HPH 10A • Sauria terrestria, Amphisbaenia

Foreword V
Dedication VII
Table of Contents XVII
Acknowledgements IX
Institutional Abbreviations XI
Abbreviations used in Figures XV
Historical Overview 1
Dating the Fossils 3
Osteology 6
The Orders of the Squamata 8
Superorder Squamata OPPEL 1811 9
Order Sauria MCCARTNEY 1802 11
Suborder Eolacertilia ROBINSON 1967a 12
Family Paliguanidae BROOM 1903 13
Paliguana BROOM 1903 13
Palaeagama BROOM 1926 13
Saurosternon HUXLEY 1868 15
Family Kuehneosauridae ROBINSON 1967a 15
Kuehneosaurus ROBINSON 1967a 16
Cteniogenys GILMORE 1928 16
Icarosaurus COLBERT 1966 18
Kuehneosuchus ROBINSON 1967b 19
Family Fulengidae CARROLL and GALTON 1977 19
Fulengia CARROLL and GALTON 1977 19
Eolacertilia incertae sedis 20
Lacertulus CARROLL 1981 20
Litakis ESTES 1964 20
Suborder Lacertilia OWEN 1842 21
Infraorder Iguania CUVIER 1807 23
Family Iguanidae GRAY 1827 23
Iguana LAURENTI 1768 23
Aciprion COPE 1873c 24
Anolis DAUDIN 1802 25
Basiliscus LAURENTI 1766 29
Crotaphytus HOLBROOK 1843 29
Ctenosaura WIEGMANN 1828 31
Cyclura HARLAN 1824 31
Cypressaurus HOLMAN 1972 32
Dipsosaurus HALLOWELL 1854 32
Erichosaurus AMEGHINO 1899 32
Geiseltaliellus KUHN 1944 33
Harrisonsaurus HOLMAN 1981 34
Holbrookia GIRARD 1851 34
Laemanctus WIEGMANN 1834 35
Leiocephalus GRAY 1827 35
Leiosaurus DUMÉRIL and BIBRON 1837 39
Oplurus DUMÉRIL and BIBRON 1837 39
Paradiplosaurus FRIES, HIBBARD and DUNKLE 1955 39
Parasauromalus GILMORE 1928 40
Phrynosoma WIEGMANN 1828 42
Pristiguana ESTES and PRICE 1973 44
Sauromalus DUMÉRIL 1856 45
SceloporusWlEGMANN 1828 45
Swainiguanoides SULLIVAN 1982 48
Urosaurus HALLOWELL 1854 48
Uta BAIRD and GIRARD 1852 48
Iguanidae incertae sedis 49
Family Euposauridae CAMP 1923 49
Euposaurus JOURDAN 1862 49
Family Arretosauridae GILMORE 1943a 50
Arretosaurus GILMORE 1943a 50
Family Agamidae GRAY 1827 52
Agama DAUDIN 1802 52
Clamydosaurus GRAY 1825 53
Mimeosaurus GILMORE 1943a 53
Stellio GRAY 1845 53
Tinosaurus MARSH 1872 54
Uromastyx MERREM 1820 56
Agamidae incertae sedis 56
Family Chamaeleonidae GRAY 1825 56
Chamaeleo LINNAEUS 1759 56
Anquingosaurus HOU 1976 56
Chamaeleonidae incertae sedis 57
Infraorder Gekkota CUVIER 1807 59
Family Ardeosauridae CAMP 1923 60
Ardeosaurus MEYER 1860 61
Eichstaettisaurus KUHN 1958 62
Yabeinosaurus ENDO and SHIKAMA 1942 63
Family Bavarisauridae KUHN 1961 65
Bavarisaurus HOFFSTETTER 1953 65
Palaeolacerta COCUDE-MICHEL 1961 67
Family Gekkonidae GRAY 1825 67
Gekko LAURENTI 1768 67
Aristelliger COPE 1861 68
Cadurcogekko HOFFSTETTER 1946 68
Coleonyx GRAY 1845 68
Cyrtodactylus GRAY 1827 70
Geckolepis GRANDIDIER 1867 70
Gerandogekko HOFFSTETTER 1946a 70
Hemidactylus CUVIER 1817 70
Lygodactylus GRAY 1864 71
Paroedura GÜNTHER 1879 71
Phelsuma GRAY 1825 71
Phyllodactylus GRAY 1828 72
Rhodanogekko HOFFSTETTER 1946a 72
Sphaerodactylus WAGLER 1830 72
Tarentola GRAY 1825 73
Thecadactylus GOLDFUSS 1820 73
Gekkonidae incertae sedis 73
Family Pygopodidae GRAY 1845 73
Infraorder Scincomorpha CAMP 1923 74
Superfamily Lacertoidea FITZINGER 1826 74
Family Teiidae GRAY 1827 74
Subfamily Polyglyphanodontinae GILMORE 1942b, n.comb. 75
Polyglyphanodon GILMORE 1940 76
Adamisaurus SULIMSKI 1972 78
Cherminsaurus SULIMSKl 1975 79
Darchansaurus SULIMSKI 1975 80
Erdenetesaurus SULIMSK1 1975 80
Haptosphenus ESTES 1964 81
Macroephalosaurus GILMORE 1943a 83
Paraglyphanodon GILMORE 1940 85
Subfamily Teiinae MACLEAN 1974 87
Tribe Teiini PRESCH 1974a 87
Ameiva MEYER 1795 87
Cnemidophorus WAGLER 1830 88
Dicrodon DUMÉRIL and BIBRON 1839 91
Leptochamops ESTES 1964 91
Meniscognathus ESTES 1964 91
Peneteius ESTES 1969a 92
Tribe Tupinambini PRESCH 1974a 92
Tupinambis DAUDIN 1802 92
Callopistes GRAVENHORST 1838 93
Chamops MARSH 1872 95
Diasemosaurus AMEGHINO 1898 96
Dracaena DAUDIN 1802 96
Teiidae incertae sedis 97
Family Gymnophthalmidae MACLEAN 1974, n.comb. 97
Family Lacertidae BONAPARTE 1831 97
Lacerta LINNAEUS 1758 98
Dracaenosaurus POMEL 1846 100
Eolacerta NÖTH 1940 102
Eremias WIEGMANN 1834 102
Nucras GRAY 1838 103
Plesiolacerta HOFFSTETTER 1942a 103
Pseudeumeces HOFFSTETTER 1944 104
Lacertidae incertae sedis 104
Superfamily Scincoidea OPPEL 1811 105
Family Scincidae GRAY 1825 105
Subfamily Scincinae GRAY 1825 105
Chalcides LINNAEUS 1758 105
Contogenys ESTES 1969c 105
Eumeces WIEGMANN 1834 106
Gongylomorphus FITZINGER 1843 109
Paracontogenys SCHATZINGER in GOLZ and LILLEGRAVEN 1977 111
"Scelotes" sp. 111
Subfamily Lygosominae MITTLEMAN 1952 111
Ablepharus LICHTENSTEIN 1823 111
Egernia GRAY 1848 112
Leiolopisma DUMÉRIL and BIBRON 1839 112
Mabuya FITZINGER 1926 112
Trachydosdurus GRAY 1825 113
Subfamily Feylininae CAMP 1923 113
Subfamily Acontinae GRAY 1845 114
Scincidae ? incertae sedis 114
Sauriscus ESTES 1964 114
Family Dibamidae BOULENGER 1884 114
Scincomorpha incertae sedis 114
Superfamily Cordyloidea FITZINGER 1826 115
Family Paramacellodidae nov.fam. 115
Paramacellodus HOFFSTETTER 1967b 115
Becklesius nov.gen. 116
Pseudosaurillus HOFFSTETTER 1967b 118
Saurillodon nov.gen. 118
Saurillus OWEN 1854 121
Family Xantusiidae BAIRD 1859 121
Xantusia BAIRD 1859 123
Lepidophyma DUMÉRIL 1851 123
Palaeoxantusia HECHT 1956 123
Family Cordylidae GRAY 1837 126
Gerrhosaurus WIEGMANN 1828 126
Pseudolacerta DE STEFANO 1903 126
Infraorder Anguimorpha FÜRBRINGER 1900 129
Superfamily Anguioidea FITZINGER 1826 129
Family Xenosauridae COPE 1886 129
Exostinus COPE 1873c 130
Restes GAUTHlER 1982 132
Family Dorsetisauridae HOFFSTETTER 1967b 132
Dorsetisaurus HOFFSTETTER 1967b 132
Family Anguidae GRAY 1825 134
Subfamily Anguinae GRAY 1825 134
Anguis LINNAEUS 1758 134
Machaerosaurus GILMORE 1928 135
Ophipseudopus KUHN 1940a 136
Ophisauriscus KUHN 1940a 137
Ophisaurus DAUDIN 1803 137
Anguinae incertae sedis 144
Subfamily Anniellinae NOPCSA 1928 144
Anniella GRAY 1852 144
Apodosauriscus GAUTHlER 1982 144
Subfamily Glyptosaurinae MARSH 1872 144
Tribe Melanosaurini SULLIVAN 1979a 145
Melanosaurus GILMORE 1928 146
Arpadosaurus MESZOELY 1970 146
Odaxosaurus GILMORE 1928 147
Peltosaurus COPE 1872 149
Proxestops GAUTHIER 1981 151
Xestops COPE 1873c 151
Tribe Glyptosaurini SULLIVAN 1979a 153
Glyptosaurus MARSH 1871a 153
Eoglyptosaurus SULLIVAN 1979a 155
Helodermoides DOUGLASS 1903 155
Paraglyptosaurus SULLIVAN 1979a 156
Placosaurus GERVAIS 1848-1852 158
Subfamily Gerrhonotinae MCDOWELL and BOGERT 1954 163
Gerrhonotus WIEGMANN 1828 163
Paragerrhonotus ESTES 1963a 165
Gerrhonotinae incertae sedis 165
Subfamily Diploglossinae BOCOURT 1873-1879 165
Diploglossus WIEGMANN 1834 165
Celestus GRAY 1838 166
Eodiploglossus GAUTHIER 1982 166
Superfamily Varanoidea CAMP 1923 168
Family Necrosauridae HOFFSTETTER 1943 168
Necrosaurus FILHOL 1876 169
Colpodontosaurus ESTES 1964 171
Eosaniwa HAUBOLD 1977 172
Parasaniwa GILMORE 1928 172
Provaranosaurus GILMORE 1942a 174
Family Helodermatidae GRAY 1837 174
Heloderma WIEGMANN 1829 175
Eurheloderma HOFFSTETTER 1957 177
Paraderma ESTES 1964 177
Family Varanidae GRAY 1827 179
Varanus MERREM 1820 179
Iberovaranus HOFFSTETTER 1969 182
Megalania OWEN 1860 182
Palaeosaniwa GILMORE 1928 183
Saniwa LEIDY 1870 184
Telmasaurus GILMORE 1943a 187
Varanidae incertae sedis 189
Lacertilia incertae sedis 189
Anhuisaurus HOU 1974 189
Changjiangosaurus HOU 1976 189
Conicodontosaurus GILMORE 1943a 189
Costasaurus nov.gen. 190
Cremastosaurus COPE 1873c 190
Diacium COPE 1873c 190
Dibolosodon AMEGHINO 1898 191
Dicarlesia HUENE 1931 191
Durotrigia HOFFSTETTER 1967b 191
Haplodontosaurus GILMORE 1942a 191
Iguanavus MARSH 1872 191
IlaerdesaurusHOFFSTETTER 1966a 192
Isodontosaurus GILMORE 1943a 192
"Lacerta" ambigua LARTET 1851 192
"Lacerta" castellanensis DE GREGORIO 1925 192
"Lacerta" philippiana LARTET 1851 192
"Lacerta" ponsortiana LARTET 1851 192
"Lacerta" pulla MEYER 1859 192
"Lacerta" rottensis MEYER 1856 193
Lisboasaurus SEIFFERT 1973 193
Meyasaurus VIDAL 1915 193
Paruprionosaurus GILMORE 1938a 194
Qianshanosaurus HOU 1974 194
Sauromorus POMEL 1953 194
Teilhardosaurus SHIKAMA 1947 195
Sauria? incertae sedis 195
Order Amphisbaenia GRAY 1844 196
Family Oligodontosauridae ESTES 1975 196
Oligodontosaurus GILMORE 1942a 196
Family Amphisbaenidae GRAY 1865 197
Omoiotyphlops DE ROCHEBRUNE 1884 197
Family Rhineuridae VANZOLINI 1951 197
Rhineura COPE 1861 197
Dyticonastis BERMAN 1976 200
Jepsibaena VANZOLINI 1951 201
Lestophis MARSH 1885 202
Macrorhineura MACDONALD 1970 202
Ototriton LOOMIS 1919 202
PseudorhineuraVANZOLINI 1951 205
Spathorhynchus BERMAN 1973 206
Rhineuridae incertae sedis 206
Family Hyporhinidae BAUR 1893 206
Hyporhina BAUR 1893 207
Family Bipedidae TAYLOR 1951 207
Family Trogonophidae GRAY 1865 207
Amphisbaenia incertae sedis 208
Taxa removed from the Squamata 209
Changisaurus microrhinus YOUNG 1959 209
Changlosaurus wutuensis YOUNG 1961 209
Chilingosaurus chingshankouensis YOUNG 1961 209
Colubrifer campi CARROLL 1982 209
Crythiosaurus mongoliensis GILMORE 1943a 209
Cuttysarkus mcnallyi ESTES 1964 209
Diacium unipedale COPE 1873c 209
Enigmatosaurus botti DE STEFANO 1903 210
Habrosaurus dilatus GILMORE 1928 210
"Lacerta" eocena OWEN 1884 210
Lacertulus bipes CARROLL and THOMPSON 1982 210
Macellodus brodiei OWEN 1854 210
Naocephalus porrectus COPE 1872 210
Neorotodactylus leonardii CASAMIQUELA 1980 210
Nordenosaurus magnus HOLMAN 1973b 210
Protolacerta patagonica CASAMIQUELA 1975 211
Rajaurisaurus estesi SAHNI and SRIVASTAVA 1976 211
Saurophis crassus BRUNNER 1957 211
Tylosteus ornatus LEIDY 1872b 211
Varanus lemoinei NOPCSA 1908 211
Familial, Generic and Specific Synonyms 212
Geological and Geographical Distribution of Lizards and Amphisbaenians 214
Literature Cited 231
Index Gernerum 247

Although the fossil record of many groups of fossil reptiles and amphibians has been summarized, not only in this series but in other studies as well, such summaries have often been more complete for extinct groups than for those whose close relatives live today. Among the latter, the lizards are notable in having a widely scattered literature on fossil forms. While lists of fossil lizards have been given by (e.g.) KUHN (1939, 1961b, 1963b) and ROMER (1956) no detailed critical revision of the known material has been made, although the general summaries by HOFFSTETTER (1955, 1962c) have been extremely valuable.

Some explanation of my approach is in order. When originally asked by Dr. OSKAR KUHN to contribute to this series, I decided that a simple literature review would be insufficient; a more useful approach appeared to be to see as much of the important fossil material that was significant and available, and to evaluate it in as critical a manner as possible. In this way maximum information content for both herpetologists and paleontologists would be obtained. A second decision was to omit from the present volume the Cretaceous marine lizard groups Dolichosauridae, Simoliophidae, Aigialosauridae and Mosasauridae. This omission seemed logical for several reasons. Although often considered as platynotan lizards, these families may actually be convergent on that group. The number of taxa involved is moreover sufficiently large so that including them here would substantially increase the size of the present volume, which has already exceeded the usual limits in this series. In addition, the morphological and ecological problems involved in dealing with these groups invite participation of relevant collaborators, so that volume 10B, Sauria maritima, will appear with the participation of DALE RUSSELL (Ottawa) and JACQUES GAUTHIER (Berkeley).

The present work has been in progress for about fourteen years. I have necessarily made extensive revisions of material that was written in earlier years, and in general, opinions expressed are my most recent evaluations. It has not been possible to be entirely consistent, however, and relevant older literature and many scattered records continue to appear. This is, nevertheless, the most complete list of fossil finds of lizards available, but obviously there are still omissions. So far as current finds are concerned, I believe that there are few not included. Unpublished works have been cited if they have been accepted for publication; some of these have come to me for review, others have been kindly provided by colleagues working with lizard fossils or systematics. Material available to me up through the first half of 1982 is included. In a few cases, I have noted studies in progress to indicate important materials or current workers. I have discussed each genus and species and have tried to give an adequate definition wherever possible. In most cases, the generic definitions include citations for the data given; if these are not cited, then the definition is either by the original author or by me. In all cases, I have given definitions for higher categories, and wherever possible have expressly stated the shared derived characters of the group. Polarities are seldom justified, however, because of space limitations, and generally (if not otherwise cited) are based on outgroup comparison made by me (or by others cited) during the course of this study. These character states and polarities are provisional, however, and are offered mainly as indications of what has been observed or directions that might be taken in future.

The descriptive sections that are included in the discussions of many species are mainly given for those represented rather well in the record, are otherwise especially significant or are unique extinct groups. Little or no description has been included for Holocene genera and species, and original bibliographic citations for these are not included in the References Cited.

So far as taxonomy is concerned, I have included all synonymies for extinct genera and species known to me or that have been suggested by my own work. I have not, however, always included complete synonymies for Holocene genera and species, nor have I given comprehensive lists of the various names applied to family or higher taxonomic units, principally because most of these are living groups and such synonymies are available in the published sections of Das Thierreich or are otherwise available.

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