R i r i \ h Ì h l i r r r d i I ' i l c o D k , l o s ci rS r í r i s n i i J
ACTINOPTERYGIANSFROM THE MIDDLE TRIASSIC
OF NORTHERN ITALY AND CANTON TICINO (SVITZERLAND):
ANATOMICAL DÉSCRIPTIONSAND NOMENCLATURAL PROBLEMS
CRISTINA LOMBARDO
R1i-etl Jnt
À.ì èrrlr:
ìrriropr.rrsirns,
1\ 2AAl:deetE.lse?k"tbú 2.J.2oal
l r l.rdinirn, r.dcsc!iption,
Ticino-Swirzerland)
(F;g.l).Thisìocrlityyicldedrr;ch
faunamadeup of morc thin 3000 spcc;mcnsof fishes,
belongingto 1s species,
and three specimcns
of rhe
^,.r!r,n,. \,cnsoDodt\criii, rulh b$c di nuo\ì$(mphri b.n
(Timori
sauropterygianLariosaurus
E Renesro l9lì1,
.oD'enÍi fhr.iicn,i
drlh Krlksrhiclcùon. (crlci.e di Mcidc,
L i n i t u \ ú p ú n r c ) d i c r . 1 r 1 r Ì r . ( v i s a i ù v, ù . r c ) . d i M ù i d . ( c r n 1990iTinroricr .1. 1985jîntor; 1990î,l990biRencsro
ÌiÌTiri,r,. Svi,/rrr), Ali J(iDoRùitji iinon po.oronosciuri/11o&?i
1993).The fish fauna consistsof five gcneraof basrl
ld6 úd1,ú, (RijpprlÌ).lrru lror,l (lilsînú Criv€lli, l3l9) e /'eràr
actinopterygians
înd sevcnof pr;mir;vc neoprerygirns
l,J.r/t,/l?d (D(r.kc. lssr), rpú.ir iriruire su ùrtcriJlc provcni.nk
(Lombardo
1997)
but rhe former exceedthe l:uterin
dìsli S.nti di I'.flcdo. pii. sorìmnxlc ddh lirmznnc di Pslcdo
vrùìrr. I nuori ri,rurrn.rri hr r, (rseùno di efldruù. h prìnr
nunber of specimens,rvith the cxccption of thc
do,trslfttr 'r.\cri,rone $rdnlct
,11A|ok?maù^ bdlùtiì c ta
neopttrygianProhalecitesponoi
(Bellon;, I 357) (Tintori
rico{n,zion. inloiii.r .onlpldr d i Pdlil,r,,1bàlr;r h prscizi d.l
1990a),
represented
by
thous:rnds
of specinrcnsioùnd
rîurido r,, n.l liir\\ico Nlcdio,prrccdcnrùn.ik ncsr in dirus
alsoin massmortrlity laycrs.Someoî rhe fish species
'nnr.. !icnc ilrrc\ì.oifcflit1. Ohrc î1lcdc\crizbniinntomiche vcn! . r . d n . u $ i r l c u n in o i n u n r c ( ^ ip m b l r m ir c h t i v ir l h n o r ì c n c h r u r i rhrt constnutc rhe Ca'del Frate faunr ivere îlrcidy
itlorr ì Jisli rúori prcc.dcnri,comepurch dìsrribuzion. rnrishfi
descr;bed
on the basisof old mrterirl collecrcdin rhc
ù dd 8d,ùc /./&t^
ch. \ì ùrnolr òdu\ivi dd ljdinico. fnfinc
Perledo
rre:,
rlong thc Ersrerncorst of Como Lrke.
\icnr d.s.i,k, un nuo\o qrDrr. irpsren.!r. rì Pù1.ìdirorin.
The Perledo-Varenna
Formarion hîs bccn known
sincethe last century for irs fossil finds, .rndin particu/rrf,?.r. Thr ic,núpr.rvliinr All,netuloù^b.Iatii l\ijppell),
/i,,, bîrii (lilìù)
CrivcÌli, l3l9) înd Pfl.ir!,r ,,1tLpr'J(Deecke,
lrr for irs verrebrare
faunr (Balsamo
Crivellil8l9i Bclr8$e),$ fú kno\n ftonr thc Pcrlc.loMenbs (uppe! menìbù oÍ rh.
lott; 1857jBrssani1886iDecckc1889iDc Alessandrì
PÚrcdo,v.ùcnnrri).m.ÍnÌ),irc húe rede$ribdùn rh( b$n olwdt1910).The studyof rhc bcttcr-presened
nraterial
from
pft\ÈnrJ mtrt.riJlconrir{ íionr rhc hc Lldinnn Kllk\chieh,onc of
'ficino,
Ca'dcl
Frrtc
provides
good
a
correlation
rvith
thc
ci dd rÌùc (viiìjiir. vfsc) î.d Msidc (c!úor
sNirzùhnl). Thc* nrs rinJ\ in rl,. Nrlkschielcrzoncfic pfriculdy impor
Perledofauna (Tintori cr al. 1985iTinror; & Renesro
hnr trccru'c rhs rLlouh( iirr d(rilcd rnùoni l dc$riprion ol
1990;Lombîrdo1997;Tintori& Lombrrdo1999).
)lbLt,i,loùt' I'd!.trii ù) ú. tomplù. inronicrl aronrion of rDcr
The comparisonbcrwccn rhc mrrerial from rhe
/dl6!r"/l?rì,hc pr6cn.. of,hc ùùrid lrb in dÈ MiddhTrirsic.
rwo units hassrressedrhrt the so-called"Perledof:u:nr'
\'hi.h hft prevnùlv b.cn querioncd, ts ùJiÍirmrd. Bcsid$ thc
rn.ronìi.rl Jc!rrit'nr\, rÌì. cotrìplì.r.rl rnd problcmrÌicrlnomùEh
acruallyconsistsof at leastthreediffercn! îsqtro,rss:
'u!c ùlo d bv pr.rnùrs iurhoÍi ld rh. \p..i.s ol rlr. Perlc,loM.ùF
rhe youngestone. coe.al"rrh L.i Jcl Frrre. i' mo'r
berir dncuxcd. Thr *r.rigrrphicildis!iburìoD of rh. gcnorP.rlirl,r
probrbly fron rhc PcrledoMembeathe uppermostp;rrt
n qur\tn .d, rì rlì. Tri.r$ic spccid pfufioù\lv isrrib.d b rhi\ senus
ot the Perledovrrennr formlion (for Jiscusion on
ùr hù. ùr\nlúrd
nor rcltrt.,l r, n. l,iluih1 r reN scnusot prn(istratigr:ìphf scc Tinrori 1998; Tintor; & Lombîrdo
1999).The two sitessharedrc besr rcprcscnrcdspccics
in both: Perleítlus dholEis, A olepidotus bellouií and
F"to trottii, whìch re to be describedin this p:rper,
Inlroducllonanclhilorical remarks
rogether with Ptobalecites
porroi (Tintori 1990i) and
The ìocrliryof Cr'dcl Frare(Visgiù,Varese,Italy) Pehopleurus
nuptìalis(Lombardo 1999),as well as rarer
bclongsto rhc Kalksch;eferzone,
spccicssuchas,4rearc1ep
rhe upperfncmberot
is naooptera (Lombardo1997i
Meridc Lincstonc (L.rreLrdinìan),which crops out
Tintor;& Lonbîrdo 1999).
bcrwccnValccresio(Iriì;a) and the Mcridc rrca (Canron
Thc comparisonwith the Perledom:rterirlis diffiùrIknf):
'r\onon,\:
noN Lr\on.
Diprdinrok) di S.i.n,c d.ìh T.trù Univ.rsni d0gliSrudidi Mihno, rìa MrnsiJcrlìi 3+, I-:0lll
Mìlino, Ihlvi c-nni1:clomhrJo(' tin.n
.1.{3
I'is.I
All,l.ùhtrcb.
,ttii lRitppcll).R.ronriÒn. scllc bú: l0 nrm.
Allolepidotushrs r narrorvrnd subvcrticalpreoperculum. Prrrsemionorids,such as W,tsoaolzs,
Pataseníonottsand Osprz,show r wide preoperculum,
cvcn i{ in somegcncra,svh asPatacottopbows,Prctne
tonnùd rnd Phakln'ona.the preoperculL,m
i. n.rrrow
Thonasinotus,
preopcrcuStezsiororrs
havc
a
lacobxlus,
îhe gents Allolepi.totu rvas erectedby Deecke lum rn.rduof 'cvcr.rl.l.mrnr.: 'hr por'criorunr i. n.,r
(1.s89)on Perledonrrter;il of rhc spccicsA. xotboso- row end contained the preoperculumsensory canaì.
noilcs. \lìrh rhis name Deeckemeant alì those forms \lhen in Prrrscrnionotidrcrhc prcopcrculun L fragwnh dcepihnk scrlesrnd squrrishvcnrraloncs
(Deecke mentedìnto smallerelements,the antero-dorsal
ele1889: ll3) in ordcr ro distinsuishthem from Herments would correspondto suborbitrls, rccording ro
úalepi.totu nd Lepìrlotas.He did not trke inro consid- Lchrmn (1es2)rnd Lchmaner rl. (lsse), and rhe prcer:Ìtionthe skulìpîftern.
opercularsensorycrml ran nearthe posteriornrarginof
Sinccrhc drra rclrrive ro skull pattern were lacking.;r rv.ì.Jifficuh ro csr.rblirh
rhe.ysrcm,rtrc
po"rrion
As in rllprrasenionotids,
rhe elementcorrespon,
of this fonn. ft rvrs usually.rnributed to Eugnathidae ding to the suborbitJ in Allolepìdotusdocsnot ovarlap
(voodwardls95i Lehmîn1966).P:ìftcrson
(1973)dis
rhe preoperculum(Paaerson1973).
rgrcerl on kccpìng Allolcpidotas wirhin rhis family
The number and arrangementof intrrorbitah nr
togctìrcrwirL othcr Trirssicgenerasuchas Eoeagzatlzs Allolepídotusis similar to thosc shovn by Watsonolus
nd Siroeugaatbu,though all were poorly known gen(Olscn 198a):thcrc are four in{morbnalsnrakìngrhe
crr. All of rhcnr"cr. ril.g.rri.l ro rhc,,,oreprirniîive ventral:nd posteriormarginof the orbit, with the Iol
grade group, the Parasenrionotidae,
togerher with thc
and lol much biggcrrhrn Io2 and I04. A vcll-expànded
frnilics Tungusichrhyidac,
ProD,ccosom;nidae
rnd Para, posterior region characterizesIol. A posrcriorly
ccnrrophoridre(Prttereon197.1:
28.1).
cxprndcd infnorbitrl in the postero-ventralcorncr of
Accordingto Pittcrson (1971)prrrscnionorids the orbit, however, is present also in rnany halecodo not have rn indcpcndcntqurdratojugal,bur Olsen
morphs and pholidophorids, but not in the
(le8a) founda smrllandsqunrish
quadratoiugrl
in Wat- parasemionotidsWrltson
al4s, Pdîasenionot,s^nd Stcnsoaol,is.In contrast,Allolepidotushrs:r splint-like sidrotas(Lehmrn19s2jOìsen1984).
qurdnrojugrl, s;nilrr ro thrr found in repriostear(lliley
The antorbitrl of ,4llolqpirlotas
hrs r widc posrcri1976;Arrît;3 Et Schultzel99l), S?ntoaotas
(Schacffer or resionrnd r moderrrelylons rosrrrlproce". rs in
& Dunklc t9s0), Ddpdian lPrrrerson 1973), parasemionotids
(Lehmîn1952iPrrtcrson1975;Olscn
Tetrasonolepìs
(Thies l99l), H,/edrd (SchrefferE{ Pît1e84), rmiiforms ^nd opbioPsis (Bàrrram leTs). An
terson r9S,+),Prcbdlecite'nd Patalepidotn (Tìrfori
rnrorb;trlwith long rostralprocess;sknown rlso;n
1990,ì,
1996).
(Patterson1975),n:ìcrosemiids(B:ìrrrîfn
Acentrophorus
Type-species:,r/l,/qrlurd
r.//.Dt (Rùppdl).
Dilriburnnì:
ividdl.Trnrsic:
L,r Lldiniú ol Perlcdo
MdÈ
b{ (r,ùrdovùùlr hnrrnnt ùnl of Kitklchiclwùn.(Múid.
Lin,c\k,nú)
orcr d.ùìIe (vi{Liùvscsc)ùrdMùidc(Tr,cH).
PLATE I
A) A ,lèt\liú"! l,.lkrùi (Rúppdl): ,ic holotlpt .r1rl P, bckft prepar*ion. Sole bilr l0 ùnri B) Allol!tuLrs'I'dlotii (Riippcll):
McsNIo
P669.S.rlÈ b.n, 20 Dnì (:) ,41/ol.tnlúrcbdlouii (Rùppoll):MCSNIO P655.S.rlc brr: 20 nrnr; D) Allohpihna bdloùìi Bnppúll):McsNlO
P615r.S.ilc bîr: r0 mmi [),llollpnlrù^ b. oîîii (Rùppell):MCSNIO P6-19ScrlcbÚ:20 nrnr.
ì / , , r i 1 h/ , ' r . , r ! ( , k t r r , , r ' t r , r r r . , , j .
1977),Seníonot"s(Olsen E Mccune t99l), Lepidote'
(Weîz t967) ^îd Huletùa (Schaeffer& Patterson1984),
though in theselatter forms this bonc is rubelike.
The rostral bone of,4/lolEidotas ìs trìangdar; in
parasemionotìds
it is tube-like, a characterconsidered
primitive. A rostrrl bone similarto that ol Allolepidotus
is present in Hulexia (Schaeflert{ Pitrerson 1984),
Catatus,Anblysenius and amiiforms (Làmbers19e2);
many pholidophorids,roo, have such a rostral bone
(Nybelin 1966; Pitterson 1975; Z^mbelli 1975, 1977,
1986).
The,ysremrricporirionof rhisgenu',r uncenrin:
rhere are too few availablcdara abour basalneopterygiansfor r comparison.Moreorea they often concern
such characters,as endocranialand endoskeleletal
patterns,which arenot availableon Ca'del Fratespecrmens.
For these rcasons,I prefer to leave this genus rs
Neopterygii incertaesedk. The systematicposition of
rhis andother relatedgeneracanbe clearedup only rfrer
.tudy of rhe mo.r primirivcTriasiicncoprerygians.
Together wirh Alolepidotk' iothosonoides,
Deeckedescribedrlso the specres
Allolepidotnsrrcppel&. The latter was previously describedby Belloni as
Pbolidophorns
ruEpell; (Bellorti in Sroppani1857:428)
on thebasisofa plastermouldtharconstirures
tLe holotype, the deposirorycollecriono{ rhe original specimen
beingunknown.Deeckeattributedthis species
to the
genusAllolEìdotus on rhe shapeof scales,deeperthan
wide on the lateralregion of rhe trunk and rhombic on
donrl andventralpattsof the body (Dcecke1889:117,
pl. V fis.s). Deeckedid nor describethe skull,that
according to him is:" ..similar to that of Lepidotus"
(Deecke188e:117). De Alesandri(1e10)agreedwith
Dceckein arrributingthis speciesto the genr:s,4//o/qptdotas ^ccord;ng to the skull strucrure, fin and scate
shrpe.He rscribedto thc.rmc 'pecie,rnotherspecimen
storedat the MuseoGeologicodell'Universitàdi Torino,
that it ;s no longeravailible.The casrof the Lolotypeof
A olePidot,'trEpe|i. storedat the Naturmuseumund
ForschunginstitutSenckcnbcrgin Frankfurr (SM
P1266),rnd a specimenof this speciesstoredat the
SeoizìoGeologicod'Italiain Rome(1918P), exduda
rhis speciesfrom the genus, based on the different
arrangmcnr
of thc skulrbones(pers.obs.).
I 1 3 ,p l .V I I I , l i g . 7 .
A olEi.lot^ belott lBellori) Oro Fn.), Dc Ahsudri, pp.
l 2 l l 2 l , p l , V I I I .f i g . L
rgta Pholidopbon6
ckrio,ii De Aìcsrndri, p. 127,pl. VIII, tis.s.
t9tÒ PhÒl;dÒpho$obl'neuJBdloÍi (rm tdre), De Ahsrndri. p.
r 2 9 - B r , p l _v l l I , f i 8 , r 0 .
1997 AlloL?Aattr notbotonoids Deeeke,l,onrbùdo pp. 9e-113.
teta
1999 A|ole?idar". nothúonoitu
Drc.ke Tinro.i & Lonrbxrdo,
Diagnosis Gnended fÌom Dc Alc$.ndri rer0): snill-\izcd
neÒp,erysiin(up to l0 cm of $andùd lcnBrh);spindhahipcd body
wnh "hùmp",nÌeior ro doral tin (juveniÌcsslcidff)r donsitc úpps
ind towf ilws vnh snr conicaleerhi nùrow preorbin rcsìon ol
Íronhì boncs;foui infiioónils vnh inlnorbiúl I beìns Lùsu nrn
orhersiopercùlumrectanBul.r.ndrwìceasdeep,srh€rubopercutunr;
elÒngÍe rnd rrirngulrr irterop4culumi skull bon.s srrÒnglt ornrmened by eÍoinè tubùdes and rid8e5which fol|o( couAc ofscnsóry cîn.ls; do6al ind r.il fins rhnsuh. viri wdr devetopedhinsins
fulcÌa; squîmi,ìon of l5 rrns!è^e rov! of rhick srl$, po*riorly
. { n E d . d c c p f l h r n w ' d eo n L l r n r r r . r i o , p r r " f r n r , \
M a l s / | a l4: . x r , h o l o ' y p e fr, o m t { 1 . J , , v r . n r , r " , r l n ì .
. r o r e dn s è r i t i o C e o l o e i c od l u l ' r i n R o m e ,M L q N I O t b i r h À o :
P669-627,P632rb,P631b, P68,1ù.P685,P688,163t,P690íiom thc
K.lkschiefcrzoneÒl Cr del F. e (VissiiL,Vareie)iSM 1237l/b,SM
P1256,SM P1244from thc PerledÒ-Virentu
Iormrtion; MCSN 31:6
froo rhe KilkschieferzoncofMùid. GI-CH)iMB 75ez (hhelled$
Alklepi.loú, aeppelh' Îrcn thc Pùlcdo-vronnr Ì Ò'm..""n.
Disribution: Middlc Triasìc UppÚ Lrdìrin Òf thc Pùkdo
Mènber (lenedo v:Ernr Formdion) ind ol ,hc Killtschicidzft
(MerideLinéfón€) ofcn deltnÈ (vi88ìù,varNc).nd M.ridr (cin-
Oescdption
5Àa1l The rostral bone is a small,trimgular clcment.on whichporesof rhe erhmoidrlco,nmis,ure
rre
visible on the ventral part (Figs. 3, 4AB, 5ABD, Pl.
1BD). The nasalboneshrve an irregularsLapeand are
dorso-venrrallyelongated(Figs. 3, 4AB, SABCD, Pl.
IBCD). They meet medially.Length of nasrl bones
equalabout1/3 of frontal boneslength.
The frontal bones are broad, wirh an expanded
posterior region and a narrower înrcrior one (Figs. 3,
4AB, SABCD, Pl. lBCD). The interfrontalsururc
showsinterdigitationsat about halfJength.There is an
embaymentar the level of rhe otbit, where the frontal
bonesconracrrhe supraorbitalbones.The supraorbnal
sensory canal mn close ro the larerJ margin in rhe
frontal bones and exrendsinto parietal bones (Figs.
4AB,sABCD).
Allolepidotusbelloltii (Rùppell)
! i g r . 2 - 6 ! P L1
l3a9 Pnlde,,,r.,r d,rú,ri H.eckel, p.500.
1357 Seùiorot,i bellorii BcllDnì,Bellori in Stoppani,p, 425.
t37] Pbotidopbow c"rioiii (Heck.l). Bellori, p.3r (nrnusipt
1339 Alkhpìdob\ noîlne,,oler Dccckc,p-l l8r pl. VI, fig-9.
188e Prolnlopho%obl','s6 Bellori, Deeckc,p. l24,pl.VI,lig.6.
t895 AllolctklÒù^ ,Òthoto,nnl4 Deecke, \voodwlrd, pin IIl.
19ra Allalcpnlrr!: nothú.,t ,uleJDecckc,De Al.s.ndri, pp. 116-
The prrietal bones are squarish,with irrcgulrr
margins.Interparietalsuture is nor clerrly derectable,
bur it seemsslightlywîly Medirn and posteriorpit lines
ar€visible(Figs.aAB, 5BCD,Pl. IBCD).
The dermopteroticsare trapezoidal,wnh rn irregular dorsalmargin.The surfrceof the bonesis strongly
irresulrrfor the pre.enceof mrny circulrr opcning'.
The infraorbitalsensorycanalran alongthe ventralm:rrgin,enre.insrheexrrdscaspular
(F;gs.l,4AB,5ABCD).
The extrascapulars
are large,trrpezoidarcrcments,
deeperthan the parietals.The supratemporalcomnis
151
s
MìddleT/;dst actìî oPterygìan
sure is paralleledby a series of small spines (Figs.
SABCD,Pl. 1CD).
Two etongate,rectangularsupraorb;talbones are
present;they forfn rhe dorsalmarginof the orbit (Figs.
l, 4AB, 5ABC,Pl. lBD).
The dermosphenoticsare small elemen* whose
mrrgins rre not clearlydetectable,owing to rhe stateof
(Figs.3,aAB, sAB). The
presenrtionof all specimens
infrrorbitalbonc' .rrefour element.of differentsize..
They border the posrero-ventralmargin of the orbit
(Fi8s.l, 4A, SABCD,Pl. IBD): Io1 is large,with a rhin
bony lrrnella dorsal to the infraorbital sensorycanal
Inrrrorbir.rl
2 rs smrllerrhrn lol. The bonei. rectLrbe.
ventrrlly
meetsthe supramaxillaThe rhird
rrngularand
infraorbial bone is the bisgestelementof rhe se.ies;it
is posteriorlycxprndedand n forms lhe Posterovenr.al
cornerof the orbir. Many spines,following the courseof
the posterior ends of the infraorbital sensory crnal
brrnches, characterizethc surface of this bone. The
foutth elemenris the smallesrand is deeperthan wide.
Al"ng rhe dor.rl edgcof .rll element.ot the 'eries;re
derecrrbleporesof rhe infraorbitalcanal.The antorbital
showsr moderrteìylong anreriorprocessand a broad
poster;orre8;on(Figs.l, 4AB, sABCD, Pl. 1BD).
The prcoperculurnis subvcrticaland naruow.The
surfrce of the bone is irregular,with small spinesthat
follow thc courseof preopercularsensorycanal.Pores
are scatteredalong the longiludinal axis of the bone
(Fiss.5ABCD, Pl. 1CD). The operculumis higherthan
brord and twice as deep rs the suboperculum(Figs.I,
4A,5ABCD, Pl. IBCD). The dorsalmarginis cuned
whiìe rhc ventral one conractsthe suboperculum
i. rectanguthrough.,con"er.uture.The "ubopercuìum
lrr and about twice as broad as high. At the anterovenual corner of the suboperculumthereis a long rnd narrow triangularinteroperculum.It showswell-developed
grnoineridgeson irs postcriormargin(Figs.4A,5ABC,
6, Pl.1D).
The maxillais rarherIong and narroq with à peglike internalheadrhat articulateswith the premaxillaand
slighdy wary oral and dorsal mars;ns (Figs. 5ABCD).
The mrxillary posterior bord€r showsa notch (Figs l,
5ACD, Pl. 1C). The oral border of the maxilh n almost
strright and bears about thirty snall, conical teeth.
There is a lrrge supramaxìlla(Figs. 3, aA, 5ABCD, Pl.
IBCD), extendins frorn the postero-dorsalcorner of
the maxillafor almost the 2/3 of its length.There is a
srrong,ttiangularprernaxilla;it bears8-9 conicalteeth,
somewhatlargerof rhoseborne by the maxilla(Figs.3,
4A, 5ABD, Pl. IBCD).
On most of the specimensrhe qurdrate,few elementsof gillarchesand the splint-likequadratojugalare
visible(Figs.l,5ABCD, Pl. 1D).
The lower jaw ìs a slrong element,dighdy longer
tfranthc maxilla (Figs.SABCD, PI. lC). The oml margin berrea scriesof conicalteerhsimilar to thoseborne
'fn
Fis.4
A(olepùlotrs
bellottii(\'rppèll).Rsronrionof rhèlkullin
htúl (^) rnd in dorsal(B) ù€ws.
by the maxilla. The ptecise number of teeth ì. "ncleat
since the oral border of the maxilla always covers them
Pores of the sensory mandibular canal are visible on ventral margin of lower jaws (Figs. 5ABCD, Pl. lBcD).
Up to 10 branchiostegal nys were counted (Figs.
3,4A, 5ABD, PL ICD). Th€re is a median, rectangular
gulaa with a V+haped pit-line visible on the middle of
rhe bone (Fiss.5AD, Pl. 1D).
The posttemporalis a large element posteriorly
placedto the exrrascapular.
The two posttemporalsmeet
Pores
of the main laterrl line are
a
short
suture.
along
visible on the ventral margin of the bone (Figs aAB,
sABCD, Pl. lD). The supracleithrumis qùite hrge,
dorso-ventrallyelongatedand parrially coveredby thc
operculum(Fiss.3, 4,A.,5ABCD, Pl. lD) Therearetwo
posrcleirhra;the uppermostone is the biggest,rectangular,anddorso-ventrallyelongared.Belowthis is a second
by many grnoine ridges
squarishelement,characterized
arrangedperpendicularlyto the posreriormarginof rhc
bone (Fiss.l,4A, sABc, Pl. ICD). The clenhrumis
only partiallyvisible,sinceir is coveredby the opercular
region (Fiss.3, 44, 58, 6).
Fins. In all specimensthe pectoral fins are only
partiallypreserved,showingtheir proximalbases(Fig 6,
Pl. IBCD). Up to 14 lepidotrichiawere counted.The
lepidotrichiaare narrow, and well-developedfringing
fulcraborder the anteriormarginof the fins Pelvicfins
are madeup at leastof l0lepidotr;chia They ire placed
anteriorly to the dorsalfin. On the anterior border of
152
Exsc
l1g.3,
-Allollpnhtr btllatit (Rùppcll).Skullboncsfu prèsded in sp(imcn| A) MCSNIO p645ij B) MCSNIO p66oiC) MCSNTO p6s/:
D) NICSNIO P6s5.Scrleùm:5 nrnì.
thescfins rhcrcis r ser;es
(Fig.3, Pl.
of fringing-fulcrr
1BC).
The dorsalfin is rriangulrr and placedat the middle of the bodv (Figs.l,6, Pl. IABE). It is mîdc up of
ir lcasrl6 srronqlepidotrìchia,
with longproximalbases
and short distal scgnrcnrs.Each ray branchesar lcisr
rhreerirnes,beconing very thin rr the distalend.There
are Nvo to thrcc bisal fulcrr and a seriesof well-dcvclopedfringing fulcn alongrhe anteriormargino{ rhe fin.
Thc rn,rlf;n ;s similarin shapcandsrructure
to the
dorsalonc,but it is snraller(Figs..l,6,Pl. IBCE). Two
to thrcc brsrl fulcrr precedethe fin that is mrde up of at
lenstI I ìepidotrichir,
whichbnnch threetimes.
Thc crudrl fin;s hemihereroccrcîl
wìth a short
body lobc. It is decply forkcd and it is madeup of 21
tays, which brrnch ar lcrst four times. Thc axirl body
lol,e shorvss;x-scvenlrrge basrl fulcrarnd rhc firsr dor,
salrry is borderedby fringing fulcra.The rentral lobe is
preced.dby: couplcof brs.rlfulcrr rnd .hou\,ì "crie\
of fringing fulcralessdevelopedthrn thoseborneby the
dorsallobe(Figs.l, 6, Pl. 1BCÈ).
Sq"antatio,.Sc le co\eringis nì.dc up of ls rrans\rr,u rosî ol rhick sc,:Ies
with strr.rrcdpo.reriorm:r(Fiss.
Pl.
3,6,
IABCE).Scalcs
aredeeperthanwide
sins
on the lrt€ril rcgion of rhe rrùnk; rhe;r depth dccrciscs
graduallytowrrds the posterior part of rhc body. The
scalesof the first two rows bchindthe cle;thrum:rre
much deeperthan widc; thcir posteriormargrnrs scrrrted. Thc sizesof scrlesdeoeasetow:rrdsthc dorsrl rnd
ventral part of the body, rhough they iever teconìc
widcr rhan deep.The scalesbearingrhc hreral l;ne are
slightly deeperthan arc thc orhcrs.The dorsarscrlesxrc
asdeep:swidc;r dorsalscalerow between
the postrcmponl rnd rhe dorsrl fin showsthe porcs for thc prsage
of rhe dorsallateralIine (Pl. lD). Posreriorro rle dorsil
fin, scalesarc snrller and rhombic, with two or rhrce
serrationson posterior mrrgin. The scalesot ventml
region bctwccnpecroraland pelvic fins of th. """y -.
vcry small.Posteriorlyto thc anrl fin the ventralscales
are equal in size to rhc orhere.The axiîì body lobc is
coveredby snrll rhombic scnles(Fiss.l, 6, Pl. lBcE).
Taxonomichislory
The taxonomichisrory of A olepi.lotu'bellotíi is
very cornplicrtcd.
Heckel(18a9)quotedthis spccrcs
as
Pahconi*u',u,ionu on the br.i' of .: ,pccimcnco,ning
fron Perledo;however,he gave ncithcr an illusration
M ì.Idlc Trid$ ì c dctìropte1Aidns
353
#
l1:. "
tll l.rtJi\. t, 1l''rr.RJ$!l
\ p c . i r r c nM ( \ N I o l b r _ . r c . r l ab r ' : l 0 n r m .
nor a picturc of it. Afrcrrvards,on the basisof a plaster
crstot Heckeìsspecnnen
(oncestoredat MuseoCivico
di Stori:rN:rrurrlcdì Mihno), Bclloni (1873)gaver
dcscriprìonof thc spccics,in his Caralogomanoscrxto
deipescìfossilì lcl MuseoCi.uìcod; Mìlano \which was
ncvcrpubl;shcd),
but hc asoibedit to the gcnusPloliThc same spccics hrs bccn nrmed by Rùppell
Scnìorotts bcllottii, wirhour giving any published
desÙiptìon,
on thc basisof a singlespecimen.
Se'rlozons bellottii \as nor bccn cven circd, before the formal
desoiptiongivenbv Bellottibothon Rtlppcll\spccinrcn
antlanotherspecinrcr
fronì rhc Curionicollecîion.
In a
lootnote,Bellottis,rotethat this description
wrs nade
togcthr with Rùppcìlduringî vìsit to Mil:ìno (18s7:
125).Accordingro rhc InrcrnrtionrlCodcof Zoologìcal
Nomenclrrure
Riippellhasto be considered
the author
of the spccies,even if hc cl not fornrally dcscr;beit
(Art.50.l.l).
Deccke (188e),in his rcvisionof rhc Perledo
tauna.
brscdonly on spccimens
ofthe Rúppellcollection
(sceibo!c), crecredrnorher species,
Allolepidotus
nothoso,rotler,without rcrlisingthe conspccificiry
of rhe
spccimcnsLc rscrìbcdro Lis new speciesand those
descrìbcdt Pholídophorus
cqrìonii andSmionotusbet
1oÍii. ln r succcssilc
srudy,De Alessandri
(1910)agr;n
did not noricc thc simil:ìritybctwccn thc diffcrenr
species
erccteilon thc brsisof thc rwo collccrons,even
rhoughìrc had thc possibilitvro rccessto rhe marer;al
Èom Mil:rno :rnd Fr:rnkfurt.As:r consequcncc,
hc
rcdcsclibcd rll thcsc specicsrs scprrrre raxr: Póo1i
,lophau arìonìì, Seniorotus belloxíi nd Allolcpìdotus
,othoso,ùoidcs.
Conccrning Pbolidopbotusc"tio";i, De
Alcssrndr;
contirnrcd
rhe mxononicrssexmentof Belìotti, providintr nes,description
:rnda pìctureof the
cxstthîî constitutcdrhc holotypcof rhc spccics(bur
rvhich rvas losr durirc rhe Vorld \lar Il). However
judgingfrom the dnwing in thc Bclloni'smrnuscripr
rnd frornthc picturcin Dc Alcssandri's
prpcr( I910,trr
VIII, fig. s), ir crn be confìdently excludedfrom the
gen\lsPbolídopborus
úd ît the sînre rinrc confirnrcdas
belonging
to the species
currentlvunderstudv
In his revision of thc Pcrlcdo n'rtcri,rl, Dc
Alcssrndri (1910) redescribedttre spcciesSe'riorotzr
,e11on;i,attr;but;ngit to the genus,4llolepirlotrrbec:rusc
of the skullpatternand shrpe.The specincnon *hich
Dc Alcssandribascdhis dcscriptionand revisionof
AllolEidotus belloxìi belongsto the Curioni collcction.
D c A l c ' s . r n d rc! o n s i d c r c di r r o b c r h c , p c c i r r i n
desoibedby Belìoai(1357)(th;sspecimen
's stillîviilrblc, bcingstorcdrt Scoizio Gcologìcod'halia in Romc
(aa31P)). Finrlly,De Alesandri c;redrnd descrilxd
Allolepidotu notbotortorderbut he bel;evedthis spccics
wrs lackingin thc collccrionof rhc Musco Civico dì
M;lano, beingclearlydiffetent from the others.
Thc dcscriptionsand thc picturcsof thc holotypcs
oî Pholidopbuuscariozii (De Alessandril9l0: l2z pl.
VIII, fig. 5) and thc comp;rrison
rvith the holotypcsof
(SM
Atlotepidotus,otùosonoides
1237i, b, pcrs. obs.)
^nd AlolEidotu' beltottii (a31 Iì par. obs.) conf;rnr
the correspondence
between thcsc rhrcc nominrl
species.To rhe srrne speciesseemsto belong also the
specirnenthat hadbcendcscribcdand figurcdby Dccckc
$ Pholidophoruoblngur (Deecke1889:12a,pl. VI, fig.
6) andstoredat the Naturmuseunund Forschunginsritut Senckenberg
in Frankfurt(SM 1256ipers.obs.).
\liLhoutr doubt.thc C.i dcl Frrrc'peci,rrcn'
cor
respondboth to the holotype of A//olepidotus
notbosono,les rnd thrr of Allolepidotusúei1orii. ìn prevrous
studiesof the C:' del Fratematcrirl(Lomtardo 1997i
Tintori & Lombrrdo 1999)rhis specicswrs circd rs
AllolEidotus nothosomoides,
for thc conrprrison hrd
beenpossibleonly wirh the holotypeoî tha! species.
The presenceof the hoìorypeol Allolepìdotus
betlottìiin
154
thc Curioni collcction at Scrvizio ceologico in Rome
wrs discovcredlatcr. Moreovcr,rhis specimcnwas not
prepared
andthe skullpatternwashidden(Pl. 1A): only
aftcr prcprr:rtion brs it been possibleto confirn the
with the
determin:uionand to verify the correspondence
Ca'deì Frarc spccimcns.Alolepidotusbettottiiis thcreforero be considered
the "rlid nrmeot the "pecies.
Halecomorphi
F:ìmily Caturidae,Owen (1860)
GcnusF,ro Gistì, 1848
Diagnosis (fro \vùodsrd l39s:236-290):Tiunk clonS cfusiforn,. ti\rerntrl bone! leebly omimcnted wirh ùbùculîions o.
rusúi tÈtlì rch.ivel) hrse and specedor rhe denúry, smallerand
i1ùn dosdr'únnsùl on,h. mÍ8in ot thc upps ji$1 ninu'c and
dnìo$ snnular on rhe inner bone\, in more rhrn one scriesovcr rhc
Nhoh ol,hc sptcnirtiprcopcrcutu,îsn,oorhand nùlow; subopùcu,
lu'n of nodeúÈ sizc, lboú hilf rs ltrryer the ncirly.ectnsuh.
ú l r c u t u , , r n . t \ . , h , , h o , , s . c n d i n Fp r o . s . r , h r n , c r o - u p c r i o r
rnsle.O\\ificriion in ,he sher,hoI the ftro.hord obsmd only in,hè
l.rsN, spccìcrìo$ificd ribs sÌcndci Fukn bisrirl. Nclldevdopedon
rll thc fiis dftp, rh. poddhl!, on shìch rhcy rc lc.blc. Pcdo6l
n,uch scecding thr pelric iins in sizc,hù thc hr.r kll devolÒpèd!
dorsil.id ùrlfiis rrirnsuhr, the fornEr ùi in8 opposneor immcdi'
rdr behindrhc pdrn iinr; ciudil fin fo.lcd. sciles,hick,wnh i nrroNorcrhpfcd bordù,rn innr rib, ud r fcebh peg androcke rtic
uh,bni rupeli.irl $noinè rtuoÒ,hÒn rhc rnt.rnr hlltof erh srle,
p6sìnsonpùtolrh.bodrirtorÌinsvùsrusrerndrs,uhtionspos
rcriÒrly;principrlfhnk scrhs rf.ll ind rh.i onlv in p.n, dccpcrrhan
brord, scvcrrls.rie\ of v.nrrl scrks much brord€r rhtrndcep;pordrricuhÌ sùlcs hrgci ro cnhrsed scàleson thc dos ridgeor in tùe
rqlion dt thc inùr. LlRnl lìnc inconspicuous.'
.|ypè-spacies:
l:no onbonoù !J iAsr$iz, 13l:l .14)
oislibution: Middh Trirsic of Ikly (cn dd FÌ.e, Pùlcdo)i
Lo\'( Junsic oI f:'rshnd (Dorershner Lyme ResisiLf,ice$ershnei
vlr*icksh;c: vhnblr Yorkshirc) ind Fnncc (vilzr Lozèrcj NoF
mrndic: L Cxind: Uppú Junsic of lÌrnc. (Ain: Cdn) rnd G(mnr (BrÌsil: llichsift rd solnloÍcn).
The genusFrro waserectedl,y Agassiz(1833-a4)
x Eugnatbustor narerìalthe Lower Lias of Lyme Regis
and Kimnrerìdgirnof Bavaria.Sincethe nrme Eagatúrs
had been used ro designatea different animal in 1833
$flhne & Moy-Thomas l9a0), the gener;cname,Faro,
proposedby Gistl in 1848,hasbe€nreconsidered.Agassiz includcdìn the gcnusfa3zatlas fisheswith rn elongatebody,,strongdentition and scalesmore or lessornamenrcd and pectinatcd. \loodwrrd ( | 8es) distin guished
six speciesfrom the Ensl;sh Lower Lias (Eugnathns
oúbosto,ùus,
E. philpotae,E. ninor, E. senatus,Eugnatùus
ahus nd Erytathus hastìngsìae)
úd three from the
Kimmeridgianof Barar;a(Erynatbuslongienatus,Eug
ndth"s nicrcl.?irlotus, Eugnatùnstatínanxs). The gents
wasreportedby De Alessandri(1910)in the Latel,adinirn of Scistidi Perledornccordìngto De Alessandri,the
genusis representedin this unir by two species:Eag
ndîb"s bernesì nd Eugnatbust/oÍti (De Alessandri
l 9 l 0 :9 l - 9 9 ) .
Lateron,,Earohasbeenreponedìn rhe Upper Lias
tt
fim tdtit (Blsno Crirelli, 1319).Rr*or
s k u l l i nh k n l ( A ) r n d d o . ú l ( B ) v i e w r .
ion oI rhr
with the speciesE,gnatbasbtou"olal,ali (Arambours
1e3s) and in the Kimmeridgirn of Cerin (Eugnatbus
praelongus,îhollìère 1873; Sainr-Sainc1949). \lcnz
(1965)successively
Futononnandica,
€recredthe specìes
from rhe Upper Lias of Norrnrndy.
The specimenof Ca' del Fratecorrespondsìn thc
generalshapeof thc body and in the skull patternto one
of the two speciesdescribedby De Alessrndri on
Perledo material ()9t01 97-99)t Furo ttottiì. Tho\tsh
incompletelypreserved,this spec;menshowssomeskull
featuresthat are typical of the genusFrro and gcncrrlly
of caturids.There are asymmetricJparietrl bones
(wh'ch arealso presentin Futo ottho*omus,Fwo ninot
andFuro elongatus(Saint-Seine1949:fig.69)) showing
an ìnterparietalsurure srrongly interdigirarcd,visiblc
alsoin Fwo nornandica (Wenz1967:fis.71).The surure
bctweenparietalandfrontalbonesh irreSulrr,lvith a lateral processrnterior to rhe surure with thc frontrl
bones,which is alsovisiblein CaturasandAnb$senìrs
(Lîmbers 1992r150).The presenceof irregularparietrl
bonesis reportedalsofor\Udt'on"l"s (Olseî 1984ifig.2)
so rhat rhis charrcrer is considercdprinitivc for the
(Lrmbers1992:151).Two largcsuborhalecomorphs
bitrls are typical for Caturidae(Venz 1967:165);they
rre presenr,for instrncc, iD Farc onhonom"s (rJ/oodw.rrdt89s1andCara'erlo/rearRrynerl94l r. The preoperculum is straight rnd subvcnicrl rnd is scparrtcd
from the skull roof by the upper suborbnal,rs in Faro
nomandícd (\tenz 1967: 166).
Thcscfertures,togetherwith r squrmationsimilar
MiL
e liìd$E
adììopî"rr9;ahs
355
nasal(Figs.7AB, 8, PL 2A). The {rontrl bonesarerathcr
short with a narrow mterior region and . brorder Poe
terior one. Their anterior nrargin is irregulrr rnd the
interfrontalsuturcis:lmost srraisht(Figs 7AB, 8, Pl.
2A). The supraorbitalscnsorycanalruns rìons rhc lxteralmarginof rhc bones,enteringrhc pirietîl bones.
s'ith irrcguÌar
The parietalboncsrre quadrangular,
suturcslighhyinterdignrting
marginsrnd interparietrl
There is r short întero-latenl processon the xnrenor
prrt of the bones; on the;r surfacerhc pores of thc
supraorbnrlsensorycanrl rrc visible (Figs.7AD, 8, Pl.
2A).
The trapezoidrl dernoptcroticsare clongrte
At the levelo{ thc contîct betwccn
antero-posteriorly.
thc parietaland the {rontrl the dermopreroticsproduce
a short antcrior process.Thc poresof the suprrtcnporal canrl are placed along the ìnfcrior nr:rrginof thc
bones(Figs.7,4B,8, Pl. 2A). The extr:rscnpul,'.tt ti- /i,? ónd;i (BrlsùÌo Cri\rlli. lsr9). Skull oI spccimen
!ig.8
rngularandlrrge(Figs.7AB, 8, Pl. 2A).
MCSNIO P.rí6. Scrl. brr: l0 nrnr.
Thcre are two lrrge suborbitalboncsrnil r third,
much smaller,placcdbetweenrhc first two. The dorsal
clenrent,the largcronc, lies below thc dcrmopterotrc:tt
to thrt dcscribcd by Voodward in the diagnos;s of the
Thc shaPc
is slightly quadranguhawith round DrarsiDs.
gcnus, makes possiblc thc ;utriburion of the Ca'deì
of the secondone is moreircgulrr rnd it contrctsthe
Fratc specimcn to the gcnus Irro.
preopcrculumat its posterior matgin (Figs 7A, 3, Pì
2A).
Furo trottii (Bals.mo Crivclli, 1819)
small(Figs zAB,8, Pl.
The dermosphcnotics:rre
l : i g \ ./ _ 9 .P l z A
pores
infraorbnrl
sensorycanalatcvist2A); rhe
of the
ble along thcir posterior and postcroventralmargiDs.
liJSe /,fnld,i óa{tr orLrrn$ Crnclli. p. 7
Thc infrrorbiul serics is badly prescrvcd:there is r
(lhli'ro Crndli). B.llotri in Sropprni,p.
Sc"i."dtJ.rù"
tliT
dorso ventnlly elongateelcnrcntborderìngthc posteri13t5 tr?nloùJrr?rtr thl\rn Crn clli, \voodwrrd,pi( lll, p 64
or prrt of the orbit rnd two to thrcc lrrger elenrenrs
rîort (Brlsinú Crnclli), D. Alcsind.i, pp 97_99,
l91o lrsrrr,r
making rhc vcntrrl orb;trl re:ìion. A nanos', slightly
curvedelemenrwith tracesof the porcs of the infrroriigs.89-91.
199l r1,,.nnii (hl\rnù Cri\ clli).lînìhùdo, pp l20_110.
bìralsensorycanal,ìs intcrpretedas thc ànrorbiral(F;ss.
letr, li,n rrorii (Bihnn Crirelli), Înkri e Lonrbùdo, pp. 495_
7AB, 8, Pì.2A).
50.t.
The preopcrculun is subverricrl,*ith r narrowcr
Malerial:MCSNIo P.156fronì .lìc (ilkshicf.ùon. (Mclidc
dorsalregionand a venrrrlone sligLdycxpmdedand
Linlcronc, U|Pc! t.r.linnn) ot ci dd lÌIc (vissiù. vrrcsc) The
forwardly direcred.Thc surfaceis smooth (Figs.74, 8.
hoknrp.. ror.d r Nlus(, cnì.o diSrorii NÍùnlc dìMihno. hrd
regionis lrrge,wirh a senrcrrcuPl. 2A). The opercular
brctr lorr. dufins\i'orl{l \V.r il.
The subopOishibution:Uppc! Lidi'ìiin {l thr Scnridi I'cllcdo (P{ledolar outline.Thc operculumis qurdr:rnguìrt.
vÌrnnl lirnn,iù,
ùìd ot,ùc Krlkschicf.rone (Meridr Lin.ionc)
crculumis abour1/l of the depthof thc operculumrnd
o f c l i . l l i r r e ( v i s s i i ' ,V i r c s o ) .
shows n short antero-dorsalprocess(Figs. 74, I, Pl.
Diagnosis (oncndùl ironr D. Ale\srndri l9l0): Dody elon
shouldhavcbcen
The triangolrrinteroperculurn
2A).
grcjù.ildnt
iron l bonN inr.riùllt con$icrcd ind cnhrgcd Poe
on thc shb.
only
rs
impresion
bcing
represented
short,
kn*rnd t'loidIntorbnil nrrùwr ìor8 irw\ vnh
rutul'rnull'".*
rntcri"r
visible
onll
rhc
n.rrrow
suboóihls
$i,h
$rlhr
oDe
ber*.tn
is
Of the m.rrrll,r
rloDc coni.rl ,o.rhr No lrrg.
rl.n; \1u11bonN on,iDr.Dt.d by Jrnr'lv spictd ruberclsj (ilos
region becausein rhe only îvliì:ìble specimcnir rs Pîr
{leúpù rhin bro bchindcl.irhrun1.brùidcÌ rhrn decpin nlid_Pos,o_
tially covercdby gul:r bone and brrnchiostegalrays;its
r n r r u g i o ro f t u n l . n r n N v c i , n l l ) i d t e p i r n l b o d y l o b t i N d l
orrl
marginbcarsn seriesof strong coni€altecrh (Fìss
d.vcl()trd hinginir lulcli on mternr nnrgin ofrlllins.
7A, 8, PL 2A). Venrralto the antcriorproccssof thc
visible,berringrccthsmrìlmax;lh,r smdl premaxilla;s
Description
is rcprcscnted
by î sinslcincomPlete cr th:ìnthoseborncby mîx;lh (Fìgs.74, I, Pì.2A)
This species
Thc lower jaw is a strong clement,whosc vcntr;rl
rvìrhr SL of t0.acm.lts skullhckspan of thc
specnren
rnarginis dighrly concave(F;es.74,8, Pl.2A). Thc
ofothers.
dcrnrlbonesandshowsdisphcemenr
antcro-venralregionis chrncteriz-edby thc prescnccof
dorso-vcntrally
A
lrrgc
bone
rectangular,
SÉ,ril
oval opcningsrnd alongrhc vcntnl marsin rhc Poresof
is
intcrprctcd
rs
rnd visiblefrom the inside,
elongrtcd,
156
Df
Iìi*,)
- /r,r r?rr
(llrlînrd (r'ir.lli, lslt). Por.rnr pr* o{ thr bodl o{ spccimcnMCSNIO P.ls6.S.ilc bir: l0 DD.
have bcen countcd;
Howevcr,;rbout22 lepidotrichia
thc nundibulrr crnrl rrc visìblc.Thc oral bordcr of thc
le.r'rrhrccri,rr^. Tht.rnrcnornr.rrgin
e.rchbr.rnche'.rr
boncbearsconicrltecrhtriggcrthanthoscof the muilla.
Thcrc is.r l;rrgcsìrieldlìkc qulrr.rvith a smooth of thc body lobc is borderedby a seriesof lons ind welì
developedbrsrl {ulcra; rhrcc basrl fulcrr prcccdethe
rrfrcc (lìigs.I, Pl.2A). Up to nine eìongatebrrnventrrl lotre.Thereareeìongîteand narow lringing luìchiostesrlrrvs hrrc bccn countcd.Their surfaccis
crr on thc n:arginsof both dorsrl rnd ventraìlobes.
srooth (Figs.7A.8.Pl. 2A).
Sqtanation. Thc singlc spccimcnof thh species
All thc skullboncs,if not othcrw;scspec;f;ed,
îre
ornrmcnrcdbv srrongrnd rounileildcnscly,rrr:rngcd showsonly the scalecovetingof the posterìorand caudal reg;onof rhe trunk. For rhis rcrson it is impossible
grnoinetubcrclcs(Figs.s, Pì.24).
Thc posncmporrls
arcrcctrngulrlthcscelcmcnts ro stîte the nunber of trrnsversescrle ros's. On thc
s;zeînd shrpeare ,rnteriorreÍtionof thc rrunk only fcw sc:rlcsrrc visibìcr
rrc incomplcte,
so thît their prec;se
they aredeeperthanbroadimnrcdiatclybchindthc suprunclerr(Figs.zAB, s. Pl.2A). Thc suprrcleithrunis:r
bur rheydecrcrsc
rr szc
lrrcc,subrcctrncular
clcncntrvnhsnoothsurfrcc(Figs. acleithrumrnd postcleithrum
7A, s. Ì' Ì.lA). The postcìeithruùis rlso r lrrgerectrn- posteriorly(Figs.9, Pl. 2A). Thc scilesof the lîterîl
posterror
sìrh.r genrlypccrinrted
m:rrgrn. regionof the fl:rnk,rt the levelof rhc dorsalrnd anrl
sulrrclcmcnr.
(F;gs. fins, are broadcrthm decp.The dcpth of thc scrles
The clcithrunis sclì<leveloped
rnd sickle*haped
decre:rses
on the dorsaland ventr:rlrcgions.Ar thc brsc
7 4 , 8 ,P l . 2 A ) .
of rhc anrl fin, scrlcsrrc nrrrorv.Aìl scrlesh:rvesen:ued
A,rr. Thc lins .rrc prcscrvcdonly in prrr, rs the
posteriormargins,cxccprùosc covcringthe body lolre.
req;onof the
spccinenis hcking rhc ;rntcto-ventrîl
with ,""1;..,"'.
by rhombicscrles,
trurrk;úc pcctoralrnd pchic finslc thelcforcnot pre- The ìrtter;scovered
gitudinrlrxcsrthey coverthc dors:rllobeof thc caudrl
The dorsrl fin l,rcksthc most :rnteriorlepi- fin for halfof its lensth(F;gs.e, Pl. 2A).
dotrichi.r:
it h:rs;ule:rstl5 r;rys.rvirhloneproximrlbascs
(Fiss.e, Pl.2A).Fdnsinsfulrnd shon disrrlsegmcnts
Taxonomichistory
cn rre nor prcserrcd.bur thcìr prcscnccis dcduciblc
lron thc impression
lelt on the rock. The rnrl fin ìs
llrc
BrlsrmoCrivelli(1819)erectedtle specics
opposirerhc dorsrlonc.k is sm:rlìcr
th:rnthe dorsalfin .rot.ii on Perledo mrrcrirl. Thc xuthor iscribed rhis
rnd ;t ìls rt lcastll raysrthc distrl cnd of thc fin is
speciesto the genusLePidot"' brî he pror;ded nc;rhcr
(Figs.
nrde oi delic.rre
rnd shorteìcmcnts
8, Pì.2A).
deraileddescriptionnor an illustr:uionof thc specrnen.
A hrgc dorsrl lobc chrmctcrìzcs
thc caudalfin
Bellofti(1857)studiedagainthe speci,ìenol Bîìsanìo
(figs. 9, Pl. 2A). It is not possibìeto stîre rhe exrcî Crivcllibut hc considcrcd
it nrorcsimilarto thc genus
numlxt of rhc lepidotrichi:r
bccruscof r fold that Senionotus5ecttse of thc shrpc of rhe body rnd rhc
div csrhc dorsrlregiorof rhc fìn fron:thc "cntralone. positionof the fins (Belloniin Stoppani18s7,ll). Dc
M ;thllc Ttia$ i c dcrùt opî eD'sid,'
(re r0) n,adeanotherrevisionof thisspecies:
Alessandri
he included it into the gcnusÀzgr,rtlrasbascdon the
ornîmentrrionof rhc skull boncs,thc scrrrtionon the
postcrior marginsof thc scrlcs and the narrow ventral
l9l0:99). Hc rlso Producedlhe
scrles(De Alcss,rndri
firsr pictr.rrcof the holotype.
Thc comprrison betwecnd,c specimenfron Cr'
del Fr:uernd thc holotypeof Farorortii is possibleonly
and the
of De Alessandri
by mcrnsof thc dcscription
losr
durfor
rhis
litter
h:ìs
been
pictureof rhc specinren,
inE rhc world \ftr Il. Ncvenhelesrhc t"o.pecimen.
ro corrcspondin body shrpe,skull patternrnd
seerns
ornimentrt;onof thc bones,scalesshapc.positionof
rhefins:rndshrpcof rhc crudrl fin.
Thc other speciesof Pcrlcdo rttributed to this
genus,l}o /tcnrcsi, is representedby r singleincornrvithacrrshedskull;theinclusionofthe
pletcspccimen,
C:r'deì Ftrte spccincn in thìs speciescrn be excludedon
wcllvisibìe
on the photogr:rph
thebrsisofthc c:rudrlfin,
ofthe holorypc(De Alessrndri1910,r:ìvVfig.2),and
rhc conclusnclylcssdcvclopedfulcraon the anterior
mrrgin of the rn.rl iin and on both lobesof drc caudrl
Bcrg,1940
Ordcr Perleidiformes
Brough, 1931
Frmily Pcrleidid;re
GcnusPc,/cnlasDe Alessendrì,1910
Dlagnosis (cnrùnl.d lron Dc Ahssndri lel0): Sn,rll ,o
n,odioùFsizcdf.'l. itùÍ inh$ Nnlì fu\ilom bodt; ri\ilh vnr
rrro{ rntrrnr fugion rnd crpnndcdPorrrior onq Pcg like rccth on
onl iisr ùd lrr{{ onó on pilrrl boDsi propdcolùm (ith vide
dorîl 'úsn)ni NidL opcrculrr rúsion Nith dPrrculunrslisùrly snilltr
rhln rhc 'uùopd.ulur: (ildrl fìn \yith 6 or z tpiriil rrlsi lcPi
doùid1n of rll li,I Nni lms ptu\iNl bÀrs Lrtcril mnk scilcs
nudcrr,clr high ind decr.r dÌn s'idc; rll$ilN sùnted.
Iype-.pecies: Pùhrl,t tlblcp; (D€c.k., I 33r)
D s l r i b u t i o nM: i d d 1 . î i r s ì c : U È p . r l , i d i n i ! n o fP . r l c d o M c n !
bù (l'.rlcdo VRnN FornÌrnn, úd of Kilkschiefcrzone(Meridc
LinrAiÍ,nc) ofCi dd l'îrc {Vissii,-VÙ.rc)rrd M.ridt (CÍton Tici_
wrs crcctcdby De Alessandri
The gcnusPerleidrrs
(l9lo) on nrîter;rl coming from Perledo,previoudy
dloicP; by Deecke(1889).Thc
descrìbcdrs Sanriozotar
of Cr'del Frrte
ncw finds in rhc Krlkschieferzone
ro
the
type speciesand
allos,edro a cerr:rin:utriburion
rhe first cornplcrern:rtomical resrorrtion of it This
rcrverls the incornpatibility of many specicsso far
ascribcdro the gen'.s Peieiclns.Severalauthors hnve
crccted specics of Pcrleìdusfron Errly Triassic of
Spìtzbcrgen(Perleirlaro'oorlodrliStensió,1921),MrdîPìlctcau, l9l4i P?r/ei
gascar(l'erlei.l$ natlascatíensis
d"s pieeter i Lehrnan, l9s2), Grecnltnd lPeieidus
*oscliezvs Srensiò,19.12),Angola (PerleìdusIutousis
Teixeirî, 1947),China (Pe eidusjangtze,r,r Su, 1981)
rnd Crnrda (ct. PerlcitlusSchaefferet Mansus, 1976)i
from Middle Tr;es;c of Spiî' (Pertei.tsssìsa tcu', Pct
leidaslidi Rek!l].,1972),Souù+vesternTurkcy (Perici
sp.,
&s sp., Beltrn ct xl., le79), of Fnncc (Pcrleidas
Mazin E Mîrt;n, 1983) înd the Uppcr Trirssicof
Morocco (cf. Perlclas Mrrt;n, 1982)
ro thc genus
Thc attributionof rll thesespecies
hrs beenmrdc only on the skull pattern,of primirivc
rype,and often on very poorly prcscrvedmaterirl Nevenhcless,
the revision,nostly b;bl;ognphicrl,of erch
spec;esreverlcda differenrsrructureof the caudrl fin, r
dirgnosticfeaturefor thc family Perlcididrc The rcprc"cp:rxirl"fin raysscnsu
sentativesof this frmily poscss
Hurchinson(Hùtchinsonl97l) andGrrdincr(Gxrdi,,cr
1989;Tintori 1990b).Since
1988iGrrdiner& Schacffcr
h;rs
orhertLanthe typespccics
noneof thc cncdspec;es
"cprxial" rays, rhc xrtribution o{ tLcse speciesto the
gems Perleídusis questioncd;rs is the prcscnceof thc
gcnusin tle ÈarlyTri:rssic.
smrtcdrvhenSrcnsiostudThis rnisinterpretrrion
ied this genus,crcctingthe specicsPcicìd's uood-^aYtl
on mnterialfrom thc E:rrlyTrirssicof Spitzbergen.Pcr
Ieidu uoodwardi ts a spccicsof renarkablcsìze (up to
30 cm of SL); is skull presentsr long rnd narrowrosrrrl
nisalbones(Stcnsiòle2l:259,
boncandlargetrianeular
regionis cìcrrly
fig.8l). The prttcrn of the erhnroidal
diffcrcnt from thrr found in Peieidr abalepn trnd ìn
pe{e;d;forns (CleithtolepsGîrdiner l98si Dùrelofu f'J
Tintoid 1990;Tbordúpte,rsTintori Et Srssit99ai PcboP'
etleìdusBiirgin 1992iCabdnellìaÎnrori & Lombrrdo
1996),which showa largepcntrgonrlrosrralbonc rnd
dorso-ventrallyelongrtcd or T+hrpcd n:rsrìbones.In
paperthe ethoìoidàlclementsrre cireit.vith
Stens;ò's
different names:nasnlbonesas antorb;rxls,thc rosrral
andtherntorbitrlrs ro*
boneasnrsal-postrostralbonc
scnsory
canals,well r;sible on
bonc.
The
course
of
tral
rhe skull resrorrtion oÍ Perleidusuooh"ardi (Srcns;o
l92l: 259, fig.8l) provesthe wrong ìntcrprctitronor
theseelemenrs,rs has:rlrerdybeensrrcsscdby P;rrrcrson
by rhc inffris rccognizable
(197s:507).The:ìntorbital
crml, whichin thiselcmentbifurcrtcsto
orbhalsensory
onc, and the eùmoidrl comntiv
ioin the supraorbìtal
uootl-^attli the clc
sure. ID the restorat;onof Perleidus
ment inre.prcred :rs the anrorbitîl is crossedby thc
suprrorbitalcanal,bcing obviouslythc nasrl Lrone Pcr
Ieiduswooduartlì hts ventrnlscrlcslonger than dccp,irr
conrrastwith the rypicrl rhombic shrpc of the ventnl
scalesof Perler&r; the scalesof the lateralregìonof thc
trunk ate rhonrbic,:rshigh asdeep(Stensiòlez l : pì 33)
":bbreviatchcteroccrcrì"rnd
Thc cnudalfin ;s defincdns
it is clerr rhrt the axialbody lobe nukes rhe dorsxlmxrl92l: pl. 3l).
gin of the fin (Srensiò
Subscquentauthors kept considerrt;onrhc work
of Stensiò,owing to his xuthor;ty,rnd rhey ;gnoredrhc
the nttribuof Deeckernd Dc Alessandri:
descriprions
tion of thc different speciesro rhe genusP./.t1,6 w.s
nr:deon the b.sir of rhe.hull prtt.rn rnd bv comprri-
35S
l:;*. tC - lù1!ùlBnltuhtt! ll)clckr, l$89). Rcsor iùn. Sc.l. bf: l0ninr
son rvìth rhc spccicscoming from the Spitzbergen,
nrhcr rhrn rvìthrhc typc spccics
of Pcrledo.
Fbr exrnrplc.Pe,/cirlasnadagatcatieais(Pìr ereau
lel{) shows:r he'niheterocerc:rl
tril, visiblein the prper
of Pivctacu(1934:48, fig. 32, pl. VI, fìs.4) rnd on thc
orìgin:rlnrrteri:rlstored rt MuséumNîtion:ìl d'Histo;re
Namrclledc Paris(MAE 6521 MAE 1371rb;MAE
16,+;ìbi
MAE llo,+bi MAE 1023b;MAE Il00; MAE
125)iit:ìlsoshovsxn crhnoid rcgionsimìhrto rhatof
Perlcidus-^ootl-^anti(Srensiól92l: 259, fig. 81) rnd of
Petleidu*oschiensìs
(P;reteauI 934: 46, fig. 30-3I ) . The
othcr spccics fron Midisrscir, Pcrteirluspioeted"l
(LchDìan1952),;srcprcscntcd
by fcw specimens,
often
incornplctc,
so thrr rlrcdcscription
of rhc crudrl rcgion
is not givcn. The skull shows eleo,entsdifferent {rom
thoseof the other species,e.e.,spitaculatplatesof var;rblc sh:rpesrnd the presenccof a postcrior parietal
(Lchnranles2).
Petleidus
:,anltzensu (Su l98l) is representedby r
singlc spccìmcn,lackingthe posreriorprrr of rhe body.
The skullis chrrrcterized
with a veryshort
by:r maxill:r
postorbitrlrcgion.rnd r rvcrkly convcxorrl bordcr;
tceth arc d;ffcrcnt frorn thosc typical of Perleididae:
theyarcconicaìon theanreriorpartof naxilì: andblunt
rvirhr smrll rcrodincc:rpon rhc posteriorpart of thc
PktJsiiern nìnus Rtorgh, 1939in the skull paaern,the
shrpe of upper and lowc jrws, thc prcopcrculunr,:ìnd
the caudalfin, clearly hemihercrocercal.This spccics
wasattributedto the genusPerleidas
only on rhe bas;sof
rhc pancrn of dcrmJ boncsof rhc chcckrnd the postcrior serrationof the scales.
Thc spccicscoming fronì thc SpanishM;ddlc Triassic, Pe eídus giganteus and Pe eidus úai (Belnn,
1972)werealreadyrttributedby Bùrgin(1992)to d;fferent gcnera,Colobodas
înd Pf,.roL?ú, respect;vely.
A
recentrevisionof the Alcover faunacon{ìrmsthe aariburion oî Perleùlussìsdnteusto Colobodu, while Perleidris zrai is more likely to belong ro rhc Ncoptcrygii,
owing ro rhc shrpe of thc preoperculumand thc presenceof rn inreroperculumbone (Lombùdo, pqs. obs.).
Perleidus sp. from Amélie Les Br;ns (Eastern
Pyrenees,
Maz;n& Maftin l98l) is represented
onìy by
few mrssivereerh,with r pointcd trrnsluccntcusp. Thc
attr;bution to the gen\lsPetleìduswas rnadeonly by
comprrison wirh rceth of Peìcidss stoschìensk,
baing
sp.tron
clearlyinsufficientfor determ;nat;on.Perleìdus
rhc I-rdinirn of Turkcy (Bchrn ct rl. )e7e) xnd cf. PerIeidus rron rhe Uppcr Trirssic of Morocco (Mîrtin
1982) are representedby disarticulatedskulls rnd fcw
sc.rlcrow,. in.rdequ.rtc
for r properdererminrrion.
TÈerefore,Peieidusaholepr seemsto be rhe only
Perlcilas lutornsìs Tcixcir:r, 1947 n<l Perteidas v id spccics of thc genus Perleidus:this impies t
1clr,aai Schacffcr, 1990 from rhe Lower Triassic of
remarkablechangein the geographicrnd strrtigrrphic
Angolr cle:rrlydo not belongto this gcnus:"....thc distribution
of thegenus,
till now considered
cosmopolr:ppcrnosr principrl crudrl r:ry nrcersrhc longcsrscrlc irn rnd prcsenrrhroughour rhe Triîssic (Battail et al.
row in thc rsirl lobc rnd there rre no "epaxirl" fin
1987;Behan1988).Its presence
shouldbe rcstrictcdto
rays...:both species
h:rvebeenidentified:sperleidids
on
the Late Ladinian of the Southern Alps (Lombardo
rhc basìsof dcrnnl skull prncrn rnd rhe squxrnar;on' r99s).
(Antuncsct il. l9e0r25).
The diagnosisof the genus has been emended
sinceir was forn,crly mrdc on thc brsis of Lowcr TrinsThc spccicsfron Lowcr Triassicof Crnada, cl
sic mater;althrt crnnot be ascribedto rhis scnustor rhc
Pcrle;drr(Schreffer& Minsus, 1976)wrs dlreadyques
with
mentroned
reasons.
tioned by Bútsin (1992) for ;ts resemblance
MnL e Tna*t acuaoprerystans
.159
Perleldusa|tolepls(Deecke,1889)
I r i A \ . 2 .l 0 1 : 1P, l . 2 B C
BrlloÌÌì ir StÒppini.p1g.419.
1357 Lqnlr\" tnù6Btllúri,
1st7 PLÒli.lopltort'ob!',tstJ BolloÌÌi. Bollori in Sroppini,pp. 4281ss9 Sotn,tauB ixl4,s Dùckc, Dccckcpp- 120-l2li bÌ Vl, fig.
10.
Oeccke,Vood!r.d, prd lll, p- 57.
1se5 5tùn,,rrs.bolqú
Dte.kc,Schrll*icn, pp- :24-25.
t9at toionr6,rlro|Ìi'
t9]10 tutunlB nLahù lD.tck.), Dc Al.$rndri. pp, 49-51,hr ll,
fis.2.
t91a H.tuúfutùlura pftnùu (B.llori). De Alesrndi, pp. l0l-105,
Pcl
t91A Pl)alnlo?l)or"i
obhnB,r (l'rop!re) Bcll{fii, Dc Alessin{lri,pp.
l l 9 - l l l , p l ,v I l l , f i s . 9 .
t92a PalenL6.tlolqn (D.t.kr), Srollty.pig. 65.
192]1 Pùhnlt\ rhol.tn (D.tckc). Stcnsiò,pp, 256'257,figs.73 79.
t-'t
H d , ' r l , t t , ] 'ú , :Ì , . r , , , t / , . ' r { f l l ' r " ' . \ ' r n \ i " , p p . 2 1 6 . 2 r 7 . l i L \ .
tett
Pù!.nlrc rltÒlctis lDee.kc), Pir€rciu,p- lrl.
te52 Pùl.id6,ùolctì' (oceckc),Lchnlrn, p. ll7.
t96(, Pùllil6.lblqì'
lD&ckc), Lchrrn in Piv.rc.u, pp. I l4-l15,
1990
l9el
r9r5
1997
1999
Polùl !altuh?^ lDccckc),Tlntori. p, l9l,ùyl,
A,r/.r.16l/',, Ilúrsnr,pp.60 61,Iis. 59.
Prl.i16.lblqn
lDeccl,c),Búrsin,pp- 306-3ll, li8s.2-6.
I'o!ù!$ nlù,la?nlDèt l,c), Lombirdo, pp.5l-75, fi8s,36-55.
tuldlu\tholqs
lDttckc), Tir roÌi & Lombrrdo.pp.l95 501.
M a t e a l : M C S N I O P + 6 1 - + 6 1I .' t o 1 . l t 9 9 l ' 1 6 , P 6 7 6 6 3 1 ,
16s6, P637. 169lrb, P692rb. MCSN 1007, 1003,5004rh, s005,
I'lNlUZ 1 .1960.I.1961,T.1793.
50O6rb,5007ùbcd,
Dislribulion:Llrr Lldiniin of sùti di Pùl.do (Perlcdo-vàren
nr lirnnÌnx) îid oi K.lkschicfcúone(Mride Limesore) ofCr'dcl
I:ri,c (vissìi,-vA) rnd ìv.rid. (cirron Ticino CH).
Dagnosis (drcrdcd fron De Alessrndri 1910): Mediúnsi2rdrpccicsof 1.20nN of mrriDun, lensrii op{cùlu m slishryrnrlld ,hii $,borùculun,i sircle suborbnrlt Jen,nion nrde ot pes-like
tc.rlì oD o'rl mùrsinoi mrxilh rnd dcnÌÍ_r rnd blún, R.,h on pilÍrl
scilc rows. lacnl turk sàres
boD.\; \qurn rbn of rz ùins6c
tlropd rhîr \' 0. ciudil fin of l8 l.pidorichir with 6 'ep:xirl" nysj
bonc onnnr.trhtion !.presentcd br'' sinoinc tubcr.les rnd short
FiE lt
- Púlèilrt dbolepú lDccckc. l83e). Reron,ion ol thc skull
in hterd (A) md dosrl (B)
parietrlbones(Figs.10,1lAB, 12ABC,Pl. 2BC).
The parietal bones rre hrge and squarish;the
interparietalsutureis wary, as is the anteriorand latcral
margin of the bones, where rhey contact the frontal
bonesand the dermopterotics.Poresof the supraorbitrl
sensorycanalarevisiblein the anteriorhalf of the bones
( F i g ( .1 0 .I I A B . I z A B C .P ' . 2 B ) .T h e d e r m o p t e r o r i c s
are rrapezoidal,elongateelements.The suprrtemporal
sensorycanalran along thc rnedianpart of the bones,
(Figs.10,11A8,12ABC,PL
whereporesaredetectable
Description
5Érll The rostralboneis lrrge and subpcntrgonal; 2BC).
The extrascrpularsare recrangularwnh rounded
rhc ìateralbordcr presentsa narialnotch. Its surfaceis
by grnoincridgcs(Figs.10, l1AB, 12BC, posterioredge.The mediansutureis short and strright.
ornamented
Poresof the supratemporrlcommissureare visible on
on spccimens
l
Pl. J87.T\e ornlnentrrionis diffcrcnL
diffcrent ontogeneticscrges,being stronger on larger the whole length of the bones.The supratemporalsenThe nasalboncsrre irrcgularrnd elongated sory canrl enteredrhe dermopteroticanteriorlyànd rhc
specimens.
posteriorly(Figs.10, 11AB,12ABC,Pl.
Jonorrntrrlly. They present.rn anteriorerprn.ion. post-temporal
with the openingfor rhe nrrirl notch belo* The poste- 2BC).
The dorsal marg;n of the orbit is bordered by
r i o r b o r d e r . . L g h r l yc o n v e xr n d o u t l i n e t' h e r n t e r i o r
rhree
suprrorbital bones;the first two are rectangular,
ran
nrarginof rhc orbir. The supraorbitrlsensorycanal
lasr
one is biggerand trianSular(Figs.10, l1AB,
the
alongthe lengthof tLc bonc (Figs.10, I lAB, |2ABC,
12A, Pl. 2C). The dermosphenoticsare very th'n and
Pì.2B).
elongareelemennplacedon the postero-dorsalmrrgin
The frorrrl boncs are broad and rectangular;the
border in contactwith the supraorbitalbonesis slighrly of the orbit rnd posteriorlyin contact with suborbital.
Poresof the infraorbnalsensorycanalare visibleon thc
convex.Thc ìntcrfronrrl suturc is straight,exceptfor an
indentationin the posterior hal{.The supraorbitalsen- lengthof the bones(Figs.10,11A8,12A,PL2BC).The
infraorbital series;s made of two largc crescenticelesory canal entered at the întero lateral corner and
menrs,vhich outline the latero-ventraledgeof orbnal
rerchcdthc posrcriorm:rrginof the bones,endingin the
160
Na
B
openinq.The elemcntin contactrvith rhe dermosphenotic is largcr rhrn rhe secondone; borh hrve setrated
ventr:rlbordcrs. A sm:rllrntorbitrl reachedrne rosrrl
bone (Fies.10, llA, I2ABC, Pl.2B).Thercis r single
suborbir:rl:n is quire largeand rcctangularand contacts
rhc dermosphenoticrnreriorly.The postcrior border is
serrltcd(Figs.10,114, l2ABC, Pl. 2BC).
The preoperculurnis very hrge, with an expanded
dorsal reeion :rnd a nrrrow ventral one. Th. p"'.e.or
edgecontrcts rhc dennohyal;thc porcs of preopercular
rn.oru crn.rlrre rrr.rnguJin r linerlong rhc posterior
border of rhc bone. A well-devcloped
infraorbital
proccss
is present(Figs.10,114, l2,{BC,Pl. 2BC).The
opercuìumis dccpcrthrn wide; it conrrcrsrhe subopercuìun with a slìghtly convcx suture. The rectanguhr
subopcrcuìumis bigger than rhe operculum (Figs. 10,
l lA, l2ABC, llA, Pl. 2BC).
The upper jaw is formcd by r srrong maxillaand a
srall prem*ilh, bcrring 4 or 5 teerh similar in shape
and sizc to thoseborne by rhc mrxilla. Thk latter hasa
narrowrnterior regionand rn exprndedposrorbnalone
(Figs.10, llA, I2ABC, Pl. 2BC). The oril margrn,s
straight; tcerh are ;rbsentin rhc posrerior part of the
fis. r?
- Pùlc;.lrs rlralqi'
(Dcecke,
1339).Skùllboncsispturoncd
'n rpcdmen\: A) MCSNIO
P60s;B) MCSNIO P607iC)
M.\NIO t60.1.iq ( I.r,J t
bone. The dentition is madeof 20-25srrongteeth,whh
the rypicalpegJikeshape;the first 10are similarin size;
reethbecomesmallerin the posteriorpart of thc maxìlIi (Figs.10,114, 12ABC,Pl. 2B).Thc palare
is dentigerous,bearingteethofdifferenr size.Largemd srourrc.rh
predorninate.The lower jaw is clongrre, with straight
oraì and ventral mrrgin. Anterior te€rh rrc sinìilar in
shapeand sizc to rhoseof the 1nteriorpîft ot rhe ma\illa; posreriorlythey becornehrgcr. Pores of the
mrndibularsensorycanri rre visibleon the ventralprrr
of the dent;rr.7(Figs.10, I lA, I2ABC, PI. 2B).
Up to 6 triangLrlar
bnnchiostegalraysrverecounted (Fig.12C).There;s a singlemedianovrl gular(Figs.
12CP
, l.2B).
The posttenrporalsare two large and tr:rpczoidal
elements,separared{rom erch other by r couple of
\crle' lFiss. 10. llAB, l2AB. PL 2BCt. The posrerior
edgeis roundedand lrterally are detectablethc poresof
the supratemporal
sensorycnnal coming trom rhc
extrascapular.
The supracleihrum k elong:rtcddorso
ventrallyand contactsthe cleithrum rr rhe lerel of the
suturebeween operculurnand suboperculum(Figs. lo,
114, I2ABC, l3A, Pl. 2BC). The anteriorrrrrsrrrrs
Mi ddlc Tiase. a. u nopt.rrszùs
361
DI
Scl
opì
fis.lJ
Pe .ì1116
nlral.p^ (Doccl(t,lsse). A) MCSNIO P60abrB) ciudrllin Ò{MCSNIO P5oln.Scilc bir: 5 n,n1.
alwryshiddcn by rhc operculum.Postcleithrumis a large
elemcnrornamentedby tuberclesand shon ridges of
ganoine(Figs.10,llA, l3A, Pl. 2BC).The cleithrumis
partially visiblc, bcins coveredby the suboperculum.
The posterior part is convexwnh ornnmentarionmade
of short ridgcsof ganoine(Fì8s.| 0, I lA, 12A8, Pl. 28).
Fias. Pectoralfins hrve 8-13 lepidorrichia.Each
ray consistsof a long proximrl baseand shorter distal
segments.Each branchesat least twice. The anterior
mrrgin of rhe f;n bearsa seriesof strong frìnging fulcra
(Figs.10, llA, PI. 2BC). The pelvicfins aresmallerthrn
rhe pecronlsr rhey have 7-t3 lcpidotrichia,s;milar in
structureto thoseof rhc pecrorals
(Egs. 10, 1lA, Pl.
2C).
Thc dorsalfin is in the posteriorhalf of the body
It is triangular
andcons;sts
of I2-17lep;dotrichia.
Distrl segncnrs branch rr least twice. Two or rhree basal
fulcraare present,and therc is r serieso{ fringing fulcta
alongthc rnrcriormrrginof the fin (Figs.10, 134, Pl.
2BC). The analfin is smrll rnd triangularand is madeof
7-r0 lepidorrichir.A coupleo{ basalfulcra and a series
of fringing fulcra on rhe rnterior margin of rhc fin are
present(Figs.10,Pl.2C). On spccimen
P 605ther€are
two largepreanrlscutcs,with senatedposterioredgc.
The caudalfin is almostsymmetrical,wìth a short
axiaìbody lobc.Thcrc arerbout 28 lepidotrichia;6-7 of
rhcm rrc "epaxial".The nys rre scgmentedand they
branchat leastthree times.Both the dorsaland ventnl
marginsbearfringing fulcra (Figs.2, 10, l3B, Pl. 2C).
Squanation.'fhescalecoveringconsisrso{ 37 vert;cal scalerows.The scalcsof the anreriorregion of the
trunk arc deeper than wide; they bccome gradually
srnallertowards the posrerior pan of the body. The
scrlesof thc mtcrolaterrl {lank region rre rccrrngular;
thcir dcprh decreases
towardsboth rhe dorsalrnd venrral resion where rhey bccomc rhombo;dal (Figs. t0,
13A, Pl. 2BC). A mid-dorsalridge scaleis present.At
the baseof the dorsJ fin, scalesare smalland wìderthan
deep(Fiss.10, llA, Pl. 2BC). Scnlcsof thc posrerior
part of the body rre rhombic, only slightly wider thrn
deep.All scalesare serrated.
Taxonomlc hlstory
Perleidusaholepiswas ercaed by Deecke;n 1889
asSenionotusaholepison mrrerialcoming from Pcdcdo.
que'rionedrhe rs\ignmcnL
Schellwein
ol rhisspecies
ro
the g€nusSezrbrotus,stressingthe primitive prtrcrn of
rhe skufl (Schellwein1901t24-25) and De Alessandri
erectedthe new genusPoleidzs for it. The speciesh;rd
actuallybeendescribed,bur not figured in 1857by Bcllo:li ^s Lepidot"s'eratu' (Bellorti 1857:419) ànd Proli
dophowsoblungn (Bellorri t8s7: 428). Complri,ons
with the counterpart (rhe only avrihble prn) of the
holotype of Lepidotus setatas and with the drawing of
the holotype oÍ Pùolidophotusoblungusmadc for Bellot,
Li' prperconfirmthe .rtrriburion
ot rhcsc,pccimenrto
For Perlcidn alnlepts,a. well as for m.rny'pccie'
coming from Perledo,there exisrs considerabletaxonomicconfusion.Deecke(188e),revisingthc Pcrlcdo
marerial,erectedthe spec;esPerleid"sattoleprìwithout
comparing his mrterial wirh that of Bellotti. Decckc
consideredrhe prper of Belloni not vrlid, beingwithout
dr.wings(De€cke1889:110-111).
Lîrer on,De Alessandri (1910)r€-studied
the originalmaterialof Bellotti,
but he did not understandthatP abolEis, L. senatusand
P oblmgaswere the samespecies:he just erectedr ncw
gcnusfor the Deeckes species.I think thrt zerle ar ser
ntus, as slell as Perleídusoólaaszs,could be considered
a nonen oblitan. ft has never been used aftcr Dc
Alessrndri,while subsequentauthorsalwaysmentioned
162
Iris. l.l
- Dr,nrì {r,r,$ g.n. n. sp. n. Resontìor. Scàlebù: 5 mm.
only PerleidusaLolepis,except for Bùrgin (1995) who
finr nored rhe nomencbtorirlproblems.sincemore
than 50 yearshas passcdwithout mentioningPerleidu'
seratus or Perleirlusoblungus, one can considt Petleidus
Itolqprr (ahhough"junior synonymous")to be the only
Danlniagen.n.
Diagnosisràsfo!specie!.
Typ€-speci€s:D,,rDn Jptdz sen. n. \p. n.
Etymology: dcdi.r.d ,Ò chnluc. Dmini, cun.or of Civico
Mu*o lfulbrico di sioria Nrunk dilndunoOlon4 for his brsic rnd
conr.i' cortribution to rhe studrof the locrlny ofctdèlFdrc.
Typelocality:Cn dd Iì& (vissii'-vr), Ihlyi Ilre Ldinid-
Remarks
This speciesis attributedto Perleidiformes
on the
skull pattern,wìth the largerosrralbone separatingthe
nisalbones,the maxillashowingan enlrrgedpostorbnal
region,rnd rhe preoperculumdorsaUyexpanded(Bùr
gin 1992;Tìntori & Lombrrdo 1996).Also, the structurc of caudalfin, with the presenceof"epaxial"fin rays,
and the lateralscalesdeeperthrn widc on the ànt€rior
rcgion of the body are typical of this group (Grrdiner
1988;jGardiner& Schreffer1989;Bùrgin 1992:TinÌor].
tr Lombardo 1996).Brsìcrlly,somc peculirr characters
do not allow attributing it to any known genus:this
genusis in facr diffcrcnt from any orher perl€idiform
taxon in the stronglyserratedscalesand in the srructure
of tLe fins. These rre chrracterizedby long prox;mal
basesand by distalsegm€ntsthat areshorte. but alwiys
muchlongerthrn broad.In addnion,the caudalfin rays,
in otLer generr usurlly scgmentedbeginningfrom the
baseof the fin, show very long proximrl brses.
Éiymology: frcm rhe L.tir vord ftxnr', rtf€rcd ro thr
,hony ippcú.nce of rhe $ale coy€rins.
P668rb (hólÒtypÈ), rro
hc
Matarial: MCSNIO
K.lk$hiefcu ore of G' dd lìrte (visgii'-vA); McsNIo P6l8ab,
P666!b, P667,P669,P67A,P671^b,P6721b,P673îb, 167a, ló75!b
fron rll€ Krlkrbiè{ezone otct dd Fn.c (viseiù vA) Groredin,ie
civico M!$o Insubrico di Sbrià Ndunle di In.lùno olori) rnd
McsN I,5008Írom rie KJkschic{e.zóneÒt Meridc (cin'on Ticino'
CH) Gtorcd in the MuroCin,onrle di storìi NÍunh diLuslno).
Type-localily:Cr'dd Fúre (viseiù-v4,lhlÍ hr L.di',ian.
DisFibution: Upper Lldini.n {Middh Trià$ic) ot
KJksl'iele.zó.e (Mùide l.úlk) ol cr dd Fr.e (visBiìrvA) .nd ol
Mddè (crnrÒnTicino-cH).
Diagnosisrsnill p{hidiform wirh elonsrÈ tùsilùnn body:
opercularlegion vidci p.opèrcùlùn wnh n.Íow vcnral resio! rnd
Ro
Na
Danlnlasplnosagen.n. sp, n.
F i g s -l 4 - 1 7 ,P ì .2 D - I
1995 O/Jioptn cl.l.?bn,t Bùrgin,pp,8l8-820, fi8,,2-14.
199/ Pcrld,rd sp. Lonìbrrdo.pp, 76 73, fis. t6-
B
F ; 9 .1 5 - Daùinia spinùd Aen.n, sp. n Rsrontion Òf the sku
hteral (A) and in do.s,l (B) liews.
363
Itl ì.Lile Trìasic dcti"optelr\ia$
median pit line is present.The interparietalsuture is
srtaight(Figs.14,l5AB, 1óBD).
shapc
The dermopteroticshavea sub-recràngular
pr€sence
of
is
irregulrr
owing
to
the
rnd their surface
small openingsand pores of the sensory cînrl, well
detcctableon the ventral marginof rhe elemen* (Figs.
Description
14,1sAB,16ABD,Pt.2F).
S,èrii.The rostral bone is large rnd pentagonal,
Extrascapulars
are triangularand narroq with an
with thc rypicrl pcrleidiformoutline.The ethmoidal irregularoutlinedue to rhc presence
of tuberclesand
perpend;cularly
arrangedto the
commissure;s well vìsiblcon the ventralmarginof the
short ganoine ridges
(Figs.
ganoine
cresn
14,
clemcnrrnd is linedby short
posteriormarginof thc elements.The two elements
l5AB, l6AC, Pl.2F). Thc nrsrl bonesconsistof large are in contactby meansof a short suture (Figs. la,
sub-rccrmguhrelenents:the suprrorbit.l scnsorycanal 15AB,lóABD, Pl. 2F). It is impossible
to locrrer dcrentersthe postero-dorsîlcorncr,wherethe nasalbones
mcct the frontal bones.Thc surfrccis smooth (Figs.14,
The scriesof infraorbitalsis mrde up of one or
lsAB, l6AC, Pl. 2F). The frontrl boncsrre wide,with
two denderand elongrtc elementsborderingrhe dorsrl
rn exprnded postcrior region and r narrower antenor
mrrgin of the maxilla,followed by r larger crescenrc
one.Thc posteriormarginis roundcd.The supraorbital bone on the postero-ventralcorner of the orbit. Pores
(Figs.14,15A8,l6AD, Pl.
sensory crnal hrs rn S-shapedcoursei it enters the
rre detectable
on allelemenrs
anrcro-vcntralcorner of the bonesrnd rcachesthe parietal bones.Thc intcrfrontal suture is stnishr. The surThe shapeof the preoperculumis typical of per
f:ce of these clcmcn* is only feebly ornamentedby
leidiforms,with an enlargeddorsrl regionanda natrow'
(Figs.
Pl.
14,
15AB,
16ABCD,
rugaein a few specnrens
er ventnl one. Its ventral margin embrrcesthe dorsal
2DF). Thc prrictal bones are large and squrrish; they
margin of the maxilla.The preopercularsensorycanaì
rcccivethe sensorycrn,rl from the frontal t onesand a
ran along the posterior margin of the bone (Figs. 14,
!\prndcd donil onc Ni'h dcleloped infr.orbitll pro.ess; lons .nd
pÒi .d k.rh on bodì upperrnd lovcr irNsi sqùlnlion ol 36 or 37
rrnsrsc scrler"*'; x.l*,lc€pd thrn widc in antro'ldcnl resion
of trurkt nl sùlcs dccph rcú.r0d: rnil rnd do6rl fins with hldd
b$.t c!ù,hl finofibou,27l.p o.icìriris.reD epùial ùúdrl fin
Fr
la
,,
li*.t6
I) nrid 'pntúì sti. n. sp. n. Skullborcs s prerned irspecinrens:A) MCSNIO 1662;B) MCSNIO P669iC) MCSNIO P6l0i D)
MCSNIO P6l2b. Scrlcbrr: ì mm,
16.{
I ir. |
- / r . r x , / i . . , r ' . . , q L ì r . \ p . . r t r i Ì u l p . r r u ' ' l ; ' J l c r n d bodyof MCSNIO P6lri. S.ilc brr:5 mm.
l5A, l6AD, PL 2DF).Thc opercular
regionis widcrnd
rccransulrr,with r rounded postcrior mnrgin.The
suture betrveenopcrculumrnd suboperculumis alwrys
difficultro derecr,
owingro rhethinness
of thebones,so
th.rrthc 'h,rpc.rnd,izeof erchelemcntrcmrinsuncerrain.On spccinrcn
MCSNIO P667a sm.llantero-dorsal
processof rhe suboperculum
gives the position of
boundarybetweenthe rwo bones(F;gs.164, Pl. 2F).
The surfrcc of thc opercularregion is ornrmentedby
concentricstriaeat ìeaston the do.srl ma.gin.
Thc m:rxilh shorvsrhc rypical perleidiform outline, wiù rn cxprndcdpostorbitalregion and r slender
anrcriorone.The or;ìì ffrrgin bcrrs scverrlrows of long
and pointed tecrh; on rhe anterior tip of the nraxillary
(premrx;lhry))teeth:ìreslighrlylonser(Figs.14, l5A,
l6AD, Pl. 2DF). Thc mand;bleis a strongwedgclike
elemenrbcaringrccthsimilarin shnpcrnd sizero those
of thc mrx;lh. On its posteriorprrt r lrrge angularis
present,;rndthe ventralrcgion showsthe courseof thc
mrndibuhr scnsorycrnal (Figs. 14, l5A, 16AD, Pl.
2DF).
The palamlregion bearsconicalrccth hrger than
thoseborne by nrrxilh and lowq jaws,rnansed 'n sev-
st.;ae(Figs.14,l5A, l6AC, 17,Pl.2EF);on a fewspccimensrherc are rwo rows of minute denticles:rrnngcd
along the antero-ventralmrrgin of the elemenr.On the
dorsalareathe courseof rhe lareralline sensorycanalis
visible.The postcleìrhrumis subtriangular,
with r
roundedposteriormarginand a straightvcnrralonc; the
ornamentrtionis sirnilir ro thar shownby the supracleithrum. The cleithrumis wide rnd sicklelikc, bur irs outlineir not clerr.osrng ro rhe vrre of pre.erv.:rion.
Firs. All fins are characterizedby :r rcmrrkable
development:paired fins are long, whilc med;anfins
show a brord base.owing ro the delicacyof rll fins,
their distal endsare poorly preserved.The pcctoralfins
haveabout ten lepidotrichir, wirh long proximal brses
andshorrdisralsegments.
They branchrt lcrst twicc and
rherearestrongfringingfulcrr rlong rhe rnteriorm.rrgin. Pelvicfins rre mrde of long and slenderrays;thcy
are placcdrr rhe I2rh scal€row înd opposircrhc dorsal
fin. Fringing:ndbasalfulcrî îre nor visible(Fiss.l,r, 17,
Pl. 2DE).
The dorsalfin is placedrr rbour rhe 18 20th scale
row rnd shows a brord base;;r has :r leîsr 15 lcpi
dorrichir, with very long proximal brscs.An ovrl scute
and 2-3 strong basalfulca prccedethe fìn. The anrl fin
At lqrst fivc rriangularbranchiostegrlrays have is locatedat abour rhe 20rh scalerow and it snowsrne
bcencounted;theirsurf:rce
is smooth.
the dorsalone; h hasrr lcrsr 15 rays
Behindthc exrrrscapulars
rhereare two postrem- andtwo or threebasalfulcrr. In fronr of the fin thereare
porrls w;th r roundedposteriormrrgin; rhe supraorbitrl two or more pre-analscutes:on specimcnMCSNIO
sensorycrnal rin on the venrralregion of eachctemenr ó73a the larger one shows a sub-ovaloud;ne, with :r
rnd rcrchcsthe supr:rcleithrum
behind(Fìss.14, l5AB,
round,:d.rnLcrior
mrrgin.:ndrn irregul.rr
po,trriorone.
l6A. 17,Pì.2EF).Therc is a largeovalsupracleithrum Ganoineridsesendingwith r tip ornrncnr rhc surface
with orn.rnrcnr.rriun
th.,rconsi'r'of delit.rre
conccnrric (Figs.14, 17,Pl.2DE). The c.udal{in has25,27rrys,
ILATE 2
A) lrrJ r?r', (B1lúno Crirelli, ls39): MCSNIO Pas6:B) Prkid6 dhalEn (Dccck., l33e): McsNIo P6o2tC) Pcleidb altok\ lDtcc^.
rss9): MCSNIO lrtorii D) D"tiÌh spnnn {e\ \. sp, n,: 'h. holÒrtpoMCSNIO P66et E) Dd|ntd Vkúa Fn n. sp-n-: MCSNIO P67]li E)
Dd,nd *nrM s.n. n. \p. r.: MCSNIO P662.
lt l
È r . .
i ,{*"Ì ' ;
;
n
166
with sevcn "epaxial"rays; lepidotrichiadre segmented
and even distal elementsdre clongate.Both the dorsal
rnd ventrrl lobes show strong and long basalfulcra;
fringingfulcraarenot visiblc(Figs.13,16,Pl. 2DE).
Squanatiox. Squamation consists of about 37
rînsverse scile rows; all scJcs are characrerizedby a
distinct;velyscruircdposteriormrrgin. Scrlesaredeeper
rhrn brord on rhc rntero-lateralregion of the flank,
while scrlcs of the ventrrl rnd dorsal region arc lower
but, os'ing ro rhc deepserrat;on,ir is impossibleto identify tbcir preciseourlinc. Beginningfrom the dorsalfin
and towards thc posterior part of the body, scrles
decrcrsein depth,becominghnceolateon rhe axialbody
Iobe. Therc rre large basalscutesin front of borh the
dorsaland venrr.l lobe of the caudalfin and one or two
prc-rnal scuteswirh largc denticleson thcir posterior
margin.Thc surfaceof the scalcsis smooth; the lateral
scalcsbehind the cleirhrunr show 6-7 long dent;cles
whose numbcr dccreasestowards the Posterior,dorsîl
rnd vcntrrlregionof thc body (Figs.14,17,PI 2DE).
recognisableon specimcnT 4964 for its pentagonal
shapeand the ethmoidalconmissurethat runs through
ir. The maxillaL not narrow and eìongate,but is madc
enlrrscdrcsionànd: nrro"cr rnterior
ol r posLorbirrl
one.The interopcrculumis absent.The axialbody lobc,
accordingto Bùrgin, extendsabout hrlf way rlong the
lengthof the dorsallobe of the caudalfin (Bùrs;n 1995,
pg. 819); the specimensfrom Ca' del Frate and Meridc
show "epaxial" fin rrys, excluding rh;s gcnus fronr
of rhese.p..;-,.' to
The .rrtriburion
neop(erygians.
the genus Opfioprs rnd in particuh. to the Perlcdo
speciesseemsto havebeenmrdc by Bùrgin on rhe brsis
of rhe genenl shapeof the body and rhc fins, thoughthc
'ome difference'
in m"'t mcrisaurhorhimselfstresse.
holorype of
(Bùrgin
l99s:
819).
The
tic characiers
re10, rav
(figured
De
Alessrndri
by
ophiopsisleptutu
III, fig. a) has been losr, but a cast of ir, storcd at the
in
Senckenberg
Nîturmuseumund Forschunginstirut
Frankfun (P 12a3),shows clerrly some of rhe typical
ncopterygianfearurcs,suchasa nrrro* and venicrl prcoperculumand the interopcrculum(p*s. obs )
Discusslon
belongingto this
The size of mosr specimens
speciesis very smrll; they reachabout 35 mm of toril
show the srnc sizeand
length.Atmost all specimens
come from rhc samebedd;ngplanc. As for other taxa
comìng from Cr'dcl Fratc, which rre clusteredin sizechsscs,(Lonbardo 1997;Tintori 1990a;Tinrori èt Lombrrdo 1999).thcy probablyreprcscntthe sameontogeby the
mostlikclyiuvenile:
thìsis supported
ncricst:rge,
presencein rhe collectionof r singlespecimenof larger
s;ze(MCSNIO P61sab).This kind o{ srmplingmaybe
relatedto thc scasonalityof thc Ca' del Fnte dcpositional environmcnt(Tintori 8. Lonrbîrdo 1999).
Some specimcnscoming from tbe Middle
of Meride,storedat MuseoCrntonale
Kalkschicferzone
di Storia Naturale of Lugano and at Palàontologische
Institur und Museumder Univcrsitàtin Zùrìch,are
rr tnburedtc rh.rr"rmc 'pecier.The .rlc of preser";tion
o{ rhescspccimensallowedcomparisonthem with the
rnd. .trLh. ..rmerime.que.tìonCr'del Frlc spcc'nrens
madeby Bùrsin (1995) Búrgin
ing the determination
describedthesespecimensts Opbiopsi cf. teptutxs,bur
rhe obsenatìon of sone skull eìen,ents(especirllyon
T 4964rnd MCSN 3014)suchasrheopercuspecimcns
Irr region,shrpeand sizeof preoperculum,rostraìbone,
fron the genus
thcscspecimens
rnd maxilla,excìudes
Opl'iop';. nd morc cencr.lly, from : neoprerygirn
group (pers.obs.).The preoperculumis not narrow,as
in neoprerygirns,but it showsa wide dorsalregionand
along the posterior margin pores of the preoperculrr
sensorycanalare dercctrblerconsequendy,this rrea is
not occupicdby the largesuborbir:rlsmentionedby Bùras the Pregin (199s:819). The elemenrinrcrprered
mrxillary is, on thc contrary,a hrge rostral bone well
Ackna@ledqenkni\.
G. Adh ind M. wilson did rhe Íniciìind ìirsunric rt ins:
I inì veryB.Íèfúl tor thci! hclprnd Pr.cio{s su88c$nrnsMintthinks
80 to A. înrorifor thc lonsrnd siruì.ting dncusìons,whichSrcit
ly mp'"'rd 'h.. prpcr v\ ',nnte rhrnk' r." '- rll 'h ir.:'J'inr'
vho save tinrncirl suppon {or fiold work borh r cr dd F-nt ind
Mùide; ,hè cirico Musco l."ub.icÒ dì s,orià Ndurft di Llduno
Olon( thc Múnicipihy of Vissii' rnd th. MuscoClnk,nrlc di Srorir
Natunle di Lùg.no: spccill thtrnksgó to c Drnini rnd M lcltrci
'fh,n",
s" ro rlì rho'c *ìr- h.lprJ 'n collc'rinr rl, 'p..iN(n' f n.
cirlly D. Moriiio. C. Ptr.orind M.Tentoro{rhe Mrrco I'rlcontolosiund
có lî Roccn'di Mon{rl.onc- I m indehtedto ,hc Nruflruru'ì
Fó6chùnsìnrirut Scnckcnbùs(Fnnkf un im Mrìn), PrLionblosn.h.
In*no, und Museun dèr Univcsitir (Zuri.h), Musóum NdnDd
dHn,onc NJruElle (Prit. Muscum fùr Nrurkundc (Berli't rn{-ì
sen izio ccoìosico d Lrlh (Rome)for úces,o thcir coll(rions: I sifr
cùcly.húkG. Plodowski,H, Fulrei G Amriiind M Ans.l.llì Thc
phoroS.rpi! wercmidc b) S. Ren€*o
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