Bollettino della Societa Paleontologica ltaliena, 44 (l), 2005, 25-34. Modet

A new perleidiform (Actinopterygii, Osteichthyes) frtofNorthern Italy

Cristina LovtBARDo & Francesca BnnrraBlLLASCA

KEY WORDS - Perleidiformes, Triassic, Adaptive radiation, Durophqg'.

ABSTRACT - Endennia licia n. gen. n. sp., a new genus of perleidiform.from the Zorz,

described. The nev, genus is charicterized bt, a .fusiform outline with a strong skull, b1

peculiar kind of denition, w,ith long and cvlindrical marginol teeth, v'ith.flattened apex, c'kind of dentition, unique among ptilttdtlorms, is indicatiie that Endennia licia n. gen. n. s

or haid exoskeletons, such as imall cruistaceans or mollrrscs and echinoderms, u'hich v'€rt

nev, taxon adds .further data to the knowledge qf the perleidiforms, mainb' knov'n.fi'om Ec

widespread, also v,ith specialized.forms, during the Late Triassic.

RIASSUNTO - [Un nuovo perleidiforme (Osteichthyes, Actinopterygii) dal Triassicc

descritto un nuovo genere e una nuova specie di pesce.fossile appartenente ai perleidiformi

Calcare di Zorzino-(Norico, Triassico Suprriore). Il nuovo genere d caratterizzqto da un p

robusto, dalla.fusiotne parziale di alcuni elementi del tetto cranico e da un tipo di denta

mascella, ^oidtbola dotati di denti lunghi e cilindrici dalla punta leggermente appiattita e

tipo di dentattlra, unico tra i perteidiformi, porta a ritenere che Endennia licia n. gen. n.

p'rovvisti di esoscheletro sottile quali'i crostacei, anche se non si esclttde la possibititd che I'o echinodermi, la cui presenza-d segnalata nella stessa unitd stratigrafica, anche sotto.fa

Questo nuovo taxon occresce li conoscenza del gruppo dei perleidiformi, molto diffi,

degli ultimi anni rivelano essere presente, qnche con.forme molto specializzate, durante

C. Lombardo, Dipartimento di Scienze della Terra "A. Desio",

I ta ly ; c r i s t ina . lombardo@unimi . i tF. Brambil lasca, Dipartimento di Scienze della Terra "A. Desio

I t a l y .

INTRODUCTION AND GEOLOGICAL SETTING

In the last th i r ty years the Calcare d i Zorz ino(Zorzino Limestone, middle Norian , Late Triassic) hasyielded an extraordinary rich ichthyofauna: at least 25genera have been already described or are under study,but probably more than 50 genera are represented(T in to r i , 1981, 1983, 1995a, b , 1996a, b ; T in to r i &Renesto, 1983; Tintori & Sassi, 1992). A few taxa arerepresented by hundreds or thousands of specimens,like pholidophori ds (Zambelli , 197 5, 1978, I 980, I 98 I a,b, 1986), others by dozens and in some cases by fewor single specimen. Apart from the numbers, this faunais important because it records the first major radiationo f neop te ryg ians (among these , pycnodon ts ,semionotids, macrosemiids and pholidophorids), whenpaleopterygians were still important, especially at thetop of the trophic hierarchy, with the "chondrosteans"

genera Birgeria and Saurichthys and the "subholosteans"

Thoracopterus and Gabanellia (Tintori, 1990a; Tintori& Sassi, 1992; Tintor i & Lombardo, 1996; Tintor i ,1998a).

Reptiles are also very important for the presence ofendemic genera and of the oldest pterosaurs so farknown. Although the Zorzino Limestone deposited inan intraplatform basin, most reptiles are terrestrial withland (Langobardisaurus), arboreal (Megalancosaurus)

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or flyin g (Eudimorphodon) special rzations (Wild , 197 8;Renesto, 1994,2000). These animals probably l ivedon more or less wide islands situated around the basins;they are usually small-sized, very lightly built, and oftenhave hollow bones: their preservation requires a shortpost mortem transport in superficial oxic and warmwaters. Some others could have spent part of their lifein water, especially for feeding, such as the thalattosaurEndennasourus, the armoured placodont Psephodermaand the phytosaur Mystr iosuchus (Renesto, 1992;Renesto & Tintori, 1995; Gozzi & Renesto, 2003).

Invertebrates are also well represented by severalcrustacean genera (Pinna , 197 4; Garassino & Teruzzi,1993, Basso & Tintori, 1994) echinoderms (echinoids,ophiuroids, crinoids and a single asteroid) (Blake et al.,2000), cnidarians and scleractinia. Gastropods are rareand bivalves are common, mainly as remains of f ishpredation, and they contributed to constitute the benthicfauna, rich in individuals but poor in species.

The Zorzino Limestone is inferred to have beendepos i ted in a mar ine bas in assoc ia ted w i th ear lyMesozoic rift ing (Jadoul et al., 1992, 1994). The basinopened within a wide and thick carbonate platform,the Dolomia Principale Formation, that extended overallof the western margin of the Tethys. Because of thewidespread shal low-water environment, connectionsbetween the basin and the open sea were probably only

26

through very long tidal channels, explaining the poorlydiversi f ied benthic fauna (Renesto & Tintor i , 1995).On the other hand, the restricted environment alloweddifferentiation of a largely endemic vertebrate faunai n c l u d i n g b o t h m a r i n e a n d t e r r e s t r i a l s p e c i e s .Superficially waters were well oxygenated, allowingnekton to thrive, and also at the margins of the basincond i t i ons a t bo t tom were favou rab le t o l i f e .Preservat ion is usual ly excel lent as a lmost a l l thespecimens are complete and fully articulated owing toan anoxic bottom and rapid sedimentat ion (Tintor i ,1992). Thin laminat ion of the foss i l i ferous levelsprovides further evidence of an undisturbed bottom atthe cen te r o f t he bas in . The foss i l s f ound a l l owreconstruction of life assemblages of both superficialwaters and benthic settings at the margins of the basin,as well as those of nearby islands, where terrestrialrepti les could l ive (Blake et al., 2000).

Abbreviations are as follows: ad, adnasal; lnrg, an-gular; ant, antorbital ; c l , c lei thrum; de, dental; dpt,dermopterotic; exsc, extrascapular; ifo3, infraorbital3, mx, maxil la; na, nasals; op, operculum; pa, parietal;p c l , p o s t c l e i t h r u m ; p m x , p r e m a x i l l a ; p o p ,preoperculum; ppa, postparietal; pt, posttemporal, ro,rostral, SC, scales; scl, Supracleithum, So, Supraorbitals;sop, suboperculum.

I n s t i t u t i o n a l a b b r e v i a t i o n s : M P U M : M u s e oPaleontologia Universitd degli Studi di Milano, Milano,I ta ly ; CMISNIO: Civ ico Museo Insubr ico d i Stor iaNaturale, Induno Olona (Varese, Italy).

Fig. | - Endennia l icia n. gen. n.sp.: restoration of the skull, a)

in lateral and b) in dorsal view.

Bollettino della Societa Paleontologica ltaliana, 44 ( I ), 2005

PALEONTOLOGICAL DESCRIPTION

Order PEnIEIDIFoRMES Berg, 1937

Family PEnlptuDAE Brough, l93l

Diagnosis

Endennia n. gen.

As for the only known species.

hp, species - Endennia licia n. gen. n. sp.

Etvmologv - From Endenna(Zogno, Bergamo, Lom-bardy), the name of locality where the holotype hasbeen found.

Tvpe locality - Zogno-Endenna (Bergaffio, Italy).

Age - Middle-late Norian (Late Triassic).

Geographical distribution - Bergamo Prealps.

Endennia licia n. gen. n. sp.P l . l ; F igs . l -6

Diagnosls (based on a combination of characters) -

Medium-sized perleidiform with fusiform body; skull-roof mainly formed by a single expanded parietal plate;total fusion of post-parietal with dermopterotic bones;supraorbital series made of a dozen elements of differ-ent size and shape arranged in rows; operculum l15 asdeep as the s[boperculum. Long marginal pencil-l iketeeth with a flattened cap and big and powerful, partlybicuspidate, crushing teeth disposed in several rowson palatal surface. Squamation consists of 42 scalerows, deeper than broad only in the anterior part of theflanks, ventral and posterior scales broader than deep;posterior margin of the scales slightly serrated, exceptfo r the vent ra l rows. Cauda l f in composed o f 32lepidotrichia; 8 epaxial rays.

Etl tmology From the Lat in word l ic ia, whichmeans comb, for the morphology and arrangement ofmarginal teeth.

Holotype - Specimen MPUM 8434 from Zogno-Endenna site (lower beds). Complete and articulatedspecimen of 160 mm in total length.

Type locality - Zogno-Endenna (Bergaffio, Italy).

Paratypes Specimen MPUM 8435 ( incompletejuvenile specimen) from P8 bed and 8436 (only skullr oo f l f r om P l 0 bed , a l l Zogno2 s i t e ; spec imenCMISNIO 267 from Endenna ( lower beds), (almostcomplete and articulated, in 195 mm of total length).

Age and horizon - Uppermost Calcare di Zorzino:middle-late Norian (Late Triassic).

Descript ion The rostral bone is a very large,subpentagonal element making the snout blunt in outline.The nasal bones are rounded elements irregular in shape;along the anterior margin there is the notch for theanter ior nar ia l opening. Each e lement contacts the

\ -.ifg I ant.. V \ ,

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Fig. 2 - Endennia licia n.MPUM 8434. b) Specimen

parietal bone at the anterolateral corner, where thesupraorbital sensory canal enters them (Figs. l-2).Theparietal bones are fused medially in a single large plate;it is characterrzed by large posterior and anterior regionsand nalrower median one, corresponding to the dorsalmargin of the orbit. The posterior outl ine is stronglyconvex. The two supraorbital sensory canals run veryc lose to each o the r med ia l l y . The bony p la te i sornamented by ganoine r idges and small tubercles,arranged mainly around the pores of the sensory canal(Figs. I -3). The postparietal and dermopterotic bonesof each side are fused to a single, wide L-shaped plate.The anterior, sharply pointed region of this elementembraces the postero-lateral margin of the parietal bone,and an anterior and a posterior pit-line are positionedon the postero-ventral corner (Figs . l-2). The mediansu tu re be tween them i s s t ra igh t and sho r t . Theextrascapular bones are sub-trapezoidal; they meet thepost-parietals*dermopterot ic bones along a straightsu ture ; owing to the s ta te o f p reserva t ion o f thespec imens , i t i s no t poss ib le t o s ta te i f t he twoextrascapulars were in contact. The supratemporalcommissu re , t yp i ca l l y T -shaped , w i t h pa tches o f

2 7

ganoine affanged around each pore, (Figs. l-2) is wellvisible. The posttemporal bones are small and roundedelements separated by the elements of the first scalerows (Figs. l -2).Of the infraorbital series only twofragmented elements have been detected: the trapezoidallarge antorbital and the IO3 (?), long and crescentic-shape, abutting the postero-ventral margin of the orbit(Fig. 2). The supraorbital series is very peculiar: it ismade of a dozen elements of different size and shape(Figs. l, 3). The anteriormost element, contacting thenasal bone, is much larger than the others and can bere fer red to as adnasa l in per le id i fo rms: i t has aquadrangular shape and it is followed posteriorly by amosaic of tesserae seemingly arranged in 3 rows. Aclear suture between maxillary and premaxillary bonesis not detectable; the powerful maxilla, with an expandedposterior region, bends slightly upwards. The anteriorpart juts out, being inclined anteroventrally-postero-dorsally; the oral margin is quite straight, hemmed byabout twenty teeth. On the anterior part of the maxilla,7 teeth are relatively longer, pointed and projectingslightly forwards; the others, more backwardly directed,decrease in length and show a different shape, becomingsmaller, stouter and blunt (Figs. l, 2, 4a). The apex ofthe anteriormost teeth is characterized by a slightlyflattened acrodine cup. Teeth different in size and shapeare visible on palatal bones, arranged in few (4?) rows:the anteriormost ones are smaller and pointed; medially,teeth are bicuspidate while the posterior ones are veryla rge and co inp le te ly f la t tened (F igs . 4b , c ) . Thepreopercular bone is backwardly oriented, with straightposterior margin; its outline is not clearly detectable inany specimen, owing to bad preservation of postorbitalregion, but it seems to be slightly enlarged in the dorsalregion; a wide infraorbital process is present. (Figs. I -

2 ) . T h e o p e r c u l u m i s m u c h s m a l l e r t h a n t h esubope rcu lum, be ing abou t I l 5 as deep as t he

Fig. 3 - Endennia l icia n. gen. n.sp.: the skull roof as preserved

in specimen MPUM 8436. Scale bar - 5 mm.

C. Lombardo, F. Brambillasca - A nev, perleidiform.from l{orthern ltallt

gen. n. sp. : skul ls . a)CMISNIO 267. Scale

The holotypebar - l0 mm.

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2 8 Bollettino della Societa Paleontologica Italiana, 44 ( I ), 2005

suboperculum; the dorsal margin is rounded while thesuture between operculum and suboperculum is quitestraight. The large suboperculum is quadrangular, witha rounded ventra l out l ine. (F igs. l -2) . In specimenCMISNIO 267, the lower ju* is visible in its antero-ventral region: i t is a strong element, sl ight ly bend

upwards, bearing at least 6 long and pointed teeth alongthe anterior oral margin. These teeth are similar in size,shape and affangement, decreasing in size posteriorly,to the anteriormost ones borne by the maxi l la. Thedentary is thickly ornamented by tubercles (Fig. 2b).It is not possible to state if branchiostegal rays werep resen t . The c le i t h rum i s c rescen t i c - shaped andornamented by a series of strong ridges running alongits anterior margin. The postcleithrum is subrectangular,deeper than broad, while the supracleithrum is oval-like, showing the same ornamentation of the cleithrumand it is clearly pierced by the sensory canal in its dorsalregion: its pores are very conspicuous and sulroundedby a r ing of tubercles (Fig. 2). The skul l bones areornamented by small ganoine tubercles mainly arrangedat the margins of each element.

Fins - The fan-shaped large pectoral fins are com-posed of at least l5 lepidotrichia: each of them con-sists of a long proximal segment followed by a seriesof much shorter, quadrangular distal ones. The firstray carries a series of strong fringing fulcra. At leasttwo spiny basal fulcra have been identified (Pl. l, fig.I ; Fig. 5a). The pelvic fins are placed at the level of thel7'h longitudinal row of scales; each of them consistsof at least 9 lepidotrichia, similar in structure to thoseof the pectorals. The anterior margin of the fins showsa series of fringing fulcra preceded by a couple of ba-sal ones (Pl . l , f ig . I ) .

The median fins are relatively small and triangular.The dorsal fin rs located at about the level of the 27'hrow of scales; it is composed of at least l3 lepidotrichiaand a series of small fringing fulcra has been observedalong the anterior margin. The anal fin, similar in shapeto the dorsal one, is composed by at least 9 lepidotrichia(Pl. l , f ig. I ; Fig. 5b) and i t is placed at the 27'h scalesrow . I n spec imen CMISNIO 267 the ana l f i n i spreceded by a couple of modified shield-shaped scale,much larger than the others, identified as the pre-analscales.

The externally almost symmetrical caudal fin con-sists of 32 lepidotrichia, with a very short axial body-lobe. There are 8 epaxial rays, with the first two seg-mented but not branched. Lepidotrichia with the samepattern are present at the base of the ventral lobe of thefin. All the other rays branch at least three times. Strongfringing fulcra and at least two basal fulcra are presenton edges of both dorsal and ventral lobe, even if thoseof the ventral lobe are much smaller (Pl. l, f igs. l-3;F ie. 6) .

Squamation - The squamation consists of 42 trans-verse scale rows, slightly deeper than broad only onmiddle-lateral region of the trunk. They decrease indepth both posteriorly and dorso-ventrally: in the dor-sal region they become rhombic,, as deep as broad. Amid-dorsal row of lanceolate scales is present. Scalesof the second horizontal row, below the mid-dorsalridge, show the openings of the dorsal lateral l ine. Thescales covering the posterior part of the body, thosebehind the dorsal f in and the caudal peduncle, and inthe ventral region of the trunk are broader than deep.The region between the pectoral f-rns is covered withvery smal l rhombic scales (Pl . l . f igs. l -3 ; F igs. 5b,c) .

Fig. 4 - Endennia licia n. gen.maxillary dentition. b) Same8435: the skul l with palatalbars - 5 mm.

n. sp. a) MPUM 8434: detail of thespecimen: palatal teeth; c) MPUMand premaxil lary denti t ion. Scale

gen. n. sp. : MPUM 8434. a) The r ightf-rn and detai l of the ventral scales. c)

bars - 5 mm.

2 9

gen. n. sp. : caudal f - rn of the holotypel 0 m m .

D I S C U S S I O N

SvsTEMATIc ' s

E n d e n n i a l i c i a n . g e n . n . s p . b e l o n g s t oPer le id i formcs for the general skul l pat tern, wi th aver t ica l ly or iented preoperculum, a maxi l la wi th anexpanded poster ior region, and a media l pentagonalrostral. Also the advanced fins structure is characteristico f p e r l e i d i f o r m s , a n d i n g e n e r a l o f t h e s o - c a l l e d"subholosteans" sensu Brough ( 1939). , wi th an equalratio between radials and lepidotrichia; these latter havelong unsegmented proximal elements and shorter distalones, as in neopterygians. The body lobe of the tail isvery short and the caudal f-rn is characterized by epaxialrays, inserted dorsally to the notochord. Perleidiformeshave been recent ly cons idered as a he terogeneousassemblage (Mutter, 2004): nevertheless, at momentwe find sti l l preferable to use the "classical" definit ionof prev ious authors (among others, see Hutchinson,1973: Gard iner , 1988; Gard iner & Schaef fe r , 1989;Btirgin, 1992; Tintori & Lombardo, 1996; Lombardo& Tintor i , 2004), s ince we bel ieve that the uniquecombinat ion of dermal skul l pattern, structure of themedian and caudal f ins and the lateral scales deeperthan wide on the anterior region of the body, representsa n e f f e c t i v e t o o l t o i d e n t i f y u n e q u i v o c a l l y t h erepresentatives of this group.

Among Per le id i formes, the character is t ics shownby the new genus match those of the family Perleididae( i nc lud ing the gene ra Pe r l e i dus , Mer idens ia . ,Aetheodontus, Ctenognathichthvs, Peltoperleidus.,Dipteronotus, Danin ia., Btirgin, 1992; Tintori,, I 998b;Lombardo, 2001 ) with the operculum generally smallerthan the subopercu lum, the preopercu lum dorsa l l yexpanded and narrower ventrally and the body coveredby thick but smooth ganoid scales, scales of the middle-lateral part of the trunk moderately deeper than broad.

C. Lornbordo, F. Brambillasca - A nev'perleidiform from l{orther"n ltalv

t , t , '

-r'ist

Fig. 6 - Endennia l icia n.MPUM 8434. Sca le bar :

Fig. 5 - Enclennia licia n.pectoral f rn: b) The analMid-dorsa l sca les . Sca le

All the scales are thick and smooth and show posteriorserration, except for those covering the caudal pedun-cle and the bel ly region, where they show a straightposterior margin. The art iculat ion between the scalesof the same vertical row seems to have been relativelyloose: in specimens MPUM 8434 and CMISNIO 267. ,wh ich are comple te and ar t i cu la ted , sca le cover ingappears "d is turbed", wi th the scales of most of thelongitudinal rows showing l itt le or not at all dorsal andventra l over lapping (Pl . l . , f igs. 2-3) : th is was prob-ably due to a small antero-dorsal art icular processes,as v is ib le on MPUM 8434.

3 0 Bollettino della Societa Paleontologica ltaliana, 44 ( I ), 2005

Endennia l icia n. gen. n. sp. shares with Perleididaealso the number of epaxial fin rays (six or seven), thepresence of a stout dentition on palatal bones and thepost-temporals separated by a couple of scales, even ifat least the first two characters are present in most ofthe perleidiform taxa.

Endennia n. gen. shows a medium-sized fusiformbody, covered by thick scales with serrated posteriormargin in the posterior flank, slightly deeper than broadon the antero-lateral region of the trunk; all the fins arerelatively small. The surface of skull bones is denselyornamented with ganoine ridges and tubercles and thedentition is well developed, both that of the oral marginsand the palatal elements. For the body out l ine, theoperculum/suboperculum size ratio and the shape andarrangement of anterior marginal teeth, Endennia n.gen. is reminiscent of some species belonging to theperleidid genera Peltoperleidus and Ctenognathichthvs,both from the lower Ladinian of Monte San Giorgio(Biirgin, 1992). With Peltoperleidus macrodontusEndennia n. gen. shares the pattern of the opercularregion, wi th the operculum much smal ler than thesuboperculum, and the fusion of part of the skull roof;moreover , P . macrodontus shows e longate andprotruding marginal teeth, as Endennia n. gen. does.Actual ly , pat terns of dent i t ion of the two taxa arediss imi lar , , being the teeth of P. macrodontus notdifferentiated, only decreasing in size posteriorly; themarginal dentition of Endennia n. gen. is on the contrarycharacterized by teeth of diverse morphologies: longand pointed anter ior ly and shor t and b lunt in theposterior region of the jaws. Concerning the fusion ofelements of the skull roof, P macrodontus shows fusedpost-parietals (Btirgin, 1992) but in Endennia n. gen.the fus ion is between the two f ronta ls , as wel l asbetween each par ie ta l + dermoptero t ic . A lso thesquamation pattern in P macrodontus is quite differentfrom that of Endennia n. gen.: in fact there are lesshorizontal scale rows and the lateral scales are muchdeeper than broad.

Concerning Ctenognathichthys bellottii, it is verysimilar to Endennia n. gen. in body outline and positionof the fins, with the anal and dorsal ones located farbehind the middle of the body length (Btirgin, 1992:Tintor i , 1998b). Also Ctenognath ichthys shows apeculiar dentition, made of large, fang-like teeth on theoral margin of both upper and lower jaws and smallerteeth on coronoids elements (Btirgin, 1992, 1996). Inspite of this, there are remarkable differences betweenthese two taxa: the marginal teeth of Endennia n. gen.are very e longate, but they are not as long as inCtenognathichthvs bellottii; besides, Endennia n. gen.has large and differentiated crushing teeth on palatalbones, lacking in Ctenognathichthys. The shape ofpreoperculum and consequently the orientation of theopercular region are other features that sep arateEndennia gen.n. from Ctenognathichthys bellottii. Bothgenera show a broad and massive preopercular bone,but in Endennia n. gen. it is dorso-ventrally elongateand shows a s t ra igh t pos te r i o r marg in , wh i l e i nCtenognathichthys bellottii it is clearly bent anteriorly.ln Endennia n. gen. both the upper and lower jaws aremuch longer and, different from Ctenognathichthys

bellotti i ,the maxil la has teeth also in its middle portion.Endennia n. gen. shows a skul l roof where the twoparietals are fused and an operculum which is about I I5 of the depth of suboperculum: in Ctenognathichthvsbe l l o t t i i pa r i e ta l s a re c l ea r l y sepa ra ted and theo p e r c u l u m i s o n l y s l i g h t l y s m a l l e r t h a n t h esuboperculum. Also appearance of the scales is verydifferent: the scales of the anterior part of the flank ofCtenognathichthlts are ornamented with irregular smallconcavities and naffow grooves horizontally arranged,while in Endennia gen. n. all scales are smooth.

Other perleidids show specialized long marginalteeth: Felberia excelsa (Lombardo & Tintori, 2004) and" DipteronotLts " ornotlts ( Btirgin, 1992), but thesespecies are clearly different in having a deep bodiedhabitus and short gapes, provided with marginal teetho n l y a n t e r i o r l y . M o r e o v e r , t h e i r s k u l l s l a c k t h esupraorbitals arranged in rows and in the opercularr e g i o n t h e o p e r c u l u m i s a t l e a s t a s d e e p a s t h esuboperculum. Marginal teeth are different from theinner ones, but dentit ion of the same region does notshow differentiation, since teeth change only in size.

So Endennia licia n. gen. n. sp. shows a combinationof characters, such as the rows of supraorbitals andthe ventral scales broader than deep, not found in anyknown taxon of Perleididae. In particular, the new genusshows a peculiar dentit ion, different from that of theother representatives of the family. In most of them, infact, adapted to a semi-durophagous diet, teeth borneby the oral maigin of both upper and lower jaws arepeg-like while palatal ones, inegularly arranged on bonyelements, are generally very small, domed and providedwith an acrodin cap. Crushing teeth generally show amorphology similar to that of grasping ones, being onlylarger. There are only few exception: for example, theEar ly Tr iass ic fo rms, such as C le i th ro lep is ,Hydropessum and Cleithrolnpidina, show slendertoothless, or almost toothless, lower ja* (Hutchinson,1973); the Ladinian genus Ctenognathichthys apparentlylacks crushing palatal teeth (Bt irgin, 1996), whi leFelberia shows long marginal teeth, with spatulate apexand blunt inner ones (Lombardo & Tintori, 2004). InEndennia licia n. gen. n. sp. outer and inner teeth arecompletely different; besides, there are morphologicald i f ferences even among teeth borne by the sameelements. The anteriormost teeth of maxil la are longand conical, while posteriorly teeth are short and blunt.On the palatal elements, the dentition consists of teethwhich are pointed anteriorly, bicuspidate medially andve ry l a rge and comp le te l y f l a t t ened pos te r i o r l y ;moreover, they are arranged in quite regular longitudinalrow s .

T h e p o s s e s s i o n o f a n u n i q u e c o m b i n a t i o n o fcharacters makes it indispensable to erect the new taxonEndenn ia l i c i a n . gen . n . sp . w i t h i n t he f am i l yPerleididae.

FpEorNc sTRATEGY

Concerning mode of feeding, Endennia licia n. gen.n. sp. can be compared with the neopterygians, owingto its fully-durophagous dentit ion. Even if in the Middle

C. Lombardo, F. Brambillasca - A new perleidiform from Northern ltaly

EXPLANATION OF PLATE I

bars : l0 mm.Figs. l -3I23

Endennia licia n. gen. n. sp. Scale- The holotype MPUM 8434.- Specimen CMISNIO 267.- Restoration.

P l . l

3 2 Bollettino della Societa Paleontologica ltaliana, 44 (l ), 2005

Triassic some "subholosteans" have reached a partiallytriturial dentit ion (Biirgin, 1996, Tintori, 1998a), highlyspec ia l i zed durophagous ac t inopteryg ians becomecommon and diversif ied only in the Norian, especiallywith semionotids and pycnodonts (Tintori, 1998a). InSargodon tomicus, for example, palatal teeth are largeand hemispherical and they are arranged in more orless regular rows; pointed teeth are aranged anteriorlyand laterally to the hemispherical ones. Sargodon alsohas incisiviform teeth, hamming premaxilla and dentary.For these characteristics, this semionotid is presumedto have fed upon hard-shelled organisms, grasped withthe chisel-l ike teeth and crushed with the grinding innerones (T in to r i , I 983 ) . Pycnodon ts a re even morespecial ized in durophagy, both in body shape, which isdeep and laterally compressed, and mouth morphology:prehensi l chisel- l ike teeth are on the premaxi l la anddentary, while the crushing teeth are arranged in regularrows on vomerine and prearticular bones (Tintori, l98l ;Nursall, 1996). The deep and flattened body of thesefishes allowed them to move in complex environmentand they probably were rather s luggish swimmers,f eed ing on ha rd -she l l ed i nve r teb ra tes , co ra l s o rencrust ing organisms. The body shape of Endennial ic ia n. gen. n. Sp., being quite fusiform, completelycovered by thick ganoine scales and furnished withrather small f ins,, does not suggest a similar mode ofl i fe. Nevertheless, in spite of i4he heavy squamation,the seemingly loose articulation between the scales couldhave improved the mobil ity of this fish, making it ableto pu rsue mov ing p reys . I t m igh t be the re fo rehypothesized that the anter ior teeth were used fors e i z i n g s m a l l s w i m m i n g o r g a n i s m s s u c h a scrustaceans, which were subsequently crushed withthe grinding teeth. Anyway, we do not exclude otherpossible food among the invertebrates provided withharder mineralized parts, as molluscs or echinoderffis,common i n t he Zo rz ino L imes tone . These l a t t e rcons t i tu ted a source o f food tha t was prev ious lyunderexploi ted by Middle Triassic act inopterygians,because of the primitive pattern of their skull, whosemobility was l imited by the cheek bones jointed to themaxilla. On the contrary, the new arrangement of theskull bones of the neopterygians, with the consequentlyimproved kinesis and the develop of a more powerfulmuscu la r sys tem, a l l owed them to ach ieve moresuccessful trophic adaptations. Endennia licia n. gen.n. sp. was able to reach full duroph agy like the moreadvanced neopterygians, even if in a different way: itwas not able to protrude the ethmoidal region, but theprojecting marginal teeth and the differentiated crushingones p robab l y a l l owed i t t o improve i t s f eed ingcapability.

CONCLUSIONS

After their extraordinary radiation during Anisian andL a d i n i a n , " s u b h o l o s t e a n s " w e r e t h o u g h t t o b e anegligible presence in Late Triassic faunas: their onlyNorian representative was considered, for a long time,Peltopleurus humilis from Seefeld (Kner, 1867).

As new localit ies and new specimens were found,it became clear that this group was more widespread

and diversif ied than once supposed; some Late Triassic"subholosteans" achieved also high special rzations, bothin denti t ion and body morphology, in spite of theiranatomical constraints and just before their last declinet o e x t i n c t i o n ( T i n t o r i & S a s s i , 1 9 9 2 : T i n t o r i &Lombardo, 1996).

One of these specialized forms is Thoracopterus,known f rom the Upper Tr iass ic of Europe (Bronn,1858; Bassani , 1895; Gr i f f i th , 1977; T intor i & Sassi ,1992). The representatives of this genus achieved thecapabil ity of gliding owing to the very long pectoraland pelvic fins (relate to standard length), associatedto a hypobatic caudal fin (Tintori & Sassi, 1992). Thiskind of spec ialization, achieved independently, is foundin ex tan t Exocoet idae, te leos tean " f l y ing f i shes"widespread in most temperate and tropical seas (Bertin,1958; Bruun, 1935; Hubbs , 1932). The adaptat ion togl ide achieved by the representat ives of the fami lyThoracopteridae could have been used to feed uponother fishes, escape from predators, or may be relatedto long distance migrat ions (Tintor i & Sassi, 1992).Probably in order to l ighten the body weight, NorianThoracopterus lacks the scale covering, but the trendto become lighter is clear also in other "subholostean"

taxa . A new genus found in the ear ly N or ian o fLumezzane Member of Garza Valley (Brescia Prealps;Brunetti et al., 200 I ), currently under study, is in factcharacter i zed by the absence of scales, except fewdorsal and ventral rows between median and caudalfins; moreover, it shows the same anal fin modification,r e l a t e d t o s e x u a l d i m o r p h i s m , s e e n i n o t h e rpeltopleuriforms and perleidiforms (Biirgin 1990, 1992;Lombardo 1999).

D i f fe ren t ly f rom most o f o ther per le id i fo rms,generally small and moderately fusiform fishes coveredwith heavy ganoid scales, Gabanellia agilis, from theNorian of the Zorzrno Limestone, is characterized by as lender body, about 25 cm long, a very th in scalecovering and a large, externally symmetrical and falcatetai l . These features, in associat ion with a powerfuldentition made of conical, radially striated, uneven teeth,clearly indicate that Gabanellia agilis was a predator,good swimmer, probably able to keep high speeds forlong d is tance, chasing smal l f ishes in open waters(Tintori & Lombardo, 1996). It is worthy to note thatneither in the Zorzino Limestone fauna, nor in otherTriassic ones, there are large pelagic actinopterygians;among this group, the first large swimmers specializedfor chase in open seas are the Jurass ic Caturus ,Hvpsocormlts and Pachycormus, all neopterygians fishes(Webb, 1984).

Among pe l t op leu r i f o rms , new taxa have beenrecently found both in the Zorzino Limestone (CL, pers.obs.) and the Lumezzane Member; moreover, in thefauna of Garza Valley, a single specimen temporarilyreferred to the genus Nannolepis (Griffith, 1977) wasfound.

Therefore, the finding of Endennia n. gen., with itsadaptation to strict durophagy, confirms once more boththe considerable presence of "subholosteans" duringthe Norian and their extraordinary versat i l i ty, whichal lowed them to compete wi th the more advancedneopterygians.

C. Lombardo, F. Brambillasca - A nev, perleidiform.from ltlorthern ltaly 3 3

ACKNOWLEDGEMENTS

The authors wish to thank G. Danini (lnduno Olona) for theloan o f spec imen CMISNIO 267 and S . Renes to f o r t hephotographs . The f ie ld -works in Zogno-Endenna s i tes ,supported by a 40o/o grants from Ministero Istruzione Universitde Ricerca, with the contribution of Bergamo Museum of NaturalH i s to r y "E . Ca f f i " , we re d i r ec ted by A . T in to r i , whosesuggestions also greatly improved this paper. Earlier version ofmanuscript was improved by R. Mutter and C. Sorbini.

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Manuscript received 05 July 2004Revised manuscript accepted 03 February 2005