FIELDIANA Geology NEW SERIES, NO. 44 Marine from the Triassic of the Reptiles Tre Venezie Area, Northeastern Italy Olivier Rieppel Fabio Marco Dalla Vecchia as January 31, 2001 Publication 1511 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY JMJ18 2W2 Information for Contributors to Fieldiana General: Fieldiana is primarily a journal for Field Museum staffmembers and research associates, although manuscripts fromnonaffiliatedauthorsmaybeconsideredas spacepermits. The Journal carries a page charge of$65.00 per printed page or fraction thereof. Payment ofat least 50% of pagechargesqualifiesapaperforexpeditedprocessing, whichreducesthepublicationtime. Contributions from staff, research associates, and invited authors will be considered for publication regardless ofability to pay page charges, however, the full charge is mandatory fornonaffiliated authors ofunsolicited manuscripts. 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Fabio Marco Dalla Vecchia Museo Paleontologico Cittadino di Monfalcone Via Valentinis 134 1-34074 Monfalcone (Gorizia) Italy Accepted March 15, 1999 Published January 31, 2001 Publication 1511 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY © 2001 Field Museum ofNatural History ISSN 0096-2651 PRINTED IN THE UNITED STATES OF AMERICA GEOLOGY LIBRARY Table of Contents 9. ICymbospondylus sp.; isolated neural arch and proximal head of a rib 7 10. IShastasaurus sp.; isolated dorsal ver- tebral centrum 7 Abstract 1 11. Placodus cf. P. gigas; posterior pala- Introduction 1 tine tooth plates 8 Systematic Paleontology 2 12. Cyamodontoidea indet.; incomplete Ichthyosauria 2 neural arch 9 Sauropterygia 8 13. Cyamodontoidea indet., carapace frag- Discussion 20 ment 11 Acknowledgments 22 14. Cyamodontoidea indet.; nuchal region Literature Cited 23 of a carapace 12 15. ICyamodus sp.; partial palatine bone .. 13 16. Nothosaurus sp.; rib fragment with iso- lated tooth 13 List of Illustrations 17. Nothosaurus sp.; incomplete femur 13 18. Nothosaurus sp.; fragmentary mandible 14 19. Nothosaurus sp.; neural arch 14 21.. MLoicxaolsiatiuersusinstp.h,epTarretiaVlensekzeileetoanrea 23 222102... NNNooottthhhooosssaaauuurrruuusss cssfpp...;;N.ppaarrgttiiigaaallntsskkeuuullsll; partial 1156 3. Mixosaurus sp., neural spine 3 skull 17 4. ICymbospondylus sp.; isolated dorsal 23. Nothosaurus cf. N giganteus; lower vertebral centrum 4 jaw fragments 18 5. ICymbospondylus sp.; isolated dorsal 24. Nothosaurus cf. N. giganteus; vertebrae 19 vertebral centrum 5 25. Nothosaurus cf. N giganteus; distal 6. ICymbospondylus sp.; isolated dorsal end of clavicle 19 vertebral centrum 5 26. Cymbospondylus sp.; dorsal vertebral 7. ICymbospondylus sp.; isolated dorsal centrum 20 vertebral centrum 6 27. Lariosaurus calcagnii; humerus 21 8. ICymbospondylus sp.; partial dorsal 28. Sauropterygia indet.; partial skeleton .. 21 vertebral centrum 6 29. Sauropterygia indet.; thoracic rib 22 m Marine Reptiles from the Triassic of the Tre Venezie Area, Northeastern Italy Olivier Rieppel Fabio Marco Dalla Vecchia Abstract Ichthyosaurian and sauropterygian remains are described from Triassic deposits in the Tre Venezie area of northeastern Italy. The taxa recorded include Mixosaurus, Cymbospondylus, IShastasaurus, Placodus, ICyamodus, an as yet unnamed cyamodontoid placodont, Nothosau- rus sp., and Nothosaurus cf. N. giganteus. The temporal distribution of these fossils ranges from the earliest Anisian to the middle Carnian. Placodus (from the lower upper Anisian Calcare di Recoaro Formation of Vallarsa and Recoaro) is recorded for the first time (on the basis of diagnostic material) from the Alpine facies of the Middle Triassic. Collectively, the sauropterygian fauna from the Tre Venezie area resembles that of the eastern Alpine Triassic more closely than that of the southwestern Alps. This may indicate two separate colonization events ofthe Alpine Triassic as intraplatform basins developed along the northwesternTethyan margin. Introduction of the European platform before the late Anisian. It is at the Anisian-Ladinian boundary that the A number of taxa of marine reptiles (Thalat- fossil record of marine reptiles becomes richer in tosauria, Ichthyosauria, and Sauropterygia) have the Alpine Triassic. A well-corroborated phylog- been collected over the years from Triassic de- eny on which paleobiogeographic reconstructions posits in theTre Venezie area ofnortheastern Italy can be based is available for the Sauropterygia (Trentino-Alto Adige, Veneto, and Friuli-Venezia only at this time (Rieppel, 1999a). Given the pa- Giulia). The localities yielding Paleozoic andMe- leobiogeographic reconstruction of the late Ani- sozoic reptiles in these areas have been cata- sian western Tethyan realm by Marcoux et al. logued by Sirna et al. (1994), along with an iden- (1993) and the phylogenetic relationships of the tification of the faunal elements found at the re- Sauropterygia (Rieppel, 1997, 1998a), alternative spective localities (Fig. 1). Some of the material scenarios can be reconstructed for the invasion of has been described in greater detail, such as thal- the Alpine Triassic by Sauropterygia. A detailed attosaurcaudal vertebrae, nothosaur("Paranotho- comparison of the faunas from the Germanic Tri- saurus") vertebrae, and a tooth-bearing palatine assic (Muschelkalk) and from the southern Alps fragment of a cyamodontoid placodont (Dalla (Monte San Giorgio and equivalent deposits in Vecchia, 1993). Other material has remained un- northern Italy; Perledo) led Rieppel and Hagdorn described and forms thebasis ofthepresentstudy. (1997) to conclude that sauropterygians invaded The occurrences of Sauropterygia and other the southern Alpine realm through a southern marine reptiles in the Alpine Triassic range from gateway (Burgundian Gate), which linkedthe An- & the early Anisian (Sirna et al., 1994; Rieppel glo-Germanic Basin with the developing southern Hagdorn, 1997) to the Rhaetian (Pinna, 1990). branch of the Neotethys. In view of their eastern Lower Anisian records are exceedingly rare, not affinities (with Chinese taxa such as Sanchiaosau- so much because of a sampling bias, but rather rus, Keichousaurus, and Hanosaurus: Rieppel & because thecharacteristic intraplatformbasinhab- Lin, 1995; Rieppel, 1998a, 1999b), sauropterygi- itats did not develop along the southeastern shelf ans might alternatively have reached the eastern FIELDIANA: GEOLOGY, N.S., NO. 44, JANUARY 31, 2001, PP. 1-25 Fig. 1. Localities in the Tre Venezie area (NE Italy) yielding Triassic marine vertebrates. 1, Ambruseit Creek, ArtaTerme(UdineProvince); 2,ClapdiVal, Forni di Sotto(UdineProvince);3,PiandelleStreghelocalityofBivera Mountain, Forni di Sotto (Udine Province); 4, Leno Creek, Specchieri (Trento Province); 5, Recoaro, (Vicenza Province);6,FuseanearTolmezzo(UdineProvince);7,FusCreek,MoggioUdinese(UdineProvince);8,Mt.Tersadia, ArtaTerme (Udine Province); 9, Lavaz Creek, Dogna (Udine Province). Alpine realm along the southeastern continental Giulia Region. Lower Pelsonian (Cuccense sub- shelf ofEurope, bordering on the western Tethys zone), lowermost upper Anisian. in the northeast and on the western branch ofthe The specimen (mfsn 19385) was collected by Neotethys in the south. In view of these alterna- C. Rosenfeld in the bed ofAmbruseit Creek near tive scenarios, a more detailed analysis ofmarine the village of Piedim (Arta Terme). Specimen reptiles from the Triassic ofthe southeastern Alps mfsn 19385 comes from alternating black silt- is likely to provide important new information on stones and gray nodular limestones with subor- the paleobiogeography of this region. dered black marly limestones. This interval is sit- uated below and in continuity with the Bivera Formation (lower Illyrian; Farabegoli et al., 1984) and is attributed to the Dont Formation (Carulli Systematic Paleontology et al., 1987). On the basis of ammonoids, the age can be determined as early Pelsonian, earliest late Ichthyosauria Blainville, 1835 Anisian (P. Mietto, pers. comm.). Mixosauridae Baur, 1887 The specimen comprises a total of 15 caudal Mixosaurus Baur, 1887 vertebral centra, ofwhich 3 are very incompletely preserved (Fig. 2). All vertebral centra are weath- Mixosaurus sp. ered to a sagittal or parasagittal plane, exposing — their deeply amphicoelous and notochordal struc- Material Museo Friulano di Storia Naturale ture. The length ofthe centra ranges from 12.5 to (mfsn), Udine, no. 19385; a partially disarticulat- 6 mm, and their height varies from 15 to 14 mm; ed string of 15 caudal vertebral centra (of which the height/lengthratio varies from 1.12 to 2.3. As- 3 are poorly preserved) associated with neural sociated with the caudal centra are neural arches arches (Fig. 2). — carrying tall neural spines. The basal portions of Locality and Horizon Ambruseit Creek, the neural arches are sutured but not fused to the Arta Terme, Province of Udine, Friuli-Venezia vertebral centra. Thebases (pedicels) oftheneural FIELDIANA: GEOLOGY AniFsiiga.n.2.ScaMliexobsaaru=rus20spm.m(.mfsn 19385) from Arta Terme, Ambruseit Creek, Province ofUdine, lowermost upper arches are generally expanded; one well-pre- and convex posterior margin. Elongation and cur- served and complete neural arch is remarkable in vatureoftheneural spines arecharacteristic ofthe that it shows a greatly expanded pedicel, which middle tail section of Mixosaurus (Repossi, appears to carry distinct pre- and postzygapo- 1902). Only a single elementis preserved thatcan physes (Fig. 3). Repossi (1902, pi. IX, fig. 9e) be identified as an incomplete chevron. figured the neural arches oftail vertebrae ofMix- Mixosaurus has a cosmopolitan distribution & osaurus cornalianus, which bear pre- and post- during the Middle Triassic (Callaway Massare, zygapophyses but show much less expanded ped- 1989; Mazin & Sander, 1993; Sander & Mazin, icels. The neural spines are slender and elongate. 1993), with five species currently recognized. They slant slightly posterodorsally and show a These are Mixosaurus atavus (Quenstedt, 1852), slight curvature, resulting in a concave anterior from the lower Muschelkalk of southern Germa- ny; Mixosaurus cornalianus (Bassani, 1886), from the Grenzbitumen horizon (Anisian-Ladini- an boundary) of southern Switzerland and north- ern Italy; Mixosaurus nordenskioeldii (Hulke, 1873), from the Middle Triassic of Spitzbergen and British Columbia; Mixosaurus natans (Mer- riam, 1908), from the Middle Triassic of north- western Nevada (Sander & Bucher, 1990); and Mixosaurus maotaiensis Young, 1965, from the lower Middle Triassic of China (Kweichou Prov- ince). Early Triassic remains ofMixosaurus have been reported from British Columbia (Callaway & Brinkman. 1989), while intheAnglo-Germanic Basin, Mixosaurus extends back to the Gogolin Fig. 3. NeuralarchofMixosaurussp. (mfsn 19385). beds (?Bithynian, lower Anisian) of Gorny Slask Scale bar =10 mm. (Huene, 1916). Coming from the Pelsonian (low- RIEPPEL & DALLA VECCHIA: TRIASSIC MARINE REPTILES FROM ITALY Fig. 4. ICymbospondylus sp. (mfsn 15275); isolated dorsal vertebral centrum from Clap di Val, Forni di Sotto, Province ofUdine, upper Ladinian. A, Anterior view; B, posterior view; C, right lateral view; D, left lateral view; E,dorsal view; F, cross-sectional area. Scalebar = 20 mm. Abbreviations: dia, diapophysis (fortheribarticulation); nc, neural canal. ermost upper Anisian), the specimen mfsn 19385, Sotto, Province ofUdine. "Formazione delle cal- from the Carnian Alps, is the earliest record for careniti rosseegrigie," Longobardian, upperLad- the genus in the Alpine Triassic. The incomplete- inian. ness of the fossil renders comparison with other The vertebrae were collected at the locality of species ofMixosaurus difficult, but the broad ex- Clap di Val, near the village of Forni di Sotto pansion of the pedicel of the neural arch appears (Udine). Specimen mfsn 15275 comes from the to be a unique character ofthe Friulian specimen. UA5 interval of "red and gray calcarenites For- mation" (Pisa, 1966), dated to as early as middle Longobardian (late Ladinian: P. Mietto, pers. Shastasauridae Merriam, 1902 coram.), from which a rich ammonoid fauna was Cymbospondylus Leidy, 1868 collected as well. It is a nearly complete centrum, lacking only a small splinter in the ventrolateral ICymbospondylus sp. anteriorpart (Fig. 4). In posteriorview itis slimghmt- — ly deformed by lithostatic pressure. It is 23.6 mm Material Museo Friulano di Storia Naturale, long, 42.8 high (1.81 height/length ratio), and mm Udine, no. 15275; isolated dorsal vertebral cen- the horizontal diameter is 46.0 (anterior sur- mm trum. Museo Paleontologico of Portogruaro (Ve- face) and 39.5 (posterior surface), respective- nezia), uncatalogued specimen; isolated dorsal ly. The anterior and posterior ends ofthe centrum vertebral centrum. — are deeply amphicoelous and notochordal, but the Locality and Horizon Clap di Val, Forni di articular surface is only weakly inclined in the FIELDIANA: GEOLOGY