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ZOOLOGICAL SCIENCE 151 415-424 (1998) @ 1 998 ZeoFogica Sloc]et yof Japan
Platycopi caompacta n, sp., the Second Species of
Platycopioida(CrustacCeoap:epoda) inthe lndo-Pacific
Region, Remarks Development, Feeding,
with on
Swimming, Zoogeography
and
Susumu Ohtsuka'kH,oYoungSoh" HiroshiUeda2
and
"Fishen'esLabonatory,HiroshirnaUniversit5y-,8-1Minato-machi,
lakehara,Hiroshima725-O024,Japan
2Marine Bioiogica Sltatio nE,hime Universit yN,akajima-cho,
Onsen-gun, Ehime 797-4502,Japan
ABSTRACT-Piatycopiacompactanew$pecies(Copepoda/PlatycopLoLda)isdescribedfromthehyperbenthic
zone inthe Tokaralslandss,outhern Japan.Thisisthe second species of the order inthe lndo-Pacific
region, and the eighth species of the genus .The presen tnew species i sdi$tinguisha bfireom the closely
related P. inornata from the Bahamas by < 1> the formatio oni allobasis in the antenna; <2 )two unequal setae
on the basai exite of the maxil1ule; (3 )the intermaxilliped aplroces swith only 3 pair sof prominence sat tip ;<4)
the fusion of the cexa and basis in ieg 3; and (5 )the fusion of the $econd and third endopod segments in
female Iegs 3-5, Copepodid stages II IV,, and Vl are compared with those of P, orientaiis collected from
Okinawa, and diffe rin the developmenta lpattern sof the antennute and legs .The presence of an
intermaxMipedaplrocessand gutcontent analysis revealed thatPIatycopiiascarnlvorous. The armature oi
iegs 2-5 oi the hyperbenth igcenus Ptatycopi ias stouter than that of cavernicolous platycopioi dwhsl,ch
indicate sthat Pla tycop imaay penetrat iento 1oose sediment, whereas the iatt egrroup might be more adapted
to pelagi ctif eT,he horizonta Ldistribut oito tnhe genus Ptatycopia apparently exhjbits a tul lTethyan track.
INTRODUCTION have found another species of P/atycopia ,which belongs to
thehyperbenthiclcavernicobuscopepodorderPIatycopioida,
Hyperbenthicandcavernico)ouscopepodshaveprovidedin the Namsei lsland ss,outhern Japan .This is the second
signlticant new informati oabnout the morphology, zoogeog- species ot the order in the lndo-Paci triecglon and the eighth
raphy, ecology and phylogeny of the maxiliopodan subciass speciesofthegenusworidwide.Thepresentpaperdescribes
Copepoda, Some ca)anoid gener asuch as Exurneiia, the new species with remarks on jts development, teeding ,
Metacalanus ,Pararnisophria ,and Stephos are broadly dis- swimming, and zoogeography,
tribute din both the hyperbent izcone aRd marine caves and
iBnodxischaatl e)feta uanL,i s1t9 i9sci0r; niBloawrimtaien s,b e1t9w7e6e; n Jtahuem et waon dre aBlomxss h<aclfl,, MATERIAL AND METHODS
1995a: Ohtsuka et at, ,1993b,1994 ;Rier aet a/, ,199t) ,Re- Copepods were colFected with a dredge (mout h50 cm wide, 15
cently numerous new famLlies ,genera a,nd species of cope- clmsl haingdhs ;,msesohut hsaizteK 5y umsmh)u, aJta 6p7a nm< 3dee'p=tOh Oo,ff8 K2u'cNh,i1n2o9 "l5s3[,a9n2 dT',oEk)a,roanJune
pods have been discovered from anchialine caves <e.g,,
1, 1997 ()oc at)ime 0815-0847) .Sediment was stjrre din sea-water
Fos$hagen and 11if f1e9,85, t988, 1989, 1991, 1994; Jaume several times and the supernatant was flFter ewidth a planktQ nnet
and Boxshal l,ri995a, b1,996a,b ;Ohtsuka et aL, 1993a,b: (meshsize/O,lmm),CopepodsweretixedinlO%neutralizedforma-
Rocha and l[if f1 e9,91) .These have shed new "ght on phylo- iin,'sea-waterimmediatelyafterfilterlng.
g1e9n8e9t; icH or,e la1t9io9n4sh)i psand bwritohuing he taicnht oqcuopeesptoidon tohrde etro t(aBl iootxfy sthheail, s(e1 um99T h1aen)d ,t ylpne sstpeictimue tnCseh,ib aa ,rTee rdmeipneosliotge y dfion 'tIhoe wN saHtuuyrsa la nHdi sBtoorxy sMhua-ll
Messinia nSalini tCyrisi isn the Mediterranean Sea abovt 5.5
Myr BP (Jaum aend Boxsha) 11,996b),
Durln gou"nvestigation of hypefbenth iccopepods we
'CorrespondJngauthor/ Tel , +81-846-22-2362;
FAX, +81-846-23-O038,
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DESCRIPTtON face; coxal endite with 3 setae: ba$al exite (arrow jen dFig,
2E) lobate ,bearin g2 setae of unequal length p;roxima )basal
Order Platycopioi Fdoasshagen ,1985 endite elongate, bearin g3 terminal spinulo$e setae: distal
Famil yPlatycopiid Saaers ,1911 basal endjte probably represented by a knob with 2 unequal
Genus Platycop iSaars, 1911 setae; both rami completely incorporated int obasis ,bearing
Platycopia compacta new species 4 setae in total,
(Fig s1,-6} Maxill (aFl 2gF.) stout; 2 praecoxa l2, coxa[, and 1 basal
Materialexamined,-Holotype:adultiemale(bodylength endites bearin g4, 1 , 3, 3, and 3 setae, respectively, one seta
O,76 mm), di$sected and mounted on 14 glas sslides, CBM each on second coxal endite and on basal endite fused to
ZC 4035, Paratype a/dult temal e{bod lyength O.8 1mm), whole segment, Firs etndopod segment completely fused te basis,
specimen, CBM ZC 4036. Non-types c:opepodid V femabe represented by 1 spinlform seta; second to fourth segments
(bod ylengt hO.74 mm}, dissecte dand mounted on 4 glass each bearin g1 spiniform seta; thir adnd fourt shegments with
slides; copepodid V male Cbod lyengt hO,62 mm}, dissected 1 and 2 setae, respectlvely, in addition to spiniform seta.
and mounted on 2 glass s[jdes; copepodid 1 11(bod lyength MaxiIlipe d<Fi g3,A) with praecoxa and coxa incompletely
O,47 mm), dlssecte dand mounted on 1 glass slide, coaiesced: coxa with 2 spinulose, subterminal setae; basis
Description.-Adu) tfemale (holoty Bpoed)y .(Fi gIA,,B) bearing2innersetae,oneofthembeingrudlmentary;endopod
compact. Prosome about 2,8 times as )ong as urosome; 6-segmented ,setal formula 1, 2, 2, 1, 1, 4. IntermaxiUipedal
cepha!osomeandfjrstpedigeroussomiteincompletelyfused, proces s(Fi g1B,,F) with 3 pair sof anterior)y directed ,acute
without distin acrtthrodial membrane: fif tphedigerous somite prom]nences at tjp.
produced posterio rilnyt oround lobe reaching beyond middle Leg 1 {Fi g3.B) wjth 2-segmented rami; coxa unarmed,
ot firs atbdomina) somite; several developin ogva observed in produced into acute outer process subtermjnally; basis ex-
posterio rpart of prosome in dorsa land Iater avilew, Rostrum panded, bearing plumose setae near base of endopod, First
(Fi gt,C} sharply pointed ,ventra)iy directe dwi,thout filaments endopod segment with longitudina glroove on anterior sur-
at tip U,rosome (Fi g1D,) 5-segmented ;genita lsomite small, face along basa lseta, segment produced at outer dista lcor-
almost concealed beneath dorsal extension of fift phedigerous ner, Firs etxopod segment uRarmed, prominen tat outer distat
somite; paire dslit-lik egonopores (arrow ien Fdig .1 D) located corner; second with acute promlnenc eand row of setules along
ventraliy, slit-lik ef;ir stto thir dabdominal somites with poste- outer margln and 6 setae, outermost more heavil cyhitinized
rior margin finel sytriated; thir dabdominal somite protruded than others. Leg 2 (Fi g3.C) with 3-segmente drami, Legs 3
dorsoposteriorl yinto w-shaped plat epartl cyovering anus; <Fi 4gA,) and 4 (Fi g4.B) with second and third endopod seg-
fourth abdom]nal somite prominen tventroposteriorly, Caudal ments eot tul lsyeparate on posteri osrurface; both legs simi-
rami (Fi g1D,,E) symmetrical, each w]th trjangula drorsa lpro- lar b,ut dMerin gin i(i c)oxa and basis separate in leg 4 and
cess; seta l rudimentary, seta ) )slender, setae 111- Vpllumose; incomplete ltyused in le g3; (2 t)hir dendopod segment bear-
setae IV and V with finel pyunctat eproxima lsurface, ing 5 spiniform setae jn leg 3 but4 in leg 4; <3 s)econd exopod
Antennui e<Fi g2.A} 23-segmented ,Firs stegment with $egment Iackin ignne ;seta in [eg 3, Leg 5 {Fi g4C,) with ex-
two longitudina rlidges and elongate, heavil cyhitinized ante- panslonalong]nnermarginofbasis;secondandthirdendopad
rior seta, Armature and fusion patter ans toliow s1:-1 mlaly be segments almost fused, wjth suture visible on both suriaces,
incorporate din firs stegment, which ls not traceabl beased on Spine and seta formul aof legs 1 to 5 shown in Table 1.
elements; IV-Vl-6 + ae; Vll-2 + ae; Vill-1 short spjniform Male unknown.
seta; IX-2 + ae: X-1 + 1 short spLniform seta; Xl-2 + ae; Copepod[d ll l{C"1 )(Altho ulegghs 3-5 are damaged in
XlI-1 ; XiI1- 1+ 1 short spin]torm seta; XIV-2 + ae; XV-1; the only individua li, tyvas identit iaesd CiH on the basjs of the
XVi-1 + ae; XV)1-; XVIII-1 ; XiXn; XX-1 : XXI-1 + ae; number of urosomal somites, the number of ramal segments
XXIi-O: XXIII-1; XXiV-1; XXV-1 + ae; XXVI-2; XXVII- ot 1eg 2 and the presenc eof leg 5, See Ohtsuka and Boxshall,
1 + 1; XXVII]" + ae, 1994).-Urosome 2-segmented. Antennule 15-segmented
Antenna (Fi g2.B) with unarmed coxa and basis .Endopod (Fi 5g},; armature as toHows :1" + ae (Fi g6.A); 2ab; 3-1
with fif ssetgment completely fused to basi sto form allobasis; +ae; 4-O; 5-1; 6--O; 7-1; 8-1+ae; 9-O;1O---1; 11-1;
second and third segments incomplete icyoalesced, bearing 12-1 + ae; 1 3-2: 14-1 + 1; 1" + ae (se eDevelopment
3 and 7 setae, respectively, Exopod indistin c7t-lsyegmented; in Discussion )An.tenna wjth basis compietely fused to tjrst
setal fermula 1, O, 1, 1. 2, 1, 4. endopod to form allobasis. Basal exite of maxillule with 2 se-
Mandibu)ar gnathobas (eFi g2,C) bearing 1 monocusped tae ot unequal Iength ,Maxi11ipe dwith onIy singl eseta on coxa;
tooth and 4 bicusped teeth ,Mandibular palp (Fi g2D,) with basis as in adult female ;endopod 4-segmented; setal fo rmula
unarmed basLs. Endopod unisegmented, with 7 termina lse- of 1, 1, 1, 4, lntermaxillip perdoacles swith 2 pair sof promi-
tae .Exopod 4-segmented ,tir stto tourt hsegments, respec- nences at tip .Leg 1 with 2-segmente drami; $econd exopod
tivel wyi,th O, 2, 1 ,and 2 setae. segment bearing only 5 setae. Leg 2 (Fj g6C.) with 2-seg-
Maxi)lu l(eFi g2,E> well developed p;raecoxa larthrite ex- mented rami; firs texopod segment with sLngle outer spine,
panded, bearing 6 stout, multicusped spines and 6 setae, 2 Legs 3-5 mi$sing.
setae of which being originated from center of anterjor sur- Capepodid V (CV f>emaLe,-U rosome 4-segmented. An-
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New Specie osf P?atycopia 417
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Fig, 1. Piatycopia compacta, adult temale Cholotyp <eA)) ,Habitus, dersal; (B )Habltus, lateral ,inte;maxillipeda lprocess arrowed: (C )Rostrum;
(D )Urosome, ve ntral, gonopores arrowed; (E )Anal somite and caudal rami, dorsal /(F )intermaxillip epdraolcess ,viewed tror nventral side. Scale
bars in mm.
tennule 22-segmented (Fi g5), ;armature as follow s1/-8 + 2 21-1 + 1; 22-4 + ae (se Deevelopment in Discvssion) .Basai
ae (Fi g6.B}: 2-1 short spin[form seta: 3-2; 4-1 short exite of maxillule with 2 setae of unequal length .Maxillipe das
spinitorm seta; 5-2 + aesthetasc; 6-1; 7-t + 1 $hort in adujt temale, lntermaxiHipeda plrocess with 3 pair soi promi-
spinitorm seta; 8-2; 9-1 ; tor1 + ae; 1 1-O; 12-1; 13-O; nences at tip ,Legs 1 and 2 as in adult female ,Leg 3 vvith 2-
14-1;15-1+ae:16-O;17-1;18-1/19-1+ae;20-2: segmented endopod and 3-segmentedexopod; armature as
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Fig. 2, Platycopi acornpacta, adult temale (ho[oty p(eA >)A,ntennule; (B )Antenna; (C )Mandlbular cutting edge; (D )Mandibula rpalp ;(E)
Max"lule, basal exite arrowed; {F )Maxill aS,cale bars in mm.
in adu[t female ,but second and thir dendopod segments not rated from coxa; 2･segmented rami; firs tendopod unarmed,
separated, without arthrodial membrane, Leg 4 with 2-seg- second bearin g6 spines; flr setxopod segment with slngle
mented endopod and 3-segmented exopod; armature as in outer spine, second segment bearin g4 tlange dspines and 3
adult female, but second and thir dendopod segments not short spines.
separate. Leg 5 {Fi g6,D) with intercoxa $lclerite not fu11 ysepa- CopepodidV(CV)male,-Uro$ome,antennule(Fig.6B),
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New Specie osf Ptatycopia 419
B
O.05B,C
Fig. 3.Ptatycopia compacta, adult iemafe Cholotyp (eA) },Maxjllipe d(;B )Leg 1 , anterior; {C >Leg 2, anterior. Scale bars in mm.
basa lexite of maxiilule, intermaxi))ip epdraolcess a,nd legs 1 - a distin pcrtocess .Althoug hsome other congeners may also
4 as in CV female. Leg 5 (Fig 6.E) with intercoxa lsclerite not bear a process, i twas not mentioned in their originai descrip-
tuMy sepafated from coxa: 2-segmented rami; second endopod tLens.
segment with 6 short spines; second exopod segment bear- The presen tnew species closeiy resemb]es Platycopia
ing 4 tlange dsplnes (2 m[ssing) and 2 short spines. inomata ,P. perplexa P,. robusta Andronov ,1985, and P, sarsi
Etymoiogy.-The specific name "compacta" <Latin M,S. WUson, 1946 in :(1 >3-segmented rami in Iegs 2-4
compactus meaning plump) is derive dfrom the compact (symplesiomorp h(y2 ))t;he fir setxopod segments ot Iegs 3
prosome ot the new species, compared with that of P. on'entalis and 4 ot adults bearin g1 inner seta (symplesiomorp hFey-).
fir sctol)ected from Japan, male P. perptexa and P, robusta both lack an inne rseta on
Comparison.-The mouthparts and Ieg 1 of specles ot the fir setxopod segment of leg 5 (synapomor pwhh"ye) P,
the genus Piatycop ieaxhibit no remarkable differences while inornata and the new speckes carry one <sympleslomorphy>,
the armature and iusLo npatter nof legs 2-5 are the important Since only males are knewn in P, sarsi, i tis impossibi teo
characters fo rdlstinguish ibnegtween species (Wils o1n94,6; compare the female legs 5. The presen tnew species i sdls-
Ohtsuk aand Boxshall ,1994) .The presenee or absence of tinguishabl efrom P. inornat ian :(1 )the basis and endopod of
theintermaxillipedaLprocessisal$oimportant,Some$pecles the antenna being completely fused (separ ain tPe, inornata);
such as P. inornata Fosshagen, 1972 and R petplexa Sars, {2 )the 2 setae on the basal exite ot the maxi"ule being of
191 1 are furni3he wdith i t<se eSars ,191 1 ; Huys an c tBoxshal1, unequal length (almo setqual in P. inornata )<;3 )the
1991> ,whereas P, orientalis Ohtsuka and Boxshal l1,994 Iacks intermaxiLlipedalprocesshaving6spinousprominencesatits
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O.05
Fig ,4. Piatycopi caompacta, adult female (hDlotype> .L(egA )3, anterior ;(B} Leg 4,amteriD r(/C )Leg 5, afiterior, Scale bar in mm,
ti p(8 in P. inornat aHiuys and Box$hal l1,99I, tig,2,1.1 <24);) Platycopia sarsi was poorl ydescribe don the basis ot a
the coxa and basis in leg 3 being fused <separ aitn eP. single adult male and two copepodids by Wiison (1946 T)h.e
inornata) {;5 )the second and thir dendopod segments i nlegs followi nfgeature sdiHe rbetween P. sarsi and the new spe-
3 and 4 being incompletely fused {separa tine P, inornata); cies: (1 t)he proximalmos tspine on the praecox aarlthrite of
and (6 t)he second and thir dendopod segments of the female maxMule is bifurca tien P, sarsibut tricusped in P, compacta;
leg 5 being almost completelyfused (separa itn eP. inornata). (2 t)he outer subterminal cerner of the coxa of leg "s round
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New Species of Piatycopia 42t
Table 1, Seta and spine formul oaf legs t to 5 of Ptatycopi caom- rate from the compound segment IV-V tat CVI.
pacta n. sp. Roman and Arabic numerals indicate spines and setae, Developmenta lpattern soi antennules, maxilljpeds and
respectjvely,
legs can be compared between P, compacta and P, otientatis
coxabasisexopod123 endopod123 as describe bdy Ohtsuka and Boxshal (l 1994). In C1l 1the num-
berof segments oftheantennutes is15inP.compacta and
Leg 1Leg o-oo-oO-OOo--1olO--1OI-1OI--O1]l1-,o1,4II-OIFOOi-lO"]iO[,,22,tO-liO-Oll1l8,2j,n2P,orientaiis.The dista1l3 segments <ancestsrega-l
2Leg ments XVI-XXVII Io)f P. compacta have the same armature
34LLeegg jill--Ollil-lO-ll[l"li[l,l2,[,OO2--l1i1D0-l-lllill,l2,,32,3 as in R orientaiis, [n CV the segmentation and setation of the
5 IFIil-llli,1,[[o-elo-Lil,11,u antennules are the same in both species except to rthe pres-
ence o{ an aesthetasc on the ancestrai segment XIV and in-
completeseparationbetweenthefirst(pre ¢ursoroWl-VII)and
second {VII sIeg)ments in R o"'entatis, ln CVI (adu fletmale}
ln R sarsi but acutely pointe din P, compacia, The )ate the ancestral compound segment 11 1{o rIV)-V a)nd segment
copepodid female describe dby W"son (194 6p)robabl ycor- Vl lare separate i nP. compacta and fused in P. orientatis,
responds to the fif tcohpepodid stage because of the 22-seg- The differenc ebsetween the armatures of CVi ot both spe-
mented antennule, 3-segmented rami of [egs 2-4 ,and 2-seg- cies are as foilow san: aesthetasc is presen ton the compound
mented ram[ of leg 5 (cf O.h,tsuka and Boxshal l1,994) ,1n P, segment IV-V lof P, compacta but absent on the part corre-
compacta the titt hcopepodid stage of both sexes bears a 2- sponding to IV-Vl on the compound segment liFVI Iot P,
segmented endopod in leg 5, in which the second and thlrd orienta tis t;he ancestral segment Vl1 is separate from the com-
segments of the adult are not fuHy separated, Leg 5 ot the CV pound segment IV-Vl in P, compacta but incorpQrated in P,
femaie of P. saisidiffers slightly from that of P. compacta: the orientalis; the ancestral segment Xll lbears 1 long and 1 short
proxima )outer spine on the second endopod segment and spinitorm seta in P, compacta, but only a single long seta in
the proxima linner spine on the second exopod segment are P. on'entatis, ln addltion, the appearance ot aesthetascs at
reiatively more eLongate than those of R satsi. the molt fror CnV to CV1 differ isn [P. compacta an aesthetasc
appears on segments )V-Vl ,iX ,and XiV, whereas in P,
DISCUSSION orientaiis only one aesthetasc i sadded on segment IX,
The developmen tot maxillipeds has so far been unknown
Development in the P)atycopioi diancludin gP. orientalis (Ohtsu kanad
The segmentation and setation ot the antennules of P, Boxshal l,1994), in P. compacta the numbers of the maxil-
compacta are depicte din Fig ,5. The most proxima lantennu- lipe dendopodal segments in CV and Cl1 lafe 6 and 4, respec-
Iar segment ot female CVI is interpret aesd IV-V lbased on tivel y,Therefor etha tin CIV may be 5, i fthe development
Huys and BoxshaH (1991 )tit:he setatlon pattern provides no foliows the pattem proposed for the Calanoida by Ferrari
evidence oHhe tate of the 3 most proximal segments". AI- (1 995) ,the penultimate endopod segment ot Cl1 ctan be iden-
though the fourt hcopepodid stage (CIV i) sunknown, the tifie ads a precurso orf the third to fif tshegments of the adult
setation of the dist a1l3 segments are unchanged from ClI Ito <altho uFgerhra r<i199 5r)egards the fir setndopDd segment
CVI. At the las tmolt from CV to CVl, an aesthetasc is added as a iobe of the basis ,vve tollo wHuys and Boxsha] l(1991):
int othe ancestral segments IV-Vl I,X ,and XIV ;a seta is added maxillipedsprimitivelybear6-segmentedendopodratherthan
to the segment X. The fir sctompound segment of OV con- 5-segmented) a;t the molt from Cll lto CIV the third segment
sjsts oi the ancestral segments IV-Vl1 t,he Ias tot which is sepa- of the adult may f[rs atppear to result in 5-segmente dendopod;
spiniform setax $etaaesthetasc
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IV.LJIVlIVilLIXXXlXllX:11xlvrvXUIXVIIrv111XIXxxxxLxx11xx1[1XXNxxvxxv)XXViLX)CVIII
Fig. 5. Schematic illustrat oifa anntennuLes of Pia tycop icoarnpacta. <I IthIi)rd copepodid stage: (V )tit tehopepodid stage <bot shexes); (Vl)aduit
female. Roman numerats lndicate the ancestral segment scheme proposed by Huys and BoxshaU (1 991).
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Fig ,6, Platycopia compacta, (A )First segment of antennuie of third copepodid stage: (B )Firs stegment ot antennule of tit tchopepodid stage
male/ (C )Leg 2, third copepodid stage; {D )Leg 5, fif tchopepodid stage temale ;(E )Leg 5, fif tchopepodid stage ma[e. Sca} ebars in mm.
the fourt hsegment of the adult could be added at the molt Table2, Pattern ot development of legs 1 to 5 ot Platycopi caom-
from CIV to CV, ln the Calanoid athe adult endopod segmen- pacta n. sp. Number of exopod segments + number of endepod seg-
tatio nis comp)eted at CIV (se eFerrar i1,995; Ferra raind mseenctos,nd Nanudmbtehri rjdsn egpmaernetns.tlheegse sindicat eisncomple tseeparation of
Markhaseva ,19g6) while in P, compacta presumably at CV.
The completion ot the endopod segmentation may be acce)- 1leg 2leg 3leg 4ieg 5
erated in the Ca)anoida, Copepodidll l 1+1 2+2 missing missing mjssing
The setation of the maxil)iped endopod of R compacta is V 1+1 3+3 3+2 3+2 2+2
completed probably at CV as well as the segmentation, Per- Vl 1+1 3+3 3+{3) 3+(3) 3+{3)
haps the setal numbers of the fir satnd sixth endopod seg-
ments of the adult are the same from Cll lto CVI. Fo)lowjng
the patter nof the Caianoida (Ferr a1r9i95,; Ferrar iand separations of these segments are delayed in legs 3 and 4. 1n
Markhaseva ,1996} ,the setal formul aof the endopod of CIV contrast, simuitaneous additions durin gthe las tmolt are sus-
may be 1, 2, 2, 1, 4. pended in the endopods of legs 2-4 in P, orientalis, which
The devebpmental patter onf 1eg st-5 of P. compacta Ls results in 2-segmented endopods in these Iegs .1n Cl1 [aproxi-
shown in Table 2 to compare i twith the common patter ont mal outer spine on the f[rs texopod segment ot leg 2 appears
copepod Ieg ontogeny proposed by Ferra r(i1988 S)i,mutta- in P. orientatis (Ohtsuk aand Boxshall, 1994) ,but not in P,
neous additions are suspended in both rami of leg 1 here as compacta,
well as in P. orientalis. The developmenta lpattern of legs 2
and 5 of P, compacta almost colncldes with the common de- Feeding and swimming
velopmental patter anlthough the second and thir dendopod Littl ies known about the biologyan decology ot the order
segments of legs 3 and 4 are not tult syeparated. Probably Platycopioid a(Huy sand Boxshal l,1991>, rwatycopi baears
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New Specie osf Ptatycopia 423
well-developed maxiliulary praecoxa larthrjtes, maxillae with anchialine caves in Bermuda (Fosshag aendn liif f1e9,85,
stoutspinitormsetaeandspines,andanintermaxillipedalpro- t988). Since there are numerous anchialine caves and other
cess in some species, al1 of which probabl ypla ya role ln grasp- types of marine caves in the Nansei island ss,outhern Japan
ing and hoidLn gprey ,This is supported by gut content analy- (Hayam iand Kase, 1993) ,undescribed taxa ot the
sis which indicat etdhat the CV female ot P, compacta had Platycopioi cdoauld be discovere idn these regions.
preyed upon copepoctids oi other copepods The distribut oifo nthe genus PJatycop iia sshown in Fig.
The strong spines and splniform setae of Iegs 2-5 ot 7 on the basis of Huys and Boxshal l<1991 O)h,tsuka and
Piatycopia imply that j tmay penetrat leoose sediment but not Boxshall ( 1994), and the presen tstudy. This patter nevidently
lnterstit iasp[aces. The Ieg armature ot the hyperbenthic colncides to a ful lTethyan track {circum-tro pthircoauglh the
Platycopiiasmuch stronger than thatof the cavernicolous entire region of theformerTethysSea)as definedby Stock
AntrisocopiFaosshagen,1985 and AlanocopiaFosshagen, (1993w)ho, inslstetdhatstygobionts thatdlspiasyuch a dis-
1988,which suggests thatPlatycopiia$more closely associ- tributionaplatterannd havepoordispersaaibilitiesmust have
ated with sediment than are the cave-dwelling genera (cL, exisied i npLace ever since the existence of the Tethys Sea
Bowman and Genzalez ,196D, which are more adapted te a (ca t,20-20 Myr BP). Althoug hthe genus Platycopi ias not a
pelagi clife. real $tygobiont, the di$tributi oonfs many hyperbenthic
copepod$ seem to coincide with the ful lTethyan track,
Zoogeography For example, hyperbenthi ccalanoid copepods such as
The order Platycopioi dwaas the fir sgtroup to dlverge Paramisophria ,Pseudocyciops , Stephos, and the
trom the main lineage of the subclass Copepoda and retains Ridgewayiidae are djstrjbut iend the shallow water$ in the
numerousextremelypleslomorphiccharacters,Piatycopioids Caribbean t,he eastern Atlant iacnd, lndo-Wes tPacif ircegions
are restricted to the hypeFbenth izcone and anchiaiLne caves (Boxsh aetlatl,, 1990; Fosshagen ,1968a,b, 1970a,b; Jaume
I nshallow waters (Huy sand Boxsha" ,1991), AI Ieight spe- and Boxshal l1,995a; Ohtsuka and Hjromi ,t987; Ohtsuka et
cies of the genus Ptatycopi aare distribut eexdclusively in the aJ. ,1993a,1996 ;Rier aetaL, 1991> .ln addition, some of these
hyperbenth izcone in the Atlant iacnd indo-Paci fOicceans taxaaccommodatebothhyperbenthicandcavernico)ousspe-
(Huy sand Boxshall ,1991; Ohtsuka and Boxshall ,1994; cies (Ohtsuk aet aL, 1993b,1994; Yeatman, 1969). Such a
present study), whlle Antrisocopia Fesshagen, 1985 and faunlsti csimilarity between in the hyperbenthi czone and
NanocopiaFosshagen,1988 are monotypic inhabitantosf marine caves strongly supports theabove assumption,
Fig ,7, Distribut iootn the genus Platycopi abased on Huys and Boxshal l<1991 O)h,tsuka and Boxsha] l(199 4a)nd present study. 1, P,
compacta, n, sp.; 2,P, orientalis Ohtsuka and Boxsha[I ;3, P. perpiexa Sars and P. pygmaea Sars; 4, P. perplexa /5, P. fobusta Andronov; 6, P.
sa rsi Wilson and P.tumida Wilsen/ 7, P, inornata Fos$hagen.
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424 S. Ohtsuka et at.
ACKNOWLEDGMENTS Fosshagen A, 11if fTeM {1994 A> new species of Erebonectes
(Copepo dCaa,lanoida )ffom marine caves on Caico sIslands,
We would lik eto express our sLncere thanks to Dr GA Boxshall We$t lndie sH,ydrebiolog l2a92t293 /1 7-22
for his revision of the early draf tA.cknowledgments are also due to Hayami 1 ,Kase T (1993 S>ubmarine eave Bivalvi farom the Ryukyu
Dr F.D. Ferrari and an anonymous referee, We are gratef utjo the lslands;systematicsandevolutionarysignificance,BullUn]vMus,
captain and crew of Toyoshio-maru ,Hiroshima Universi ttyor their Univ To kyo 35/ 1" 33
cooperation at sea. Ho JS {1994 )Origi nand evolution Df the parasit iccyclopoid cope-
pods, lnt J Parasitol 24/ t293-1300
Huys R,BoxshaHGA (1991C)opepod Evolution.The Ray Society,
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on Lanzarote C,anary Islands S,tygologi 5a: 33-41 Ohtsuka S, Hirom iJ (198 7C)alanoid copepods corlected fro mthe
Fe r ramrenj tFsD {of1 c9oBpBe pD)oedve pbopsmte-nmtaa)l cpiaitlitpe eI;de niagns ls n.Cumrbeurssta cote raa nm5aa4l: 2s5eg6-- nHeoanrs-hbvo,tJtaopman .in T Paunbalb See Btao yM aorn Btihoe l LPaabe i3f2 i:cc2ea1 s9t- 2o3t2 the middle
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trom the barrie rfeet iR Belize ,with comments on appendage 781 57-67
F e rrdaer viFeDl, oMpamreknhat s,Pervaoc B EiLo lS{o1c9 9Wa6 s)Pha r1k0iBu/ s1k8a0r-e2n0w0ishnera ea, new O ht scuopkeap oSd,s ,Roe tHhJe SH, yBpeorxbsihoanyl clGiAda e(, 19 9c3obl)Le cAte dnew f rfoamm itlhy eo fd ceaeiapn-oisdea
genu$ and species et calanoid copeped (ParkiLd naeew, family) hyperbenthi ccommunity in the northeasterm Atlanti cS,arsi a7B/
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F o s sPmharsao,gce B4n i, AoP lsS(eo1uc9d Wo6acs8y hacMla )r1oi0pn9ie]d 2ab 6e(i4oC-lo2o8pg5iec paioln vdCeaas,ltaignaotiid oifannr )stohm et hBea hBaa-- O h ts<tiuCoko na Spso,f e Bcpoopoxep sdoPhdaLlada/lt ;yGc Aostp ai(goei1s.d9a )9 J, f 4PCr)/roaum tstythcea ocNpeoia rano tBriih eoPnMat4ac/liisf ,1i 5cwn1ie,t-wh1 d6se7pseccireisp-
FFo os s ssthmShhahaaa$aem, grgaBees sann8i , h. 3A AaaS5B ma/o( (ar5t 11s9s7t97j -o0 w6a6mia38t4-)2hbl : MiM>3raave9rTmi- aiinAr6nnkrgee2sl ebbtj ieooolnl[lo iogPdigaa clrea(ca lCaimiolninvpsveoeespstpihtogridaig t aaaciCt,toaunitl sheaiainennn stot hihTed,e aBB S)aafchhroaaot--mt, O Oh htt s$bt6duuea0ekktb1m aawl e1eiSSr0es,,nhs 5 BmFa- eool1anxrsc7tjssa 2ehothlfeaaa [ltlgn ilhGeodrA ni ge,dA e ec,Rn noeoSpweeoer hpH g ao{SeHdJnYCPe o(;r<a pa191 c.9eBo94puc)9 lao P6]Tl dNhhaC)yaarnalteuo/l Hgesa ei{nnnseRoe wtiitMdidu cgass pe)r ewwe,Lciliatoaehytnsi idotin hs<Ddhetai Z petesesh}-eotrlo)m
mas. 12, StephLda e(Copepo dCaa,lanoida )from the Bahamas Okinawa, $euthern Japan. Sarsia 81: 247-263
Fo $ swShiaatrghs erinea m4Aa1 r:{ks317 9e-n47 80S btMea)pFhionse sbiinuoatlusog iW cianlavliee syatndig aSt. ia oricnnt sitchuse BSaahras-, Ri e ra(L aCTn,od p(Vewivpeeoss dtFCae ,a M/rleCnadiitn tioJel-rdM r afa(f)no1ema9 na 9) O1 s.cu)Sbemtaareinnpoehloo sgmc aaiAivageqaut eaofiti iMca asj1po0,r: c3na1o l7v,s--
mas. 15. Ricigewayi {aCopepod aC,alanoida )and two new gen- 323Recha
Fo $ sehraag oefn cAal(a1no9i7ds2)M tarroimn tehbei Boalhoagmiacsa.] iSnavresslt a4j4g:a t2i5o-n5s8intheBahamas Co pCeEpFo,d a1 1CiCt yfTceMl oCp1o9i 9dfr1a o))Smp aenlcheioaliitneho nciadvea sea, onne wth e fBaamhialm yaoi
F o s sSn1hea7awr. gsP eiolnra ad4AetBr,y : cl5ooiftpi -Ci f6oTifepMde ea (pe(oC1do9ap,8 e 5PTpl)woaotd yCnaceaw,olpa g]neoonii eddtraar a,)ot morf o mmCa artilnhaeen o Bicaahdvaa em$aands ,o na S a r s lnAGesrwOicah tn{i ydv 1MS.p9aa et1r1ohfs) iPNd alae7ate6tup: yr-1cv w63oa71pt-:i e1p1a r7e-Cr51pa6Xleaxan,o ni. dfgareonm, atnhde Nspo.r wAe gFeimaarnk acbolaest.
Fo s sBheargmuedna .A, Sa11risfif ae7TM0: 3(415-93858 8)A nevv genus of Platycopioida S t o cAk mJpHh i(p1o9da9, 3SJo) Nmaet rHeimsal r2k7a: bl8ee7 d-i8s1t9ribut ipoanttern isn stygobiont
(Copepoda)fromamarimecaveonBermuda,HydroblWoill$oogni, MaS1 6(71,9'4 6T)he specles of Platycopia Sars <Copepoda,
168: 357-361 CalanQida )S.mithson Misc Col l106: 1-16
F o s sChaalgaenno Ai.d aI )wLlitfh f TetMwo (ne1w98 s9 pe}BcioehsoY firnoa m,a anne wanc hgieajnlunes ( cCavoep ien pthoeda/ Y e a twmltahn dHeCsc r(ip1t9io6n 9oAf) raend eusncursiupatlion $p eofc icmeepenp.od ,J Te Rninaegsesweae pAaca amadr kSi,ci
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339-346
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