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Multequina ISSN: 0327-9375 [email protected] Instituto Argentino de Investigaciones de las Zonas Áridas

Cei, José M. Comparative studies of supraocular lepidosis in (reptilia) and its relationships with an evolutionary Multequina, núm. 16, 2007, pp. 1-52 Instituto Argentino de Investigaciones de las Zonas Áridas Mendoza, Argentina

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COMPARATIVE STUDIES OF SUPRAOCULAR LEPIDOSIS IN SQUAMATA (REPTILIA) AND ITS RELATIONSHIPS WITH AN EVOLUTIONARY TAXONOMY ESTUDIOS COMPARATIVOS DE LA LEPIDOSIS SUPRA-OCULAR EN SQUAMATA (REPTILIA) Y SU RELACIÓN CON LA TAXONOMÍA EVOLUCIONARIA

JOSÉ M. CEI †

las subfamilias Leiosaurinae y RESUMEN Enyaliinae. Siempre en Iguania Observaciones morfológicas se evidencian ejemplos previas sobre un gran número de como los inconfundibles patrones de especies permiten establecer una escamas supraoculares de correspondencia entre la , Leucocephalidae, peculiaridad de los patrones , . A nivel sistemáticos de las escamas específico la interdependencia en supraoculares de Squamata y la de los géneros Iguana, posición evolutiva de cada taxón Cercosaura, Brachylophus, considerado en los cladogramas Conolophus, puede llevar a propuestos por Estes et al. (1988). postular pretéritos acontecimientos Aparte del significado biológico paleogeográficos. También amerita general de estos hallazgos, incluso énfasis la llamativa separación, para discutidas orientaciones según este criterio morfológico, en- taxonómicas, la lepidosis tre Iguania y , la supraocular llega a refrendar una uniforme lepidosis de centenares de decisión sistemática con su , o la excepcional fisonomía evidencia. Así, en Iguania, la familia de Autarchoglossa, en sus ramas tan , propuesta por Frost individualizadas de et al. (2001), aparece sostenida (; Teiioidea; hasta en el detalle de su división en Scincoidea) o .

MULTEQUINA 16: 1-52, 2007 1 Palabras clave: diagnosis, INTRODUCTION escamas supraoculares, pa- The selection of significant somatic trón específico lepidosis, morphological characters (shape, Pleurodonta, Squamata, size, structure) will play an essential taxonomía role in providing a systematic herpetological arrangement, at least SUMMARY to level: furthermore it can Former observations on apply also to more elevated ranks characteristic patterns of supraocular of the herpetological classification. scutellation in different specific or We are pointing out here some supraspecific taxa of Squamata are interesting aspects of the so-called extended and analyzed in this general lepidosis, or cutaneous scale critical review. A relationship covering of Reptilia, mainly in between a defined morphological Lacertilian . More exactly, supraocular lepidosis in a particular the differentiated scales bilaterally taxon and its evolutionary position located in the supraocular region of in a classificatory system, such as the the head, bounded inside by the cladograms obtained by Estes et al. orbital semicircles, and by the (1988), was tested and confirmed. ciliary eye border outside (Figure Supraocular scales of Squamata are 1). Given the noticeable position of not morphological elements at these supraocular scales in the head random, but can be assumed as scutellation of Squamata, their morphological scale characters, presence has generally been useful in systematic comparative reported in many diagnostic examinations and able to strengthen descriptions of species. some discussed new systematic Nevertheless, no true emphasis was assessments, such as the family status ever placed on their peculiar of Leiosauridae and its subfamilies taxonomic significance: either as in Pleurodont Iguanians, proposed an important generic characteristic by Frost et al. (2001), as well as in structure of the dorsal head several other cases. lepidosis, or as distinctive supraocular patterns in separated systematic groups, from different Key words: Squamata, genera to families and higher supraocular scutellation, categories. A preliminary paper was specific lepidosis pattern, provided on this matter (Cei, 2003), diagnosis, Pleurodonta, dealing with supraocular scales in taxonomy species and genera of Iguania, as

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3 Figure 1. Sketched fundamental morphology of supraocular scutellation in Squamata, Pleurodonta: 1. supraorbital semicircles, 2. supraocular scales, 3. intermediate supraorbital scales and 4. superimposed superciliaries Figura 1. Esquema fundamental de la morfología de la escutelación supraocular en Squamata, Pleurodonta: 1. semicírculos supraorbitales, 2. escalas supraoculares, 3. escalas supraorbitales interme- dias y 4. superciliares superimpuestos well as in some far-away taxonomic , besides a preliminary categories, such as Scleroglossa paper (Cei, 2003), deal with Teiid lizards. Thus, considering the circumorbitals in one row or in two remarkable number of past and rows between supraoculars and recent species descriptions, a median head shields, as emphasized detailed review and discussion of in Frost‘s (1992) taxonomic such a noticeable but still revision of groups of disregarded anatomical peculiarity lizards that points out could be a very interesting job, discrimination among the genera postulating a distribution pattern , , not at random but in agreement Plesiomicrolophus and Tropidurus. with the phylogenetic subdivisions Likely in Etheridge‘s (1970) of well-known modern comparative research the classificatory essays. systematic differences in supraciliaries were suitably applied Some isolated reports about to discriminate and specific supraocular scutellation in Uranoscodon from other members

MULTEQUINA 16: 1-52, 2007 3 of the Tropidurus groups. However, Museo de Zoologia, Universidade apart from these interesting but de São Paulo, Brasil; MCZ: limited citations, a whole zoological Museum of Comparative , discussion and critical valuation of Harvard University, Cambridge, lacertilian supraocular scutellation USA; NMNH: National Museum is yet wanting to our knowledge. of Natural History, Smithsonian The present work will attempt to Institution, Washington, USA; reach better and conclusive FMNH: Field Museum of Natural information on such a suggestive History, Chicago, USA; MNHN and little analyzed subject. Paris: Museum Nationale Histoire Naturelle, Paris; UNNE: MATERIAL AND METHOD Universidad Nacional del Nordeste, Corrientes, Argentina. Given the aim of this work, its outcome is a detailed representation Given the Figures including the of the postulated relationships total samples drawn that support our between the supraocular present report and discussion, a scutellation patterns and the separated list of the examined taxonomic categories of Squamata. specimens would be prolix and Our comparative graphic unnecessary. Moreover the general documentation, exposed in Figures, localities of samples were was mainly obtained from careful opportunely indicated in the Figures. observations and drawings of When it was necessary, preserved specimens, or in some morphological details of the lizards case from careful adaptations of were drawn under a dissecting controlled reproductions by reliable microscope. Peter‘s Dictionary authors. The available materials (1964) was a reference for above all have been the specimens morphological and anatomical of a personal collection (JMC-DC: nomenclature, as well as the several José M. Cei Diagnostic Collection), Etheridge‘s papers, i.e. his occasionally complemented by past Ctenoblepharis adspersa studies of species from the redescription (1995). The drawings Herpetological Collections of some of the Figures have been all more or Museums or Scientific Institutions less slightly magnified: we believe whose list and acronyms are that individual magnification of following. IBA-UNC: Collections symbols in every identified drawing of the Instituto de Biología , scale is useless to our purposes. Universidad Nacional de Cuyo, Besides the specimens documented Mendoza, Argentina; MZUSP: in the Figures, hundreds of specimens

4 José M. Cei corresponding to the patterns have phylogenetic indented taxonomy been observed and compared, for exhibited in the valuable conclusive improving the truthfulness and statements by Estes et al. (1988) significance of our samples. was considered as a suitable model Among the most important (Figure 2), together with the literature supporting the harvesting, reassessment of Iguania by Frost et arranging and discussion of our al. (2001). Such taxonomy has been representative samples, we wish to followed here for comparison with highlight the useful, good or our detailed morphological outstanding papers or reviews by observations, far away from any Anderson (1999), Angel (1942), aprioristic personal opinion. Then, Blarc (1977), Brygoo (1971, 1978), without disregarding other Dixon (1973), Donnelly and Myers interesting systematic contributions, (1991), Greer (1970), Grismer et as the recent paper by Schulte et al. al. (1994), Haas and Werner (1969), (2003), in full agreement with our Halliday and Adler (2002), findings, we recognize the taxon Hoogmoed (1973), Kluge (1974), Pleurodonta and its included Leviton et al. (1992), Mertens families as in the mentioned Frost (1958, 1959), Pregill (1992), Ruibal et al.’s (2001) arrangement. On the (1964), Savage (1963), Tayer contrary, the opposite general taxon (1956), Van Demburg (1978), Iguanidae (sensu Boulenger, 1895) Vanzolini et al. (1965, 1980) and postulated in the paper by Schulte Wiens (1993). et al., based on bio-molecular and morphological researchs, is fully discordant with our results. RESULTS In the following 27 Figures, Analytical review of comparative specific supraocular scutellation reports on supraocular lepidosis in patterns belonging to genera and Squamata families of the major taxa listed in As pointed out in the the indented taxonomy are reported. Introduction, the aim of this Evidently it was impossible to research was to put in evidence a check the thousands of living correlative relationship between a species of lizards, but significantly classificatory system and a representative drawings of enough significant set of data on generic and specific taxa, are supraocular scutellation in provided. Squamata reptiles. The

MULTEQUINA 16: 1-52, 2007 5 Iguania Chamaeleonidae

PLEURODONTA Iguanidae Leiocephalidae Leiosauridae Leiosaurinae Enyaliinae Opluridae Polychrotidae Tropiduridae

Scleroglossa GEKKOTA

AUTARCHOGLOSSA Scincomorpha Lacertoidea Xantusiidae Lacertiformes Teiioidea

Scincoidea Scincidae

Anguimorpha Helodermatidae Lanthanotidae

Figure 2. Squamata: phylogenetic indented taxonomy according to Estes et al. (1988). The taxon Iguania was adapted in accordance with Frost et al. (2001) Figura 2. Squamata: taxonomía filogenética indentada según Estes et al. (1988). El taxón Iguania fue adoptado de acuerdo con Frost et al. (2001)

6 José M. Cei Two opposed major systematic patterns has been strengthened by categories stand out in the their direct control in many other Squamata indented taxonomy: agamid specimens, or by other Iguania and Scleroglossa, sustained careful documentary reports by by anatomical, mostly osteological reliable authors. Agamid patterns characters. Acrodonta and of Figure 3 have been compared Pleurodonta are a subdivision of and confirmed in other 15 African Iguania, likewise countersigned by and Indian taxa of the Agama, osteological differences. We can in 15 Asiatic taxa of Draco, in 4 anticipate that a fairly simple Indian taxa of Calotes whose supraocular scutellation belongs to irregular supraocular scales appear Acrodonta, but a remarkable noticeably bulky, in 70 Australian variety of differentiated taxa of the genera Goniocephalus, supraocular scales is shown by the Amphibolurus, Tympanocryptis, Pleurodonta families. Chelosania, Dipariphera, Agamidae and Chamaeleonidae Physignathus and Lophognathus. are two large and widespread Chamaeleonidae, a specialized, Acrodont families. With more than mostly arboreal family, with more 50 genera and 400 species, than 130 species but only 6 genera Agamidae are found in the African, are distributed in Africa, Euro-Asiatic and Australian Madagascar, Arabia, India and Sri continents, with an extraordinary Lanka. The protruding eyes of morpho-ecological diversification. , covered with scaly But a relatively uniform covering lids, are unique: in Figure 4, the of rudely roundish or enlarged supraoculars of Chamaeleo and scales appears in their supraocular from Madagascar are region, the circumorbital emphasized. A fundamental semicircles being scarcely evident analogy between the Chamaeleo and the supraciliaries moderately pattern and the agamid supraocular prominent or sharp-pointed, with scutellation is evident from its some important exceptions like the general lining of irregular scales, smart or spiny supraciliaries of distinct supraciliary border and Lyriocephalus, Phrynocephalus almost indistinguishable inner and Draco. In Figure 3 several circumorbital semicircles. The examples of these lizards are given, examples of Brookesia taxa in significant enough for the multiple Figure 4 point out the peculiar species of the Acrodonta generic feature of dermal cranial taxa. Representativeness of these ossifications in these advanced

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G H I Figure 3. AGAMIDAE. A. Agama planiceps (Angola SW Africa); B. Laudakia rupta (Iran); C. Trapelus pallidus (Arabia); D. Uromastyx asmussi (Irán); E. Phrynocephalus maculatus (Arabia Saudita); F. Draco maculatus (India); G. Japalura tricarinata (Nepal - India); H. Diporiphera reginae; I. Amphibolurus vadnappa (S Australia) Figura 3. AGAMIDAE. A. Agama planiceps (Angola SW Africa); B. Laudakia rupta (Iran); C. Trapelus pallidus (Arabia); D. Uromastyx asmussi (Irán); E. Phrynocephalus maculatus (Arabia Saudita); F. Draco maculatus (India); G. Japalura tricarinata (Nepal - India); H. Diporiphera reginae; I. Amphibolurus vadnappa (S Australia)

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Figure 4. CHAMAELEONIDAE. A. Chamaeleo bifidus (E Madagascar); B. Chamaeleo cephalolepis (I. Comore-Madagascar); C. Ch. lateralis (Madagascar); D. Chamaeleo cucullatus (NE Madagascar); E. Brookesia decaryi (Madagascar); F. Brookesia thieli (Madagascar); G. Brookesia ebenaui (Madagascar); H. B. perarmatz (Madagascar); I. Brookesia stumpffi (Madagascar) Figura 4. CHAMAELEONIDAE. A. Chamaeleo bifidus (E Madagascar); B. Chamaeleo cephalolepis (I. Comore- Madagascar); C. Ch. lateralis (Madagascar); D. Chamaeleo cucullatus (NE Madagascar); E. Brookesia decaryi (Madagascar); F. Brookesia thieli (Madagascar); G. Brookesia ebenaui (Madagascar); H. B. perarmatz (Madagascar); I. Brookesia stumpffi (Madagascar)

MULTEQUINA 16: 1-52, 2007 9 chamaeleontids, leading to a moderate, regular circumorbital striking masking of their semicircles inside. In the same supraocular scale coating, here Figure 5, midmost and inferior illustrated for several species of rows, representative of the family uncommon lizards from the Polychrotidae, are shown: isolated Madagascar. Polychrus with few neotropical The partition of the Pleurodonta species and Anolis with more than evolutionary stock is remarkable. 230 species, spread with several Also remarkable are our findings characteristic groups from south about a corresponding association eastern United States to West between different supraocular Indies, northern Mexico, scale patterns and the different southwards through Central and families listed in the indented to about 20ºS taxonomy exposed in Figure 2. latitude. A true “Polychrotidae” These families have been pattern (sensu Frost et al., 2001) is discussed in a serial issue of recognizable in the Figure 5, herpetological contributions: from bearing impressive large the paper by Frost and Etheridge circumorbital semicircles and few (1989) to the successive works by very variable internal, small shields, Frost (1992) and Frost et al. (2001). surrounded by many minute scales, Without any intentional attempt and very diminished supraciliaries of phylogenetic references, every outside. The foreseeable specific family shall be analyzed now with variation in the multiple Anolis is regard to its supraocular lepidosis. emphasized in Figure 5 by some taxa from Cuba and Colombia. In Figure 5, upper row, Leiocephalidae is only limited to Two families with few genera the genus Leiocephalus from West and species are registered in the Indies. Formerly related to upper and inferior rows of Figure Tropiduridae, its family status was 6. In Opluridae from Madagascar established by Frost et al. (2001). and Comore islands, the monotypic In its different insular taxa, the well genus Chalarodon and some differentiated supraocular lepidosis species of keep a rather prevailing in Pleurodonta is simple supraocular pattern maintained, with minor specific reminiscent of Acrodont differences. A longitudinal central scutellations. Both very similar row of large supraocular scale examples of Oplurus in this Figure stretches is evident between a exhibit irregular supraocular scales narrow row of supraciliaries and decreasing from their almost

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Figure 5. LEIOCEPHALIDAE. A. Leiocephalus psammodromus (W Indies - Caicos Isl.); B. Leiocephalus stictigaster (Cuba); C. L. barahonensis (Republica Dominicana); POLYCHROTIDAE. D. Polychrus acutirostris (Pernambuco, Brasil); E. Anolis isolepis (Guantanamo, Cuba); F. Anolis alutaceus (Guantanamo, Cuba); G. A. cyanopleurus (Guantanamo, Cuba); H. Anolis spectrum (Matanzas, Cuba); I. Anolis biporcatus (Colombia) Figura 5. LEIOCEPHALIDAE. A. Leiocephalus psammodromus (W Indies - Caicos Isl.); B. Leiocephalus stictigaster (Cuba); C. L. barahonensis (Republica Dominicana); POLYCHROTIDAE. D. Polychrus acutirostris (Pernambuco, Brasil); E. Anolis isolepis (Guantanamo, Cuba); F. Anolis alutaceus (Guantanamo, Cuba); G. A. cyanopleurus (Guantanamo, Cuba); H. Anolis spectrum (Matanzas, Cuba); I. Anolis biporcatus (Colombia)

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Figure 6. OPLURIDAE. A. Oplurus cyclurus (Madagascar); B. Oplurus fierinensis (Madagascar); C. Chalarodon madagascariensis (Madagascar). PHRYNOSOMATIDAE. D. Prhynosoma modestum (New Mexico, USA); E. Prhynosoma douglassi (Montana, USA); F. Prhynosoma cornutum (Texas, USA). CORYTOPHANIDAE. G. Corytophanes cristatus (Costa Rica); H. Basiliscus plumifrons (Costa Rica); I. serratus (Mexico) Figura 6. OPLURIDAE. A. Oplurus cyclurus (Madagascar); B. Oplurus fierinensis (Madagascar); C. Chalarodon madagascariensis (Madagascar). PHRYNOSOMATIDAE. D. Prhynosoma modestum (New Mexico, USA); E. Prhynosoma douglassi (Montana, USA); F. Prhynosoma cornutum (Texas, USA). CORYTOPHANIDAE. G. Corytophanes cristatus (Costa Rica); H. Basiliscus plumifrons (Cos- ta Rica); I. (Mexico)

12 José M. Cei indistinct circumorbital semicircles Sceloporus orcutti and S. olivaceus. to their equally indistinguishable Interesting to remark is the finding supraciliaries. In Chalarodon of these kinds of supraocular madagascariensis a quite uniform scutellation here reported for other supraocular layer of small rounded genera of such heterogeneous scales is better delimited by narrow family, like Uma, Uta, Holbrookia, circumorbital semicircles and a with analogous geographic recognizable border of very small distribution. Likewise related but supraciliaries. Corytopharidae from peculiar supraocular lepidosis may Mexico and Central America be pointed out for the arboreal likewise include only three genera Urosaurus (Figure 8), so carefully with few species each. Irregular reviewed by Wiens (1993) for the supraocular scales are broader in same arid environment. Their Corytophanes and Laemanctus, variable six broad longitudinal small in Brasiliseus. Circumorbital supraocular scales, the other semicircles and supraciliaries, adherent rows of minor scales as sharp-pointed in Corytophanes, well as the shape and size of stand out in this family. Striking in circumorbital semicircles and central row of Figure 6 is the supraciliaries, support an undifferentiated supraocular unmistakable generic identity. surface of the horny skull roof of Similar valuation can be made for some Phrynosoma, odd genus of the related Petrosaurus species the family Phrynosomatidae, inhabiting mainly the same exposed in the following Figures. Californian territories. Even in Several genera, found from south Figure 8, inferior row, the western North America southward supraocular lepidosis in to Panama, were assembled in Crotaphytidae can be observed. Phrynosomatidae. Collared lizards and Leopard In Figure 7 the mentioned lizards ( and “Pleurodont pattern” is Gambelia) also from the subdesert recognizable for the about fifty taxa rocky or sandy of of the spiny Sceloporus, southwestern North America are distinguished by their expanded unmistakable for their nearly circumorbital semicircles and smooth supraocular covering by prominent superciliaries, very minute circumorbital surrounding 5-6 broad supraocular semicircles and sprinkled by some scales. A somewhat diverging insignificant scales. morphology can be observed, as in

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Figure 7. PHRYNOSOMATIDAE. A. Sceloporus merriami (Texas, USA); B. Sceloporus poinsetti (Texas, USA); C. Sceloporus clarki (Arizona, USA); D. S. magíster (S California, USA); E. S. orcutti (S. California, USA); F. Sceloporus olivaceus (Texas, USA); G. Uma notata (S. California, USA); H. Uta stamburiana (S. California, USA); I. Holbrookia texana (Texas, USA) Figura 7. PHRYNOSOMATIDAE. A. Sceloporus merriami (Texas, USA); B. Sceloporus poinsetti (Texas, USA); C. Sceloporus clarki (Arizona, USA); D. S. magíster (S California, USA); E. S. orcutti (S. California, USA); F. Sceloporus olivaceus (Texas, USA); G. Uma notata (S. California, USA); H. Uta stamburiana (S. California, USA); I. Holbrookia texana (Texas, USA)

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F G H Figure 8. PHRYNOSOMATIDAE. A. Urosaurus ornatus (Arizona, USA); B. Urosaurus nigricaudus (Baja California, USA); C. Urosaurus graciosus (California, USA); D. Petrosaurus mearnsi (California, USA); E. Petrosaurus thalassinus (California, USA). CROTAPHYTIDAE. F. Crotaphytus insularis (S. California, USA); G. Crotaphytus collaris (Texas, USA); H. Gambelia wislizenii (Baja California, USA) Figura 8. PHRYNOSOMATIDAE. A. Urosaurus ornatus (Arizona, USA); B. Urosaurus nigricaudus (Baja California, USA); C. Urosaurus graciosus (California, USA); D. Petrosaurus mearnsi (California, USA); E. Petrosaurus thalassinus (California, USA). CROTAPHYTIDAE. F. Crotaphytus insularis (S. California, USA); G. Crotaphytus collaris (Texas, USA); H. Gambelia wislizenii (Baja California, USA)

MULTEQUINA 16: 1-52, 2007 15 Two families of Pleurodonta the isolated insular taxa as the with a common general supraocular Polynesian lizards (Brachylophus lepidosis are documented in Figure fasciatus, B. vitiensis) or the 9. Belonging to the neotropical peculiar Conolophus from the Hoplocercidae, with a dozen Galapagos Archipelago. species, the three queer genera The important family Hoplocercus, and Liolaemidae, exemplified in show always very Figures 10-11, was also established distinct circumorbital semicircles in the cited assessment by Frost et and supraciliaries, sharp pointed in al. (2001): it comprises the genera Enyahioides. Supraocular scales are Ctenoplepharis, Phymaturus and undifferentiated, but with three from southern South central quadrangular rows in America. As stated by Etheridge Morunasaurus. Similar lepidosis, (1995) in his redescription of though with visible tendency to Ctenoblepharis adspersa from polymorphism is shown by the desert landscapes of , the interesting taxon Iguanidae (sensu unresolved relationships of such Frost et al., 2001), whose genera highly derived arenicolous reptiles hold no more than two species each. with Phymaturus and Liolaemus Minor differences among the are emphasized as differential South American Iguana, characters by their small, subequal Cercosaura and Dipsosaurus supraoculars. That is congruent depend upon the shape and size of with our drawing in Figure 10. supraocular scales, whose more or Other drawings illustrate several less regular rows are encircled by species of the rock-climbing genus evident supraciliaries, and behind Phymaturus, all similar enough for by circumorbital semicircles, very the scarcely differentiated enlarged in Iguana. They show the supraocular scutellation. differences between the almost Notwithstanding, the species of the undifferentiated probably primitive patagonicus group (those drawn patterns of the insular Pacific here are: indistinctus, patagonicus, Brachylophus and Conolophus, and somuncurensis, payuniae) have the more differentiated lepidosis in more enlarged supraorbital scales Iguana iguana from neotropical along the circumorbital semicircles. forests. Evolutionary factors of Liolaemus assembles about 160 primeval distribution in geological taxa, mainly Andean and Sub- times are likely related with the Andean in many cases likely of actual conservative conditions of Pleistocene or post-glacial

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Figure 9. HOPLOCERCIDAE. A. Hoplocercus spinosus (); B. (Rio San- tiago, Ecuador): C. Enyalioides heterolepis (Ecuador). IGUANIDAE. D. Iguana iguana (Brazil); E. Cercosaura similis (Central America); F. Dipsosaurus dorsalis (South California, USA); G. Brachylophus fasciatus (Orchid Isl., Polynesia); H. B. vitiensis (Orchid Isl., Polynesia); I. Conolophus subvristatus (Galapagos Isl.) Figura 9. HOPLOCERCIDAE. A. Hoplocercus spinosus (Brazil); B. Morunasaurus annularis (Rio Santia- go, Ecuador): C. Enyalioides heterolepis (Ecuador). IGUANIDAE. D. Iguana iguana (Brazil); E. Cercosaura similis (Central America); F. Dipsosaurus dorsalis (South California, USA); G. Brachylophus fasciatus (Orchid Isl., Polynesia); H. B. vitiensis (Orchid Isl., Polynesia); I. Conolophus subvristatus (Galapagos Isl.)

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G H I Figure 10. LIOLAEMIDAE. A. Ctenoblepharis adspersa (Peru); B. Phymaturus indistinctus (Chubut, Argentina); C. Phymaturus patagonicus (Chubut, Argentina); D. Ph. somuncurensis (Rio Ne- gro, Argentina); E. Ph. payuniae (Payunia, Argentina); F. Ph. punae (San Juan, Argentina); G. Ph. mallimaccii (La Rioja, Argentina); H. Ph. verdugo (South Mendoza, Argentina); I. Ph. antofagastensis (Catamarca, Argentina) Figura 10. LIOLAEMIDAE. A. Ctenoblepharis adspersa (Peru); B. Phymaturus indistinctus (Chubut, Argentina); C. Phymaturus patagonicus (Chubut, Argentina); D. Ph. somuncurensis (Rio Ne- gro, Argentina); E. Ph. payuniae (Payunia, Argentina); F. Ph. punae (San Juan, Argentina); G. Ph. mallimaccii (La Rioja, Argentina); H. Ph. verdugo (South Mendoza, Argentina); I. Ph. antofagastensis (Catamarca, Argentina)

18 José M. Cei speciation. Its more characterizing , the purpose of feature is, alongside the very our recent review (Cei et al., 2004), evident circumorbital semicircles, as well as (Cei et al., the longitudinal row of 5-6 broad 2003), are the most undifferentiated supraocular scales, not as irregular members of the subfamily. All the and confused as in the Phymaturus recognized species of patagonicus group (Figure 11). In Pristidactylus were analyzed in few eastern Brazilian arenicolous detail in our revision, adding three taxa, as Liolaemus occipitalis, a analogous forms to the four diverging pattern of Argentine entities reported in undifferentiated scales, perhaps Figure 12. Pristidactylus primitive, may be pointed out. nigroiugulus from Chubut exhibits Before moving to the complex an acrodont-like pattern, somewhat family Tropiduridae of which some modified in P. scapulatus and P. taxa as Liolaemidae and araucanus, but equal in P. Leiocephalidae have already been achalensis, P. casuhatiensis from referred to (Frost and Etheridge, Argentina and P. alvaroi from 1989), we must pay attention to the Chile. However in P. fasciatus, Pleurodont lizards Leiosauridae, as from the arid scrub of central well included in Polychrotidae prior Argentina, a lepidosis very similar to the remembered reclassification to those of the other two genera is by Frost et al. (2001). found, evoking probably evolutionary genetic relationships. Parted into two subfamilies, In Figure 12 a unique, just Leiosaurinae and Enyaliinae, a mentioned, iguanian supraocular number of genera and a variety of lepidosis is shown, both in patterns pertain to this unusual Diplolaemus and , with iguanian systematic entity, to which a central rounded bouquet of larger Figures 12 and 13 are dedicated. In scales in a very homogeneous field Leiosauridae, three peculiar genera of diminished scales. In neotropical are known, Pristidactylus, Enyaliinae the here commented Diplolaemus and Leiosaurus, of extant conditions of morphological very ancient lineage: Leiosaurus is supraocular uniformity appear found fossil in Pliocene layers of altered enough. In Brazilian Argentina, Pristidactylus down the homogeneous pattern right in the Miocene sediments of of E. perditus is replaced by a Chubut (Argentina). condition almost repeating that of Pristidactylus fasciatus or

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Figure 11. LIOLAEMIDAE. A. Liolaemus occipitalis (Rio Grande do Sul, Brazil); B. Liolaemus wiegmanni (Tucuman, Argentina); C. L. nigriceps (Antofagasta, Chile); D. L. chiliensis (Concepcion, Chile); E. L. ruibali (San Juan, Argentina); F. L. kingi (Santa Cruz, Argentina); G. L. silvanae (S. Cruz, Argentina); H. L. fitzingeri (Chubut, Argentina); I. Liolaemus elongatus (Neuquen, Argentina) Figura 11. LIOLAEMIDAE. A. Liolaemus occipitalis (Rio Grande do Sul, Brazil); B. Liolaemus wiegmanni (Tucuman, Argentina); C. L. nigriceps (Antofagasta, Chile); D. L. chiliensis (Concepcion, Chile); E. L. ruibali (San Juan, Argentina); F. L. kingi (Santa Cruz, Argentina); G. L. silvanae (S. Cruz, Argentina); H. L. fitzingeri (Chubut, Argentina); I. Liolaemus elongatus (Neuquen, Argentina)

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Figure 12. LEIOSAURIDAE. Leiosaurinae. A. Pristidactylus scapulatus (Mendoza, Argentina); B. P. araucanus (Neuquen, Argentina); C. P. nigroiugulus (Chubut, Argentina); D. P. fasciatus (Mendoza, Argentina); E. Diplolaemus bibroni (Santa Cruz, Argentina); F. Diplolaemus darwini (S. Cruz, Argentina); G. Leiosaurus catamarcencis (San Juan, Argentina); H. L. paronae (Mendoza, Argentina); I. Leiosaurus belli (Chubut, Argentina) Figura 12. LEIOSAURIDAE. Leiosaurinae. A. Pristidactylus scapulatus (Mendoza, Argentina); B. P. araucanus (Neuquen, Argentina); C. P. nigroiugulus (Chubut, Argentina); D. P. fasciatus (Mendoza, Argentina); E. Diplolaemus bibroni (Santa Cruz, Argentina); F. Diplolaemus darwini (S. Cruz, Argentina); G. Leiosaurus catamarcencis (San Juan, Argentina); H. L. paronae (Mendoza, Argentina); I. Leiosaurus belli (Chubut, Argentina)

MULTEQUINA 16: 1-52, 2007 21 Diplolaemus (Figure 13), as in E. considered as in the brasiliensis, E. iheringi and cited Frost’s paper (1992). Its likewise in . More fundamental feature is a complete intriguing is the genus , supraocular lining of imbricate whose specific patterns can look keeled scales, together with evident alike to that of Urostrophus vautieri keeled circumorbital semicircles, (i.e. Anisolepis grillii), superficially and variable supraciliaries, showy to that of the Corythophanid and sharp-pointed in Uranoscodon, Basiliscus plumifrons (i.e. superimposed in Stenocercini taxa. ), and The other prevailing Tropidurus strikingly alike to those of the shows in the similar genera genera Urosaurus and Petrosaurus Microlophus, Plesiomicrolophus of the family Phrynosomatidae (i.e. and Tropidurus a general unfolding Anisolepis longicauda). of longitudinal rows of 4-6 broad Remodeled as subfamily by supraocular scales, of different size Frost (1992), Tropiduridae was and flanked by minor, rounded, confirmed as a family by Frost et irregular scales. A remarkable al. (2001) without making clear the variation is even observed in the definitive position (subfamily or evident circumorbital semicircles tribe) of the diversified Stenocercini which are more moderate in tribe. The supraocular lepidosis in Stenocercini taxa. Always Tropidurus appears to be the most particularly distinct in our drawings significant pattern of Iguanian is the interparietal scale and its Pleurodonta (Figures 14, 15, 16). “eye”, more or less developed (crf. Two strongly diverging types are Frost et al., 2001: p.21). The distinguishable. The type of the unifying tendency of Frost`s genus Uranoscodon from tropical assessment (1992) led to extend South America is present too in the the generic Tropidurus name to Stenocercini tribe, with some taxa known as Stenocercus guentheri from and Plica, now Tropidurus Ecuador, Ophryessoides caducus strobilurus, T. plica and T. umbra. from Northern Argentina, A third Plica, now Tropidurus Proctotetrus pectinatus and P. lumaria from Venezuelan Tepui, doellojuradoi from central was well documented by Donnelly Argentina, the last three lizards and Myers (1991).

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Figure 13. LEIOSAURIDAE. Enyaliinae. A. Enyalius brasiliensis (Espirito Santo, Brazil); B. Enyalius iheringi (Sao Paulo, Brazil); C. E. perditus (Brazil); D. Urostrophus vautieri (Sao Paulo, Brazil); E. Urostrophus iheringi (S. Paulo, Brazil); F. U. gallardoi (Cordoba, Argentina); G. Anisolepis grillii (S. Paulo, Brazil); H. Anisolepis undulatus (S. Paulo, Brazil); I. A. longicauda (Corrien- tes, Argentina) Figura 13. LEIOSAURIDAE. Enyaliinae. A. Enyalius brasiliensis (Espirito Santo, Brazil); B. Enyalius iheringi (Sao Paulo, Brazil); C. E. perditus (Brazil); D. Urostrophus vautieri (Sao Paulo, Brazil); E. Urostrophus iheringi (S. Paulo, Brazil); F. U. gallardoi (Cordoba, Argentina); G. Anisolepis grillii (S. Paulo, Brazil); H. Anisolepis undulatus (S. Paulo, Brazil); I. A. longicauda (Corrien- tes, Argentina)

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Figure 14. TROPIDURIDAE. Tropidurini. A. Uranoscodon superciliosus (Northern S. America); B. (Arica, Chile); C. (Ecuador); D. Tropidurus hispidus (); E. (S. Paulo, Brazil); F. T. etheridgei (La Rioja, Argentina); G. T. spinulosus (Formosa, Argentina); H. T. spinulosus (Cordoba, Argentina); I. Tropidurus guarani (Asuncion, ) Figura 14. TROPIDURIDAE. Tropidurini. A. Uranoscodon superciliosus (Northern S. America); B. Microlophus peruvianus (Arica, Chile); C. Microlophus occipitalis (Ecuador); D. Tropidurus hispidus (Venezuela); E. Tropidurus torquatus (S. Paulo, Brazil); F. T. etheridgei (La Rioja, Argentina); G. T. spinulosus (Formosa, Argentina); H. T. spinulosus (Cordoba, Argentina); I. Tropidurus guarani (Asuncion, Paraguay)

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G H I Figure 15. TROPIDURIDAE. Tropidurini. A. Tropidurus semitaeniatus (Bahia, Brazil); B. Tropidurus pictus (male; Salta, Argentina); C. T. pictus (female; Salta, Argentina); D. Tropidurus hygomi (Sergipe, Brazil); E. Tropidurus bogerti (Bolivar, Venezuela); F. Tropidurus amathites (Bahia, Brazil); G. Tropidurus strobilurus (NW Brazil); H. Tropidurus plica (Para, Brazil); I. Tropidurus umbra (French ) Figura 15. TROPIDURIDAE. Tropidurini. A. Tropidurus semitaeniatus (Bahia, Brazil); B. Tropidurus pictus (male; Salta, Argentina); C. T. pictus (female; Salta, Argentina); D. Tropidurus hygomi (Sergipe, Brazil); E. Tropidurus bogerti (Bolivar, Venezuela); F. Tropidurus amathites (Bahia, Brazil); G. Tropidurus strobilurus (NW Brazil); H. Tropidurus plica (Para, Brazil); I. Tropidurus umbra (French Guyana)

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Figure 16. TROPIDURIDAE. Stenocercini. A. Stenocercus ornatus (Guayaquil, Ecuador); B. Stenocercus trachycephalus (Bogota, Colombia); C. S. guentheri (Ecuador); D. S. varius (Pichincha, Ecua- dor); E. Ophryoessoides caducus (Jujuy, Argentina); F. O. iridescens (Esmeralda, Ecuador); G. Stenocercus roseiventris (Salta, Argentina); H. Stenocercus pectinatus (Rio Negro, Argen- tina); I. S. doellojuradoi (Santiago del Estero, Argentina) Figura 16. TROPIDURIDAE. Stenocercini. A. Stenocercus ornatus (Guayaquil, Ecuador); B. Stenocercus trachycephalus (Bogota, Colombia); C. S. guentheri (Ecuador); D. S. varius (Pichincha, Ecua- dor); E. Ophryoessoides caducus (Jujuy, Argentina); F. O. iridescens (Esmeralda, Ecuador); G. Stenocercus roseiventris (Salta, Argentina); H. Stenocercus pectinatus (Rio Negro, Argen- tina); I. S. doellojuradoi (Santiago del Estero, Argentina)

26 José M. Cei Critical reports and annotations (Rhinogecko misonnei, on so extensive Iguania stem of the fasciata), or sharp-pointed species followed indented taxonomy are (Phyllurus platurus, finished. We must comment now on ciliaris). Sometimes growths or the next major embranchment, tubercles appear on the supraocular Scleroglossa, then parted into two surface, as in Phyllurus cornutus, natural lineages: Gekkota and Ph. caudiannulatus, Nephrurus Autarchoglossa. The families asper from Queensland and attributed to Gekkota are the like, Nephrurus wheeleri from W ecologically versatile Gekkonidae, Australia. Besides Australian Diplodactylidae, Eublepharidae and examples, these investigations were the unusual -like fossorial extended to several other samples Pygopodidae, reviewed by Kenge from Africa, Middle East India, Iran (1974) in a prolix fashion. Obviously and America. Our research on it is impossible to offer here Gekkonidae, Diplodactylidae, proportional scutellation samples for Eublepharidae has thus been carried taxa having some 320 species and out on about a hundred and twenty 88 genera, such as the almost specimens. cosmopolitan Gekkonidae, or for Strongly dissimilar are the more the 120 species and 15 genera of than 30 species and 8 genera Diplodactylidae from New Zealand belonging to the snake-like and Australia, or the 22 species and Pygopodidae from Australia and New 6 genera of Eublepharidae, scattered Guinea. These really unusual lizards across the SW of North America, are represented in Figure 18, pointing Central America, Africa and out their enlarged cephalic shields Southern Asia. However the and the extreme reduction of their examples of Figure 17 are supraocular scales: only one in Aclys, efficacious in expressing the general and Paradelma, two in simplicity of supraocular Delma, Pygopus and Pletholax, scutellation of all these Gekkota, where evident supraciliaries are given the modest specific or generic shown. A most impressive reduction variation in their uniform small is found in Lialis and scales even reduced to short spots, Ophidiocephalus. Lastly, nothing can or almost absent. Also the be added concerning the evolutionary circumorbital semicircles have been relationships of Pygopodidae with restrained to a nearly concealed the other Gekkota, anatomical structure. Very different in size and considerations being the decisive shape are the supraciliaries, mostly factor in this classification. linear and narrow, in some notched MULTEQUINA 16: 1-52, 2007 27 ABC D

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Figure 17. GEKKOTA. GEKKONIDAE. A. Cyrtopodion heterocercum (Iran); B. Rhinogecko misonnei (Pakistan); C. Briba brasiliana (Minas Gerais, Brazil); D. Phyllopezus pollicaris (Pernambuco, Brazil); E. turcicus (Mediterranean region); F. Hemidactylus persicus (Saudi Arabia); G. Gonatodes concinnatus (Napo, Ecuador); H. Stenodactylus grandiceps (Jordan, Middle East); I. Homonota underwoodi (San Juan, Argentina); J. Homonota whitii (Cordoba, Argentina); K. Homonota darwini (Payunia, Argentina); L. Homonota fasciata (Cordoba, Argentina). EUBLEPHARIDAE. M. Coleonyx brevis (Texas, USA); N. Eublepharis macularius (Pakistan); O. Eublepharis turcmenicus (Iran). DIPLODACTYLIDAE. P. Phyllurus platurus (South Australia) Figura 17. GEKKOTA. GEKKONIDAE. A. Cyrtopodion heterocercum (Iran); B. Rhinogecko misonnei (Pakistan); C. Briba brasiliana (Minas Gerais, Brazil); D. Phyllopezus pollicaris (Pernambuco, Brazil); E. Hemidactylus turcicus (Mediterranean region); F. Hemidactylus persicus (Saudi Arabia); G. Gonatodes concinnatus (Napo, Ecuador); H. Stenodactylus grandiceps (Jordan, Middle East); I. Homonota underwoodi (San Juan, Argentina); J. Homonota whitii (Cordoba, Argentina); K. Homonota darwini (Payunia, Argentina); L. Homonota fasciata (Cordoba, Argentina). EUBLEPHARIDAE. M. Coleonyx brevis (Texas, USA); N. Eublepharis macularius (Pakistan); O. Eublepharis turcmenicus (Iran). DIPLODACTYLIDAE. P. Phyllurus platurus (South Australia)

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Figure 18. PYGOPODIDAE. A. Aclys concinna (W Australia); B. Aprasia aurita (SE Australia); C. Aprasia parapulchella (SE Australia); D. Aprasia repens (SW Australia); E. Delma australis (W Aus- tralia); F. Delma elegans (W Australia); G. Lialis jicari (New Guinea); H. Ophidiocephalus taeniatus (N Australia); I. Pygopus nigriceps (Australia); J. Pygopus lepidopodus (SW Aus- tralia); K. Paradelma orientalis (Queensland, Australia); L. Pletholax gracilis (SW Australia) Figura 18. PYGOPODIDAE. A. Aclys concinna (W Australia); B. Aprasia aurita (SE Australia); C. Aprasia parapulchella (SE Australia); D. Aprasia repens (SW Australia); E. Delma australis (W Austra- lia); F. Delma elegans (W Australia); G. Lialis jicari (New Guinea); H. Ophidiocephalus taeniatus (N Australia); I. Pygopus nigriceps (Australia); J. Pygopus lepidopodus (SW Australia); K. Paradelma orientalis (Queensland, Australia); L. Pletholax gracilis (SW Australia)

MULTEQUINA 16: 1-52, 2007 29 Drastic morphological changes, border are visible, the external facing examined Iguania supraocular narrower; almost rudimentary in the patterns, shall now be commented two adjacent supraocular rows of upon for the successive branches of Cercosaura; that in Lepidophyma Autarchoglossa, foreseen in our flavimaculatum, the reduction of the indented taxonomy, beginning with supraocular scales is extreme, to “a Scincomorpha and its noticeable lines narrow fleshy ridge above the eye” Lacertoidea and Scincoidea. Without (Savage, 1963). any doubt, observing their cephalic Leaving so exceptionally modified lepidosis, some evolutionary trends scale features, the specimens reported are difficult to understand at first sight in the inferior drawings of Figure 19 between the night lizards Xantusidae introduce us to the great American and the Lacertiformes Lacertidae and stem of Lacertiformes, the Teiioidea, Teiioidea of Lacertoidea, as well as to whose taxon Teiidae belong the the families Scincidae and Cordylidae, reproduced species of the genus of Scincoidea. Studied formerly by Tupinambis, still justifying a Savage (1963), who firstly assigned systematic definition as subfamily. them to Gekkota, these viviparous, The specific variation in the pattern secretive reptiles, with their genera of Tupinambis anticipates here a Lepidophyma, Xantusia and general distribution of supra-ocular Cricosaura (Figure 19), live in scales dominant in Scincomorpha, crevices or cracks of granite flakes, in with major or minor alteration. forested regions of Mexico, Central Lacking a definition of circumorbital America, Southern and Western semicircles, 4-5 supraocular scales, United States, and there being only somewhat different in size, stretch one species confined to Cuba, the out on the eye border, edged by one or monotypic Cricosaura (C. typica). two narrower linear supraciliary rows. From Figure 19 we can note that in That simple scheme, of the large- Xantusia vigilis circumorbital sized, powerful Tupinambis shall be semicircles are absent, as in X. commented upon, more or less henshawi or X. arizonae, besides the modified, in the following Figures of other genera, and that the supraocular Teiioidea, Lacertidae and Scincoidea. scales consist of a narrow longitudinal Fossil remains of Tupinambis from row near the eye border, with no South American early Miocene to evident supraciliaries; that in Xantusia late Miocene are very suggestive riversiana (ancient Klauberina records stating the significant Savage 1957) and Cricosaura typica evolutionary lineage of Teiid forms two narrow adjacent rows on the eye in the continent (Albino et al. 2006).

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Figure 19. XANTUSIIDAE. TEIIDAE. XANTUSIIDAE. A. Xantusia vigilis (SW USA); B. Xantusia vigilis (Lower California, USA); C. X. henshawi (SW USA); D. X. riversiana (SW California, USA); E. Lepidophyma flavimaculatum (Costa Rica); F. Cricosaura typica (E Cuba). TEIIDAE. G. Tupinambis nigromaculatus (Barinas, Venezuela); H. T. teguixin (E Brasil); I. T. merianae (Bs. Aires, Argentina); J. Tupinambis rufescens (Mendoza, Argentina) Figura 19. XANTUSIIDAE. TEIIDAE. XANTUSIIDAE. A. Xantusia vigilis (SW USA); B. Xantusia vigilis (Lower California, USA); C. X. henshawi (SW USA); D. X. riversiana (SW California, USA); E. Lepidophyma flavimaculatum (Costa Rica); F. Cricosaura typica (E Cuba). TEIIDAE. G. Tupinambis nigromaculatus (Barinas, Venezuela); H. T. teguixin (E Brasil); I. T. merianae (Bs. Aires, Argentina); J. Tupinambis rufescens (Mendoza, Argentina)

MULTEQUINA 16: 1-52, 2007 31 In Figure 20, upper row, the greatest careful Dixon’s research (1973) it Teiid species are documented. Five was possible to remark the supraocular scales with much reduced evolutionary process leading to loss circumorbital semicircles and relatively of the supraocular scales in the wide supraciliaries are shown by gymnophthalmid genus : a Ameiva and Callopistes: 6-5 process whose progressive steps have supraocular scales and a narrower been already accomplished, i.e., in supraciliary row characterize Gymnophthalmus underwoodi from Dracaena. Taxa of Dicrodon and Surinam or Heterodactylus , on the intermediate row, imbricatus from SE Brazil. In Figure exhibit 3-2 larger broad scales with a 22 we can point out the reduced but very small terminal scale, reduced or still recognizable two supraocular absent circumorbital semicircles, but scales of Bachia heteropa evident overlapping supraciliaries. On (Venezuela), B. flavescens (Guyana), the inferior row of the figure, the B. pallidiceps (Panama, Colombia), repeated Teiid lepidosis accounts for and also their elimination from the 3 supraocular scales of Teius, with Bachia talpa (Colombia) to B. circumorbital semicircles and intermedius (Peru), with narrow overlapping supraciliaries. For the supraciliaries remaining in the end. numerous and so variable whiptails Differences in body, scales, (Cnemidophorus) we reported 4-3 ecological relationships and supraocular scales and developed evolutionary trends within the genus, supraciliaries: only for Cnemidophorus related to fossorial adaptation, were ocellifer the circumorbital semicircle preliminarily discussed in the cited was drawn here. paper. Corresponding to the old Now we will go on with the subdivision into Microteiids and inferior row of Figure 22 and Figure Macroteiids, the family 23, illustrating characteristic Gymnophthalmidae was established features of the African and Euro- for the many small-sized forms Asiatic great family Lacertidae, distributed with more than 150 with more than 220 species and 25 species and 35 genera in Central and genera. That is our last information South America. Figures 21-22 deal about Lacertoidea of the with those lizards having 4, 3, 2 Scincomorpha embranchment. The supraocular scales, mainly lacking striking morphological affinities in circumorbital semicircles but putting supraocular lepidosis between in evidence a more or less wide Lacertidae and their counterpart supraciliary border. Through the Teiioidea from the American

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Figure 20. TEIIDAE. A. Ameiva ameiva (French Guyana); B. Callopistes palluma (Chile); C. Dracaena paraguayensis (Mato Grosso, Brazil); D. D. guianensis (Guyana); E. Dicrodon guttulatus (Ecua- dor); F. Kentropyx viridistriga (Corrientes, Argentina); G. Kentropyx lagartica (Tucuman, Ar- gentina); H. K. pelviceps (Amazonia, Ecuador); I. Teius oculatus (Cordoba, Argentina); J. Cnemidophorus lemniscatus (Yaracui, Venezuela); K. C. ocellifer (Chaco, Argentina); L. C. lacertoides (Jujuy, Argentina) Figura 20. TEIIDAE. A. Ameiva ameiva (French Guyana); B. Callopistes palluma (Chile); C. Dracaena paraguayensis (Mato Grosso, Brazil); D. D. guianensis (Guyana); E. Dicrodon guttulatus (Ecuador); F. Kentropyx viridistriga (Corrientes, Argentina); G. Kentropyx lagartica (Tucuman, Argentina); H. K. pelviceps (Amazonia, Ecuador); I. Teius oculatus (Cordoba, Argentina); J. Cnemidophorus lemniscatus (Yaracui, Venezuela); K. C. ocellifer (Chaco, Argentina); L. C. lacertoides (Jujuy, Argentina)

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Figure 21. GYMNOPHTHALMIDAE. A. angulatus (Northern South America); B. bicarinatus (Brasil, Guyana); C. Pantodactylus schreibersi (Cordoba, Argenti- na); D. Arthrosaura kockii (Surinam, N Brazil); E. Anadia bipunctata (Merida, Venezuela); F. agilis (Guyana, N Brazil); G. Leposoma guianense (Guyana, N Brazil); H. Iphisa elegans (Northern South America); I. Microblepharus maximiliani (NE Brazil, Paraguay); J. argulus (Ecuador, Guyana); K. Gymnophthalmus multiscutatus (NE Brazil); L. Gymnophthalmus underwoodi (Surinam, Tobago Is.) Figura 21. GYMNOPHTHALMIDAE. A. (Northern South America); B. Neusticurus bicarinatus (Brasil, Guyana); C. Pantodactylus schreibersi (Cordoba, Argenti- na); D. Arthrosaura kockii (Surinam, N Brazil); E. Anadia bipunctata (Merida, Venezuela); F. (Guyana, N Brazil); G. Leposoma guianense (Guyana, N Brazil); H. Iphisa elegans (Northern South America); I. Microblepharus maximiliani (NE Brazil, Paraguay); J. Prionodactylus argulus (Ecuador, Guyana); K. Gymnophthalmus multiscutatus (NE Brazil); L. Gymnophthalmus underwoodi (Surinam, Tobago Is.)

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Figure 22. GYMNOPHTHALMIDAE. A. Heterodactylus imbricatus (SE Brazil); B. Bachia heteropa (Ve- nezuela); C. Bachia flavescens (Guyana); D. B. pellidiceps (Panama, Colombia); E. B. talpa (Colombia); F. B. trisanale (Peru, Ecuador); G. B. huallagana (Peru); H. B. intermedia (Peru). LACERTIDAE. I. Psammodromus algirus (SW Europe, NW Africa); J. Acanthodactylus erytrurus (SW Europe); K. A. pardalis (Maroc, Marruecos); L. A. boskianus (NW Africa) Figura 22. GYMNOPHTHALMIDAE. A. Heterodactylus imbricatus (SE Brazil); B. Bachia heteropa (Vene- zuela); C. Bachia flavescens (Guyana); D. B. pellidiceps (Panama, Colombia); E. B. talpa (Co- lombia); F. B. trisanale (Peru, Ecuador); G. B. huallagana (Peru); H. B. intermedia (Peru). LACERTIDAE. I. Psammodromus algirus (SW Europe, NW Africa); J. Acanthodactylus erytrurus (SW Europe); K. A. pardalis (Maroc, Marruecos); L. A. boskianus (NW Africa)

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Figure 23. LACERTIDAE. A. Acanthodactylus blanfordi (Afghanistan); B. A. grandis (Siria); C. Eremias strauchi (Armenia, Iran); D. Eremias arguta (Iran, Central Asia); E. Lacerta lepida (SW Europe); F. Lacerta defilippi (Iran); G. L. vivipara (Northern Eurasia); H. Gallotia atlantica (Canary Isl.); I. Mesalina brevirostris (Middle East); J. Ichnotropis bivittata (Angola); K. scalaris (Angola); L. Nucras taeniolata (SW Africa) Figura 23. LACERTIDAE. A. Acanthodactylus blanfordi (Afghanistan); B. A. grandis (Siria); C. Eremias strauchi (Armenia, Iran); D. Eremias arguta (Iran, Central Asia); E. Lacerta lepida (SW Europe); F. Lacerta defilippi (Iran); G. L. vivipara (Northern Eurasia); H. Gallotia atlantica (Canary Isl.); I. Mesalina brevirostris (Middle East); J. Ichnotropis bivittata (Angola); K. Nucras scalaris (Angola); L. Nucras taeniolata (SW Africa)

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Figure 24. SCINCIDAE. A. Leiolopisma coventryi (S Australia); B. Emoia nativitatis (Christmas Is., W Australia); C. Egernia striolata (E Australia); D. Egernia coventryi (S Australia); E. inornatus (N Australia); F. C. helenae (W Central Australia); G. Ctenotus dux (Central Austra- lia); H. C. allotropis (SE Australia); I. Mabuya dorsivittata (Cordoba, Argentina); J. Mabuya frenata (Brazil); K. Mabuya aurata (Lebanon, Syria); L. Mabuya maculilabris (Ghana, W Africa) Figura 24. SCINCIDAE. A. Leiolopisma coventryi (S Australia); B. Emoia nativitatis (Christmas Is., W Aus- tralia); C. Egernia striolata (E Australia); D. Egernia coventryi (S Australia); E. Ctenotus inornatus (N Australia); F. C. helenae (W Central Australia); G. Ctenotus dux (Central Australia); H. C. allotropis (SE Australia); I. Mabuya dorsivittata (Cordoba, Argentina); J. Mabuya frenata (Brazil); K. Mabuya aurata (Lebanon, Syria); L. Mabuya maculilabris (Ghana, W Africa)

MULTEQUINA 16: 1-52, 2007 37 continent sound suggestive. The relation as regards (regarding) the sixteen species and eight genera Scincomorpha branch of represented may be a sufficient Autarchoglossa. The general sample, given our multiple controls homogeneity and shape uniformity carried out on a remarkable number of supraorbital scales, before of additional specimens. In the well revised, are again almost the same known genus Acanthodactylus 3-2 in Scincoidea, such as an authentic broad supraocular scales are the “Autarchoglossa band”. rule, without apparent In all subfamilies of Scincidae circumorbital semicircles, (Greer, 1970) the supraocular pronounced supraciliaries being pattern exposed in Figures 24-25 instead separated or not from reveals no noticeable variations for supraoculars by a narrow row of either the 6, 5, 4, 3 broad diminutive scales. Comparing this longitudinal scales, or the reduced pattern with the other drawn supraciliaries known for supraocular patterns of Lacertidae, Leiolapisma, Emoia, Egernia, such as Psammodromus, Eremias, Ctenotus from Australia, Mabuya Lacerta, Mesalina, Ichnotropis, from middle East Africa and South Nucras or Gallotia, their America, Eumeces taeniolatus fundamental trends of similarity from Pakistan, Eumeces obsoletus are recognizable at first sight. from Kansas, USA, scincus However, a diligent comparison of from Irak, or Scincus mitranus from the supraocular lepidosis in Saudi Arabia. Lacertidae with the above reported supraocular lepidosis in Teiioidea The scarce exceptions, (Figures 19, 20 and 21) may point registered in Figure 25, were drawn out the independent systematic for (Angola), Chalcides identity of both these major taxa, (South Europe, Canary Islands), surely in accordance with different Malaconthas holomelas, evolutionary lines, in spite of any Typhlosaurus lineatus (S. Africa), indubitable morphological Tiliqua occipitalis (Australia), convergence of their supraocular Ablepharus kitaibelii (SE Europe). scutellar patterns. All the evolutionary diversifications (scale reduction, It is now the turn of our shape, etc.) likely originated from comments on supraocular lepidosis some primitive common strain in Scincoidea families, Scincidae under adaptive environmental push and Cordylidae - , (ecological, climate changes, thus completing our analytical fossorial life, etc.).

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IJKL Figure 25. SCINCIDAE. A. Eumeces taeniolatus (Pakistan); B. Eumeces obsoletus (Kansas, USA); C. (Irak, Arabia); D. Scincus mitranus (Saudi Arabia); E. Sepsina copei (Angola, W Africa); F. Sepsina bayonii (Angola, W Africa); G. Chalcides chalcides (S Italy); H. Chalcides viridens (Canary Isl.); I. Malacontias holomelas (Madagascar); J. Typhlosaurus lineatus (S Africa); K. Tiliqua occipitalis (Australia); L. Ablepharus kitaibelii (SE Europe, Turkey) Figura 25. SCINCIDAE. A. Eumeces taeniolatus (Pakistan); B. Eumeces obsoletus (Kansas, USA); C. Scincus scincus (Irak, Arabia); D. Scincus mitranus (Saudi Arabia); E. Sepsina copei (Angola, W Africa); F. Sepsina bayonii (Angola, W Africa); G. Chalcides chalcides (S Italy); H. Chalcides viridens (Canary Isl.); I. Malacontias holomelas (Madagascar); J. Typhlosaurus lineatus (S Africa); K. Tiliqua occipitalis (Australia); L. Ablepharus kitaibelii (SE Europe, Turkey)

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EF GH

IJ KL

Figure 26. CORDYLIDAE. A. Cordylus vittifer (Angola); B. Cordylus giganteus (S Africa) ; C. Cordylus polyzonus (S Africa); D. Pseudocordylus microlepidotus (Cape Prov., S Africa); E. Platysaurus guttatus (S Africa); F. Platysaurus capensis (S Africa); G. Chamaesaura aenea (Transvaal, S Africa); H. Chamaesaura anguina (Cape Prov., S Africa). GERRHOSAURIDAE. I. Gerrhosaurus major (S Africa); J. G. nigrolineatus (S Africa); K. Zonosaurus maximus (Madagascar); L. Tetradactylus tetradactylus (S Africa) Figura 26. CORDYLIDAE. A. Cordylus vittifer (Angola); B. Cordylus giganteus (S Africa); C. Cordylus polyzonus (S Africa); D. Pseudocordylus microlepidotus (Cape Prov., S Africa); E. Platysaurus guttatus (S Africa); F. Platysaurus capensis (S Africa); G. Chamaesaura aenea (Transvaal, S Africa); H. Chamaesaura anguina (Cape Prov., S Africa). GERRHOSAURIDAE. I. Gerrhosaurus major (S Africa); J. G. nigrolineatus (S Africa); K. Zonosaurus maximus (Madagascar); L. Tetradactylus tetradactylus (S Africa)

40 José M. Cei Cordylidae and Gerrhosauridae, reporting before for scincids as some considered as separate families in Eumeces, Scincus or Chalcides, all recent classifications, are lizards lizards showing a tendency to living in African and Madagascar minimization of legs and to having territories. Exemplified in Figure snake-like features. Of course, some 26, we remark the impressive adaptive Anguid characters are similarity between their recognizable in drawings of Figure supraocular lepidosis and the above 27, as the strengthening of analysed scincid pattern. The supraciliaries or the 7, 6, 5, 4 broad supraciliary borders of these supraocular scales. A snake-like Scincomorpha are wider and more form is besides visible in the Euro- developed than supraciliaries of Asiatic Anguis, in Ophisaurus Scincidae. In the large, armoured gracilis or O. apodus from SE Asia, Cordylus species, such as C. in the several Diploglossus and giganteus, C. cataphractus, C. Ophiodes from South America and warreni, C. polyzonus, the West Indies. supraocular region is subject to In Estes et al.’s indented sourrounding quills, horns, sharp- taxonomy, Xenosauridae, points of a strong dermal sclerosis. Anguidae and Varanoidea are The broad longitudinal scales of so included under the taxon peculiar reptiles differ from 6 to 4 Anguimorpha. But, as these authors in number (mostly 4). An analogous stated, “the Xenosauridae and remark can be made for other genera Anguidae cannot be demonstrated of Cordylidae and Gerrhosauridae, to be more closely related to each such as Platysaurus, Gerrhosaurus, other than either is to Varanoids”. Zonosaurus, Tetradactylus or the Therefore, the authors placed unusual snake-like Chamaesaura xenosaurs, anguids and varanoids whose residual legged species in an unresolved triatomy, and they exhibit deeply keeled scales. mentioned that a metataxon To all appearance the “scincoid” Anguioidea wants sufficient reason model of supraocular scutellation is for being formally recognized. a prevalent one in the Scleroglossa The impoverished family taxon. Indeed, if we examine a Xenosauridae adds to its strange, sample of Squamata Anguimorpha, disjoint distribution (China, the great family Anguidae, i.e., with Mexico) some peculiarities of its its more than 100 species and 13 supraocular scutellation, being thus genera, we get back to the kind of in accordance with the sentence supraocular pattern we were not long ago stated here on the

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E F G H

IJKL Figure 27. ANGUIDAE. A. Anguis fragilis (SE Europe); B. A. fragilis (N Europe); C. Ophisaurus gracilis (Hymalayas, Burma); D. Elgaria coerulea (Pacific coast USA, Costa Rica); E. Diploglossus lessonae (NE Brazil); F. D. hewardii (Jamaica, West Indies); G. D. monotropis (Costa Rica); H. D. warreni (Hispaniola, West Indies); I. Ophiodes yacupoi (Misiones, Argentina); J. O. striatus (S. Paulo, Brazil); K. Ophiodes intermedius (Entre Ríos, Argentina); L. O. vertebralis (Buenos Aires, Argentina) Figura 27. ANGUIDAE. A. Anguis fragilis (SE Europe); B. A. fragilis (N Europe); C. Ophisaurus gracilis (Hymalayas, Burma); D. Elgaria coerulea (Pacific coast USA, Costa Rica); E. Diploglossus lessonae (NE Brasil); F. D. hewardii (Jamaica, West Indies); G. D. monotropis (Costa Rica); H. D. warreni (Hispaniola, West Indies); I. Ophiodes yacupoi (Misiones, Argentina); J. O. striatus (S. Paulo, Brasil); K. Ophiodes intermedius (Entre Ríos, Argentina); L. O. vertebralis (Buenos Aires, Argentina)

42 José M. Cei precariousness of systematic covered by dense, irregular small categories such as Anguimorpha scales, without circumorbital or Anguioidea. In fact, the Chinese semicircles but bordered by monotypic Shinisaurus (S. overlapping ciliaries. Somewhat crocodilurus, Figure 28) displays a similar is the pattern of supraocular bare covering with suspectum and H. horridum, the particular circumorbital semicircles robust venomous lizards from sandy and supraciliaries, round a American deserts, even though size somewhat eccentric relief more and distribution of their prominent related to a Chinese decoration than round scales are distinct. In Figure to cephalic lepidosis. The few 28, inferior row, the supraocular Xenosaurus species from Mexico, covering of the African Varanus on the contrary, exhibit a very niloticus and V. exanthematicus is regular Squamata supraocular drawn, together with that of the pattern with some Pleurodontid South Asiatic Varanus monitor. The reminiscences, affording complete, uncertain boundaries of reduced evident circumorbital semicircles, supraocular semicircles, the absence opposed to moderate supraciliaries of broad scales and the very narrow and, within their boundaries, a ciliaries of the African forms can be longitudinal row of broader scales, pointed out, contrasting with the 4-5 in number, having some rows more regular small supraocular of irregular smaller scales on both scales of Varanus monitor, whose sides. supraciliaries are reduced in the In this same Figure 28, taxa extreme. In Figure 29, a selection of belonging to the heterogeneous supraocular coverings in several Varanoidea assemblage are species from the extensive presented. It is about two small, distribution of the genus is provided, probably primitive families, as from the gigantic Varanus Lanthanotidae with only one species komodensis or the differentiated scales from Borneo, or Helodermatidae of Varanus grayi from Philippines, with two species from Mexico and where circumorbital semicircles are SW United States, besides the identifiable, to the very variable remarkable Varanidae, whose more patterns of Australian taxa, where only than 55 species of the unique genus the minimized scales of supraocular Varanus spread from Africa and SE coverings and the utmost restricted Asia to Australia and New Guinea. space of supraciliaries are common In Lanthanotus borneensis, the characters. supraocular region is uniquely

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Figure 28. XENOSAURIDAE. A. Shinisaurus crocodilurus (China); B. Xenosaurus rackhani (Mexico); C. Xenosaurus penai (Mexico). HELODERMATIDAE. D. Heloderma suspectus (SW USA, N Mexico); E. Heloderma horridum (Tehuantepec, Mexico). LANTHANOTIDAE. F. Lanthanotus borneensis (Borneo). VARANIDAE. G. Varanus niloticus (E Africa); H. V. exanthematicus (S Africa); I. V. monitor (S Asia) Figura 28. XENOSAURIDAE. A. Shinisaurus crocodilurus (China); B. Xenosaurus rackhani (Mexico); C. Xenosaurus penai (Mexico). HELODERMATIDAE. D. Heloderma suspectus (SW USA, N Mexico); E. Heloderma horridum (Tehuantepec, Mexico). LANTHANOTIDAE. F. Lanthanotus borneensis (Borneo). VARANIDAE. G. Varanus niloticus (E Africa); H. V. exanthematicus (S Africa); I. V. monitor (S Asia)

44 José M. Cei A B C

D E F

G H I

Figure 29. VARANIDAE. A. Varanus komodoensis (Komodo Isl. Indonesia); B. Varanus grayi (Philippine Isls.); C. V. nebulorus (Burma, Malasia); D. V. mitchelli (N Australia); E. V. giganteus (N Australia); F. V. gouldii (N Australia); G. V. spenceri (N Australia); H. V. mertensi (SW Austra- lia); I. V. timorensis (Timor Isls., Australasia) Figura 29. VARANIDAE. A. Varanus komodoensis (Komodo Isl. Indonesia); B. Varanus grayi (Philippine Isls.); C. V. nebulorus (Burma, Malasia); D. V. mitchelli (N Australia); E. V. giganteus (N Australia); F. V. gouldii (N Australia); G. V. spenceri (N Australia); H. V. mertensi (SW Aus- tralia); I. V. timorensis (Timor Isls., Australasia)

MULTEQUINA 16: 1-52, 2007 45 DISCUSSION well differentiated supraocular scales in Pleurodontid families as We must apologize for so long, Leiocephalidae, Phrynosomatidae, often monotonous descriptive Polychrotidae and Tropiduridae. reports despite their unavoidable Impressive is the family documentary essence. As first Leiosauridae, whose supraocular sentence, we can now state that a scutellation is unmistakable, of postulated relationship between the course taking generic and specific many morphological trends of variations into account. But even Lacertilian supraocular lepidosis more remarkable is its partition into and a taxonomic evolutionary two subfamilies, Leiosaurinae and system is a reality. That is supported Enyaliinae, proposed by Frost et by drawings of our 27 Figures and al. (2001) and reproduced in the their joined analytical comparative differentiation of supraocular scales comments. We would like to add in their respective taxa. Generic that the chosen indented taxonomy and specific variations occur in from the thought over treatise by other families such as Estes et al. (1988) was a very Hoplocercidae, Corytophanidae, congruent one with the peculiar Iguanidae, Liolaemidae, already features and the systematic with a recognizable Pleurodont distribution of the many taxa brand, however with some examined in this paper. A still more primitive characters as in convincing observation can be Opluridae, and exceptional emphasized here with the divergence perhaps adaptive as in undoubted affinity of our results Crotaphytidae. with the reclassification of the Iguania by Frost et al. (2001). The The great Squamata division into families proposed in such a Iguania and Scleroglossa appears reclassification are well fully in accordance with the individualized by their supraocular evolutionary morphological status scale patterns, thus a first distinction of the supraocular lepidosis in between Acrodonta and lizards. A first separation between Pleurodonta is possible on these Gekkota and Autarchoglossa is also lepidosis characters. The almost justified. In the cosmopolitan uniform supraocular lepidosis of Gekkota stem, the families Agamid or Chamaeleonid taxa, Gekkonidae, Diplodactylidae, with their more or less rounded or Eublepharidae are distinguished by squared moderate scales, is in fact a variable expression of their at first sight dissimilar from the scantily differentiated supraocular

46 José A. Scolaro, F. Videla , S. Puig y A. Marcus lepidosis, wanting recognizable evidence a clear-cut example of circumorbital semicircles, parallel evolution, with their almost showing moderate supraciliaries similar supraocular lepidosis exceptionally prominent or sharp- patterns illustrated in our Figures pointed, and having a very 19-23. Some minor features attain heterogeneous layer of moderate to highlight the natural roundish scales, frequently characteristics of an African or rarefied. Striking is a comparison European lacertilid as well as those with the joined snake-like family, of a South American Teiid or the Australian Pygopodidae, Gimnophthalmid. whose adaptive modifications of Gymnophthalmidae and the cephalic lepidosis, formerly Teiidae, formerly considered as studied by Kluge (1974), reaffirm microteiids or macroteiids are the importance of supraocular moreover distinguished now at scales as a discriminating family level also by their significant taxonomic character. supraocular morphology. The Autarchoglossa and its peculiar case of the evolutionary subdivisions exhibit the most clear- modification of supraocular scales cut difference from Iguania because until their disappearance in the of the supraocular lepidosis of all Gymnophthalmid genus Bachia has its numerous taxa, either been reported. Scincoidea is still Scincomorpha or Anguimorpha. A more respectful of the mentioned general pattern of some wide scales Autarchoglossa model of (2 to 7) longitudinally disposed supraocular scutellation, either in along the ciliary border is repeated its cosmopolitan family Scincidae either in Lacertoidea or Scincoidea, or in its related African family as well as in Anguidae. A somewhat Cordylidae, now divided by the discordant supraocular scale creation of the related taxon reduction of the family Xantusiidae Gerrhosauridae. With unavoidably is unusual and takes us back to its distinct morphological features, probably uncertain taxonomic also Anguidae, South American and position invoked in a past careful Euro-Asiatic lizards appear not to Savage’s research (1963). deviate from the just commented The two major branches of morphological expression of Lacertiformes, Lacertidae from supraocular lepidosis in Scincoidea. Africa and Eurasia, Teiioidea from But the other taxon placed in the American continents, put in Anguimorpha in our chosen indented taxonomy, the family

MULTEQUINA 16: 1-52, 2007 47 Xenosauridae, is the rarest finding morphological or biological in the whole Scleroglossa systems that are the subject of embranchment, either for its research. Thus, in a synthetic disjoint distribution (China, sequence, the first conclusion of Mexico) or for the unusual the present study is that the supraocular scales. The probable supraocular lepidosis in the artificiality of the taxon Squamata taxa is not a structure at Anguimorpha was considered by random, but a constant, defined the same authors of the indented scale arrangement, different in taxonomy. shape and size, in accordance with We reach the end of a simple, the species or other taxa considered objective discussion of data from and their evolutionary systematic our patient comparative checking position. Second conclusion is the of hundreds of Squamata lizards. significant agreement observed The last scanty families between a classificatory system, as Helodermatidae and Lanthanotidae the indented taxonomy elicited by show a quite uniform, likely Estes et al. (1988), and the results primitive, supraocular lepidosis, of our comparative research, with roundish scales, evident leading to establishing defined supraciliaries but indistinct supraocular lepidosis patterns as circumorbital semicircles. The comparable morphological extensive family Varanidae, characters of every taxon. Third unknown in the American conclusion is that a reassessment continents, assembles a very of the Iguanian Pleurodonta taxon variable set of supraocular of Estes et al.’s indented taxonomy, coverings, from scales distinctly as in the reclassification proposed arranged to almost naked surfaces, by Frost et al. (2001), is also in mottled or variegated. A typical agreement with morphological Varanid supraocular pattern is thus results on the supraocular indefinable. scutellation of Squamata, shown in our comparative review.

CONCLUSIONS A final conclusive remark is the repeated application of the Conclusions of a scientific paper pointed out relationships between are the summarized exposition of indented phylogenetic taxonomy every confirmed novelty or and differentiated supraocular improvement concerning the lepidosis, such as in the case of original status or condition of several systematic new proposals.

48 José M. Cei The new Pleurodont family NOTE: This manuscript was still on his Leiosauridae and its two desk when unfortunately Dr. Cei died. He subfamilies must be here was internationally recognized and his remembered, according to Frost et work will remain indelible for future al. (2001). Other significant generations of herpetologists, because of systematic agreements in relation his profuse and invaluable scientific to supraocular lepidosis can be cited contribution and proposal of new ideas for the families Scincoidea, for the biological sciences and evolution. Scincidae and Cordylidae and for the families Teiioidea, Teiidae and REFERENCES Gymnophthalmidae (macroteiids and microteiids). ALBINO, A.M., S. BRIZUELA & C.I. MONTALVO, 2006. New Tupinambis remains from the ACKNOWLEDGEMENTS late Miocene of Argentina and For their valuable aid in the a review of the South selection of documentary American Miocene Teiids. J. Herpetol. 40(2): 206-213. specimens I thank very much the ANDERSON, S.T., 1999. The friendly support of Kraig Adler Lizards of Iran. SSAR (Dep. Neurobiology, Cornell Contributions to Herpetology, University, Ithaca, N.Y., USA), and Vol. 15. 442 pp. of William E. Duelman (Nat. Hist. ANGEL, F., 1942. Les Lézards de Mus. Univ, of Kansas, Lawrence, Madagascar. Mem. Acad. Kansas, USA). My acknowledgement Malgache 36: 1-193. must be here emphasized for the heavy BLARC, Ch.P., 1977. Reptiles and careful work of Valeria Corbalán Sauriens Iguanidae. Faune de (IADIZA-CONICET) in the editorial Madagascar. ORSTOM - phase of my paper, given the CNRS, París, 45. 195 pp. unfortunate conditions of my personal BRYGOO, E.R., 1971. Reptiles activity during this period. Sauriens Chamaelonidae, genre Chamaeleo. Faune de His wife Giudita and his disciples J. Madagascar. ORSTOM - Alejandro Scolaro and Fernando CNRS, París, 33. 318 pp. Videla considered the publication of BRYGOO, E.R., 1978. Reptiles this paper to be a very important Sauriens Chamaelonidae, contribution to science, and genre Brookesia et acknowledge the editorial revision complements pour la genre made by Nelly Horak, Silvia Puig Chamaeleo. Faune de Madagascar. ORSTOM - and Nora Ibarguengoytía. CNRS, París, 47. 173 pp.

MULTEQUINA 16: 1-52, 2007 49 CEI, J.M., 2003. Specific supraocular ESTES, R., K. DE QUEIROZ & J. scutellation patterns as GAUTHIER, 1988. Phyloge- significant diagnostic characters: netic relationships within a taxonomic inter and Squamata. Pp. 119-281, In: intrageneric “Finger-Print” in Estes R. and G. Pregill (Eds.), Lacertilia. Facena 19: 155- Phylogenetic relationships of 161. the families. Essays com- CEI, J.M., J.A. SCOLARO & F. memorating Charles L. Camp. VIDELA, 2003. A taxonomic Stanford University Press, revision of recognized Argentine California, USA, 631 Pp. species of the Leiosaurid genus ETHERIDGE, R., 1970. A review Diplolaemus (Reptilia, of the South American iguanid Squamata, Leiosauridae). genus Plica. Bull. Br. Mus. Facena 19: 87-106. Nat. Hist. (Zoology) 19: 237- CEI, J.M., J.A. SCOLARO & F. 256. VIDELA, 2004. An updated ETHERIDGE, R., 1995. Redescrip- biosystematic approach to the tion of Ctenoblepharis Leiosaurid genus Pristidactylus. adspersa Tschudi, 1845, and Boll. Mus. Reg. Scienze the taxonomy of Liolaeminae Naturali, Torino (Italy) 21(1): (Reptilia, Squamata, 159-192. Tropiduridae). Amer. Mus. DIXON, J.R., 1973. A systematic Novitates 3142: 24 pp. review of the Teiid lizards, ETHERIDGE, R. & K. DE genus Bachia, with remarks QUEIROZ, 1988. Phylogeny on Heterodactylus and of Iguanidae. Pp. 283-367, In: Anotosaura. Miscellaneous Estes, R and G. Pregill (Eds.), Publications 57, Mus. Nat. Phylogenetic relationships of Hist., Univ. of Kansas. 47 pp. the lizard families. Essays com- DONNELLY, M.A. & C.W. memorating Charles L. Camp. MYERS, 1991. Herpetologi- Stanford University Press, cal results of the 1990 Ven- California, USA, 631 Pp. ezuelan expedition to the sum- FROST, D.R., 1992. Phylogenetic mit of Cerro Guaiquirima, with analysis and taxonomy of the new tepui reptiles. American Tropidurus group of Lizards Mus. Novitates 3017. 54 pp. (Iguania, Tropiduridae). Amer. Mus. Novitates 3033: 1-68. FROST, D.R. & R. ETHERIDGE, 1989. A phylogenetic analysis and taxonomy of Iguania Lizards (Reptilia, Squamata). Misc. Publ. Mus. Nat. Hist. Univ. Kansas USA 81: 1-65. 50 José M. Cei FROST, D.R., A.G. KLUGE & D.M. HOOGMOED, M.S., 1973. Notes HILLIS, 1992. Species in on the Herpetofauna of contemporary herpetology: Surinam. IV. The Lizards and comments on phylogenetic Amphisbaenians of Surinam. inferences and taxonomy. W. Junk, The Hague. 417 pp. Herpetological Review 23(2): KLUGE, A.G., 1974. A taxonomic 46-54. revision of the lizards family FROST, D.R., R. ETHERIDGE, D. Pygopodidae. Miscell. JANIES & T.A. TITOS, 2001. Publications, Mus. Zoology, Total evidence, sequence University of Michigan, 1-227 pp. alignment, evolution of LEVITON, A.E., S.C. ANDERSON, Polychrotid Lizards, and a K. ADLER & S.A. MINTON, reclassification of the Iguania 1992. Handbook to middle east (Squamata, Iguania). Amer. amphibians and reptiles. SSAR Mus. Novitates 3344: 1-38. Contributions to Herpetology GREER, A.E., 1970. A subfamilial 8. Oxford, Ohio. 252 pp. classification of Scincid MERTENS, R., 1958. Bemerkungen lizards. Bull. Mus. Comp. Zool. über die Varane Australiens. 139(3): 151-184. Senck Biol. 39(5-6): 229-264. GRISMER, L.L., J.A. McGUIRE & MERTENS, R., 1959. Liste der B.D. HOLLINGSWORTH, Varane Asiens under der Indo- 1994. A report on the australischen Inselwelt mit Herpetofauna of the Vizcaino systematischen Peninsula, Baja California, Bemerkungen. Senck Biol. Mexico, with a Discussion of 40(5-6): 221-240. its biogeographic and PETERS, J.A., 1964. Dictionary of taxonomic implications. Bull. Herpetology. Hafner Publ. Co. Southern California Acad. Sci. New York. 392 pp. 93(2): 45-80. PREGILL, G.K., 1992. Systematics of HAAS, G. & Y. L. WERNER, 1969. the West Indian lizard genus Lizards and from Leiocephalus (Squamata, Southwestern Asia, collected Iguania, Tropiduridae). by Henry Field. Bull. Mus. Miscellaneous Publ. 84. Mus. Comp. Zool. Harvard Univ. Nat. Hist. Univ. of Kansas. 69 pp. 138(6): 327-406. RUIBAL, R., 1964. An annotated HALLIDAY, T. & K. ADLER, 2002. checklist and key to the The Firefly encyclopedia of Anoline lizards of Cuba. Bull. reptiles and amphibians. of the Museum Comparative Firefly Books Ltd., Ontario, Zoology 130 (8) : 476-520. Canada: 240 pp.

MULTEQUINA 16: 1-52, 2007 51 SAVAGE, J.M., 1963. Studies on the VANZOLINI, P.E. & J. VALENCIA, lizard family Xantusiidae IV. 1965. The genus Dracaena, The genera. Contrib. Sci. .Los with a brief consideration of Angeles County Mus. 71: 1-38. macroteiid relationships SCHULTE II, J.A., J.P. (Sauria, Teiidae). Arquivos de VALLADARES & A. Zoologia, 13: 1-35. LARSON, 2003. Phylogenetic VANZOLINI, P., A. M. RAMOS- relationships within Iguanidae COSTA & L.J. VITT., 1980. inferred using molecular and Repteis das Caatingas. Academia morphological data and a Brasileira de Ciencias, Río de phylogenetic taxonomy of Janeiro. 161 pp. iguanian lizards. Herpetologica WIENS, J.J., 1993. Phylogenetic 59: 399-419. systematics of the tree lizards TAYLOR, E.H., 1956. A review of (genus Urosaurus). lizards of Costa Rica. Univ. Herpetologica 49(4): 399-420. Kansas Sci. Bull. 38: 1-320. VAN DEMBURG, J., 1978. Recibido: 04/2007 Herpetology of lower California : Collected Papers. Aceptado: 07/2007 Facsimile reprints in herpetology. SSAR. 85 pp.

52 José M. Cei