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Home , Calcare di Zorzino, Colobodontidae, Dipteronotus, Endennasaurus, Thoracopterus

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

A newperleidiform (, ) frt ofNorthernItaly

Cristina LovtBARDo& FrancescaBnnrraBlLLASCA

"A. C. Lombardo, Dipartimento di Scienze della Terra Desio", Universitddegli Studi di Milano. Via Mangiagalli34. I-20133Milano, Italy; cristina.lombardo@unimi. it "A. ", F. Brambillasca,Dipartimento di Scienzedella Terra Desio Universiti degli Studi di Milano. Via Mangiagalli34, I-20133Milano, Ita ly.

KEY WORDS - , , Adaptive radiation, Durophqg'.

ABSTRACT - Endennia licia n. gen. n. sp., a new of perleidiform.from the Zorz, described. The nev, genus is charicterized bt, a .fusiform outline with a strong skull, b1 apex, c 'kindpeculiar kind of denition, w,ith long and cvlindrical marginol teeth, v'ith.flattened 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 .

RIASSUNTO - [Un nuovo perleidiforme (Osteichthyes, Actinopterygii) dal Triassicc descritto un nuovo genere e una nuova specie di pesce.fossile appartenente ai perleidiformi -(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. possibititd che I 'op'rovvisti di esoscheletro sottile quali'i crostacei, anche se non si esclttde la echinodermi, la cui presenza-d segnalata nella stessa unitd stratigrafica, anche sotto.fa molto diffi, Questo nuovo taxon occresce li conoscenza del gruppo dei perleidiformi, degli ultimi anni rivelano essere presente, qnche con.forme molto specializzate, durante

INTRODUCTION AND GEOLOGICAL SETTING or flyin g ()special rzations (Wild , 1978; Renesto,1994,2000). These animalsprobably lived In the last thirty years the Calcare di Zorzino on more or lesswide islandssituated around the basins; (Zorzino ,middle , Late Triassic) has they areusually small-sized, very lightly built, and often yielded an extraordinaryrich ichthyofauna:at least 25 have hollow bones:their preservationrequires a short generahave been already describedor are under study, post mortem transport in superficial oxic and warm but probably more than 50 genera are represented waters.Some otherscould have spentpart of their life (Tintori, 1981, 1983, 1995a,b, 1996a,b; Tintori & in water,especially for feeding,such as the thalattosaur Renesto,1983; Tintori & Sassi, 1992).A few taxa are Endennasourus,the armoured placodont Psephoderma representedby hundredsor thousandsof specimens, and the phytosaur (Renesto, 1992; like pholidophorids (Zambelli, 1975, 1978,I 980, I 98 I a, Renesto& Tintori, 1995;Gozzi & Renesto,2003). b, 1986), others by dozensand in some casesby few Invertebratesare also well representedby several or single specimen.Apart from the numbers,this fauna crustaceangenera (Pinna , 1974; Garassino& Teruzzi, is important becauseit recordsthe first major radiation 1993,Basso & Tintori, 1994)echinoderms (echinoids, of neopterygians (among these, pycnodonts, ophiuroids,crinoids and a singleasteroid) (Blake et al., semionotids,macrosemiids and pholidophorids),when 2000), cnidariansand scleractinia.Gastropods are rare paleopterygianswere still important, especiallyat the and bivalves are common, mainly as remains of "chondrosteans" top of the trophic hierarchy,with the ,and they contributedto constitutethe benthic "subholosteans" generaBirgeria andSaurichthys and the fauna,rich in individualsbut poor in species. Thoracopterusand Gabanellia (Tintori, 1990a;Tintori The Zorzino Limestone is inferred to have been & Sassi, 1992; Tintori & Lombardo, 1996; Tintori, depositedin a marine basin associatedwith early 1998a). Mesozoicrifting (Jadoulet al., 1992, 1994).The basin are also very important for the presenceof opened within a wide and thick carbonateplatform, endemic genera and of the oldest pterosaursso far the Dolomia PrincipaleFormation, that extendedoverall known. Although the Zorzino Limestone depositedin of the western margin of the Tethys. Becauseof the an intraplatform basin, most reptiles are terrestrialwith widespreadshallow-water environment, connections land (), arboreal () betweenthe basin and the open seawere probably only

/ss// 0375-7633 26 Bollettino della SocietaPaleontologica ltaliana, 44 ( I ), 2005 through very long tidal channels,explaining the poorly PALEONTOLOGICALDESCRIPTION diversifiedbenthic fauna (Renesto& Tintori, 1995). On the other hand, the restrictedenvironment allowed Order PEnIEIDIFoRMESBerg, 1937 differentiation of a largely endemic fauna Family PEnlptuDAE Brough, l93l including both marine and terrestrial species. Endennia n. gen. Superficially waters were well oxygenated,allowing nekton to thrive, and also at the margins of the basin Diagnosis As for the only known species. conditions at bottom were favourable to life. Preservationis usually excellent as almost all the hp, species - Endennia licia n. gen. n. sp. specimensare completeand fully articulatedowing to an anoxic bottom and rapid sedimentation(Tintori, Etvmologv - From Endenna(Zogno,Bergamo, Lom- 1992). Thin lamination of the fossiliferous levels bardy), the name of locality where the holotype has provides further evidence of an undisturbed bottom at been found. the center of the basin. The fossils found allow reconstructionof life assemblagesof both superficial Tvpe locality - Zogno-Endenna(Bergaffio, Italy). watersand benthic settingsat the marginsof the basin, as well as those of nearby islands, where terrestrial Age - Middle-lateNorian (Late Triassic). reptilescould live (Blake et al., 2000). Geographical distribution - Bergamo Prealps. Abbreviationsare as follows: ad, adnasal;lnrg, an- gular; ant, antorbital; cl, cleithrum; de, dental; dpt, Endennialicia n. gen.n. sp. dermopterotic;exsc, extrascapular;ifo3, infraorbital Pl. l; Figs. l-6 3, mx, maxilla; na,nasals; op, operculum;pa, parietal; - pcl, postcleithrum; pmx, premaxilla; pop, Diagnosls (basedon a combinationof characters) preoperculum;ppa, postparietal;pt, posttemporal,ro, Medium-sizedperleidiform with fusiform body; skull- rostral, SC,scales; scl, Supracleithum,So, Supraorbitals; roof mainly formed by a single expandedparietal plate; sop, suboperculum. total fusion of post-parietalwith dermopteroticbones; supraorbitalseries made of a dozen elementsof differ- Institutional abbreviations: MPUM: Museo ent size and shape arrangedin rows; operculum l15 as PaleontologiaUniversitd degli Studidi Milano, Milano, deep as the s[boperculum.Long marginal pencil-like Italy; CMISNIO: Civico Museo Insubricodi Storia teeth with a flattened cap and big and powerful, partly Naturale, Induno Olona (Varese,Italy). bicuspidate,crushing teeth disposedin severalrows on palatal surface.Squamation consists of 42 scale rows, deeperthan broad only in the anterior part of the flanks, ventral and posterior scalesbroader than deep; posteriormargin of the scalesslightly serrated,except for the ventral rows. Caudal fin composedof 32 lepidotrichia;8 epaxial rays.

Etltmology From the Latin word licia, which means comb, for the morphology and arrangementof marginal teeth. \ -.ifg I .V\, ant. Holotype - Specimen MPUM 8434 from Zogno- :-\- / Endennasite (lower beds). Complete and articulated specimenof 160 mm in total length.

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

Paratypes SpecimenMPUM 8435 (incomplete juvenile specimen)from P8 bed and 8436 (only skull roofl from P l0 bed, all Zogno2 site; specimen CMISNIO 267 from Endenna(lower beds), (almost completeand articulated,in 195 mm of total length).

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

Description The rostral bone is a very large, subpentagonalelement making the snoutblunt in outline. The nasalbones are rounded elements irregular in shape; Fig. | - Endennia licia n. gen. n.sp.: restoration of the skull, a) along the anterior margin there is the notch for the in lateral and b) in dorsal view. anterior narial opening. Each element contactsthe C. Lombardo, F. Brambillasca - A nev,perleidiform.from l{orthern ltallt 27

ganoineaffanged around each pore, (Figs. l-2) is well visible. The posttemporalbones are small and rounded elementsseparated by the elementsof the first scale rows (Figs. l-2).Of the infraorbitalseries only two fragmentedelements have been detected: the trapezoidal large antorbitaland the IO3 (?), long and crescentic- shape,abutting the postero-ventralmargin of the orbit

\_< (Fig. 2). The supraorbitalseries is very peculiar:it is l. pop\ of different size and shape I made of a dozen elements l' (Figs. l, 3). The anteriormostelement, contacting the nasal bone, is much larger than the others and can be referred to as adnasal in perleidiforms: it has a quadrangularshape and it is followed posteriorly by a mosaic of tesseraeseemingly arrangedin 3 rows. A clear suturebetween maxillary and premaxillary bones is not detectable;the powerful maxilla,with an expanded posterior region, bends slightly upwards. The anterior part juts out, being inclined anteroventrally-postero- dorsally; the oral margin is quite straight,hemmed by about twenty teeth.On the anteriorpart of the maxilla, 7 teeth are relatively longer, pointed and projecting slightly forwards;the others,more backwardlydirected, decreasein lengthand show a differentshape, becoming smaller,stouter and blunt (Figs. l, 2, 4a).The apex of the anteriormostteeth is characterizedby a slightly flattenedacrodine cup. Teethdifferent in sizeand shape are visible on palatalbones, arranged in few (4?) rows: the anteriormostones are smallerand pointed;medially, teeth are bicuspidatewhile the posteriorones are very large and coinpletely flattened (Figs. 4b, c). The preopercularbone is backwardlyoriented, with straight posteriormargin; its outline is not clearly detectablein any specimen,owing to bad preservationof postorbital region,but it seemsto be slightly enlargedin the dorsal - Fig. 2 - Endennia licia n. gen.n. sp.:skulls. a) The holotype region; a wide infraorbital processis present.(Figs. I MPUM 8434. b) Specimen CMISNIO 267. Scalebar- l0 mm. 2). The operculum is much smaller than the suboperculum,being about I l5 as deep as the

parietal bone at the anterolateralcorner, where the supraorbitalsensory canal entersthem (Figs. l-2).The parietalbones are fusedmedially in a single largeplate; it is characterrzedby large posteriorand anteriorregions and nalrower median one, correspondingto the dorsal margin of the orbit. The posterioroutline is strongly convex. The two supraorbital sensory canals run very \{ close to each other medially. The bony plate is 4i,,, ornamentedby ganoine ridges and small tubercles, t: i r,?i:Fc sr j f"""J:, ?::j,, arrangedmainly around the pores of the sensorycanal I ,'_;crr;$1 , (Figs. I -3). The postparietaland dermopteroticbones #i: €\i- t co "n. of each side are fusedto a single,wide L-shapedplate. R''','.'it,;I PA The anterior, sharply pointed region of this element k-lu'1 ,ffi \ parietal I embracesthe postero-lateralmargin of the bone, l and an anterior and a posterior pit-line are positioned dtl on the postero-ventralcorner (Figs. l-2). The median ,'""- ioo St€ suture between them is straight and short. The extrascapularbones are sub-trapezoidal;they meet the post-parietals*dermopteroticbones along a straight suture; owing to the state of preservationof the specimens,it is not possible to state if the two extrascapularswere in contact. The supratemporal Fig. 3 - Endennia licia n. gen. n.sp.: the skull roof as preserved commissure,typically T-shaped,with patchesof in specimen MPUM 8436. Scale bar - 5 mm. 28 Bollettino della SocietaPaleontologica Italiana, 44 ( I ), 2005

suboperculum;the dorsal margin is roundedwhile the upwards,bearing at least6 long and pointed teeth along suture between operculum and suboperculumis quite the anteriororal margin. Theseteeth are similar in size, straight.The large suboperculumis quadrangular,with shape and affangement,decreasing in size posteriorly, a roundedventral outline. (Figs. l-2). In specimen to the anteriormostones borne by the maxilla. The CMISNIO 267, the lower ju* is visible in its antero- dentary is thickly ornamentedby tubercles (Fig. 2b). ventral region: it is a strong element,slightly bend It is not possibleto state if branchiostegalrays were present.The cleithrum is crescentic-shapedand ornamentedby a seriesof strong ridges running along its anteriormargin. The postcleithrumis subrectangular, deeperthan broad, while the supracleithrumis oval- like, showing the sameornamentation of the cleithrum and it is clearlypierced by the sensorycanal in its dorsal region: its pores are very conspicuousand sulrounded by a ring of tubercles(Fig. 2). The skull bones are ornamentedby small ganoinetubercles mainly arranged at the margins of each element. Fins - The fan-shapedlarge pectoral fins are com- posed of at least l5 lepidotrichia:each of them con- sistsof a long proximal segmentfollowed by a series of much shorter,quadrangular distal ones. The first ray carries a seriesof strong fringing fulcra. At least two spiny basal fulcra have been identified (Pl. l, fig. I ; Fig. 5a). The pelvic fins are placedat the level of the l7'h longitudinal row of scales;each of them consists of at least 9 lepidotrichia,similar in structureto those of the pectorals.The anteriormargin of the fins shows a seriesof fringing fulcra precededby a couple of ba- sal ones(Pl.l, fig. I ). The median fins are relatively small and triangular. The rs located at about the level of the 27'h row of scales;it is composedof at leastl3 lepidotrichia and a seriesof small fringing fulcra has been observed along the anteriormargin. The anal fin, similar in shape to the dorsalone, is composedby at least9 lepidotrichia (Pl.l, fig. I ; Fig. 5b) and it is placedat the 27'hscales row. In specimenCMISNIO 267 the anal fin is precededby a couple of modified shield-shapedscale, much larger than the others, identified as the pre-anal scales. The externally almost symmetricalcaudal 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 same pattern are presentat the baseof the ventral lobe of the fin. All the other rays branch at leastthree times. Strong fringing fulcra and at least two basal fulcra are present on edgesof both dorsal and ventral lobe, even if those of the ventral lobe are much smaller (Pl. l, figs. l-3; Fie. 6). Squamation- The squamationconsists of 42 trans- verse scale rows, slightly deeperthan broad only on middle-lateralregion of the trunk. They decreasein depth both posteriorly and dorso-ventrally:in the dor- sal region they become rhombic,,as deep as broad. A mid-dorsalrow of lanceolatescales is present.Scales of the secondhorizontal row, below the mid-dorsal ridge, show the openingsof the dorsallateral line. The scalescovering the posterior part of the body, those behind the dorsal fin and the caudalpeduncle, and in Fig. 4 - Endennialicia n. gen.n. sp.a) MPUM 8434:detail of the than maxillarydentition. b) Samespecimen:palatal teeth; c) MPUM the ventral region of the trunk are broader deep. 8435:the skull with palataland premaxillarydentition. Scale The region betweenthe pectoralf-rns is coveredwith bars- 5 mm. very small rhombicscales (Pl.l. figs. l-3; Figs.5b,c). C. Lornbordo,F. Brambillasca - A nev'perleidiform from l{orther"nltalv 29

t,t,'

-r'ist

Fig. 6 - Endennia licia n. gen. n. sp.: caudal f-rnof the holotype MPUM 8434. Scale bar : l0 mm.

DISCUSSION

SvsTEMATIc's

Endennia licia n. gen. n. sp. belongs to Perleidiformcsfor the generalskull pattern,with a vertically orientedpreoperculum, a maxilla with an expandedposterior region, and a medial pentagonal rostral.Also the advancedfins structureis characteristic of perleidiforms,and in generalof the so-called "subholosteans"sensu Brough ( 1939).,with an equal ratio betweenradials and lepidotrichia;these latter have long unsegmentedproximal elementsand shorterdistal ones,as in neopterygians.The body lobe of the tail is very shortand the caudalf-rn is characterizedby epaxial rays,inserted dorsally to the notochord.Perleidiformes have been recently consideredas a heterogeneous assemblage(Mutter, 2004): nevertheless,at moment "classical" we find still preferableto usethe definition Fig. 5 - Enclennia licia n. gen.n. sp.:MPUM 8434.a) The right of previousauthors (among others,see Hutchinson, pectoral frn: b) The anal f-rnand detailof the ventralscales. c) 1973:Gardiner, 1988; Gardiner & Schaeffer,1989; Scalebars- 5 mm. Mid-dorsal scales. Btirgin, 1992; Tintori & Lombardo, 1996; Lombardo & Tintori, 2004), since we believe that the unique combinationof dermal skull pattern,structure of the median and caudal fins and the lateral scalesdeeper than wide on the anteriorregion of the body,represents All the scalesare thick and smoothand show posterior an effective tool to identify unequivocallythe serration,except for thosecovering the caudalpedun- representativesof this group. cle and the belly region,where they show a straight Among Perleidiformes,the characteristicsshown posteriormargin. The articulationbetween the scales by the new genusmatch those of the family Perleididae of the samevertical row seemsto have beenrelatively (including the genera , Meridensia., loose:in specimensMPUM 8434and CMISNIO 267., Aetheodontus, Ctenognathichthvs,Peltoperleidus., which are completeand articulated,scale covering ,Danin ia., Btirgin, 1992; Tintori,, I 998b; "disturbed", appears with the scalesof most of the Lombardo,2001 ) with the operculumgenerally smaller longitudinalrows showinglittle or not at all dorsaland than the suboperculum,the preoperculumdorsally ventraloverlapping (Pl. l., figs. 2-3): this was prob- expandedand narrowerventrally and the body covered ably due to a small antero-dorsalarticular processes, by thick but smoothganoid scales, scales of the middle- as visibleon MPUM 8434. lateral part of the trunk moderatelydeeper than broad. 30 Bollettino della SocietaPaleontologica ltaliana, 44 ( I ), 2005

Endennia licia n. gen. n. sp. shareswith Perleididae bellottii,themaxilla hasteeth also in its middle portion. also the number of epaxial fin rays (six or seven),the Endennia n. gen. shows a skull roof where the two presenceof a stout dentition on palatal bones and the parietalsare fused and an operculumwhich is about I I post-temporalsseparated by a couple of scales,even if 5 of the depth of suboperculum:in Ctenognathichthvs at least the first two charactersare presentin most of bellottii parietals are clearly separatedand the the perleidiform taxa. operculum is only slightly smaller than the Endennia n. gen. shows a medium-sizedfusiform suboperculum.Also appearanceof the scalesis very body, covered by thick scaleswith serratedposterior different: the scalesof the anterior part of the flank of margin in the posteriorflank, slightly deeperthan broad Ctenognathichthltsare ornamentedwith irregular small on the antero-lateralregion of the trunk; all the fins are concavitiesand naffow grooveshorizontally arranged, relatively small. The surfaceof skull bonesis densely while in Endennia gen. n. all scalesare smooth. ornamentedwith ganoine ridges and tuberclesand the Other perleidids show specializedlong marginal dentitionis well developed,both that of the oral margins teeth:Felberia excelsa(Lombardo & Tintori, 2004) and " " and the palatal elements.For the body outline, the DipteronotLts ornotlts ( Btirgin, 1992), but these operculum/suboperculumsize ratio and the shapeand speciesare clearly different in having a deep bodied arrangementof anterior marginal teeth, Endennia n. habitus and short gapes,provided with marginal teeth gen. is reminiscentof some speciesbelonging to the only anteriorly.Moreover, their skulls lack the perleidid generaPeltoperleidus and Ctenognathichthvs, supraorbitalsarranged in rows and in the opercular both from the lower of Monte San Giorgio region the operculum is at least as deep as the (Biirgin, 1992). With Peltoperleidus macrodontus suboperculum.Marginal teeth are different from the Endennia n. gen. sharesthe pattern of the opercular inner ones,but dentition of the same region does not region, with the operculum much smaller than the show differentiation,since teeth changeonly in size. suboperculum,and the fusion of part of the skull roof; So Endennialicia n. gen.n. sp. showsa combination moreover, P. macrodontus shows elongate and of characters,such as the rows of supraorbitalsand protruding marginal teeth, as Endennia n. gen. does. the ventral scalesbroader than deep, not found in any Actually, patternsof dentition of the two taxa are known taxon of Perleididae.In particular,the new genus dissimilar,,being the teeth of P. macrodontus not shows a peculiar dentition, different from that of the differentiated,only decreasingin size posteriorly; the other representativesof the family. In most of them, in marginaldentition of Endennian. gen.is on the contrary fact, adaptedto a semi-durophagousdiet, teeth borne characterizedby teeth of diverse morphologies: long by the oral maigin of both upper and lower jaws are and pointed anteriorly and short and blunt in the peg-likewhile palatalones, inegularly arrangedon bony posteriorregion of the jaws. Concerningthe fusion of elements,are generallyvery small, domedand provided elementsof the skull roof, P macrodontusshows fused with an acrodin cap. Crushing teeth generally show a post-parietals(Btirgin, 1992) but in Endennia n. gen. morphologysimilar to that of graspingones, being only the fusion is between the two frontals, as well as larger.There are only few exception:for example,the between each parietal + dermopterotic.Also the Early Triassic forms, such as Cleithrolepis, squamationpattern in P macrodontusis quite different Hydropessum and Cleithrolnpidina, show slender from that of Endennia n. gen.: in fact there are less toothless,or almost toothless,lower ja* (Hutchinson, horizontal scalerows and the lateral scalesare much 1973);the Ladinian genusCtenognathichthys apparently deeperthan broad. lacks crushing palatal teeth (Btirgin, 1996), while Concerning Ctenognathichthysbellottii, it is very Felberia showslong marginalteeth, with spatulateapex similar to Endennian. gen. in body outline and position and blunt inner ones (Lombardo & Tintori, 2004). In of the fins, with the anal and dorsal ones located far Endennia licia n. gen. n. sp. outer and inner teeth are behind the middle of the body length (Btirgin, 1992: completely different; besides,there are morphological Tintori, 1998b). Also Ctenognathichthysshows a differences even among teeth borne by the same peculiardentition, made of large,fang-like teeth on the elements.The anteriormostteeth of maxilla are long oral margin of both upper and lower jaws and smaller and conical,while posteriorlyteeth are short and blunt. teeth on coronoids elements(Btirgin, 1992, 1996). In On the palatal elements,the dentition consistsof teeth spite of this, there are remarkabledifferences between which are pointedanteriorly, bicuspidate medially and thesetwo taxa: the marginal teeth of Endennia n. gen. very large and completely flattened posteriorly; are very elongate, but they are not as long as in moreover,they arearranged in quite regularlongitudinal Ctenognathichthvsbellottii; besides,Endennia n. gen. row s. has large and differentiated crushing teeth on palatal The possessionof an unique combinationof bones, lacking in Ctenognathichthys.The shape of charactersmakes it indispensableto erectthe new taxon preoperculumand consequentlythe orientation of the Endennia licia n. gen. n. sp. within the family opercular region are other features that separate Perleididae. Endenniagen.n. from Ctenognathichthysbellottii. Both generashow a broad and massivepreopercular bone, but in Endennia n. gen. it is dorso-ventrallyelongate FpEorNc sTRATEGY and shows a straight posterior margin, whi le in Ctenognathichthysbellottii it is clearly bent anteriorly. Concerningmode of feeding,Endennia licia n. gen. ln Endennia n. gen. both the upper and lower jaws are n. sp. can be comparedwith the neopterygians,owing much longer and, different from Ctenognathichthys to its fully-durophagousdentition. Even if in the Middle C. Lombardo, F. Brambillasca - A new perleidiform from Northern ltaly Pl. l

EXPLANATION OF PLATE I

Figs. l -3 Endennialicia n. gen. n. sp. Scalebars: l0 mm. I - The holotype MPUM 8434. 2 - SpecimenCMISNIO 267. 3 - Restoration. 32 Bollettino della SocietaPaleontologica ltaliana, 44 (l ), 2005

"subholosteans" Triassicsome have reacheda partially and diversifiedthan oncesupposed; some Late Triassic "subholosteans" triturial dentition(Biirgin, 1996,Tintori, 1998a),highly achievedalso high specialrzations, both specializeddurophagous actinopterygians become in dentition and body morphology, in spite of their common and diversifiedonly in the Norian, especially anatomicalconstraints and just before their last decline with semionotidsand pycnodonts(Tintori, 1998a).In to (Tintori & Sassi, 1992: Tintori & Sargodon tomicus, for example, palatal teeth are large Lombardo, 1996). and hemisphericaland they are arrangedin more or One of these specializedforms is , lessregular rows; pointed teeth are aranged anteriorly known from the Upper Triassic of Europe (Bronn, and laterally to the hemisphericalones. Sargodon also 1858;Bassani, 1895; Griffith, 1977;Tintori & Sassi, hasincisiviform teeth,hamming premaxilla and dentary. 1992). The representativesof this genus achievedthe For thesecharacteristics, this semionotidis presumed capability of gliding owing to the very long pectoral to have fed upon hard-shelledorganisms, grasped with and pelvic fins (relate to standardlength), associated the chisel-liketeeth and crushedwith the grinding inner to a hypobaticcaudal fin (Tintori & Sassi, 1992).This ones (Tintori, I 983 Pycnodontsare even more kind of specialization, achieved independently, is found ). "flying specialized in durophagy,both in body shape,which is in extant Exocoetidae,teleostean " deepand laterallycompressed, and mouth morphology: widespreadin most temperateand tropical seas(Bertin, prehensilchisel-like teeth are on the premaxilla and 1958; Bruun, 1935; Hubbs, 1932).The adaptationto dentary,while the crushingteeth are arrangedin regular glide achievedby the representativesof the family rows on vomerineand prearticular bones (Tintori, l98l ; Thoracopteridaecould have been used to feed upon Nursall, 1996). The deep and flattened body of these other fishes, escapefrom predators,or may be related fishesallowed them to move in complex environment to long distancemigrations (Tintori & Sassi, 1992). and they probably were rather sluggish swimmers, Probably in order to lighten the body weight, Norian feeding on hard-shelledinvertebrates, corals or Thoracopteruslacks the scale covering, but the trend "subholostean" encrustingorganisms. The body shapeof Endennia to becomelighter is clear also in other licia n. gen. n. Sp.,being quite fusiform, completely taxa. A new genus found in the early N orian of covered by thick ganoine scalesand furnished with LumezzaneMember of Garza Valley (Brescia Prealps; rather small fins,,does not suggesta similar mode of Brunetti et al., 200I ), currently under study, is in fact life. Nevertheless,in spite ofi4he heavy squamation, characterized by the absenceof scales,except few the seeminglyloose articulation between the scalescould dorsal and ventral rows between median and caudal have improvedthe mobility of this fish, making it able fins; moreover,it showsthe sameanal fin modification, to pursue moving preys. It might be therefore related to sexual dimorphism, seen in other hypothesizedthat the anterior teeth were used for peltopleuriformsand perleidiforms(Biirgin 1990,1992; seizing small swimming organisms such as Lombardo 1999). crustaceans,which were subsequentlycrushed with Differently from most of other perleidiforms, the grinding teeth.Anyway, we do not exclude other generallysmall and moderatelyfusiform fishescovered possiblefood among the invertebratesprovided with with heavy ganoid scales,Gabanellia agilis, from the harder mineralizedparts, as molluscs or echinoderffis, Norian of the Zorzrno Limestone,is characterizedby a common in the Zorzino Limestone.These latter slenderbody, about 25 cm long, a very thin scale constituteda source of food that was previously coveringand a large,externally symmetrical and falcate underexploitedby actinopterygians, tail. These features,in associationwith a powerful becauseof the primitive patternof their skull, whose dentitionmade of conical,radially striated,uneven teeth, mobility was limited by the cheekbones jointed to the clearly indicate that Gabanellia agilis was a predator, maxilla. On the contrary, the new arrangementof the good swimmer, probably able to keep high speedsfor skull bonesof the neopterygians,with the consequently long distance,chasing small fishes in open waters improved kinesis and the develop of a more powerful (Tintori & Lombardo, 1996).It is worthy to note that muscular system,allowed them to achieve more neither in the Zorzino Limestone fauna, nor in other successfultrophic adaptations.Endennia licia n. gen. Triassic ones,there are large pelagic actinopterygians; n. sp. was able to reach full durophagy like the more among this group, the first large swimmers specialized advancedneopterygians, even if in a different way: it for chase in open seas are the Caturus, was not able to protrude the ethmoidal region, but the Hvpsocormltsand Pachycormus, all neopterygiansfishes projectingmarginal teeth and the differentiatedcrushing (Webb, 1984). ones probably allowed it to improve its feeding Among peltopleuriforms,new taxa have been capability. recentlyfound both in the Zorzino Limestone(CL, pers. obs.) and the LumezzaneMember; moreover, in the CONCLUSIONS fauna of Garza Valley, a single specimentemporarily referred to the genus Nannolepis (Griffith, 1977) was After their extraordinaryradiation during and found. "subholosteans" Ladinian, were thought to be a Therefore,the finding of Endennia n. gen., with its negligiblepresence in Late Triassicfaunas: their only adaptationto strict durophagy,confirms once more both "subholosteans" Norian representativewas considered,for a long time, the considerablepresence of during Peltopleurushumilis from Seefeld(Kner, 1867). the Norian and their extraordinaryversatility, which As new localitiesand new specimenswere found, allowed them to compete with the more advanced it becameclear that this group was more widespread neopterygians. C. Lombardo, F. Brambillasca - A nev,perleidiform.from ltlorthern ltaly 33

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