MESOLEPTOS AND THE ORIGIN OF SNAKES 



141 



Camp, C.L. 1923. Classification of the lizards. Bulletin of the American Museum of 

 Natural History 48: 289-481. 



Carroll, R.L. 1985. Evolutionary constraints in aquatic diapsid reptiles. Special 

 Papers in Palaeontology 33:145-155. 



Carroll, R.L. & Debraga, M. 1992. Aigialosaurs: mid-Cretaceous Varanoid lizards. 

 Journal of Vertebrate Paleontology 12: 66-86. 



Cope, E.D. 1869. On the reptilian orders Pythonomorpha and Streptosauria. Proceed- 

 ings of the Hasten Society of Natural History 12: 250-266. 



Cornalia, E. & Chiozza, L. 1852. Cenni geologic] sull' Istria. Giomale dell' 1. R. 

 Institute Lombardo 3: 1-35. 



Dollo, L. 1903. Les ancetres des Mosasauriens. Bulletin scicniifique de France el 

 Belgique 38: 137-130. 



1904. L'origine des Mosasauriens. Bulletin du Societe belgique de Geologie, 



Hydrologie et Palaeontologie, Bruxelles 18: 217-222. 



Estes, R., De Queiroz, K. & Gauthier, J. 1988. Phylogenetic relationships within 



Squamata. In R. Estes and G. Pregill (eds). Phylogenetic relationships of the lizard 



families. Stanford University Press, Stanford, California. Pp. 119-281. 

 Gardner, J.D. & Cifelli, R.L. 1999. A primitive snake from the Cretaceous of Utah. 



Special Papers in Palaeontology 60: 87- 1 00. 

 Gasc, J.-P. 1974. L' interpretation fonctionelle de lappareil musculo-squelettique de 



1' axe vertebral chez les serpents (ReptWtd). Memoires du Museum nalionale d'Histoire 



natu relic. Series A 83: 1-182. 

 Goldman, N, Anderson, J. P. & Rodrigo, A.G. 2000. Likelihood-based tests oi 



topologies in phylogenetics. Systematic Biology 49: 652-670. 

 Gorjanovic-Kramberger, C. 1892. Aigialosaurus, eine neue Eidechse aus die 



Krcideschiefern der Insel Lesina mil Rucksicht auf die bereils beschriebenen 



Lacertiden von Comen und Lesina. Glasnik huvatskoga naravoslovnoga drustva 



(Societas historico-naturalis croatica) u Zagrebu 7: 74-106. 

 Greene, H.W. 1997. Snakes - The evolution of mystery in nature. University of 



California Press. Berkeley. 

 Haas, G. 1979. On a new snakelike reptile from the lower Cenomaniai) of Ein Jabrud, 



near Jerusalem. Bulletin du Museum national d'Histoire naturelle. Paris (4). Section 



CI: 51-64. 



1980. Remarks on a new ophiomorph reptile from the lower Cenomanian of Ein 



Jabrud. Israel. In L. L. Jacobs fed.). Aspects of vertebrate history: Essays in Honour 

 of E. H. Colbert. Museum of Northern Arizona Press: Flagstaff AZ. Pp. 177-202. 



Hoffstetter, R. 1955. Squamates de type moderne. Trade de Palaontologie 5: 606-662. 



1960. Un serpent terrestre dans le Cretace inferieurdu Sahara. Bulletin du Sot ieti 



giologique de France 7: 897-902. 



& Gayrard, Y. 1965. Observations sur I'osteologie et la classification des 



Acrochordidae (Serpentes). Bulletin du Museum national d'Histoire naturelle. Paris 

 (series 2) 36: 677-696. 



Janensch, W. 1906. Uber Archaeophis proavus Mass.. eine Schlange aus dem Eoc.m 



des Monte Bolca. Beitriige zur Palaontologie und Geologic Osterreich-Ungams 



iiml des Orients 19:1-33. 

 Kluge, A.G. 1987. Cladistic relationships in the Gekkonoidea (Squamala. Saunal. 



Miscellaneous Publications of the Museum of Zoology, University of Michigan 173: 



1-54. 

 Kornhuber, A. 1 873. Ubereinen neuen fossilen Saurier aus Lesina. Abhandlungen der 



k. k. geologischen Reichsanstalt 5: 75-90. 2 plates. 

 Kornhuber, A. 1901. Opetiosaurus bticchichi. Eine neue fossile Eidechse aus der 



unteren Kreide von Lesina in Dalmatien. Abhandlungen der k. k. geologischen 



Reichsanstalt 17: 1-24. 

 Lee, M.S.Y. 1998 Convergent evolution and character correlation in burrowing rep- 

 tiles: towards a resolution of squamate phylogeny. Zoological Journal oj the 1 nine an 



Society 65: 369-453. 

 1999. Aquatic reptiles. In R. Singer and M. K. Diamond (eds). Encyclopedia of 



Paleontology. Fitzroy Dearbon: London. Pp. 96-101. 

 2002. Squamate phylogeny. taxon sampling, and data congruence. Hcrpelologicul 



Monographs in review. 



& Caldwell, M.W. 1998. Anatomy and relationships of Pachyrhachis 



problematicus. a primitive snake with hindlimbs. Philosophical Transactions oi the 

 Royal Society of London B 353: 1521-1 552. 



& 2000. Adriosaurus and the affinities of mosasaurs. dolichosaurs and 



snakes. Journal of Paleontology 74: 915-937. 



, & Scanlon, J.D. 1999. A second primitive marine snake: Pachyophis 



woodward! from the Cretaceous of Bosnia-Herzegovina. Journal of Zoology, Lon- 

 don 248: 509-520. 



& Scanlon, J.D. 2002. Snake phylogeny based on osteology, soft anatomy and 



ecology. Biological Reviews of the Cambridge Philosophical Society 77. in press. 



McDowell, S.B. 1987. Systematics. In R.A. Seigel. J.T.C. Collins and S.S. Novak 

 (eds.). Snakes: Ecology and Evolutionary Biology. MacMillan. New York. Pp. 1-50. 



& Bogert, CM. 1954. The systematic position of Lanthanotus and the affinities 



of the Anguimorphan lizards. Bulletin of the American Museum of Natural History 

 105: 1-154. 



Mosauer, W. 1 935. The myology of the trunk region of snakes and its significance for 

 ophidian taxonomy and phylogeny. University of California Publications in Biologi- 

 cal Sciences 1 : 8 1 - 1 20. 



Nopcsa, F. 1903. Uber die Varanusartigen Lacertenlstriens. Beitriige zur Palaontologie 

 und Geologie Osterreich-Ungams und des Orients 15: 3 1^42. 



1908. Zur Kenntnis der fossilen Eidechsen. Beitriige zur Palaontologie und 



Geologie Osterreich-Ungams unci des Orients 21: 33-62. 



1923. Eidolosaurus und Pachyophis. Zwei neue Neocom-Reptilien. 



Palaeontographica 65: 99-154. 

 Owen, R. 1842. Description of the remains of a bird, tortoise and lizard from the Chalk 



of Kent. Transactions of the Geological Society of London. Series 2. 6: 411 — 4 1 3- 

 1850a. Description of the fossil reptiles of the Chalk Formation. In F. Dixon (ed.). 



The Geology and Fossils of the Tertian and Cretaceous Formations of Sussex. 



Longman. Brown. Green and Longman, London. Pp. 378^t00. 403-404. 

 1850b. Monograph on the Reptilia of the London Clay. Part III. Order Ophidia. 



British Museum: London 



1851. .4 monograph on the fossil Reptilia of the Cretaceous formations. Part 1. 



The Palaeontological Society: London. 



1877. On the rank and affinities of the reptilian class of the Mosasauridae. 



Gervais. Quarterly Journal of the Geological Society oj London 33: 682-715. 



Radovanovic, M. 1935. Anatomische Studien am Schlangenkopf. Jenaische Zeitschrift 



fiir Naturwissenschaft 69: 321-422. 

 Rage, J.C. 1984. Hanilbttch der Paldoherpetologie. Ted II. Serpentes. Gustav Fischer. 



Stuttgart. 

 Rieppel, O. 1988. A review of the origin of snakes. Evolutionary Biology 22: 37-130. 

 & Zaher, H. 2000. The intramandibular joint in squamates. and the phylogenetic 



relationships of the fossil snake Pachyrhachis problematicus Haas. Fieldiana (Geol- 

 ogy) 43: 1-69. 

 Romer, A.S. 1956. Osteology of the reptiles. University of Chicago Press, Chicago. 

 Scanlon, J.D. 1993. Madtsoiid snakes from the Eocene Tingamarra Fauna of eastern 



Queensland. Kaupia: Oarinstddtcr Beitriige zur Naturgcschichtc 3: 3-8. 

 1996. Studies in the palaeontology and systematics of Australian snakes. Unpub- 

 lished Ph.D. thesis. University of New South Wales. Sydney. 

 & Lee, M.S.Y. 2000. The Pleistocene serpenl Wonambi and the early evolution of 



snakes. Nature 403: 416420. 

 , . Caldwell, M.W. and Shine, R. 1999. Ecology and laphonomy of the 



primitive snake Pachyrhachis. Historical Biology 13: 127-150. 

 Seeley, H.G. 1 88 1 . On remains of a small li/ard from the Neocomian rocks of Comen. 



near Trieste, preserved in the Geological Musem of the University of Vienna. 



Quarterly Journal of the Geological Society of London 37: 52-56. 

 Sorenson, M. 2000. TreeRot Version 2b. Computer program and documentation. 



Distributed b\ the author. Boston University. 

 Swofford, D. L. 1999. PAUP*: Phylogenetic Analysis Using Parsimony (*and other 



methods). Version 4. Sinauer Associates, Sunderland MA. 

 Tchernov, E., Rieppel, ()., Zaher, H., Polcyn, M. & Jacobs, L.L. 2000. A fossil snake 



with limbs. Science 287: 2010-2012 

 Underwood, G. 1967. A Contribution to the Classification of Snakes. British Museum 



(Natural History) Publication No. 653. Trustees of the British Museum (Natural 



History i. I .ondon. 179pp. 

 von Meyer, H.V. I860, Actaeosaurut tommasinii aus dem schwarzen Kreide-Schiefer 



von Comen am Karste. Palaeontographica 7: 223-231. 

 Walls, G.L. 1940. Ophthalmological implications lor the early history of snakes. 



Copeia 1940: 1-8. 

 Zaher, H. & Rieppel, O. 1999. The phylogenetic relationships of Pachyrhachis 



problematicus, and the evolution of limblessness in snakes (Lepidosauria, Squamata). 



Comptes vendues de I' Academic des Sciences de Paris. Sciences de la terre el des 



planetes 329: 831-837. 



Appendix 1 



Additions to the osteological data matrix of Lee (2002) used in this 

 cladistic analysis. Mesoleptos was added to the taxon list and could 

 be coded for the following characters. 



Axial Skeleton 



171. Centra. Not constricted anterior to condyle, i.e. condyle not 

 wider than posterior end of centrum (0): slightly constricted 

 anterior to condyle, i.e. condyle slightly wider than posterior 

 end of centrum ( 1 ); greatly constricted anterior to condyle, i.e. 

 condyle much wider than posterior end centrum (2). Mesoleptos 

 0. 



172. Vertebral articular surfaces. Vertical, condyles (if present) 

 facing posteriorly, much of the articular surface is visible in 

 ventral view (0); slightly anterodorsal, condyles facing slightly 

 dorsally, only the ventral edge of the articular surface is visible 



