmentandSexualDimorphism
Thehypothesisthatvariationintotallive、veightandshellsizeinmature
specimenscanbeattributedtosexualdimorphismhasalreadybeencOnfirmed
forNpompi]iusiOmthePhilippinesandtheFijilslands(Haven,1977a;や Vard
etal,1977:ヤVardandMartin,1980;Hayasakaet αl,1982;Tanabeet α
l,1983),
forNmα cromphα lusfromNewCaledonia〔 WardandMartin,1980),andfOrN
belα uensisfromPalau,VVestCarolinelslands(SaundersandSpinosa,1978)In
thesespecies,Inaturemalesgenerallyhavelargershellsandheaviersoftparts
団π
Ш Ｌく α Ｚ 〇
一 り
２ く Ｏ Ｘ Ｏ コ
匡 ０
工
ゝ ＞
０ ∞  ∞
の 一 × く Ｏ Ｚ 一 Ｊ 一 〇 ＯШ エ ト Ｏ Ｆ の コ Ｃ Ｏ 工 ≧ ´ Ш エ ト Ｌ Ｏ Ш ＯＺ く 卜 の 一 〇
RELATIVEWHORLTHiCKNCSS FLANKPOSITiON

112
8
:器1ミ 」
]脚 日PttNES
含
洋誕腎
習
elヽ認￡
古
曰Л
 ▲
口
 △
彗
lfi
▲
▲
.
ヽ ロ
1lo120,3o,40150160
SHELLDIAMETER(mm)
Chapter7
Figure7Doublescatterp10tsof
80nadindexltestisorovary
WeiよνSOfttissuewe18ht(%】 vs
shelldiameterintheselectedspec―
imellsofNpOmpiliusfTomthe
PhilippinesandtheFijilslands
︵ ぎ ︶Ｘ Ш ０ ２ 一 ０ く Ｚ Ｏ Ｏ
170130
thanmaturefemaleslnaddition,Stenzel〔 1964),Haven(1977a),Saundersand
Spinosa(1978),andArnold(1984)havepOintedoutthattheshellsofinature
maleshaveabrOaderandrOunderaperturethanthoseoffemalesofsimilarage
ThesedifferencesaresupportedbythespecimenscollectedfromtheFijilslands
(Fig6),althOughprincipalcomponentanalysisOfmaturespecimensfromthe
Philippinesrevealednostatisticallysignificantdifferencesinthebasicshell‐ form
ratiosbetv′ eensexes(Tanabeet αl,1983)
Indescribingshelllnaturity,theearlierliteraturehasreliedmainlyonchar―
acteristicshellfeaturessuchasablackenedandconstrictedaperture,fllsingto‐
getherofthefinalt、vOOrthreesepta,andathickenedlastseptumlnthisstudy,
weexaminedtherelativegonadweightinselectedspecimensfromtheF"ilslands
andthePhilippinesScatterplotsofthegOnadindexloVaryortestisweightsOlt
tissueweight(%)]VSShelldiameterareshownfortwosamplesinFig71nthe
femalesllomtheFijilslands,Ovaryweightincreasesabruptlyafterashelldiameter
of120mmandattainsamaximum(〜 15g)ataShelldiameterOfapproximately
140mlnlncontrast,gonaddevelopmentislnoremarkedinmales,andfullde―
velopmentOftestesdoesnotOccuruntilashelldiameterof140‑150mlnlf、 ve
considersexuallnaturityas30onthegonadindex,maleshavelargershellsthan
femalesatthispointThelargegonadindicescalculatedforthePhilippinespec‐
imensindicatethatmostofthem(especiallythefemales)haVeapproachedor
attainedsexualmaturity(Tsukahara,1985)TheaVerageshelldiameteratmaturity
forbOthsexesinthePhilippinesampleisabOut25mmlargerthanintheFijian
sampleleg"17164mmand14721mmformalesfromthePhilippines(N=15)
andFijilslands(rヾ =14),respectively,seealsoFig7,andFig12inChapterlll
llVardetαl(1977)andHayasakaetal(1982)havealSOnotedthatmatureoralmost
maturespecimensofNpompiliusfromthePhilippinesaremuchheavierand
largerthanthosefrOmtheFililslands
4.ConclusiOns
lntheFijilslandsandthePhilippines,rヾ pompiliusexhibitsadistinctsexual
dimorphisminweight,shelldimensions,andsofttissuesMaturemalesgenerally

BiometricAnalysisofNautil.lspompi]ius ■13
possessabroaderM7horlaperturethanlnaturefemalesThetwOsamplesfromthe
PhilippinesandtheFililslandsareclearlydifferentinoverall、 veight,sizeat
maturity,andtheslopesofthea1lometricrelationshipsofseverallnorphological
charactersThelnorphologicaldifferencesbetM/eenthesamplesfromthet、 voareas
aremuchgreaterthanthosebet、 veensexes、 vithinthesamearealnaddition,
ヽ4asudaandShinOmiya〔 1983)discOVeredastatisticallysignificantdifferencein
allelefrequenciesbetM′ eensamplesfromthesetwoareasFromtheselinesof
evidence,weconcludethatpopulationsofNpompiliusfromtheFililslandsand
thePhilippinesaredifferentiatedbothgeneticallyandmorphologically
AcKNOWLEDG卜4ENTSWeareparticularlyindebtedtoProfShozoHayasakaOfKa―
goshimaUniversityfortheuseofhisfacilitiesandfOrhisencouragementboth
inthefieldandinthelaboratoryOurthankstoDrAlfredoEEvangelista,As‐
sistantDirectorofthePhilippinesNationalMuseun■ ,ProfAngelCAlcala,Vice
PresidentofSilhmanUniversityinthePhilippines,andDrUdayRaj,Director
ofthelnstituteofヽlarineResources〔 IMR)attheUniversityoftheSouthPacific
(Fijl),fOrtheirassistanceandtheuseoftheirfacilitiesforplanningandoperating
ourfieldstudiesThanksalsototheothermembersoftheprojectinthePhilip‐
pinesandFilifortheirassistanceinthefieldandfortheirhelpfuldiscussions
ヽ
へ
√eexpressoursinceregratitudetOthestaffoftheIヽ 4Randヽlr ヽ
へ7ilsonValloces
inBindoy,Philippines,M′ hohelpedustrapNautilusDrNeilHLandmanofthe
AmericanMuseumofNaturalHistOryandDrAndrewSwanoftheUniversiけ
CollegeofSv√anseacriticallyreadthemanuscriptThiswork、 vassupportedby
Grantsin‐ AidforOverseasScientificResearch(No57043059for1982;No
58041055for1983,No59043050for1984)

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―
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――
――
―
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―――――
―
1983:NaturalratesOf growthandlOngevityOfNautilus
hclュ聖
cns■ s・ 2ュlcoLユ。1。 gy,9,280‑288.
―――
――
1984:N2● ■■■
塁旦growthand10ngevity:evidencefrommarlく ed
andrecapturedaninals 旦。
1,ュoo,224,99o992
‑― ―
andsPINOSA,C.1978:sexualdilnOrphisnin
ゝュ
ュ■■
1● ,from
Palau.2ュ■
Qgo■。19gy,4,349‑358.
STENZEL,H.B.1964:LivingNュ 坐Lll塾 旦
・ェュ
:R.C.MooRE(ed):
エ
ェ￡
皇二
=旦,。ュ
ェ
ュ
yュ ニ也
。Lニュ也Q221,Qュ 也
ology2ュ ニ
也
xl■ 911■,9ュ旦立
K59‑K93.
TANABE,K.,HAYASAKA,S.,sAISHo,T.,sHINOMIYA,A.andAOKI,K
1983:MOrlhologicvariatiOnOf 型ュ

也■
1聖 旦
2Qュ plll聖,fronlthe
PhilippinesandFijilslands.1ュ :HAYASAKA,S.(ed.):Studies
onニ
ュュ上
i■ uュpQュp■ 1■ 聖旦andits assOciatedfaunafromTanon
Strait,thephilippines. こュ
gQ,Lニュュ
型ュ
ニYニュE二 ,.Q,ュ ■
c二 』。
ェ旦
.
2ュQ■ f19.QgQュ 旦
ュQュ ュ12ュ ●ュ,(1)19‑21.
―
――
――
,andTsuKAHARA, 」.1985:MOrphO10gicanalysisOf
ざユ
理二
11●旦
poЩpil=菫旦
. ■
■
:S,HAYASAKA(ed:):Marineeco10gical
StudiesOnthehabitatOfNautilu,p9ュ piliuslntheenvirOnsOf
VitiLebu,Fiji.Kュ gQttL■ ュュ
」ュ
ュ
y二 塁
,ュニQ皇ュニ2■ 19ェ  旦.2皇 o■ f■o
QQQュ,■ 。ュュ
■
2ュ p.,(4):38‑49.
TAYLOR,B.E.andWARD,P.D.1983:stableisOtOpicstudies
Of 世ュ
utti■u旦 ェュ
。ェ
Qュ pLュ■
uttSowerby(NewCaredonia)andNautilus
po■ 211lЦ ttL.(Fijl).2ュl二 〇Qg02Eェニュ22■aeO。 1llnat01.221ュ QoQQg■ ,

41,1‑16.
THOMPSoN,D.1917:Qュ QェQEttL ュュ
塁E。ニュ
.1116p.canbridgeuniv.
Press,London.
TSUKAHARA, 」
.1985:HistO10gicalandhistOchenicalstudiesOf
gonadsof ゝュ
Ч上ユ■
2旦  ⊇
Qュ plll■旦frOmFij■ . ェ
ュ
:S.Hayasaka(ed.):
MarineecOlogicalstudiesthehabitatOfN奎 菫Lll彗旦29■2111聖旦in
theenvironsOfvitiLebu,Fijl. ≦ュ
g。ミLユ ュニ
」ュ
ュY.塁
o皇=
Q,ュ主Q二 ￡Qニュニ
2ュ。■』
ユQ9QQュ 量loュュ
12ュ⊇.,(4):50‑60.
WARD,P.D.andMARTIN,A.W.1980:DepthdistributionOf
ざュ聖
圭iluttp。 ュ
p■■■
ュ旦 ■
nFijiald ゝユu■ ■
■聖
ニュュQ=。 ェ
pLュ1坐 , ■
nNew
CaredOnia.y,1■ gttr,22,259‑264.
,STONE,R.,wESTERMANN,C.andMARTIN,A1977:NotesOn
aninalweight,caneralfluids,swilnmingspeed,andc。 10r
polymOrpllislnofthecepha10pOd ゝュ聖
■■■
uニ ュ
aQ二Qュ ph全 ■
坐旦
22192L19■2gy,3,377‑388.