HPH 3A2 • Temnospondyli I

Editor’s Preface V
Institutional abbreviations VI
Abbreviations used in the figures VIII
Introduction 1
Historical Overview 4
Comparative Anatomy of Temnospondyli 5
Systematic Paleontology 17
Temnospondyli ZITTEL, 1888 17
Temnospondyl classification 17
Edopoidea ROMER, 1945b 18
Edopidae ROMER, 1945b 18
Edops ROMER, 1936 18
Cochleosauridae BROILI, 1923 20
Adamanterpeton MILNER & SEQUEIRA, 1998 21
Chenoprosopus MEHL, 1913 21
Cochleosaurus FRITSCH, 1885a 21
Nigerpeton SIDOR et al., 2005 24
Procochleosaurus SEQUEIRA, 1996 24
Cochleosauridae incertae sedis 24
Eutemnospondyli SCHOCH, 2013 24
Dendrerpetidae FRITSCH, 1885a (as Dendrerpetontidae) 24
Dendrerpeton OWEN, 1853 25
Dendrysekos STEEN, 1934 26
Balanerpeton MILNER & SEQUEIRA, 1994 26
Rhachitomi WATSON, 1919 sensu SCHOCH, 2013 28
Dvinosauria YATES & WARREN, 2000 28
Eugyrinus WATSON, 1921 28
Trimerorhachidae COPE, 1882b 29
Trimerorhachis COPE, 1878b 30
Neldasaurus CHASE, 1965 33
Lafonius BERMAN, 1973 33
Unnamed clade Erpetosaurus + Dvinosauroidea 34
Erpetosaurus MOODIE, 1909b 34
Dvinosauroidea SÄVE-SÖDERBERGH 1935 34
Eobrachyopidae SHISHKIN, 1964 34
Isodectes COPE, 1896c 34
Acroplous HOTTON, 1959 35
Dvinosauroidea incertae sedis 35
Not referable to Dvinosauroidea 35
Dvinosauridae WATSON, 1919 36
Dvinosaurus AMALITZKY, 1921 36
Tupilakosauridae KUHN, 1960 37
Tupilakosaurus NIELSEN, 1954 37
Slaugenhopia OLSON, 1962 39
Batrachosuchoides SHISHKIN, 1966 39
Kourerpeton OLSON & LAMMERS, 1976 39
Thabanchuia WARREN, 1999 40
Tupilakosauridae incertae sedis 40
Dvinosauria incertae sedis 40
Dawsonerpeton ZAJÍC, 1984 40
Dissorophoidea BOLT, 1969 42
Micromelerpetidae BOY, 1972 42
Branchierpeton BOY, 1972b 42
Eimerisaurus BOY, 2002a 43
Limnogyrinus MILNER, 1986 43
Micromelerpeton BULMAN & WHITTARD, 1926 44
Nyranerpeton WERNEBURG, 2012 45
Micromelerpetidae incertae sedis 46
Xerodromes, new taxon 46
Branchiosauridae FRITSCH, 1879 46
Apateon MEYER, 1844 46
Branchiosaurus FRITSCH, 1876 48
Leptorophus BULMAN & WHITTARD, 1926 50
Melanerpeton FRITSCH, 1878 50
Schoenfelderpeton BOY, 1986 52
Branchiosauridae incertae sedis 52
Eumicrerpeton MOODIE, 1910 52
Milnererpeton HUNT et al., 2002 52
Tungussogyrinus EFREMOV, 1939 52
Additional branchiosaurid material 54
Differentiation of larvae within Dissorophoidea 54
Amphibamidae MOODIE, 1910 56
Amphibamus COPE, 1865 56
Doleserpeton BOLT, 1969 58
Eoscopus DALY, 1994 58
Georgenthalia ANDERSON et al., 2008a 60
Gerobatrachus ANDERSON et al., 2008b 60
Micropholis HUXLEY, 1859a 60
Nanobamus, new genus 60
Pasawioops FRÖBISCH & REISZ, 2008 61
Platyrhinops STEEN, 1931 61
Plemmyradytes HUTTENLOCKER et al., 2007 62
Rubeostratilia BOURGET & ANDERSON, 2011 62
Tersomius CASE, 1910 62
Nomina dubia 63
Limnerpeton FRITSCH, 1881 63
Olsoniformes ANDERSON et al., 2008a 64
Trematopidae WILLISTON, 1910a 64
Basal trematopids 64
Ecolsonia VAUGHN, 1969a 64
Mordex STEEN, 1938 64
Tambachia SUMIDA et al., 1998 65
Derived trematopids 66
Acheloma COPE, 1882a 66
Actiobates EATON, 1973 67
Anconastes BERMAN et al., 1987 67
Fedexia BERMAN et al., 2010 67
Phonerpeton DILKES, 1990 67
Rotaryus BERMAN et al., 2011 68
Dissorophidae BOULENGER, 1902 69
Dissorophinae WILLISTON, 1910b 70
Broiliellus WILLISTON, 1914a 70
Dissorophus COPE, 1895 72
Eucacopinae SCHOCH & SUES, 2013 74
Cacops WILLISTON, 1910b 74
Alegeinosaurus CASE, 1911 75
Anakamacops LI & CHENG, 1999 76
Brevidorsum CARROLL, 1964b 76
Conjunctio CARROLL, 1964b 76
Kamacops GUBIN, 1980 76
Parioxys COPE, 1878a 77
Reiszerpeton MADDIN et al., 2013 77
Scapanops SCHOCH & SUES, 2013 77
Zygosaurus EICHWALD, 1848 78
Aspidosaurinae WILLISTON, 1914a 78
Aspidosaurus BROILI, 1904 78
Aspidosaurinae incertae sedis 79
Platyhistricinae, new taxon 79
Platyhistrix WILLISTON, 1911 79
Astreptorhachis VAUGHN, 1971 80
Dissorophidae incertae sedis 80
Fayella OLSON, 1965a 80
Iratusaurus GUBIN, 1980 80
Macrerpeton MOODIE, 1909a 81
Perryella CARLSON, 1987 82
Stegops MOODIE, 1909a 82
Zatracheidae COPE, 1882b 83
Acanthostomatops KUHN, 1961a 83
Dasyceps HUXLEY, 1859b 83
Zatrachys COPE, 1878a 85
Zatracheidae incertae sedis 86
Eryopiformes SCHOCH, 2013 86
Iberospondylus LAURIN & SOLER-GIJÓN, 2001 86
Eryopidae COPE, 1882b 88
Actinodon GAUDRY, 1866 88
Eryopinae, new taxon 88
Eryops COPE, 1878a 89
Glaukerpeton ROMER, 1952 91
Onchiodon GEINITZ, 1862 91
Osteophorus MEYER, 1856 93
Clamorosaurus GUBIN, 1983 94
Eryopidae incertae sedis 94
Stereospondylomorpha YATES & WARREN, 2000 95
Stereospondyli ZITTEL, 1888 95
Stereospondyli incertae sedis 95
Peltobatrachus PANCHEN, 1959 95
Brachyopoidea LYDEKKER, 1885 95
Brachyopidae LYDEKKER, 1885 97
Brachyops OWEN, 1854 97
Banksiops WARREN & MARSICANO 2000a 98
Bathignathus DAMIANI & JEANNOT, 2002 98
Batrachosuchus BROOM, 1903 98
Bothriceps HUXLEY, 1859a 101
Gobiops SHISHKIN, 1991 101
Notobrachyops COSGRIFF, 1973 101
Platycepsion KUHN, 1961a 101
Sinobrachyops DONG, 1985 102
Vanastega DAMIANI & KITCHING, 2003 102
Vigilius WARREN & MARSICANO, 2000b 102
Xenobrachyops WARREN & HUTCHINSON, 1983 103
Indeterminate Brachyopidae 103
Ferganobatrachus NESSOV, 1990 103
Plagiosauridae JAEKEL, 1914 104
Plagiosuchinae, new taxon 104
Plagioscutum SHISHKIN, 1986 104
Plagiosuchus HUENE, 1922 104
Plagiosterninae + Plagiosaurinae 106
Plagiosaurinae JAEKEL, 1914 106
Gerrothorax NILSSON, 1934 106
Plagiosaurus JAEKEL, 1914 106
Plagiosaurine from southern China 108
Plagiosterninae SHISHKIN, 1986 108
Aranetsia NOVIKOV & SHISHKIN, 1992 108
Melanopelta SHISHKIN, 1967 108
Plagiosternum FRAAS, 1896 109
Plagiorophus KONZHUKOVA, 1955b 109
Plagiosauridae incertae sedis 110
Plagiobatrachus WARREN, 1985 110
Bear Island plagiosaurid 110
Indeterminate plagiosaurid material 110
Temnospondyli incertae sedis 111
Saharastega SIDOR et al., 2005 111
Nomina dubia 111
Genera 111
Arkanserpeton LANE, 1932 111
Euchirosaurus GAUDRY, 1879 112
Hesperoherpeton PEABODY, 1958 112
Nannospondylus OLSON, 1965a 112
Pleuronoura GAUDRY, 1879 112
Porierpeton FRITSCH, 1885b 112
Species 112
Nomen nudum 113
Unnamed temnospondyl material from the Paleozoic 114
Not Temnospondyli 115
Genera 115
Austrobrachyops COLBERT & COSGRIFF, 1974 115
Chalcosaurus MEYER, 1866 115
Enosuchus KONZHUKOVA, 1955b 115
Erierpeton MOODIE, 1912 115
Erpetobrachium MOODIE, 1912 115
Jugosuchus RIABININ, 1962 116
Moraverpeton AUGUSTA, 1947b 116
Species 116
Stratigraphy and occurrences 118
Ontogeny and life histories 120
Major clades of temnospondyls 123
Temnospondyl evolution 125
Literature cited 127
Index of taxa 146

The present volume, Temnospondyli I by Drs. R. R. SCHOCH (Stuttgart) and A. R. MILNER (London), represents the long-awaited companion to Part 3B by the same team of authors, which dealt with one major clade of temnospondyls, Stereospondyli. A third volume (Part 3A1) by Professor J. A. CLACK and Dr. A. R. MILNER will discuss stem tetrapods and the proximate sister-taxa of Tetrapoda (Tetrapodomorpha). Such has been the pace of discoveries of and research on stem amphibians and stem tetrapods in recent decades that the editor and publisher decided to issue Volume 3 of the Handbook of Paleoherpetology in three rather than two parts to do justice to the wealth of available information. It is to be hoped that these syntheses will provide important sources of data for all researchers concerned with the early evolution of tetrapods.

Although temnospondyls have been the subject of scientific study since the first half of the nineteenth century, these tetrapods received relatively little scientific attention for many years. ROMER’s (1947) monographic review of all taxa known at that time was foundational. It compiled and synthesized the widely scattered information on the diversity and morphology of temnospondyls and established a series of major grades among these tetrapods. During the second half of the twentieth century, the study of temnospondyls experienced a major renaissance. In addition to increasingly detailed studies on the morphology of previously known as well as new taxa, researchers explored the ontogeny of various temnospondyls and, using the explicitly phylogenetic methods, elucidate the evolutionary history of Temnospondyli, with special reference to their relationships to extant amphibians. The latter subject remains contentious, but most researchers now would argue for a sister-group relationship between one particular clade of Temnospondyli, Dissorophoidea, and at least Batrachia (comprising Anura and Caudata).

The authors of the present volume have long been the leaders in the study of temnospondyl morphology and phylogeny and have personally examined most of the important specimens. Most recently, SCHOCH (2013) published a major analysis of temnospondyl interrelationships, which provides the phylogenetic framework for the present review.

The authors and editor take great pleasure in thanking Dr. F. PFEIL for his continuing enthusiasm for and tireless support of the Handbook of Paleoherpetology.

H.-D. SUES