Abstract
Small terrestrial non-eusuchian mesoeucrocodylians are common components of Cretaceous assemblages of Gondwanan provinces with notosuchians and araripesuchids as flagship taxa in South America, Africa and Madagascar, well into the Late Cretaceous. On the other hand, these are exceedingly rare in Laurasian landmasses during the Late Cretaceous. Small terrestrial mesoeucrocodylians from Europe were often referred to the genus Theriosuchus, a taxon with stratigraphic range extending from the Late Jurassic to the late Early Cretaceous. Theriosuchus is abundantly reported from various European localities, although Asiatic and possibly North American members are also known. It has often been closely associated with the first modern crocodilians, members of the Eusuchia, because of the presence of procoelous vertebrae, a widespread key character diagnosing the Eusuchia. Nevertheless, the relationships of Theriosuchus have not been explored in detail although one species, Theriosuchus pusillus, has been extensively described and referred in numerous works. Here, we describe a new basal mesoeucrocodylian, Theriosuchus sympiestodon sp. nov. from the Maastrichtian of the Haţeg Basin, Romania, suggesting a large temporal gap (about 58 myr) in the fossil record of the genus. Inclusion of the new taxon, along with Theriosuchus guimarotae, in a phylogenetic analysis confirms its referral to the genus Theriosuchus, within a monophyletic atoposaurid clade. Although phylogenetic resolution within this clade is still poor, the new taxon appears, on morphological grounds, to be most closely related to T. pusillus. The relationships of Atoposauridae within Mesoeucrocodylia and especially to Neosuchia are discussed in light of the results of the present contribution as well as from recent work. Our results raise the possibility that Atoposauridae might not be regarded as a derived neosuchian clade anymore, although further investigation of the neosuchian interrelationships is needed. Reports of isolated teeth referable to a closely related taxon from the Upper Cretaceous of Romania and France, together with the presence of Doratodon and Ischyrochampsa, indicate a previously unsuspected diverse assemblage of non-eusuchian mesoeucrocodylians in the Late Cretaceous European archipelago.
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References
de Andrade MB, Bertini RJ (2008) A new Sphagesaurus (Mesoeucrocodylia: Notosuchia) from the Upper Cretaceous of Monte Alto City (Bauru Group, Brazil), and a revision of the Sphagesauridae. Hist Biol 20:101–136
Benton MJ, Clark JM (1988) Archosaur phylogeny and the relationships of the Crocodilia. In: Benton MJ (ed) The phylogeny and classification of tetrapods. Systematics association special volume no. 35A Amphibians, reptiles, birds, vol. 1. Clarendon Press Oxford, UK, pp 295–338
Bojar AV, Grigorescu D, Ottner F, Csiki Z (2005) Paleoenvironmental interpretation of dinosaur- and mammal bearing continental Maastrichtian deposits, Haţeg basin, Romania. Geol Q 49:205–222
Brinkmann W (1992) Die Krokodilier-fauna aus der Unter-Kreide (Ober-Barremium) von Uña (Provinz Cuenca, Spanien). Berl Geowiss Abh 5:1–123
Brochu CA, Wagner JR, Jouve S, Sumrall CD, Densmore LD (2009) A correction corrected: consensus over the meaning of Crocodylia and why it matters. Syst Biol 58:537–543. doi:10.1093/sysbio/syp053
Buckley GA, Brochu CA (1999) An enigmatic new crocodile from the Upper Cretaceous of Madagascar. Sp Pap Palaeontol 60:149–175
Buckley GA, Brochu CA, Krause DW, Pol D (2000) A pug-nosed crocodyliform from the Late Cretaceous of Madagascar. Nature 405:941–944
Buffetaut E (1979) Revision der Crocodylia (Reptilia) aus den Gosau-Schichten (Ober-Kreide) von Österreich. Beitr Paläontol Österr 6:89–105
Buffetaut E (1982) Radiation évolutive, paléoécologie et biogéographie des Crocodiliens mésosuchiens. Mém Soc Géol Fr num sp 60:1–88
Buffetaut E (1983) The crocodilian Theriosuchus Owen, 1879 in the Wealden of England. Bull Brit Mus Nat Hist Geol 37:93–97
Bunzel E (1871) Die Reptilfauna der Gossau-Formation in der Neuen Welt bei Wiener-neustadt. Abh Geol Reichs 5:1–18
Buscalioni AD, Sanz JS (1988) Phylogenetic relationships of the Atoposauridae (Archosauria, Crocodylomorpha). Hist Biol 1:233–250
Clark JM (1986) Phylogenetic relationships of the crocodylomorph archosaurs. Ph.D. dissertation. University of Chicago, Chicago, p 556
Clark JM (1994) Patterns of evolution in Mesozoic Crocodyliformes. In: Fraser N, Sues HD (eds) In the shadow of the dinosaurs: Early Mesozoic tetrapods. Cambridge University Press, New York, pp 84–97
Company J, Pereda-Suberbiola X, Ruiz-Omeñaca JI, Buscalioni AD (2005) A new species of Doratodon (Crocodyliformes: Ziphosuchia) from the Late Cretaceous of Spain. J Vertebr Paleontol 25:343–353
Cuny G, Buffetaut E, Cappetta H, Martin M, Mazin JM, Rose JM (1991) Nouveaux restes de Vertébrés du Jurassique terminal du Boulonnais (Nord de la France). N Jb Geol Paläont Abh 180:323–347
Delfino M, Codrea V, Folie A, Dica P, Godefroit P, Smith T (2008) A complete skull of Allodaposuchus precedens Nopcsa, 1928 (Eusuchia) and a reassessment of the morphology of the taxon based on the Romanian remains. J Vertebr Paleontol 28:111–122
Folie A, Codrea V (2005) New lissamphibians and squamates from the Maastrichtian of Haţeg Basin, Romania. Acta Palaeontol Pol 50:57–71
Galton PM (1996) Notes on Dinosauria from the Upper Cretaceous of Portugal. N J Geol Pal Mh 2:83–90
Gmelin JF (1789) Caroli a Linné systema naturae per regna tri naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. GE Beer, Leipzig
Grigorescu D (1992) Nonmarine Cretaceous formations of Romania. In: Mateer NJ, Chen PJ (eds) Aspects of nonmarine Cretaceous geology. Ocean Press, Beijing, pp 142–164
Grigorescu D, Csiki Z (2000) Sedimentology taphonomy and paleoecologic reconstruction of the Tuştea nesting site from the Upper Maastrichtian of the Haţeg Basin (Romania). In: Frey E (ed) 5th European Workshop on Vertebrate Palaeontology, Kalsruhe, Volume of Abstracts, p. 33
Grigorescu D, Weishampel D, Norman D, Şeclăman M, Rusu M, Baltreş A, Teodorescu V (1994) Late Maastrichtian dinosaur eggs from the Haţeg Basin (Romania). In: Carpenter K, Hirsch KF, Horner JR (eds) Dinosaur Eggs and Babies. Cambridge University Press, Cambridge, pp 75–87
Grigorescu D, Garcia G, Csiki Z, Codrea V, Bojar AV (2010) Uppermost Cretaceous megaloolithid eggs from the Haţeg Basin, Romania, associated with hadrosaur hatchlings: Search for explanation. Palaeogeogr Palaeoclimatol Palaeoecol 293:360–374. doi:10.1016/j.palaeo.2010.03.031
Joffe J (1967) The ‘dwarf’ crocodiles of the Purbeck formation, Dorset: a reappraisal. Palaeontology 10:629–639
Karl HV, Gröning E, Brauckmann C, Schwarz D, Knötschke N (2006) The Late Jurassic crocodiles of the Langerberg near Oker, Lower Saxony (Germany), and description of related materials (with remarks on the history of quarrying the “Langenberg Limestone” and “Obernkirchen Sandstone”). Clausthaler Geowiss 5:59–77
Kusuhashi N, Hu YM, Wang YQ, Hirasawa S, Matsuoka H (2009) New triconodontids (Mammalia) from the Lower Cretaceous Shahai and Fuxin formations, northeastern China. Geobios 42:765–781
Lapparent de Broin F, Murelaga X (1999) Turtles from the Upper Cretaceous of Lano (Iberian Peninsula). Est Mus Cien Nat Alava Num esp 14:135–211
Lapparent de Broin F, Murelaga Bereikua X, Codrea V (2004) Presence of Dortokidae (Chelonii, Pleurodira) in the earliest Tertiary of the Jibou Formation, Romania: Paleobiogeographical implications. Acta Pal Rom 4:203–215
Larsson HCE, Sues HD (2007) Cranial osteology and phylogenetic relationships of Hamadasuchus rebouli (Crocodyliformes: Mesoeucrocodylia) from the Cretaceous of Morocco. Zool J Linn Soc 149:533–567
Lauprasert K, Laojumpon C, Saenphala W, Cuny G, Thirakhupt K, Suteethorn V (2010) Atoposaurid crocodyliforms from the Khorat group of Thailand: first record of Theriosuchus from Southeast Asia. Paläontol Z. doi:10.1007/s12542-010-0071-z
Makádi L (2006) Bicuspidon aff. hatzegiensis (Squamata: Scincomorpha: Teiidae) from the Upper Cretaceous Csehbánya Formation (Hungary, Bakony Mts). Acta Geol Hung 49:373–385
Marinho TS, Carvalho IS (2009) An armadillo-like sphagesaurid crocodyliform from the Late Cretaceous of Brazil. J S Am Earth Sci 27:36–41
Martin JE (2009) Le crocodile de l’Allier: Diplocynodon, sa systématique, son écologie et sa place dans l’évolution des alligatoroïdes primitives. Thèse de doctorat, Université Lyon 1, p 429
Martin JE, Benton MJ (2008) Crown clades in vertebrate nomenclature: correcting the definition of Crocodylia. Syst Biol 57:173–181
Martin JE, Buffetaut E (2005) An overview of the Late Cretaceous crocodilian assemblage from Cruzy, southern France. Kaupia 14:33–40
Martin JE, Delfino M (2010) Recent advances on the comprehension of European eusuchians. Palaeogeogr Palaeoclimatol Palaeoecol 293:406–418. doi:10.1016/j.palaeo.2009.10.021
Martin JE, Csiki Z, Grigorescu D, Buffetaut E (2006) Late Cretaceous crocodilian diversity in Hateg Basin, Romania. Hantkeniana 5:31–37
Matheron P (1869a) Note sur les reptiles fossiles des dépôts fluvio-lacustres crétacés du bassin à lignite de Fuveau. Mém Acad Imp Sci Belles Lettres et Arts de Marseille 1–39
Matheron P (1869b) Note sur les reptiles fossiles des dépôts fluvio-lacustres crétacés du bassin à lignite de Fuveau. Bull Soc Géol Fr 26:781–795
Michard JG, de Broin F, Brunet M, Hell J (1990) Le plus ancien crocodilien néosuchien specialisé à caractères “eusuchiens” du continent africain (Crétacé inférieur, Cameroun). C R Acad Sci Paris Ser II 311:365–371
Nobre PH, Carvalho IS (2006) Adamantinasuchus navae: a new Gondwanan Crocodylomorpha (Mesoeucrocodylia) from the Late Cretaceous of Brazil. Gond Res 10:370–378
Nopcsa F (1915) Die Dinosaurier der siebenbürgischen Landesteile Ungarns. Mitt J der Kgl Ung Geol Reichs 1–24
Norell M, Clark JM (1990) A reanalysis of Bernissartia fagesii, with comments on its phylogenetic position and its bearing on the origin and diagnosis of the Eusuchia. Bull Inst R Sci Nat Belg 60:115–128
Nydam RL, Cifelli RL (2002) A new teiid lizard from the Cedar Mountain Formation (Albian–Cenomanian boundary) of Utah. J Vertebr Paleontol 22:276–285
Ortega F, Gasparini Z, Buscalioni AD, Calvo JO (2000) A new species of Araripesuchus (Crocodylomorpha, Mesoeucrocodylia) from the Lower Cretaceous of Patagonia (Argentina). J Vertebr Paleontol 20:57–76
Ősi A (2005) Hungarosaurus tormai, a new ankylosaur (Dinosauria) from the Upper Cretaceous of Hungary. J Vertebr Paleontol 25:370–383
Ősi A, Clark JM, Weishampel DB (2007) First report on a new basal eusuchian crocodyliform with multicusped teeth from the Upper Cretaceous (Santonian) of Hungary. N Jb Geol Paläont Abh 243:169–177
Owen R (1879) Monograph on the fossil reptilia of the Wealden and Purbeck Formations. Supplement no. VI, Crocodilia (Hylaeochampsa). 7pp. and pl. II of Suppl. V. Palaeontogr. Soc. (Monogr.), London (1874). Supplement no. IX. Crocodilia (Goniopholis, Brachydectes, Nannosuchus, Theriosuchus and Nuthetes). p 19, 4 pls. loc. cit. (1879)
Pereda-Suberbiola X (1999) Ankylosaurian dinosaur remains from the Upper Cretaceous of Laño (Iberian Peninsula). Est Mus Cienc Nat Álava 14(Número especial 1):273–288
Pereda-Suberbiola X (2009) Biogeographical affinities of Late Cretaceous continental tetrapods of Europe: a review. Bull Soc Géol Fr 180:67–71
Pereda-Suberbiola X, Galton PM (2001) Reappraisal of the Nodosaurid Ankylosaur Struthiosaurus austriacus Bunzel from the Upper Cretaceous Gosau Beds of Austria. In: Carpenter K (ed) The Armored Dinosaurs. The Indiana University Press, Bloomington, pp 173–210
Pol D (2003) New remains of Sphagesaurus huenei (Crocodylomorpha: Mesoeucrocodylia) from the Late Cretaceous of Brazil. J Vertebr Paleontol 23:817–831
Pol D, Norell MA (2004) A new crocodyliform from Zos Canyon, Mongolia. Am Mus Novit 3445:1–36
Pol D, Apesteguía S (2005) New Araripesuchus remains from the Early Late Cretaceous (Cenomanian- Turonian) of Patagonia. Am Mus Novit 3490:1–38
Pol D, Gasparini Z (2009) Skull anatomy of Dakosaurus andiniensis (Thalattosuchia: Crocodylomorpha) and the phylogenetic position of Thalattosuchia. J Syst Palaeontol 7:163–197
Pol D, Turner AH, Norell MA (2009) Morphology of the Late Cretaceous crocodylomorph Shamosuchus djadochtaensis and a discussion of neosuchian phylogeny as related to the origin of Eusuchia. Bull Am Mus Nat Hist 324:1–103
Pouech J, Mazin JM, Billon-Bruyat JP (2006) Microvertebrate biodiversity from Cherves-de-Cognac (Lower Cretaceous, Berriasian: Charente, France). 9th International Symposium on Mesozoic Terrestrial Ecosystems and Biota, Abstracts and Proceedings Volume: 96–100
Prasad GVR, Lapparent de Broin F (2002) Late Cretaceous crocodile remains from Naskal (India): comparisons and biogeographic affinities. Ann Paleontol 88:19–71
Rabi M (2008) New discovery of the Late Cretaceous ziphodont crocodyliform, Doratodon from the Santonian Csebánya Formation of Hungary. 6th Meeting of the European Association of Vertebrate Palaeontologists, Spišská Nová Ves, Slovak Republic. Volume of Abstracts, 82
Rage JC (2002) The continental Late Cretaceous of Europe: toward a better understanding. C R Palevol 1:257–258
Rogers JVI (2003) Pachycheilosuchus trinquei, a new procoelous crocodyliform from the Lower Cretaceous (Albian) Glen Rose Formation of Texas. J Vertebr Paleontol 23:128–145
Schwarz D, Salisbury SW (2005) A new species of Theriosuchus (Atoposauridae, Crocodylomorpha) from the Late Jurassic (Kimmeridgian) of Guimarota, Portugal. Geobios 38:779–802
Schwarz-Wings D, Rees J, Lindgren J (2009) Lower Cretaceous Mesoeucrocodylians from Scandinavia. Cretac Res 30:1345–1355
Seeley HG (1881) The reptile fauna of the Gosau Formation preserved in the Geological Museum of the University of Vienna. Q J Geol Soc Lond 37:620–704
Sereno PC, Larsson HCE, Sidor CA, Gado B (2001) The giant crocodyliform Sarcosuchus from the Cretaceous of Africa. Science 294:1516–1519
Sereno PC, Sidor CA, Larsson HCE, Gado B (2003) A new notosuchian from the Early Cretaceous of Niger. J Vertebr Paleontol 23:477–482
Sereno PC, Larsson HCE (2009) Cretaceous crocodyliforms from the Sahara. Zookeys 28:1–143
Turner AH (2004) Crocodyliform biogeography during the Cretaceous: Evidence of Gondwana vicariance from biogeographical analysis. Proc R Soc Lond B 271:2003–2009
Turner AH (2006) Osteology and phylogeny of a new species of Araripesuchus (Crocodyliformes: Mesoeucrocodylia) from the Late Cretaceous of Madagascar. Hist Biol 18:255–369
Vasse D (1995) Ischyrochampsa meridionalis nov. gen. nov. sp., un crocodilien d’affinités gondwanienne dans le Crétacé supérieur du Sud de la France. N J Geol Pal Mh 8:501–512
Weishampel DB, Jianu CM, Csiki Z, Norman DB (2003) Osteology and phylogeny of Zalmoxes (n. g.), an unusual euornithopod dinosaur from the latest Cretaceous of Romania. J Syst Palaeontol 1:65–123
Weishampel DB, Csiki Z, Benton MJ, Grigorescu D, Codrea V (2010) Palaeobiogeographic relationships of the Haţeg biota – between isolation and innovation. Palaeogeogr Palaeoclimatol Palaeoecol 293:419–437. doi:10.1016/j.palaeo.2010.03.024
Whetstone KN, Whybrow PJ (1983) A “cursorial” crocodilian from the Triassic of Lesotho (Basutoland), Southern Africa. Occ Pap Mus Nat Hist Kans 106:1–37
Wu XC, Sues HD (1996) Anatomy and phylogenetic relationships of Chimaerasuchus paradoxus, an unusual crocodyliform reptile from the Lower Cretaceous of Hubei, China. J Vertebr Paleontol 16:688–702
Wu XC, Sues HD, Sun A (1995) A plant-eating crocodyliform reptile from the Cretaceous of China. Nature 376:678–680
Wu XC, Sues HD, Brinkman DB (1996) An atoposaurid neosuchian (Archosauria: Crocodyliformes) from the Lower Cretaceous of Inner Mongolia (People’s Republic of China). Can J Earth Sci 33:599–605
Young MT, de Andrade MB (2009) What is Geosaurus? Redescription of Geosaurus giganteus (Thalattosuchia: Metriorhynchidae) from the Upper Jurassic of Bayern, Germany. Zool J Linn Soc 157:551–585
Zhang WT, Chen PJ, Palmer AR (2003) Biostratigraphy of China. Elsevier Science, Beijing, p 599
Acknowledgments
We are grateful to Silvia Burnaz (Museum of Deva) for the loan of some as yet undescribed specimens. We appreciate the sharing of unpublished information and the useful discussions with Marco Brandalise de Andrade (University of Bristol), Diego Pol (Museo Paleontológico Egidio Feruglio), Daniella Schwarz-Wings (Museum für Naturkunde, Berlin), Francisco Ortega (Universidad Nacional de Educación a Distancia), Christopher Brochu (University of Iowa) and Eric Buffetaut (Centre National de la Recherche Scientifique). We wish to say thanks to Sandra Chapman (Natural History Museum of London) and Daniella Schwarz-Wings for the access to the type material of T. pusillus and T. ibericus, respectively. The helpful suggestions of four anonymous reviewers significantly improved the quality of an earlier version of the manuscript. JEM was funded by the Laboratoire SPE UMR 6134 (Université de Corse). ZCs was supported by grants CNCSIS-UEFISCSU 1930 PNII-IDEI/2009 and RO0023/2009 (Romanian Academy of Sciences).
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Martin, J.E., Rabi, M. & Csiki, Z. Survival of Theriosuchus (Mesoeucrocodylia: Atoposauridae) in a Late Cretaceous archipelago: a new species from the Maastrichtian of Romania. Naturwissenschaften 97, 845–854 (2010). https://doi.org/10.1007/s00114-010-0702-y
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DOI: https://doi.org/10.1007/s00114-010-0702-y