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Phylogenetic Position of the Crocodylian Megadontosuchus Arduini and Tomistomine Palaeobiogeography

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Phylogenetic Position of the Crocodylian Megadontosuchus Arduini and Tomistomine Palaeobiogeography Phylogenetic position of the crocodylian Megadontosuchus arduini and tomistomine palaeobiogeography PAOLO PIRAS, MASSIMO DELFINO, LETIZIA DEL FAVERO, and TASSOS KOTSAKIS Piras, P., Delfino, M., Del Favero, L., and Kotsakis, T. 2007. Phylogenetic position of the crocodylian Megadontosuchus arduini and tomistomine palaeobiogeography. Acta Palaeontologica Polonica 52 (2): 315–328. A cladistic analysis of Megadontosuchus arduini from the middle Eocene of Monte Duello (NE Italy) confirms its tomistomine relationships, but the low number of scorable characters determines a low resolution within the tomistomine clade. However, Megadontosuchus is clearly distinct from the other Eocene European or North African tomistomines, in having a moderate elongated but robust rostrum, massive maxillary and dentary teeth and large supratemporal fenestrae. The rostrum and teeth characteristics could indicate that M. arduini had a degree of feeding specialization intermediate between Maroccosuchus zennaroi and the Eocene European tomistomines. A summary of tomistomine palaeobiogeo− graphy suggests that despite only one species with a rather restricted range survives at present, such a clade had a glorious past with a world wide distribution documented by a conspicuous fossil record that starts at least in the early Eocene. At present, a detailed knowledge of tomistomine palaeobiogeography is hindered by the lack of modern taxonomic revisions of some fossil remains and therefore by the poor understanding of phylogenetic relationships. Key words: Crocodylidae, Tomistominae, Tomistoma schlegelii, palaeobiogeography, Eocene, Italy. Paolo Piras [[email protected]] and Tassos Kotsakis [[email protected]], Dipartimento di Scienze Geologiche, Università Roma Tre, Largo S. Leonardo Murialdo 1, I−00146, Roma, Italy; Massimo Delfino [[email protected]], Dipartimento di Scienze della Terra, Università di Firenze, Via La Pira 4, I−50121 Firenze, Italy; Letizia Del Favero [[email protected]], Museo di Geologia e Paleontologia, Università degli studi di Padova, Via Giotto 1, I−35137, Padova, Italy. Introduction —currently preserved in the Gand Museum, Belgium, K. spenceri—BMNH 19633, and M. arduini—MGPD 1Z) evi− De Zigno (1880) described longirostrine crocodylian re− denced that the three forms represent probably three different mains from the middle Eocene of Monte Duello (Verona, species. Recent phylogenetic analyses (Jouve 2004; Delfino Italy) and formally erected a new species upon this material, et al. 2005; Brochu 2006, in press) showed that Dollosuchus Crocodylus arduini (originally written as Crocodilus). Since dixoni is a basal tomistomine close to Kentisuchus spenceri. then, many authors (Kuhn 1938; Brochu 1997, 2001; Kot− Recently, Brochu (in press) expressed doubts about the ge− Dollo− sakis et al. 2004) declined to include this taxon in the genus neric and specific identity for the Belgian specimen of suchus dixoni. No modern description or phylogenetic analy− Crocodylus, placing it instead among tomistomines. Mook sis have been published until now for Megadontosuchus (1955) proposed a new genus for this species, Megadonto− arduini. We present here a complete list of the remains attrib− suchus, and in the same paper erected another new tomisto− uted to M. arduini to date (Appendix 1), a redescription of all mine genus, Kentisuchus, for “Crocodylus” spenceri Buck− known material referred to this species and a cladistic analy− land, 1836. A number of papers describe or mention M. sis primarily based on Gatesy et al.'s (2004) dataset. As the arduini, accepting its specific validity (Nicolis 1882; Lydek− holotype has never been designated by de Zigno (1880) nor a ker 1886; Uzielli 1886; Fabiani 1915; Del Vecchio 1921; lectotype by Mook (1955), we choose the specimen MGPD Mook 1955; Steel 1973; Altichieri 1980; Pinna 1989; Rocca− 1Z as the lectotype (figured in de Zigno 1880: pls. 1, 2, and forte et al. 1994; Brochu 2001). Kotsakis et al. (2004, 2005) here as Figs. 1–3). quoted this species as a basal tomistomine. Brochu (1997, 2001) hypothesized that Kentisuchus spenceri, Megadonto− Institutional abbreviations.—BMNH, The Natural History suchus arduini, and Dollosuchus dixoni (Owen, 1850) are Museum, London, United Kingdom; IRScNB, Institut Royal closely related and underlined that in case of congeneric at− des Sciences Naturelles, Bruxelles, Belgium; MGPD, Museo tribution the genus Dollosuchus should have priority. Direct di Geologia e Paleontologia, Università degli Studi di Padova, observation of the type specimens (D. dixoni—IRScNB 482 Italy. Acta Palaeontol. Pol. 52 (2): 315–328, 2007 http://app.pan.pl/acta52/app52−315.pdf 316 ACTA PALAEONTOLOGICA POLONICA 52 (2), 2007 supratemporal fenestra frontal squamosal postorbital prefrontal lacrimal nasal premaxilla parietal occipital condyle supraoccipital temporal canal quadrate external naris orbit maxilla jugal 50 mm postorbital infratemporal bar quadratojugal fenestra Fig. 1.Tomistomine crocodile Megadontosuchus arduini (de Zigno, 1880), MGPD 1Z, lectotype from Monte Duello (Roncà, Province of Verona, NE Italy); middle Eocene. Skull in dorsal view. Photograph (A) and explanatory drawing of the same (B). Referred material.—MGPD 5Z, skull lacking the anterior Systematic palaeontology part of the rostrum; MGPD 4Z, anterior part of the rostrum belonging to the specimen MGPD 5Z; MGPD 6Z, fragment Eusuchia Huxley, 1875 of a right mandibular ramus, lacking retroarticular process; Crocodylia Gmelin, 1789 (sensu Benton and Clark MGPD 8Z, nearly complete mandibular ramus; probably 1988) these remains belong to the original type series. Two verte− Crocodyloidea Fitzinger, 1826 (sensu Brochu 2003) brae MGPD 24Z (cervical) and MGPD 25Z (dorsal; de Zigno Crocodylidae Cuvier, 1807 (sensu Brochu 2003) 1880: pl. 2: 6 and 4, 5, respectively). Tomistominae Kälin, 1955 (sensu Brochu 2003) Emended diagnosis.—Megadontosuchus arduini is a tomisto− mine differing from all other members of the clade because of Genus Megadontosuchus Mook, 1955 this combination of features: the robust rostrum, the massive Type species: Crocodilus arduini de Zigno, 1880; see below. maxillary and dentary teeth and large supratemporal fenestrae. Megadontosuchus arduini (de Zigno, 1880) Approximately 14–16 maxillary teeth, and approximately 14– Lectotype: MGPD 1Z, a nearly complete skull (Figs. 1–3) with lower 17 dentary teeth. Small occipital condyle. Large narial aperture jaw (de Zigno 1880: pl. 1: 1–3 and pl. 2: 1–3). and foramen incisivum. Nasals in contact with external naris Type locality and age: Monte Duello (Roncà, Province of Verona, NE without bisecting it. Large mandibular fenestra. Elongated Italy); middle Eocene. squamosal prongs reaching nearly the half of quadrate ramus. PIRAS ET AL.—REDESCRIPTION OF MEGADONTOSUCHUS ARDUINI 317 infratemporal quadratojugal fenestra orbit 50 mm palatine maxilla foramen incisivum quadrate occipital condyle premaxilla occlusal pits enlarged teeth palatine fenestra supratemporal postorbital fenestra bar Fig. 2. Tomistomine crocodile Megadontosuchus arduini (de Zigno, 1880), MGPD 1Z, lectotype from Monte Duello (Roncà, Province of Verona, NE Italy); middle Eocene. Skull in ventral view. Photograph (A) and explanatory drawing of the same (B). riorly, the quadrate ramus is very expanded medio−laterally (in Description occipital view it is nearly the half of semi cranial width at max− imum quadrate lateral expansion). The skull morphology in Skull ventral view (Fig. 2) is difficult to reconstruct precisely due to Preservation, form, and general features.—The lectotype the collapse of secondary palate posteriorly to the maxillo−pal− (Figs. 1–3) is an almost complete skull that, although dorso− atine suture; such suture is wedge−like shaped. Palatines are ventrally flattened, retains the undulatory outline (“festoon− antero−posteriorly elongated. Pterygoids and ectopterygoids ing”) of the maxillary and dentary visible in dorsal and, de− are totally crushed down. Choanae are not visible. Despite a spite the deformation, lateral view. It is very difficult to recog− marked erosion, basioccipital seems to be medio−distally nize cranial and mandibular sutures due to the preservation short. In occipital view (Fig. 3) only the sutures between squa− and historical museum preparation that focused on the mainte− mosals and exoccipitals are slightly visible. The general pro− nance of the general external shape and not of the fine mor− file is flattened and compressed. phology. The minimal distance between supratemporal fene− Dorsal view.—In dorsal view, the lectotype shows an evi− strae is approximately 10 mm. Only the right infratemporal dent festooning despite the post−mortem deformation. Exter− fenestra is visible due to the absence of the left jugal bar. The nal nares are wide but antero−posteriorly elongated, with jugal border of the right orbit is medially displaced. The right maximum width at their anterior border. It is difficult to eval− orbit appears longer, due to post−mortem deformation. Poste− uate the position of the anterior nasal tip; however, the nasals http://app.pan.pl/acta52/app52−315.pdf 318 ACTA PALAEONTOLOGICA POLONICA 52 (2), 2007 squamosal foramen magnum 50 mm exoccipital quadrate occipital condyle basioccipital tubera Fig. 3. Tomistomine crocodile Megadontosuchus arduini (de Zigno, 1880), MGPD 1Z, lectotype from Monte Duello (Roncà, Province of Verona, NE Italy); middle Eocene. Skull in occipital view. Photograph (A) and explanatory drawing of the same (B). profile suggests
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