A Cladistic Analysis of Ten Lizard Families (Reptilia: Squamata) Based on Cranial Musculature

Virginia Abdala, Silvia Moro


The cranial musculature of species of ten families of lizards (Gekkonidae, Teiidae, Gymnophthalmidae, Anguidae, Scincidae, Lacertidae, Tropiduridae, Liolaemidae, Leiosauridae, and Polychrotidae) was analyzed. Using 93 myological cranial characters, a cladistic analysis was performed. To root the trees, data of Sphenodon were added to the matrix. The cladistic analysis yielded 129 equally parsimonious trees with a fit of 426.0 (39%) and 467 steps. In the consensus tree we observed only five nodes congruent with traditional phylogenetic hypothesis of the groups analyzed: teiids + gymnophthalmids (node 128); geckos (node 136); liolaemids (node 116); genus Liolaemus (node 115); and genus Phymaturus (node 92). Thus in these groups, phylogeny seems to be better predictor of muscle morphology than ecology or diet.

They are many problematic groups whose monophyly requires further analysis (tropidurids, polychrotids, and scincids). In no case we were able to recognize differences that could be attributed to functional features.


cranial musculature; lizards; phylogeny; cladistic analysis

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Abdala V. and Moro S. (1996), «Cranial musculature of South American Gekkonidae», J. Morphol., 229, 59 – 79.

Adams L. A. (1919), «A memoir on the phylogeny of the jaw muscles in recent and fossil vertebrates», Ann. N. Y. Acad. Sci., 28, 51 – 166.

Avery D. F. and Tanner W. W. (1964), «The osteology and myology of the head and thorax regions of the obesus group of the genus Sauromalus Dumeril (Iguanidae)», Brigham Young Univ. Sci. Bull. Biol. Ser., 5, 1 – 30.

Brock G. T. (1939), «Cranial muscles of the gecko», Proc. Zool. Soc. London, 108(4), 735 – 761.

Caldwell M. W. (1999), «Squamate phylogeny and the relationships of snakes and Mosasauroids», Zool. J. Linn. Soc., 125, 115 – 147.

Camp C. L. (1923), «Classification of the Lizards», Bull. Am. Mus. Nat. Hist., 48, 289 – 481.

Costelli J. (1973), Iguanid Trigeminal Musculature and Its Role in the Phylogeny of the Iguanidae, UMI Dissertation Services, A Bell and Howell Co., Ann Arbor, MI, USA.

Cundall D. (1987), «Functional morphology, in: Snakes: Ecology and Evolutionary Biology, R. A. Seigel, J. T. Collins. and S. S. Novak (eds.), MacMillan, New York, pp. 106 – 140.

Dalrympe G. H. (1979), «On the jaw mechanism of the snail-crushing lizards, Dracaena Daudin 1802 (Reptilia, Lacertilia, Teiidae)», J. Herpetol., 13, 303 – 311.

Edmund A. G. (1969), «Dentition», in: Biology of the Reptilia. Vol. 1, C. Gans and T. S. Parsons (eds.), Acad. Press, London, pp. 117 – 200.

Estes R., de Queiroz K., and Gauthier J. (1988), «Phylogenetic relationships within Squamata», in: Phylogenetic Relationships of Lizard Families: Essays Commemorating Charles L. Camp, R. Estes and G. Pregill (eds.), Stanford Univ. Press, pp. 119 – 281.

Etheridge R. (1995), «Redescription of Ctenoblepharys adspersa Tschudi, 1845, and the taxonomy of Liolaeminae (Reptilia: Squamata: Tropiduridae)», Am. Mus. Novitates, 3142, 1 – 34.

Etheridge R. and de Queiroz K. (1988), «A phylogeny of Iguanidae», in: Phylogenetic Relationships of Lizard Families: Essays Commemorating Charles L. Camp, R. Estes and G. Pregill (eds.), Stanford Univ. Press, pp. 283 – 367.

Fanghella C., Avery D. F., and Tanner W. W. (1975), «Urosaurus and its phylogenetic relationship to Uta as determined by osteology and myology (Reptilia: Iguanidae)», Great Basin Nat., 35, 245 – 268.

Fisher D. L. and Tanner W. W. (1970), «Osteological and myological comparisons of the head and thorax regions of Cnemidophorus tigris septentrionalis Burger and Ameiva undulata parva Barbour and Noble (Family Teiidae)», Brigham Young Univ. Sci. Bull. Biol. Ser., 11(1), 1 – 39.

Frost D. R. (1992), «Phylogenetic analysis and taxonomy of the Tropidurus group of lizards (Iguania: Tropiduridae)», Am. Mus. Novitates, 3033, 1 – 68.

Frost D., Etheridge R., Janies D., and Titus T. (2001), «Total evidence, sequence alignment, evolution of Polychrotid lizards, and a reclassification of the Iguania (Squamata: Iguania)», Am. Mus. Novitates, 3343, 1 – 38.

Gans C. and De Vree F. (1987), «Functional bases of fiber length and angulation in muscle», J. Morphol., 192, 63 – 85.

Gans C., De Vree F., and Carrier D. (1985), «Usage pattern of the complex masticatory muscles in the shingleback lizard, Trachydosaurus rugosus: a model for muscle placement», Am. J. Anat., 173, 219 – 240.

Gauthier J., Kluge A. G., and Rowe T. (1988), «Amniote phylogeny and the importance of fossils», Cladistics, 4, 105 – 209.

Gnanamuthu C. P. (1937), «Comparative study of the hyoid and tongue of some typical genera of reptiles», Proc. Zool. Soc. London, 107, 1 – 63.

Goloboff P. A. (1993a), PIWE: Parsimony and Implied Weights. MS-DOS Program and Documentation Distributed by the Author.

Goloboff P. A. (1993b), «Stimating character weights during tree search», Cladistics, 9, 83 – 91.

Goloboff P. A. (1995), «A revision of the South American spiders of the family Nemesiidae (Araneae, Mygalomorphae). Part I: species from Peru, Chile, Argentina, and Uruguay», Bull. Am. Mus. Nat. Hist., 224, 1 – 189.

Goloboff P. A. and Farris J. S. (2001), «Methods for quick consensus estimation», Cladistics, 17, 26 – 34.

Gorniak G. C., Rosemberg H. I., and Gans C. (1982), «Mastication in the tuatara, Sphenodon punctatus (Reptilia: Rhynchocephalia), structure and activity of the motor system», J. Morphol., 171, 321 – 353.

Haas G. (1960), «On the trigeminal muscles of the lizards Xenosaurus grandis and Shinisaurus crocodilurus», Am. Mus. Novitates, 2017, 1 – 54.

Haas G. (1973), «Muscles of the jaw and associated structures in the Rhynchocephalia and Squamata», in: Biology of the Reptilia, C. Gans and T. S. Parsons (eds.), Acad. Press, USA, pp. 285 490.

Halloy M., Etheridge R., and Burghardt G. M. (1998), «To bury in the sand: phylogenetic relationships among lizards species of the boulengeri group, Liolaemus (Reptilia: Squamata: Tropiduridae), based on behavioral characters», Herpetol. Monogr., 12, 1 – 37.

Haines R. W. (1935), «A consideration of the constancy of muscular nerve supply», J. Anat. London, 70, 35 – 45.

Haines R. W. (1939), «A revision of the extensor muscles of the forearm in tetrapods», J. Anat. Cambridge, 73, 211 – 233.

Harvey M. and Gutberlet R. Jr. (2000), «A phylogenetic analysis of the tropidurine lizards (Squamata: Tropiduridae), including new characters of squamation and epidermal microstructure», Zool. J. Linn. Soc., 128, 189 – 233.

Howell A. B. (1938), «The phylogenetic arrangement of the muscular system», Anat. Rec., 66(3), 295 – 316.

Iordansky N. N. (1964), «The jaw muscles of the crocodiles and some relating characteres of the crocodilian skull», Anat. Anz., 115(3), 256 – 280.

Iordansky N. N. (1970), «Structure and biomechanical analysis of functions of the jaws muscles in the lizards», Anat. Anz., 127(4), 383 – 413.

Iordansky N. N. (1994), «Tendons of jaw muscles in Amphibia and Reptilia: homology and evolution», Russ. J. Herpetol., 1(1), 13 – 20.

Kesteven H. C. (1944), «The evolution of the skull. Part III, The Sauria», Mem. Aust. Mus., 8(3), 238 – 268.

Kluge A. G. (1989), «Progress in Squamate classification», Herpetologica, 45(3), 368 – 379.

Lakjer T. (1926), Die Trigeminus-versorgte Kaumuskulatur der Sauropsiden, Reitzel, Kopenhagen.

Lobo F. (2001), «A phylogenetic analysis of lizards of the Liolaemus chiliensis group (Iguania: Tropiduridae)», Herpetol. J., 11, 137 – 150.

Lubosch W. (1938), «Muskeln des Kopfes: Viscerale Muskulatur. C. Amphibien und Sauropsiden», in: Handbuch der vergleichenden Anatomie der Wirbeltiere, L. Bolk, E. Göppert, E. Kallius, and W. Lubosch (eds.), Urban und Schwarzenberg, Berlin – Wien, pp. 1025 – 1064.

MacLean W. P. (1974), «Feeding and locomotor mechanisms of teiid lizards: functional morphology and evolution», Pap. Avulsos Zool., 27(15), 179 – 213.

Mivart St. G. (1867), «Notes on the myology of Iguana tuberculata», Proc. Zool. Soc. London, 35, 766 – 797.

Mivart St. G. (1870), «On the myology of the Chamaeleo parsoni», Proc. Zool. Soc. London, 57, 850 – 890.

Moro S. and Abdala V. (1998), «Cranial myology of some species of Liolaemus and Phymaturus (Squamata: Tropiduridae: Liolaeminae)», Amphibia–Reptilia, 19, 171 – 192.

Moro S. and Abdala V. (2000), «Cladistic analysis of Teiidae based on cranial myological characters», Russ. J. Herpetol., 7(2), 87 – 102.

Nixon C. K. (1999), «The parsimony ratchet, a new method for rapid parsimony analysis», Cladistics, 15, 407 – 414.

Nixon C. K. and Carpenter J. M. (1993), «On outgroups», Cladistics, 9, 413 – 426.

Oelrich T. M. (1956), «The anatomy of the head of Ctenosaura pectinata (Iguanidae)», Mus. Zool. Univ. Michigan, 94, 1 – 121.

Ostrom J. (1962), «On the constrictor dorsalis muscles of Sphenodon», Copeia, 1962(4), 732 – 735.

Poglayen-Neuwall I. (1954), «Die Kiefermuskulatur der Eidechsen und ihre Innervation», Z. Wiss. Zool., 158, 79 – 132.

Presch W. (1974), «Evolutionary relationships and biogeography of the macroteiid lizards (Family Teiidae, subfamily Teiinae)», Bull. S. California Acad. Sci., 73, 23 – 32.

Raikow R. J. (1977), «The origin and evolution of the Hawaiian honeycreepers (Drepanididae)», Living Bird, 15, 95 – 117.

Raikow R. J., Bledsoe A. H., Myers B. A., and Welsch C. J. (1990), «Individual variation in avian muscles and its significance for the reconstruction of phylogeny», Syst. Zool., 39, 362 – 370.

Rieppel O. (1980), «The trigeminal jaw adductor of Tupinambis, with comments on the phylogenetic relationships of the Teiidae (Reptilia: Lacertilia)», Zool. J. Linn. Soc., 69, 1 – 29.

Rieppel O. (1984), «The structure of the skull and the jaw adductor musculature in the Gekkota, with comments on the phylogenetic relationships of the Xantusiidae (Reptilia: Lacertilia)», Zool. J. Linn. Soc., 82, 291 – 318.

Romer A. S. (1942), «The development of tetrapod limb musculature — the thigh of Lacerta», J. Morphol., 71, 251 – 298.

Russell A. (1988), «Limb muscles in relation to lizard systematics: a reappraisal», in: Phylogenetic Relationships of Lizard Families: Essays Commemorating Charles L. Camp, R. Estes and G. Pregill (eds.), Stanford Univ. Press, pp. 493 – 568.

Sanders A. (1870), «Notes on the myology of Platydactylus japonicus», Proc. Zool. Soc. London, 1870, 413 – 427.

Sanders A. (1872), «Notes on the myology of Liolepis belli», Proc. Zool. Soc. London, 1872, 154 – 183.

Sanders A. (1874), «Notes on the myology of Phrynosoma coronatum», Proc. Zool. Soc. London, 1874, 71 – 89.

Schulte J. A. III, Macey J. R., Larson A. and Papenfuss T. J. (1999), «Molecular tests of phylogenetic taxonomies: a general procedure and example using four subfamilies of the lizard family Iguanidae», Mol. Phylogen. Evol., 10(3), 367 – 376.

Schwenk K. (1988), «Comparative myology of the lepidosaur tongue and its relevance to squamate phylogeny», in: Phylogenetic Relationships of Lizard Families: Essays Commemorating Charles L. Camp, R. Estes and G. Pregill (eds.), Stanford Univ. Press, pp. 569 – 598.

Versluys J. (1912), «Das Streptostylie-Problem und die Bewegungen im Schädel bei Sauropsiden», Zool. Jahrb., 15(2), 545 – 716.

Wainwright P. C. and Bennet A. F. (1992), «The mechanism of tongue projection in chameleons. I. Electromyographic tests of functional hypotheses», J. Exp. Biol., 168, 1 – 21.

Wainwright P. C., Kraklau D., and Bennet A. F. (1991), «Kinematics of tongue projection in Chamaeleo oustaleti», J. Exp. Biol., 159, 109 – 133.

Wassersug R. (1976), «A procedure for differential staining of cartilage and bone in whole formalin fixed vertebrates», Stain. Tech., 51, 131 – 134.

DOI: https://doi.org/10.30906/1026-2296-2003-10-1-53-78


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