Eryops Megacephalus

Pawley, K. 2006. 247 The Postcranial Skeleton of Temnospondyls (Tetrapoda: Temnospondyli) PhD Thesis. La Trobe University, Melbourne

APPENDIX 1. TABLES OF PUBLICATIONS WITH FIGURES OF TEMNOSPONDYL POSTCRANIAL MATERIAL

TABLE 2. BASAL TEMNOSPONDYLI

BASAL TEMNOSPONDYLI AUTHOR es es es g g

r ulocoracoi als p p resacral vert resacral vert halan ubis halan tibia fibula tarsals skull mandible atlas / axis p caudal vert ribs interclavicle clavicle cleithrum sca humerus ulna radius car p p ischium rib sacral femu p ilium branchials / osteoderms scales (Milner and Sequeira, Balanerpeton woodi ** * ******** * * * * **** * 1994) (Godfrey and Holmes, Caerorhachis bairdi ** ***** * * * * **** * 1995) (Godfrey and Holmes, Caerorhachis bairdi ** ***** * * * * **** * 1995) Caerorhachis bairdi (Ruta et al., 2002) * * * * * * * * * ** * * * (Sequeira and Milner, Capetus palustris ** * * 1993) Dendrerpeton acadianum (Carroll, 1967) ** * ** ** * * * * ** Dendrerpeton acadianum (Dawson, 1882) * Dendrerpeton acadianum (Holmes et al., 1998) * * * * * * * * * * * * * * * * Dendrerpeton acadianum (Milner, 1980b) * * * * * * * * * ** Dendrerpeton acadianum (Milner, 1982b) * * * Dendrerpeton acadianum (Shishkin, 1989) * Dendrerpeton acadianum (Steen, 1934) ** * **** (Romer and Witter, Edops craigi ** * * * * * * 1942) (Laurin and Soler- Iberospondylus schultzei * * Gijón, 2001)

TABLE 3. COCHLEOSAURIDAE

COCHLEOSAURIDAE AUTHOR es es es g g

TABLE 4. DVINOSAURIA

DVINOSAURIA AUTHOR es es es g g

r ulocoracoi als p p resacral vert resacral vert halan ubis halan tibia fibula tarsals skull mandible atlas / axis p caudal vert ribs interclavicle clavicle cleithrum sca p p ischium rib sacral femu p humerus ulna radius car ilium branchials / osteoderms scales Acroplous vorax (Coldiron, 1978) * * * * * * * * * Acroplous vorax (Hotton, 1959) * * * * * * * * * * Acroplous vorax (Shishkin, 1989) * Dvinosaurus campbelli (Gubin, 2004) ** * ** * * Dvinosaurus egregius (Nikitin, 1995) * Dvinosaurus primus (Bystrow, 1938) * * * * * * * * * * * * * * * * * * * Dvinosaurus primus (Nikitin, 1995) * Dvinosaurus primus (Nikitin, 1997) * Dvinosaurus primus (Shishkin, 1989) * Dvinosaurus primus (Shishkin, 2000) * * Dvinosaurus primus (Sushkin, 1936) * * * * * * * * * * * * * Isodectes (Saurerpeton) (Milner, 1982a) ** ******** * * obtusus Isodectes obtusus (Watson, 1956) ** * ** * (Eobrachyops townendae) Isodectes? obtusus (Werneburg, 2002) * * * * * * * (Olson and Lammers, Kourerpeton bradyi * * *** ** * * 1976) Neldasaurus wrightae (Chase, 1965) * * * * * * * * * * * * * * * Thabanchuia oomie (Warren, 1998b) * * * * * * * * Trimerorhachis insignis (Case, 1911a) ** * * * * * * Trimerorhachis insignis (Case, 1935) ** * * **** * * * * * Trimerorhachis insignis (Colbert, 1955) * (Cope and Matthew, Trimerorhachis insignis ** * * * 1915) Trimerorhachis insignis (Olson, 1979) * * Trimerorhachis insignis (Williston, 1910a) * Trimerorhachis insignis (Williston, 1915a) * * * * * * * * * * * Trimerorhachis insignis (Case, 1911a) * (alleni) Trimerorhachis (Berman and Reisz, ** * ** * * * sandovalensis 1980) Tungussogyrinus bergi (Shishkin, 1998) * * * * * * * * * * * Tupilakosaurus (Shishkin, 1961) * Tupilakosaurus heilmani (Nielsen, 1954) * * * * * * Tupilakosaurus (Shishkin, 1989) * wetlugensis

APPENDIX 1: TABLES OF PUBLICATIONS 249

TABLE 5. EUSKELIA

EUSKELIA AUTHOR es es es g g

r ulocoracoi als p p resacral vert resacral vert halan ubis halan tibia fibula tarsals skull mandible atlas / axis p caudal vert ribs interclavicle clavicle cleithrum sca humerus ulna radius car p p ischium rib sacral femu p ilium branchials / osteoderms scales ? Actiobates peabodyi (Milner, 1985a) * * **** ? Eryops (Werneburg, 1993a) ** * * Acanthostomatops (Steen, 1937) * * ****** (Acanthostoma) vorax Acanthostomatops vorax (Boy, 1989) * ** * * * * * Acanthostomatops vorax (Werneburg, 1998) * * *** * * Acheloma cumminsi (Case, 1911a) * **** Acheloma cumminsi (Romer, 1922) * Acheloma cumminsi (Case, 1911a) * * * * * * * * * * * * * * * * * * * * * * (Trematops milleri) Acheloma cumminsi (Olson, 1941) * * * * * * * * * * * * * (Trematops milleri) Acheloma cumminsi (Schaeffer, 1941) **** (Trematops milleri) Acheloma cumminsi (Williston, 1909a) * * * * * * * * * * * * * ** * * (Trematops milleri) Acheloma spp. (Sullivan et al., 2000) * Alegeinosaurus aphthitos (Case, 1911a) * * ***** * * * Amphibamus grandiceps (Bolt, 1979) ** * Amphibamus grandiceps (Carroll, 1964a) * * * * * * * * * * * ** * Amphibamus grandiceps (Daly, 1994) Amphibamus grandiceps (Gregory, 1950) Amphibamus grandiceps (Milner, 1982a) * ** ** * * * * * Amphibamus grandiceps (Werneburg, 2002) ** * ******** * Amphibamus grandiceps (Watson, 1940) ** ********** * * * * ** * * (Miobatrachus romeri) Anconastes vesperus (Berman et al., 1987a) ** * Aspidosaurus (Zatrachys) (Case, 1907) * ******** * apicaulis Aspidosaurus (Zatrachys) (Case, 1911a) * * apicaulis (Berman and Lucas, Aspidosaurus binasser ** * 2003) Aspidosaurus chiton (Case, 1911a) * * Aspidosaurus crucifer (de Mar, 1966) * Aspidosaurus glascocki (Case, 1911a) * * * Aspidosaurus (Carroll, 1964a) * * * * novomexicanus Aspidosaurus sp. (Carroll, 1964a) * Astreptorhachis ohioensis (Vaughn, 1971) * 250 K. PAWLEY PHD THESIS

EUSKELIA AUTHOR es es es g g

r ulocoracoi als p p resacral vert resacral vert halan ubis halan tibia fibula tarsals skull mandible atlas / axis p caudal vert ribs interclavicle clavicle cleithrum sca humerus ulna radius car p p ischium rib sacral femu p ilium branchials / osteoderms scales Brevidorsum profundum (Carroll, 1964a) * * * Broiliellus brevis (Carroll, 1964a) ** *** **** * Broiliellus texensis (de Mar, 1966) * Broiliellus texensis (Williston, 1915b) * * * Cacops aspidephorus (de Mar, 1966) * Cacops aspidephorus (Williston, 1910a) * * * * * * * * * * * * * * * * * * * * * * * * * Cacops cf. aspidephorus (Milner, 1985b) * * * (Trematopsis seltini) Cheliderpeton latirostre (Boy, 1993) ** * Conjunctio sp. (Carroll, 1964a) * * * * Dissorophid cf. Cacops (Sullivan et al., 2000) * Dissorophus angustus (Carroll, 1964a) * * * ** *** * Dissorophus multicinctus (de Mar, 1966) * * * * * * * * * * * * * * * * * * * * * * * * Dissorophus multicinctus (Williston, 1910b) * * * * * * * * * Dissorophus multicinctus (Case, 1911a) * **** * * (Otocoelus testudineus) Doleserpeton annectans (Bolt, 1969) * * Doleserpeton annectans (Carroll et al., 2004) * Doleserpeton annectans (Daly, 1973) * Doleserpeton annectans (Shishkin, 1989) * Ecolsonia cutlerensis (Berman et al., 1985) * * * * * * * * * * * * * * * * * Ecolsonia cutlerensis (Shishkin, 1989) * Eoscopus lockardi (Daly, 1994) * * * * * * * * * * * * * * * * * * * * * Eryopoidea ?Onchiodon (Werneburg, 1993c) * * * * * * Eryopoidea c.f. Onchiodon (Werneburg, 1995) * * Eryops megacephalus (Case, 1911a) * * * * * * * * * * (Cope and Matthew, Eryops megacephalus * * * *** * 1915) Eryops megacephalus (Cope, 1888) * * * * * * * * * Eryops megacephalus (Emery, 1897) **** (Gregory and Raven, Eryops megacephalus ******** 1941) Eryops megacephalus (Gregory et al., 1923) * * * * Eryops megacephalus (Gregory, 1949) * Eryops megacephalus (Miner, 1925) * * * * * * * * * Eryops megacephalus (Moulton, 1974) **** (Romer and Witter, Eryops megacephalus * 1941) APPENDIX 1: TABLES OF PUBLICATIONS 251

EUSKELIA AUTHOR es es es g g

r ulocoracoi als p p resacral vert resacral vert halan ubis halan tibia fibula tarsals skull mandible atlas / axis p caudal vert ribs interclavicle clavicle cleithrum sca humerus ulna radius car p p ischium rib sacral femu p ilium branchials / osteoderms scales Eryops megacephalus (Romer, 1922) ***** Eryops megacephalus (Williston, 1899) * * Eryops megacephalus (Langston, 1953) * * * * * * (grandis) Eryops megacephalus (Moodie, 1910) * * * **** * (willistoni) Fayella chickashaensis (Olson, 1972) * * * * * * * * * * (Broili and Schröder, Micropholis stowi ** *** * * * 1937) Micropholis stowi (Watson, 1913) ** * *** **** * * Onchiodon (Actinodon) (Gaudrey, 1884) * frossardi Onchiodon (Actinodon) (Werneburg and Steyer, ** ** * ** * frossardi 1999) Onchiodon labyrinthicus (Boy, 1971) ** * ****** * Onchiodon labyrinthicus (Boy, 1990) ** * * * Onchiodon labyrinthicus (Witzmann, 2005) ** * ******** * * * Onchiodon lagenhani (Werneburg, 1993b) * * Onchiodon manebachensis (Werneburg, 1996) ** * Onchiodon sp. (Werneburg, 1997) * * * * * * * * * * Parioxys bolli (Carroll, 1964b) * * * * * * * * * ** Parioxys ferricolus (Moustafa, 1955a) * * * * * * * * * * * * ** Peltobatrachus pustulatus (Panchen, 1959) * * * * * * * * * * * * Peltobatrachus pustulatus (Shishkin, 1989) * Perryella olsoni (Carlson, 1987) ** * ** Phonerpeton (Acheloma) (Dilkes, 1990) ** *** ***** pricei Platyhistrix rugosus (Langston, 1953) * * * (Lewis and Vaughn, Platyhistrix rugosus * 1965) Platyrhinops (Carroll et al., 1999) ** * ******** * * (Amphibamus) lyelli Platyrhinops (Carroll, 1964a) ************* * * * ** * (Amphibamus) lyelli Platyrhinops (Daly, 1994) (Amphibamus) lyelli Platyrhinops (Amphibamus) lyelli (Hook and Baird, 1984) * * * * * * * * ** * * * (Ichthycanthus platypus) Sclerocephalus credneri (Onchiodon labyrinthicus, (Werneburg, 1993b) ** * ****** * Lusor tenellus) Sclerocephalus haeuseri (Boy, 1988) **** * * * * * ** * 252 K. PAWLEY PHD THESIS

EUSKELIA AUTHOR es es es g g

r ulocoracoi als p p resacral vert resacral vert halan ubis halan tibia fibula tarsals skull mandible atlas / axis p caudal vert ribs interclavicle clavicle cleithrum sca humerus ulna radius car p p ischium rib sacral femu p ilium branchials / osteoderms scales (Lohmann and Sachs, Sclerocephalus haeuseri * * * * * * * * * * * * * * * * * * * * * * * 2001) Sclerocephalus haeuseri (Meckert, 1993) ******* * Sclerocephalus (Werneburg, 1992) * * * * * * * * * * * * jogischneideri Tambachia trogallas (Sumida et al., 1998) * * * * * * * * * * * * Tersomius cf. T. texensis (Daly, 1973) ** * Tersomius cf. T. texensis (Daly, 1994) * * * * * * * * * * Tersomius graumanni (Boy, 1980) * * ** * * * Tersomius mosei (Daly, 1994) * * ** * Trematopid cf. Acheloma (Sullivan et al., 2000) * Zatrachys serratus (Case, 1911a) * ** Zatrachys serratus (Langston, 1953) ** * * Zatrachys serratus (Shishkin, 1989) * APPENDIX 1: TABLES OF PUBLICATIONS 253

TABLE 6. STEREOSPONDYLOMORPHA

d STEREOSPONDYLO- AUTHOR

MORPHA es es g g

r ulocoracoi als al rib al p r p resacral vert resacral vert halan ubis halan femu tibia fibula tarsals skull mandible atlas / axis p caudal vert ribs interclavicle clavicle cleithrum sca humerus ulna radius car p p ischium sac p ilium branchials / osteoderms scales Almasaurus habbazi (Dutuit, 1976) ** * *** * * Apachesaurus gregorii (Hunt, 1993) ** * ** * Aphaneramma rostratum (Nilsson, 1943) * * * * * * * * (Säve-Söderbergh, Aphaneramma rostratum * * * 1936) Aphaneramma rostratum (Wiman, 1910) ** * ** (Lonchorhynchus oebergi) Archegosaurus (Milner, 1978) ** ** (Memonomenos) dyscriton Archegosaurus decheni (Jaekel, 1896) * *** Archegosaurus decheni (Meyer, 1857) * * * * * * * * * * * * * * * * * * * * * Archegosaurus decheni (Witzmann, 2006a) * * * * * * * * * * * * * * * * * * * * * * Archegosaurus sp. (Emery, 1897) ** * * **** Archegosaurus sp. (Gaudrey, 1884) * (Dias and Schultz, Australerpeton cosgriffi * ******** * * * 2003) Banksiops (Blinasaurus) (Cosgriff, 1974) * * * * * * * * townrowi Batrachosuchoides (Shishkin, 1966) * * * (Warren and Batrachosuchus * Marsicano, 2000) Batrachosuchus (Cosgriff, 1969) * * * (Blinasaurus) henwoodi (Chernin and Kitching, Batrachosuchus concordi *** * ** 1977) Batrachosuchus henwoodi (Cosgriff, 1974) * Batrachosuchus sp. (Watson, 1956) ** (Welles and Estes, Batrachosuchus sp. * * 1969) (Bystrow and Efremov, Benthosuchus sushkini * * * * * * * * * * * * * * * * ** * 1940) Bothriceps major (Warren, 1997) * Buettneria maleriensis (Sengupta, 2002) * * * * * * * * * * * * * * * * * ** * Buettneria maleriensis (von Huene, 1940) ** * ** * (metoposaurid) Buettneria perfecta (Case, 1922) * ** Buettneria perfecta (Hunt, 1993) * * ** * Buettneria perfecta (Olsen, 1951) * Buettneria perfecta (Sengupta, 1992) * * 254 K. PAWLEY PHD THESIS

d STEREOSPONDYLO- AUTHOR

MORPHA es es g g

r ulocoracoi als al rib al p r p resacral vert resacral vert halan ubis halan femu tibia fibula tarsals skull mandible atlas / axis p caudal vert ribs interclavicle clavicle cleithrum sca humerus ulna radius car p p ischium sac p ilium branchials / osteoderms scales Buettneria perfecta (Borborophagus (Branson and Mehl, ** * ** * wyomongensis, 1929) Koskinodon princeps) Buettneria perfecta (Sawin, 1945) ************* * * * * * (howardensis) Buettneria perfecta (Metoposaurus browni, (Roy Chowdery, 1965) * * * * * * * fraasi fraasi, malerensis) Bukobaja enigmatica (Ochev, 1972) ** * (Warren and Damiani, Chigutisauridae indent. ** * 1999) Chomatobatrachus halei (Cosgriff, 1974) * * * Compsocerops cosgriffi (Sengupta, 1995) ** * ** * * (Colbert and Cosgriff, Cryobatrachus kitchingi *** ** * * * 1974) (Cosgriff and Hammer, Cryobatrachus kitchingi * 1984) Cyclotosaurus hemprichi (Kuhn, 1942) * * * * Cyclotosaurus intermedius (Sulej and Majer, 2005)** ***** Deltasaurus (Cosgriff, 1965) * * * * kimberleyensis Dutuitosaurus (Dutuit, 1976) * * * * * * * * * * * * * * * * * * * * * (Metoposaurus) ouazzoui Dutuitosaurus (Shishkin, 1989) * (Metoposaurus) ouazzoui Dutuitosaurus ouazzoui (Steyer et al., 2004) * Eocyclotosaurus sp. (Milner et al., 1990) * * Eocyclotosurus sp. indent (Cyclotosaurus randalli) (Welles, 1947) * * * * Heylerosauridae indent (Rhadalognathus boweni) Eryosuchus (Parotosaurus) (Howie, 1970) * * * * * * * * * * * * * * * * * * * * * pronus Eryosuchus tverdochlebovi (Ochev, 1972) * * * * * * * * * * * * * * * * * * * * * Eryosuchus tverdochlebovi (Shishkin, 2000) * (garjainovi) Ferganobatrachus (Nessov, 1990) * * * * riabinini Gerrothorax (Plagiosaurus) (von Huene, 1922) * * * * * * pulcherrimus Gerrothorax (Plagiosaurus) (Jaekel, 1914) ** * * pulcherrimus Gerrothorax pulcherrimus (Nilsson, 1937) * * * * * Gerrothorax Hellrung (2003) ** * *** ** *** * * pustuloglomeratus APPENDIX 1: TABLES OF PUBLICATIONS 255

d STEREOSPONDYLO- AUTHOR

MORPHA es es g g

r ulocoracoi als al rib al p r p resacral vert resacral vert halan ubis halan femu tibia fibula tarsals skull mandible atlas / axis p caudal vert ribs interclavicle clavicle cleithrum sca humerus ulna radius car p p ischium sac p ilium branchials / osteoderms scales Gerrothorax rhaeticus (Nilsson, 1946a) * * * * * * * ** * * Gerrothorax sp. (Shishkin, 1989) * (Maisch and Matzke, Gobiops desertus **** * * 2005) Gobiops desertus (Shishkin, 1991) * * Gondwanosaurus (Lydekker, 1885) ** * *** * bijoriensis Gonioglyptyus longirostris (Huxley, 1885) ** * * Khonzukovia (Melosaurus) (Konzhukova, 1955a) ** * * vetusta Koolasuchus cleelandi (Warren et al., 1997) * *** * Laidleria gracilis (Kitching, 1957) * * * * * * * * * * * * Laidleria gracilis (Warren, 1998a) * * * * (Warren and Lapillopsis nana ** ** ** * Hutchinson, 1990a) Lapillopsis nana (Yates, 1999) ** * ** * (Broili and Schröder, Lydekkerina huxleyi * ** 1937) Lydekkerina huxleyi (Broom, 1930) * **** Lydekkerina huxleyi (Parrington, 1948) * Lydekkerina huxleyi (Watson, 1919) * * ***** * * * ** Lydekkerina huxleyi (Parrington, 1948) ** **** (Limnoiketes paluditans) (Mazin and Janvier, Lyrocephaliscus euri * * *** 1983) Lyrocephaliscus euri (Nilsson, 1943) * * Mastodonsauridae incertae sedis (Promastodonsaurus (Bonaparte, 1963) ** ** bellmani) Mastodonsaurus (Wepfer, 1923) ******* **** * * * * * cappelensis Mastodonsaurus giganteus (von Huene, 1922) * * * * * * * * * * Mastodonsaurus giganteus (Schoch, 1999a) * * * * * * * * * * * * * * * * * * * * Mastodonsaurus giganteus (Werneburg, 2004a) ** Mastodonsaurus giganteus (Fraas, 1889) ** * *** **** * * (acuminatus, keuperinus) Mastodonsaurus torvus (Konzhukova, 1955a) ** * Mastodonsaurus torvus (Ochev, 1972) ** * * * * Mastodonsaurus torvus (Shishkin, 1989) * Melosauridae gen. indent. (Gubin, 1991) * Melosaurus kamaensis (Gubin, 1991) * Melosaurus uralensis (Gubin, 1991) * 256 K. PAWLEY PHD THESIS

d STEREOSPONDYLO- AUTHOR

MORPHA es es g g

r ulocoracoi als al rib al p r p resacral vert resacral vert halan ubis halan femu tibia fibula tarsals skull mandible atlas / axis p caudal vert ribs interclavicle clavicle cleithrum sca humerus ulna radius car p p ischium sac p ilium branchials / osteoderms scales (Branson and Mehl, Metoposauridae * ** 1929) Metoposaurus (Buettneria) (Case, 1931) * * * * bakeri Metoposaurus (Buettneria) (Case, 1932) **** *** *** * * * * * bakeri Metoposaurus (Metopias) (Fraas, 1889) * * * * * * * diagnosticus Metoposaurus (Metopias) (Fraas, 1896) * * * * * * diagnosticus (Colbert and Imbrie, Metoposaurus diagnosticus * ** 1956) Paracyclotosaurus davidi (Watson, 1958) * * * * * * * * * * * * * * * * * * * * * * * Parotosaurus bogdananus (Ochev, 1972) * ** Parotosaurus (Ochev, 1972) ** * * orenburgensis Parotosuchus haughtoni (Damiani, 2002) ** * Pelorocephalus sp. (Marsicano, 1993) * * * * * * * * * * * * * Peltostega erici (Nilsson, 1946b) * * Phrynosuchus whaitsi (Broom, 1913) * * * * * * * * * * * * * Plagiobatrachus australis (Warren, 1985) * * Plagiosauridae indent. (Milner et al., 1995) * * * Plagiosauridae indent. (Shishkin, 1967) * Plagiosauridae indent. (Shishkin, 1987) * * * * * Plagiosauridae indent. (Schmidt, 1931) (Plagiosternum nanum) Plagiosaurus depressus (Nilsson, 1939) * * * Plagiosaurus sp. (Piveteau, 1928) * Plagioscutum ochevi (Shishkin, 1989) * Plagioscutum ochevi (Shishkin, 2000) * Plagioscutum ochevi / (Shishkin, 1986) * * * caspiense Plagioscutum ochevi / (Shishkin, 1987) * * * caspiense Plagiosternum granulosum (Fraas, 1896) * Plagiosternum granulosum (Fraas, 1913) * ** Plagiosternum granulosum (Schmidt, 1931) Plagiosuchus pustuliferus (von Huene, 1922) * * * * * * * Plagiosuchus pustuliferus (Schmidt, 1931) * ** Platycepsion (Platyceps) (Stephens, 1887) * * * * * * wilkinsoni Platycepsion (Platyceps) (Watson, 1956) ** ** * wilkinsoni APPENDIX 1: TABLES OF PUBLICATIONS 257

d STEREOSPONDYLO- AUTHOR

MORPHA es es g g

r ulocoracoi als al rib al p r p resacral vert resacral vert halan ubis halan femu tibia fibula tarsals skull mandible atlas / axis p caudal vert ribs interclavicle clavicle cleithrum sca humerus ulna radius car p p ischium sac p ilium branchials / osteoderms scales Platycepsion wilkinsoni (Cosgriff, 1969) * * * * (Warren and Platycepsion wilkinsoni * ** * Marsicano, 1998) Platyoposaurus (Platyops) (Konzhukova, 1955b) * * * * * * * * * * * * * * * * * * stuckenbergi Platyoposaurus (Gubin, 1991) * * * * * * * * * * * * * * * * * stuckenbergi Platyoposaurus (Shishkin, 1989) * stuckenbergi Platyoposaurus (Shishkin, 2000) * stuckenbergi Platystega depressa (Nilsson, 1943) * * (Cosgriff and Zawiskie, Pneumatostega potamia ** * ** 1979) Prothoosuchus blomi (Getmanov, 1989) * ****** * * Rewana (Arcadia) (Warren and Black, ** *** ** myriadens 1985) Rewana quadricuneata (Howie, 1972a) * * * * * * * * * * * * ** (Cosgriff and Hammer, Rhytidostidae indent. * 1984) (Warren and Siderops kehli *********** * * * * Hutchinson, 1983) Stereospondyli indent. (Efremov, 1931) ** * ** (Capitosaurus) Thoosuchus jakovlevi (Efremov, 1940) * * Trematolestes hagdorni (Schoch, 2006) * * * * * * * * * * * * * * * Trematosauria indent. (Maisch et al., 2004) * Trematosauridae sp. (Shishkin, 1989) * Trematosaurus brauni (Burmeister, 1849) ** ** Trematosaurus brauni (Efremov, 1931) ** * * * Trematosaurus (Janvier, 1992) * * madagascariensis Tryphosuchus kinelensis (Gubin, 1991) * * * * Tryphosuchus paucidens (Konzhukova, 1955a) ** * * * * Uranocentrodon (Haughton, 1915) * * * * * * * * * * * * * * * * * * * * (Myriodon) senekalensis Uranocentrodon (Haughton, 1915) * * * * * * * * * * * * * * * * * * * * (Myriodon) senekalensis Uranocentrodon (van Hoepen, 1915) ** ********** * * * * * * * ****** (Myriodon) senekalensis Uranocentrodon (Findlay, 1968) * (Rhinesuchus) senekalensis Uranocentrodon (Broom, 1930) ** **** senekalensis Vigilius (Taphrognathus) (Welles, 1947) * * wellesi 258 K. PAWLEY PHD THESIS

d STEREOSPONDYLO- AUTHOR

MORPHA es es g g

TABLE 7. LARVAL TEMNOSPONDYLS

LARVAL TEMNOSPONDYLS AUTHOR es es es g g

r ulocoracoi als p p resacral vert resacral vert halan ubis halan tibia fibula tarsals skull mandible atlas / axis p caudal vert ribs interclavicle clavicle cleithrum sca p p ischium rib sacral femu p humerus ulna radius car ilium branchials / osteoderms scales Apateon gracilis (Werneburg, 1991) ** ********** * * * * ** *** (Melanerpeton gracile) Apateon (Branchiosaurus) (Boy, 1978) *** * ** cauducus Apateon cauducus (Schoch, 1992) * * * * * * * * * * ** * * Apateon dracyiensis (Werneburg, 2002) ** * ******** * * Apateon flagrifera (Werneburg, 2002) ** * ******** * flagrifera Apateon intermedius (Werneburg, 1996) * * * * * * * Apateon pedestris (Schoch, 1992) * * * * * * * * * * ** * * Apateon pedestris (Werneburg, 2002) ** * ******** * * Apateon pedestris (Werneburg, 2004b) * * * * * * * * * * * * ** * Apateon pedestris (Werneburg, 2004b) * * * * * * * * * * * * * * * * * * * * * * Apateon sp. (Werneburg, 1990) * Branchierpeton (Werneburg, 1991) * * * * * * * * * * * * * * * * * * * * amblystomus Branchierpeton cf. (Zajic et al., 1990) ** * ******** * * saalensis Branchierpeton saalensis (Werneburg, 1990) * * * * * * * * * * * * * ** * Branchierpeton saalensis (Werneburg, 1990) * * * * * * * * * * * * * * * ** * Branchierpeton saalensis (Werneburg, 1996) ** * *** **** * * * ** * Branchiosaurus ?fayoli (Werneburg, 2002) ** * * * Branchiosaurus cf. petrolei (Boy, 1971) * * * * * * * * * * * * * * * * * * * * Branchiosaurus cf. petrolei (Boy, 1974) ** * * Branchiosaurus cf. petrolei (Boy, 1978) ** ** ***** * * ** Branchiosaurus cf. petrolei (Milner, 1982a) ** ***** ** * * * ** ** Branchiosaurus (Boy, 1978) ** * *** * ** * humbergensis Branchiosaurus (Carroll et al., 1999) ** * * **** * ** * salamandroides Eimerisaurus guembeli (Boy, 2002b) ** * ** *** * * * Limnogyrinus elegans (Milner and Sequeira, (Branchiosaurus ** *** * 2003) salamandroides) Melanerpeton gracile (Werneburg, 2003) ** * ******** * * * ** Melanerpeton (Werneburg, 2004b) ** ** * **** * * *** humbergense Melanerpeton (Werneburg, 2004b) * * * * * * * * * * * * * * * * * * * * * humbergense Melanerpeton (Spinar, 1949) ** ***** *** * * * **** longicaudatum 260 K. PAWLEY PHD THESIS

LARVAL TEMNOSPONDYLS AUTHOR es es es g g

TABLE 8. TEMNOSPONDYLI INDET.

TEMNOSPONDYLI INDET. AUTHOR es es es g g

r ulocoracoi als p p r resacral vert resacral vert halan ubis halan tibia fibula tarsals skull mandible atlas / axis p caudal vert ribs interclavicle clavicle cleithrum sca humerus ulna radius ca p p ischium rib sacral femu p ilium branchials / osteoderms scales (Welles and Estes, Hadrokkosaurus bradyi ** * ** * 1969) Hadrokkosaurus bradyi (Welles, 1947) ** * ** * (Werneburg and Lysipterygium risinensis ** ***** * * * Schneider, 1996) Milnererpeton (Milneria) (Hunt et al., 1996) * * * * * * * * * * * * * * * * huberi Mordex (Limnerpeton) (Milner and Sequeira, ** * ******** *** * * * * laticeps 2003) Palatinerpeton (Boy, 1996) * * * * * * * * * * kraetschmeri Sclerothorax hyselonotus (von Huene, 1932) * * * * * * * * Temnospondyli indent. (Fastnach, 2004) * * * * Temnospondyli indent. (Godfrey, 2003) * * * * * * * * * * *

Published as: Pawley, K. and Warren, A. 2004. Immaturity vs. paedomorphism: A rhinesuchid 261 stereospondyl postcranium from the Upper Permian of South Africa. Palaeontologica africana. 40: 1-10. Copyright © 2004 Bernard Price Institute for Palaeontological Research

APPENDIX 2. LOCALITY AND REPOSITORY INFORMATION FOR RHINESUCHID SPECIMENS

SOUTH AFRICA, BEAUFORT GROUP

Cistecephalus Assemblage Zone

Beaufort West: M460, a skull and articulated anterior part of the pectoral girdle,

Dicynodon Assemblage Zone

Senekal: Uranocentrodon (Myriodon) senekalensis NMQR1438; TM75, 77-79, 85, 103, 208 and 4703, complete skeletons. Lucerne 70 farm near Graaff-Reinet, South Africa: SAM-PK-K10021, consisting of a partial skull, vertebral column and ribs, pectoral girdle, proximal limb elements and ventral scutes of a rhinesuchid, preserved in a grey mudstone block.

Lystrosaurus Assemblage Zone

Oliviershoek: Broomistega putterilli BP/1/3241, skull and anterior skeleton.

Published as: Pawley, K. and Warren, A. 2006. The appendicular skeleton of Eryops megacephalus 263 (Temnospondyli: Eryopoidea) from the Lower Permian of North America. Journal of Paleontology. 80: 561-580. Copyright © 2006 The Paleontological Society

APPENDIX 3. LOCALITY AND REPOSITORY INFORMATION FOR SPECIMENS OF ERYOPS MEGACEPHALUS

Lithostratigraphic information assigned to specimens at the time of collection is revised following Hentz and Brown (1987), and specific collection localities are defined using information from Romer (1935; 1974) and Romer and Price (1940). Most specimens come from the Lower Permian red beds of Wichita, Baylor, and Archer counties in north- central Texas; one is from Sandoval County of New Mexico.

TEXAS, BOWIE GROUP

Archer City Formation

Location one mile south of Archer City, Archer County, Latitude 33º 34’ 0 N Longitude 98º 40’ W: MCZ 1421 clavicle (Figure 10.6).

TEXAS, WICHITA GROUP

Nocona Formation

North Fork of the Little Wichita River, Baylor County, exact locality undeterminable (see Romer, 1935): AMNH 4180 scapulocoracoid; AMNH 4183 scapulocoracoid, humerus, pelvis, and femur; AMNH 4186 articulated pectoral girdle and right forelimb; AMNH 4203 radius (Figure 10.8), femur (Figure 11.6), and tibia (Figure 10.10); AMNH 4754 clavicle, ulna, radius, tibia; AMNH 23449 fibula (Figure 10.11). Geraldine Quarry, Archer County, Latitude 33º 43’ 0 N Longitude 98º 39’ W: AMNH 4198 tibia (Long Ck. of Romer, 1935); MCZ 1883 clavicle fragment (Figure 10.4); MCZ 1984 cleithrum (Figure 10.7); MCZ 1900 interclavicle (two specimens) (Figure 10.2, 10.5) and cleithra (two specimens); MCZ 7766 interclavicle (Figure 10.3); MCZ 7768 cleithrum; MCZ 7773 ilium (Figure 11.4); MCZ 7797 tibia. Rattlesnake Canyon, Archer County, Latitude 33º 40’ 0 N Longitude 98º 50’ W: MCZ 6475 fibula. Location northwest of Megargel, Archer County, Latitude 33º 28’ 0 N Longitude 98º 55’ W: MCZ 2712 scapulocoracoid. Location three miles south of Black Flat, Texas, Archer County, Latitude 33º 39’ 0 N Longitude 98º 50’ W: MCZ 7279 interclavicle; MCZ 7551 humerus (Figure 11.3); MCZ 7555 tibia and fibula; MCZ 7559 scapulocoracoid. ‘Long Ck’ (see Romer, 1935) Archer County, Latitude 33º 41’ 0 N Longitude 98º 40’ W: AMNH 4215 scapulocoracoid. 264 K. PAWLEY PHD THESIS

Upper Briar Creek, Archer County, Latitude 33º 37’ 0 N Longitude 98º 50’ W: MCZ 2565 ventral scapulocoracoid (Figure 11.2). ‘Big Wichita’ Archer County, exact locality undeterminable (see Romer, 1935): AMNH 4255 humerus.

Petrolia Formation

Mount Barry six miles southwest near Big Wichita River, Wichita County, Latitude 33º 52’ 0 N Longitude 98º 37’ W: AMNH 4211 dorsal scapula and cleithrum (Figure 11.1). Lower Brushy Creek, Baylor County, Latitude 33º 45’ 0 N Longitude 99º 00’ W: MCZ 1137 ulna. Oilfield, north of Westover and Met Survey A-963, Texas, Baylor County, Latitude 33º 33’ 0 N Longitude 99º 05’ W: MCZ 2675 interclavicle (Figure 10.1). South side of Godwin's Creek, Baylor County, Latitude 33º 33’ 0 N Longitude 99º 05’ W: AMNH 4862 ulna (Figure 10.9). Hicks Farm, half mile south of Godwin Creek, Baylor County, Latitude 33º 33’ 0 N Longitude 99º 05’ W: AMNH 4307 scapulocoracoid. Steve Pierce's Place, two miles west and five miles south of Dundee, Archer County, Latitude 33º 42’ 0 N Longitude 98º 55’ W: MCZ 1219 pelvis. Tit Mountain northeast of Dundee, Archer County, Latitude 33º 48’ 0 N Longitude 98º 50’ W: AMNH 4204 scapulocoracoid and humerus. Palo Pinto School Land, Archer County, Latitude 33º 47’ 0 N Longitude 98º 55’ W: MCZ 2092 tibia.

NEW MEXICO, CUTLER FORMATION

San Diego Canyon, four miles south of Jemez Springs, NoSee'um Quarry, Sandoval County. New Mexico, Latitude 35º 45’ 0 N Longitude 106º 45’ W: MCZ 1858 pelvis (Figure 11.5).

Published as: Pawley, K. in press. The postcranial skeleton of Trimerorhachis insignis Cope 1878 265 (Temnospondyli: Trimerorhachidae) a plesiomorphic, secondarily aquatic temnospondyl from the Lower Permian of North America. Journal of Paleontology. Copyright © 2006 The Paleontological Society

APPENDIX 4. LOCALITY AND REPOSITORY INFORMATION FOR SPECIMENS OF TRIMERORHACHIS INSIGNIS

Lithostratigraphic information assigned to specimens at the time of collection was revised following Hentz and Brown (1987), specific collection localities were defined using information from Romer (1935; 1974) and Romer and Price (1940). All specimens come from the Wolfcampian Lower Permian red beds of the Wichita, Baylor, and Archer counties of north-central Texas.

TEXAS, WICHITA GROUP

Nocona Formation

North Fork of the Little Wichita River, Baylor County, exact locality undeterminable (see Romer, 1935): AMNH 23330 leg and foot; AMNH 23302 proximal femur (Figure 22.26). Geraldine Quarry (Long Ck. Romer, 1935), Archer County, Latitude 33º 43’ 0 N Longitude 98º 39’ W: AMNH 23319 distal humerus (Figure 22.20). Rattlesnake Canyon, Archer County, Latitude 33º 40’ 0 N Longitude 98º 50’ W: MCZ 8116 ilium; MCZ 8117 interclavicle; MCZ 8128 articulated pectoral girdle (Figure 21.1); MCZ 8164 fibula (Figure 22.30); UMMP 15995 right clavicle (Figure 22.6), UMMP 16033 foot bones, mounted, UMMP 16055 clavicle.

Petrolia Formation

Hay Camp, south of Fulda, Baylor County, Latitude 33º 39’ 0 N Longitude 99º 00’ W: MCZ 840 interclavicle; MCZ 8413 3 ulnae (Figure 22.8, 22.9); MCZ 8536 articulated vertebrae and pelvis. Four mi east of Rendham, Baylor County, Latitude 33º 33’ 0 N Longitude 99º 00’ W: MCZ 8312 pleurocentra (Figure 22.2). Godwin's Creek, Baylor County, Latitude 33º 33’ 0 N Longitude 99º 05’ W: AMNH 23321 distal femur (Figure 22.27). Tit Mountain, northeast of Dundee, Archer County, Latitude 33º 48’ 0 N Longitude 98º 50’ W: AMNH 4578 interclavicle and clavicle, ilium (Figure 22.23); AMNH 23283 2 scapula (Figure 22.12); MCZ 8244 radius (Figure 22.7) and small femur; MCZ 8245 rib fragments, MCZ 8515 ilium; TMM 40031-61 left ilium (Figure 22.22); TMM 40031-62 rib fragments; TMM 40031-63 left femur; TMM 40031-64 left femur (Figure 22.24); TMM 40031-65 left clavicle fragment; TMM 40031-70 pathological left femur; TMM 40031-71 left femur (Figure 22.25); TMM 40031-72 right femur; TMM 40031-73 right femur; TMM 40031-74 proximal left humerus; TMM 40031-75 distal left femur; TMM 40031-76 intercentrum; TMM 40031-80 266 K. PAWLEY PHD THESIS

left humerus (Figure 22.19); TMM 40031-81 left humerus (Figure 22.18); TMM 40031-82 proximal left ulna; TMM 40031-84 distal left fibula; TMM 40031-85 proximal right humerus; TMM 40031-86 proximal left femur; TMM 40031-87 intercentrum; TMM 40031-88 neural arch; TMM 40031-90 left ischium fragment; TMM 40031-91 neural arch fragment; TMM 40031-93 proximal right femur; TMM 40031-94 distal left humerus; TMM 40031-95 distal left ilium; TMM 40031-96 left ischium; TMM 40031-97 right scapula; TMM 40031-98 intercentrum; TMM 40031-101 rib fragment; TMM 40031-105 distal left humerus; TMM 40031-106 distal right humerus; TMM 40031-107 right scapula; TMM 40031-108 right scapula; TMM 40031-109 distal right humerus; TMM 40031-110 right clavicle fragment. Trematops locality, west of Williams, Baylor County, Latitude 33º 40’ 0 N Longitude 98º 58’ W: MCZ 8365 rib fragments. Slippery Ck., 3 mi south of Dundee, Archer County, Latitude 33º 41’ 0 N Longitude 98º 55’ W: MCZ 8577 clavicle and interclavicle. Thrift, Wichita County, Latitude 34º 07’ 0 N Longitude 98º 37’ W: TMM 40998-39 articulated hind limb (Figure 21.2); UMCP 105153 articulated vertebrae and ribs; UMCP 105230 distal femur; UMCP 105232 distal femur; UMCP 105236 distal humerus; UMCP 105247 interclavicle; UMCP 105249 proximal femur; UMCP 105258 proximal humerus; UMCP 105259 distal radius; UMCP 174885 humerus (Figure 22.16); UMCP 174886 humerus (Figure 22.14); UMCP 174887 distal humerus; UMCP 174888 humerus (Figure 22.15); UMCP 174889 humerus; UMCP 174890 distal humerus; UMCP 174891 humerus (Figure 22.13); UMCP 174892 distal femur; UMCP 174893 distal femur; UMCP 174894 left ischium (Figure 22.28); UMCP 174895 ischium; UMCP 174896 proximal humerus; UMCP 174897 ischium; UMCP 174898 scapula (Figure 22.11); UMCP 174899 left scapula (Figure 22.10); UMCP 187000 proximal radius; UMCP 187001, proximal femur. Fulda Nr, Baylor County, Latitude 33º 33’ 0 N Longitude 99º 05’ W: UMCP 24009 femur; UMCP 24010 humerus. Hackberry Creek, 3 mi southeast of Fulda, Baylor County, Latitude 33º 40’ 0 N Longitude 99º 00’ W: AMNH 4763 six intercentra (Figure 22.3, 4, 5), six phalanges (Figure 22.31), one ilium, half neural arch (Figure 22.1), one distal fibula. Little Wichita R. south side, Baylor County, exact locality undeterminable (see Romer, 1935): AMNH 4720 femur, two humeri (Figure 22.17), three ilia (Figure 22.21), tibia.

TEXAS, LOWER WAGGONER RANCH FORMATION

Coal Ck, Baylor County, Latitude 33º 37’ 0 N Longitude 99º 07’ W: MCZ 8611 tibia (Figure 22.29).

Pawley, K. 2006. 267 The Postcranial Skeleton of Temnospondyls (Tetrapoda: Temnospondyli) PhD Thesis. La Trobe University, Melbourne

APPENDIX 5. REPOSITORY INFORMATION FOR SPECIMENS OF MORPHOGENETIC SERIES

Acheloma cumminsi (Trematops milleri)

AMNH 4205, 7150; MCZ 1413, 2524.

Buettneria (Metoposaurus) perfecta

AMNH 2994; MCZ 1919-1925.

Eryops megacephalus

AMNH 4086, 4121, 4180, 4183, 4186, 4191, 4198, 4203, 4204, 4208, 4211, 4215, 4255, 4280, 4307, 4661, 4754, 4862, 23449; MCZ 1128, 1137, 1219, 1539, 1858, 1883, 1900, 1984, 2092, 2565, 2604, 2615, 2630, 2631, 2675, 2712, 2716, 4121, 4308, 4763, 6475, 7279, 7551, 7555, 7559, 7645, 7647, 7765, 7766, 7768, 7770, 7773, 7797; NMV P202186, P202187.

Lydekkerina huxleyi

BP/1/1373, BP/1/5021, BP/1/fn 3900; UMZC T218, T243.

Rhinesuchidae

BP/1/3241; M 460; NMQR 1438; TM 75, 77, 78, 79, 85, 103, 208, 4703; SAM-PK- K100021.

Trimerorhachis insignis

AMNH 4578, 4720, 4763, 4765, 23283, 23302, 23319, 23321, 23330; MCZ 840, 8116, 8117, 8128, 8164, 8244, 8245, 8312, 8365, 8413, 8515, 8536, 8577, 8611; TMM 40031:61-110, 40998:25, 40998:39; UMCP 105230, 105232, 105236, 105247, 105249, 105258, 105259, 174885-187001, 24009, 24010; UMMP 16033, 15995.

Pawley, K. 2006. 269 The Postcranial Skeleton of Temnospondyls (Tetrapoda: Temnospondyli) PhD Thesis. La Trobe University, Melbourne

APPENDIX 6. TERMINAL TAXA: THE POSTCRANIAL SKELETON OF TEMNOSPONDYLS

(mi) indicates that the endochondral postcranial skeleton of all described specimens is morphogenetically immature.

Acheloma cumminsi (Trematops milleri) (Williston, 1909a; Case, 1911a; Olson, 1941; Schaeffer, 1941; Bolt, 1974a, 1974b, 1974c; Dilkes and Reisz, 1987). Specimens examined: AMNH 4205, 7150; MCZ 1413, 2524. Acroplous vorax (mi) (Hotton, 1959; Coldiron, 1978; Foreman, 1990). Adamanterpeton ohioensis (Milner and Sequeira, 1998). Almasaurus habbazi (mi) (Dutuit, 1976). Specimens examined: MNHN ALM43, ALM44, ALM45, ALM51, ALM62, ALM65. Amphibamus grandiceps (mi) (Watson, 1940; Gregory, 1950; Carroll, 1964a; Bolt, 1979; Daly, 1994). Archegosaurus decheni (mi) (Meyer, 1857; Jaekel, 1896; Whittard, 1928; Gubin, 1997; Witzmann, 2006a, 2006b). Specimens examined: NMV P10836, P24165, P198455, P198457. Balanerpeton woodi (Milner and Sequeira, 1994). Batrachosuchus spp. (mi) (Watson, 1956; Cosgriff, 1969; Chernin, 1977; Warren and Marsicano, 1998). Specimens examined: UCMP 80859, 80860, 140579, 140580, 140583, 140589-140592. Benthosuchus sushkini (mi) (Bystrow and Efremov, 1940). Broiliellus brevis (mi) (Carroll, 1964a; Bolt, 1974a, 1974c). Specimens examined: MCZ 1424, 3272. Caerorhachis bairdi (Holmes and Carroll, 1977; Ruta et al., 2002). Capetus palustris (Sequeira and Milner, 1993). Cheliderpeton spp. (mi) (Boy, 1993; Werneburg and Steyer, 2002). Chenoprosopus lewisi (mi) (Hook, 1993). Chenoprosopus milleri (Langston, 1953). Cochleosaurus bohemicus (Sequeira, 2004). Cochleosaurus florensis (Rieppel, 1980; Godfrey and Holmes, 1995). Dendrerpeton acadianum (Dawson, 1882; Steen, 1934; Carroll, 1967; Milner, 1980b; Godfrey et al., 1987; Holmes et al., 1998; Robinson et al., 2005). Dissorophus spp. (Williston, 1910b; Case, 1911a; Carroll, 1964a; de Mar, 1968; Bolt, 1974a, 1974c; Milner, 2003). Specimens examined: AMNH 4343, 4593; MCZ 2122, 4169, 4170, 4172, 4173, 4175, 4176, 4177, 4178, 4180, 4181, 4185, 4186, 4187, 4190, 4194. Doleserpeton annectans (Gregory et al., 1956; Bolt, 1969, 1974a, 1974b; Lombard and Bolt, 1988). Dutuitosaurus (Metoposaurus) ouazzoui (mi) (Dutuit, 1976; Hunt, 1993; Steyer et al., 2004). 270 K. PAWLEY PHD THESIS

Dvinosaurus spp. (mi) (Sushkin, 1936; Bystrow, 1938; Shishkin, 1973; Nikitin, 1995, 1997; Shishkin, 2000; Gubin, 2004). Ecolsonia cutlerensis (Berman et al., 1985). Edops craigi (mi) (Romer and Witter, 1942; Clack and Holmes, 1988). Specimens examined: MCZ 1378, 1781, 6489, 6490, 7140. Eoscopus lockardi (mi) (Daly, 1994). Eryops megacephalus (Cope, 1884, 1888; Case, 1911a; Cope and Matthew, 1915; Romer, 1922; Miner, 1925; Sawin, 1941; Moulton, 1974; Pawley and Warren, 2006). Specimens examined: AMNH 4086, 4121, 4180, 4183, 4186, 4191, 4198, 4203, 4204, 4208, 4211, 4215, 4255, 4280, 4307, 4661, 4754, 4862, 23449; MCZ 1128, 1137, 1219, 1539, 1858, 1883, 1900, 1984, 2092, 2565, 2604, 2615, 2630, 2631, 2675, 2712, 2716, 4121, 4308, 4763, 6475, 7279, 7551, 7555, 7559, 7645, 7647, 7765, 7766, 7768, 7770, 7773, 7797; MV P202186, P202187. Eryosuchus (Parotosuchus) pronus (mi) (Howie, 1970). Iberospondylus schultzei (Laurin and Soler-Gijón, 2001, 2006). Isodectes (Eobrachyops, Saurerpeton) obtusus (mi) (Watson, 1956; Milner, 1982a; Sequeira, 1998). Specimens examined: AMNH 6919, 6922, 6923, 6934, 6945, 6937, 6947, 6978; USMN 4471 (cast), 4474. Keratobrachyops australis (Warren, 1981a; Damiani and Warren, 1996). Konzhukovia (Melosaurus) vetusta (Konzhukova, 1955a; Gubin, 1991). Laidleria gracilis (Kitching, 1957; Warren, 1998a). Specimens examined: ALM 4313 (cast). Lapillopsis nana (mi) (Warren and Hutchinson, 1990a; Yates, 1999). Luzocephalus spp. (Säve-Söderbergh, 1935; Shishkin, 1980; Warren, 1998a). Specimens examined: MGUH AT1 (cast), PIN 3784/1 (cast). Lydekkerina huxleyi (Watson, 1919; Broom, 1930; Broili and Schröder, 1937; Findlay, 1968; Jupp and Warren, 1986; Shishkin et al., 1996; Jeannot, 2004; Pawley and Warren, 2005). Specimens examined: BP/1/1373, BP/1/5021, BP/1/fn 3900; UMZC T218, T243. Mastodonsaurus giganteus (mi) (Schoch, 1999a, 2000b, 2002a, 2002b). Micropholis stowi (Watson, 1913; Broili and Schröder, 1937; Boy, 1985; Schoch and Rubidge, 2005). Specimens examined: BMNH R510, BP/1/5209, 5571, UCMP 42879. Neldasaurus wrightae (mi) (Chase, 1965). Specimens examined: MCZ 1371, 1463, 2406, 2407, 2516, 2518. Nigerpeton ricqlesi (Sidor et al., 2005; Steyer et al., 2006). Onchiodon (Sclerocephalus) labyrinthicus (Credner, 1893; Boy, 1990; Werneburg, 1993b; Witzmann, 2005). Parioxys spp. (Moustafa, 1955a; Carroll, 1964b). Specimens examined: AMNH 7118, 23568, 23569, 23572, 23573; MCZ 1162, 1290, 1545, 1728, 4457, 5767, 5768, 6496, 6499, 6503, 6540, 6587. Pelorocephalus spp. (mi) (Warren, 1981b; Marsicano, 1993, 1999). Peltobatrachus pustulatus (Panchen, 1959) (Jupp and Warren, 1986). Phonerpeton pricei (Bolt, 1974a; Dilkes, 1990, 1993). Plagiosauridae (mi) (Nilsson, 1937, 1939, 1946a; Jupp and Warren, 1986; Shishkin, 1987; Hellrung, 2003). Platyoposaurus (Platyops) stuckenbergi (Konzhukova, 1955a, 1955b; Gubin, 1991; Shishkin, 2000). Platyrhinops (Amphibamus) lyelli (Carroll, 1964a; Milner, 1982a; Hook and Baird, 1984; Clack and Milner, 1993; Daly, 1994; Carroll et al., 1999; Schoch and Milner, 2004). APPENDIX 6: TEMNOSPONDYL TERMINAL TAXA 271

Rewana (Arcadia) spp. (mi) (Howie, 1972a; Warren and Black, 1985). Specimens examined: AM F54126; QM 10121, 12277, F6471. Rhinesuchidae (van Hoepen, 1915; Broom, 1930; Watson, 1962; Pawley and Warren, 2004) (Schoch, 2000a; Shishkin and Rubidge, 2000). Specimens examined: BP/1/3241; M 460; NMQR 1438; TM 75, 77, 78, 79, 85, 103, 208, 4703; SAM-PK- K100021. Saharastega moradiensis (Sidor et al., 2005; Damiani et al., in press). Sclerocephalus spp. (Broili, 1926; Boy, 1986; Werneburg, 1992; Meckert, 1993; Lohmann and Sachs, 2001; Schoch, 2002c, 2003). Siderops kehli (mi) (Warren and Hutchinson, 1983). Specimens examined: QM F7882. Tersomius spp. (mi) (Carroll, 1964a; Bolt, 1974a, 1974b, 1974c; Daly, 1994). Thabanchuia oomie (mi) (Warren, 1998b). Thoosuchus yakovlev (Damiani and Yates, 2003). Specimens examined: WAM 96.8.1. Trematosauridae (mi) (Jaekel, 1922; Säve-Söderbergh, 1936; Nilsson, 1943; Mazin and Janvier, 1983; Hellrung, 1987; Shishkin and Welman, 1994; Schoch and Milner, 2000; Steyer, 2002; Schoch, 2006). Trimerorhachis insignis (mi) (Cope, 1884; Case, 1911a; Cope and Matthew, 1915; Case, 1935; Nilsson, 1944; Colbert, 1955; Watson, 1956; Olson, 1979; Lombard and Bolt, 1988; Schoch, 1999b). Specimens examined: AMNH 4578, 4720, 4763, 4765, 23283, 23302, 23319, 23321, 23330; MCZ 840, 8116, 8117, 8128, 8164, 8244, 8245, 8312, 8365, 8413, 8515, 8536, 8577, 8611; TMM 40031:61-110, 40998:25, 40998:39; UMCP 105230, 105232, 105236, 105247, 105249, 105258, 105259, 174885-187001, 24009, 24010; UMMP 15995, 16033, 16055. Tupilakosaurus spp. (Nielsen, 1954; Shishkin, 1961; Nielsen, 1967; Shishkin, 1973). Xenobrachyops (Brachyops) allos (Howie, 1972b; Warren, 1981b; Damiani and Warren, 1996). Zatrachydidae (Acanthostomatops vorax (mi) and Zatrachys serratus) (Case, 1911a; Langston, 1953; Boy, 1989; Schoch, 1997). Specimens examined: AMNH 4560; MCZ 1170.

Pawley, K. 2006. 273 The Postcranial Skeleton of Temnospondyls (Tetrapoda: Temnospondyli) PhD Thesis. La Trobe University, Melbourne

APPENDIX 7. REINTERPRETATIONS OF PUBLISHED MATERIAL: THE POSTCRANIAL SKELETON OF TEMNOSPONDYLS

Morphologically immature specimens are noted; all other specimens are morphologically mature because they have ossified coracoid, pubis, carpus, and tarsus (Chapter 4).

Acanthostomatops vorax: The endochondral postcranial skeleton (Boy, 1989) is morphogenetically immature. The clavicle (Boy, 1989: figure 5.B) and interclavicle (Boy, 1989: figure 5.A) are inverted, anterior is to the bottom of the page. Acheloma cumminsi (Trematops milleri): The forelimb elements figured by Case (1911a: figure 34) are morphogenetically immature. Acroplous vorax: The endochondral postcranial skeleton (Hotton, 1959; Coldiron, 1978) is morphogenetically immature. Almasaurus habbazi: The endochondral postcranial skeleton (Dutuit, 1976) is morphogenetically immature. On the humerus, the process labelled as the supinator process (pr.sup) (Dutuit, 1976: figure 103) is reinterpreted as the deltopectoral crest. Amphibamus grandiceps: The specimen figured by Daly (1994: figure 18) is morphogenetically immature. Archegosaurus decheni: The endochondral postcranial skeleton (Meyer, 1857) is morphogenetically immature. Balanerpeton woodi: Some characteristics of the postcranial skeleton (Milner and Sequeira, 1994) are indeterminate due to its state of preservation. Batrachosuchus concordi: The endochondral postcranial skeleton of the specimens examined (UCMP 80859, 80860, 140579, 140580, 140583, 140589-140592) is morphogenetically immature. The postcranial elements described by Chernin (1977: figure 4) belong to a dicynodont. Benthosuchus sushkini: The endochondral postcranial skeleton (Bystrow and Efremov, 1940) is morphogenetically immature. The tibia (Bystrow and Efremov, 1940: figure 55) is interpreted as a left tibia in A, flexor; and B, extensor views. Broiliellus brevis: The humerus (Carroll, 1964a: figure 10) is morphogenetically immature. Caerorhachis bairdi: The area labelled ‘adductor crest’ (add cr) (Ruta et al., 2002: figure 12) is reinterpreted as the fourth trochanter at the distal end of the adductor blade. Cheliderpeton spp.: The endochondral postcranial skeleton (Werneburg and Steyer, 2002) is morphogenetically immature. Chenoprosopus lewisi: The endochondral postcranial skeleton (Hook, 1993: figure 3) is morphogenetically immature. Dendrerpeton acadianum: The endochondral postcranial skeleton of some specimens (Carroll, 1967) is morphogenetically immature, these specimens are readily determinable by their comparatively small size. 274 K. PAWLEY PHD THESIS

Doleserpeton annectans: In accordance with Bolt (1969) the postcranial elements are indistinguishable from those described as Cardiocephalus sternbergi (Gregory et al., 1956). The fibulae (Gregory et al., 1956: figure 30) are inverted. Dutuitosaurus (Metoposaurus) ouazzoui: The endochondral postcranial skeleton of (Dutuit, 1976; Steyer et al., 2004) is morphogenetically immature Dvinosaurus spp.: The endochondral postcranial skeleton (Sushkin, 1936; Bystrow, 1938; Nikitin, 1995, 1997) is morphogenetically immature. Ecolsonia cutlerensis: The proximal half of the humerus (Berman et al., 1985: figure 11.D) is a left rather than a right as in the distal half, and the latissimus dorsi is broken off. The ulnae (Berman et al., 1985: figure 11.E, 11.F) are also of the left side, as is the clavicle (Berman et al., 1985: figure 11.B). Edops craigi: The humerus (Romer and Witter, 1942: figure 13) is not completely morphogenetically mature, in that the proximal humeral ridge is unossified and the radial condyle of the humerus is only partially ossified. Eoscopus lockardi: The endochondral postcranial skeleton of some specimens of the hind limb (Daly, 1994: figure 8, 16) are morphogenetically immature. The interclavicle (Daly, 1994: figure 12) is inverted. Eryops megacephalus: The manus of Eryops has four digits, in agreement with Miner (1925), rather than five as described by Gregory et al. (1923). The sacral rib described by Case (1911a: figure 30) and Moulton (1974: figure 13) is more typical than that in Olson (1936a). Eryosuchus (Parotosuchus) pronus: The endochondral postcranial skeleton (Howie, 1970) is morphogenetically immature. Isodectes (Eobrachyops, Saurerpeton) obtusus: The endochondral postcranial skeleton (AMNH 6919, 6922, 6923, 6934, 6945, 6937, 6947, 6978; USMN 4471 (cast), 4474) is morphogenetically immature. Specimens were used as the source of data, rather than the reconstructions of Sequeira (1998). Konzhukovia (Melosaurus) vetusta: Konzhukovia is interpreted as possessing lateral line sulci on the skull roof, these are not obvious in the figures (Konzhukova, 1955a; Gubin, 1991). Lydekkerina huxleyi: The endochondral postcranial skeleton described by Watson (1919) is morphogenetically immature. Mastodonsaurus giganteus: The ‘radius’ (Schoch, 1999a: figure 44) is reinterpreted as an inverted left fibula in anterior, flexor, posterior and extensor views respectively. The ‘fibula’ (Schoch, 1999a: figure 48) is an inverted subadult right tibia in flexor, posterior, extensor, and anterior views respectively. Most of the specimens of the endochondral postcranial skeleton (Schoch, 1999a) are morphogenetically immature. Neldasaurus wrightae: The endochondral postcranial skeleton (Chase, 1965) is morphogenetically immature. The figure of the tibia (Chase, 1965: figure 13 H) is reinterpreted as the extensor surface of the right tibia. Onchiodon (Sclerocephalus) labyrinthicus: The tarsus described by Credner (1893: Taf XXXII, figure 10) probably does not belong to Onchiodon, as it has an astragalus and calcaneum, not known in any other temnospondyl. Pelorocephalus spp.: The endochondral postcranial skeleton (Marsicano, 1993) is morphogenetically immature. The femur (Marsicano, 1993: figure12), is a right femur in A, extensor; B, flexor; C, posterior; and D, anterior views. The right tibia (Marsicano, 1993: figure13) is in A, extensor; B, posterior; C, flexor; and D, anterior views. APPENDIX 7: TEMNOSPONDYL REINTERPRETATIONS 275

Plagiosauridae: The endochondral postcranial skeleton of plagiosaurs (Nilsson, 1939; Hellrung, 2003) is morphogenetically immature. Platyrhinops (Amphibamus) lyelli: The specimens figured by Carroll (1964a: figure 23, 24), Carroll et al. (1999: figure 4 A, B) are morphogenetically immature. Rewana spp.: Included genera are R. (Arcadia) myriadens (Warren and Black, 1985) and R. quadricuneata (Howie, 1972a). The endochondral postcranial skeletons of both species are morphogenetically immature. The right tibia of R. quadricuneata (Howie, 1972a: figure 5.K) is in flexor view. Platyoposaurus (Platyops) stuckenbergi: Platyoposaurus is interpreted as possessing lateral line sulci on the skull roof, these are not obvious in the figures (Konzhukova, 1955a, 1955b; Gubin, 1991). Rhinesuchidae: Included taxa are Uranocentrodon (Myriodon) senekalensis (Haughton, 1915; van Hoepen, 1915); Broomistega putterilli BP/1/3241; an unnamed rhinesuchid, field no. M460, and SAM-PK-K10021 (Pawley and Warren, 2004). The endochondral postcranial skeletons of Broomistega and SAM-PK-K10021 are morphogenetically immature. Siderops kehli: The endochondral postcranial skeleton (Warren and Hutchinson, 1983), is morphogenetically immature. Thabanchuia oomie: The endochondral postcranial skeleton (Warren, 1998b) is morphogenetically immature. Trematosauridae: The endochondral postcranial skeletons of Aphaneramma rostratum (Nilsson, 1943) and Watsonisuchus elongatus (Steyer, 2002) are morphogenetically immature. Trimerorhachis insignis: The endochondral postcranial skeleton (Pawley, in press) is morphogenetically immature. Zatrachydidae: Included genera are Acanthostomatops vorax and Zatrachys serratus. In Zatrachys, the scapula (1911a: figure 43) is closely resembles that of Trimerorhachis insignis (Pawley, in press), which is known from the same locality. The clavicle (Boy, 1989: figure 5.B) and interclavicle (Boy, 1989: figure 5.A) of Acanthostomatops are inverted, anterior is to the bottom of the page.

Pawley, K. 2006. 277 The Postcranial Skeleton of Temnospondyls (Tetrapoda: Temnospondyli) PhD Thesis. La Trobe University, Melbourne

APPENDIX 8. CHARACTER STATE LIST: THE POSTCRANIAL SKELETON OF TEMNOSPONDYLS

The materials and methods section (Chapter 5) outlines the selection criteria used for determining which character states were included or modified from previous analyses, and for formulating new character states. Character states in plain type (e.g. TRU VER 1) were previously published and their distribution discussed in Ruta et al. (2003). Characters from Yates and Warren (2000) are marked ‘YW’ and include the character number used by Yates and Warren (2000) (e.g. YW 51). Characters used in other previously published analyses (Sequeira and Milner, 1993; Carroll, 1995; Lebedev and Coates, 1995; Coates, 1996; Paton et al., 1999; Anderson, 2001; Damiani, 2001a; Gardner, 2001; Klembara and Ruta, 2004b; Pawley and Warren, 2004; Sequeira, 2004; Clack and Finney, 2005; Pawley and Warren, 2005) have only the numbers in bold (e.g. TRU VER 28). New characters not previously used in phylogenetic analyses of basal tetrapods are entirely in bold (e.g. INTCLA 7.). Many of these characters were also used in the phylogenetic analysis of Chapter 6 (Appendix 14). Postcranial character states subject to morphogenetic development (Chapter 4) are marked ‘m’, which means that the morphogenetic stage of the specimens should be assessed before coding the character states, as the derived state may be absent in a specimen due to its immature morphogenetic stage. Additional notes are provided to clarify character states in cases of potential confusion. Note that character state distributions refer only to the main analysis (Figure 44).

CRANIAL CHARACTERS

Skull

1. SK 1 (YW 21). Snout margins continually converging towards tip (0) tip of snout expanded so that snout margins run parallel, or are concave before tip (1). STATE 1: Archegosauridae.

Ornament

2. L SC SKU 1. Absence (0) or presence (1) of lightly sculptured area along skull midline. STATE 1: Cochleosauridae more derived than Saharastega.

Sensory canals

3. SC SK 1 (YW 17). Sensory canals: absent (0) present (1). [Repolarised and simplified from Ruta et al. (2003).] STATE 1: Nigerpeton, Dvinosauria, Cheliderpeton, Stereospondylomorpha. 278 K. PAWLEY PHD THESIS

4. SC 3 (YW 18). Infraorbital sulcus (where present): straight or gently curved (0) with a step-like or 'Z' shaped flexure between the orbit and the naris (1). STATE 1: some Stereospondyli (Lydekkerina, Mastodonsauroidea plus Thoosuchus, Metoposauroidea, Brachyopoidea).

Nares

5. NOS 3. Absence (0) or presence (1) of condition: nostrils elongate and keyhole shaped. STATE 1: Trematopidae.

Premaxilla

6. PREMAX 1 (YW 5). Absence (0) or presence (1) of alary process. STATE 1: Dendrerpetontidae, some Dvinosauria (Trimerorhachis, Isodectes), Euskelia. 7. PREMAX 4. Premaxillae without (0) or with (1) expanded anteromedial dorsal surface and marginal elongation. STATE 1: Edops, Cochleosauridae more derived than Saharastega. 8. PREMAX 5. Premaxillary tusks: absent (0) present (1). STATE 1: Nigerpeton, Acheloma plus Phonerpeton. 9. PREMAX 6 (YW 88). Rugose, medial tubercle on the palatal surface of the premaxillae, posterior to the tooth row: absent (0) present (1). STATE 1: Saharastega, Dvinosaurus, Archegosauridae plus Konzhukovia.

Septomaxilla

10. SPTMAX 2 (YW 13). Septomaxilla ornamented and part of skull roof (0) or a detached ossification inside nostril (1). [This character was re-written from Ruta et al. (2003) in order to clarify it] State 1: Dissorophoidea.

Maxilla

11. MAX 1 (YW 12). Maxilla-nasal suture: absent (0) present (1). STATE 1: Nigerpeton plus Chenoprosopus spp., Neldasaurus, Actinodontidae, Stereospondylomorpha. 12. MAX 2. Maxillary tusks: absent (0) present (1). STATE 1: some Cochleosauridae (Nigerpeton, Chenoprosopus lewisi), some Trematopidae (Acheloma plus Phonerpeton). 13. MAX 5. Maxilla not entering (0) or entering orbit margin (1). STATE 1: some Amphibamidae (Eoscopus, Platyrhinops, Amphibamus). 14. MAX 8. Absence (0) or presence (1) of condition: maxillary facial process shaped like a rectangular flange. STATE 1: derived Amphibamidae (Amphibamus plus Doleserpeton).

Nasal

15. NAS 3 (YW 10). Ventral flange of nasal: absent (0) present, forming posterior internal wall of the naris (1). STATE 1: Dissorophoidea. APPENDIX 8: TEMNOSPONDYL CHARACTER STATE LIST 279

Prefrontal

16. PREFRO 1 (YW 9). Prefrontal ventral process: absent (0) descending ventral flange (ventral process of the prefrontal- VPP) on the orbit rim (1). STATE 1: Dissorophoidea. 17. PREFRO 8 (YW 27). Absence (0) or presence (1) of prefrontal entering nostril margin. STATE 1: some Dvinosauria (Eobrachyopidae), some Dissorophoidea (Trematopidae). 18. PREFRO 9. Prefrontal not sutured with maxilla (0) or sutured. STATE 1: derived Superstereospondyli (Laidleria, Rewana spp. plus Brachyopoidea).

Lachrymal

19. LAC 1 (YW 3). Presence (0) or absence (1) of lachrymal. STATE 1: derived Superstereospondyli (Laidleria, Rewana spp. plus Brachyopoidea). 20. LAC 2 (YW 7). Lachrymal included in orbital margin, prefrontal-jugal suture absent (0) excluded from orbital margin, prefrontal-jugal suture present (1). [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Edops; Cochleosauridae; basal Euskelia (Actinodontidae plus Onchiodon plus Eryops); Stereospondylomorpha (except Plagiosauridae).

Frontal

21. FRO 4 (YW 4). Frontal excluded from (0) or contributing to (1) margin of orbit. STATE 1: Dissorophoidea (except Platyrhinops plus Amphibamus), some Superstereospondyli (Mastodonsauridae, Plagiosauroidea).

Parietal

22. PAR 2 (YW 1). Absence (0) or presence (1) of parietal-postorbital suture. STATE 1: derived Dvinosauroidea (Dvinosaurus plus Tupilakosauridae).

Pineal foramen

23. PIN FOR 1. Presence (0) or absence (1) of pineal foramen. STATE 1: Cochleosauridae (except Adamanterpeton and Chenoprosopus lewisi).

Postparietal

24. POSPAR 11. Each postparietal with a medial posterior lappet extending as a continuation of the skull roof: absent (0) present (1). STATE 1: Cochleosaurus spp.

Postfrontal

25. POSTFRO 2 (YW 1). Postfrontal-supratemporal suture: absent (0) present (1). STATE 1: Acroplous, Peltobatrachus plus Eryopoidomorpha. 280 K. PAWLEY PHD THESIS

Intertemporal

26. INTEMP 1 (YW 1). Intertemporal present (0) or absent (1). STATE 1: Dvinosauroidea (except Isodectes); Peltobatrachus plus Eryopoidomorpha.

Tabular

27. TAB 2 (YW 24). Absence (0) or presence (1) of subdermal, unornamented blade- like postero-lateral portion of tabular. [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: basal Stereospondylomorpha (Archegosauridae plus Konzhukovia plus Rhinesuchidae). 28. TAB 9 (YW 23). Tabulars: well developed rectangular to triangular bones (0) reduced to thin slivers lying against the posterior margin of the skull roof (1). STATE 1: Dvinosauroidea. 29. TAB 10 (YW 29). Terminal crest of tabular horn: absent (0) present on ventral side of tabular horn, lying lateral and parallel to the paraoccipital process (sensu Bystrow and Efremov, 1940) (1). STATE 1: Mastodonsauroidea plus Trematosauroidea. 30. TAB 11 (YW 44). Tabular - exoccipital contact in paraoccipital process, lateral to post-temporal fossa: absent (0) present (1). STATE 1: Stereospondyli. 31. TAB 12. Tabular horns: absent or posteriorly directed (0) distally enlarged and laterally directed towards a process of the squamosal, partly or fully enclosing the otic notch (1). STATE 1: some Superstereospondyli (Mastodonsauridae, Metoposauroidea). 32. TAB 13. Tabular sutured with quadrate enclosing otic notch: absent (0) present (1). STATE 1: Dissorophidae (except Broiliellus, Phonerpeton).

Squamosal

33. SQU 3 (YW 28). Squamosal with (0) or without (1) broad, concave semicircular embayment. [Repolarised from Ruta et al. (2003).] STATE 1: Dvinosauria, Plagiosauroidea. 34. SQU 6 (YW 14). Posterior squamosal margin in dorsal view: straight or concave (0) convex, bulging posteromedially narrowing the otic notch (1) convex and overhanging (falciform crest) (2). Ordered. STATE 1: some Stereospondylomorpha (Archegosauridae plus Konzhukovia, Thoosuchus plus Luzocephalus). STATE 2: some Stereospondyli (Rhinesuchidae, Lydekkerina, Mastodonsauroidea, Metoposauroidea). 35. SQU 7 (YW 22). Simple rounded or sharp edged posterodorsal margin of the otic notch (0) smooth 'gutter' bordered above and below by sharp ridges around the margin of the otic notch (1). STATE 1: Metoposauroidea. 36. SQU 8. Supratympanic flange of squamosal: absent (0) present (sensu Bolt, 1974c) (1). STATE 1: Dissorophidae. 37. SQU 9. Semi-lunar flange of squamosal: absent (0) present (sensu Bolt, 1974c) (1). STATE 1: Dissorophidae, Eoscopus. APPENDIX 8: TEMNOSPONDYL CHARACTER STATE LIST 281

Jugal

38. JUG 2 (YW 8). Jugal not contributing (0) or contributing (1) to ventral margin of skull roof. STATE 1: Cochleosauridae, Eobrachyopidae, Konzhukovia plus Stereospondyli. 39. JUG 3 (YW 83). Jugal not contacting (0) or contacting (1) pterygoid (alar process of jugal). STATE 1: Cochleosauridae, Dvinosauroidea, Konzhukovia plus Stereospondyli.

Quadratojugal

40. QUAJUG 4 (YW 32). Posterior border of quadratojugal drawn out into spike: absent (0) present (1). STATE 1: Chigutisauridae. 41. QUAJUG 5 (YW 35). Quadratojugal forms a simple corner with the quadrate in occipital view (0) a sulcus present on the quadratojugal, lateral to the quadrate condyles, so that the quadratojugal forms an overhang in occipital view (1). STATE 1: Plagiosauroidea. 42. QUAJUG 6 (YW 42). Posterior medial process of quadratojugal: absent (0) extending posterior to, and wrapping around, the dorsal process of the quadrate (1). STATE 1: Dissorophoidea 43. QUAJUG 7 (YW 43). Paraquadrate foramen on the occipital face of the quadratojugal: present (0) absent (1). STATE 1: some Cochleosauridae (Nigerpeton, Cochleosaurus spp.), Euskelia more derived than Actinodontidae.

Quadrate

44. QUA 1 (YW 41). Quadrate without (0) or with (1) enlarged dorsal process. STATE 1: Dissorophoidea.

Post-temporal fossa

45. PTF 1 (YW 46). Fossa near dorsolateral corner of occiput, roofed over by occipital flanges of tabular and postparietal sometimes bordered laterally and ventrally by ossified dorsolateral extension of opisthotic meeting ventromedial flange of tabular (0) absence of fossa (1). [This character was revised from Ruta et al. (2003) following Clack (2003a)] STATE 1: Plagiosauroidea.

Vomer

46. VOM 2 (YW 60). Vomers: form a large sheet of bone between the anterior end of the cultriform process and the posterior end of the anterior palatal fossa (0) forming a narrow bar between the anterior end of the cultriform process and the posterior edge of the anterior palatal fossa (1). STATE 1: Dvinosauria more derived than Neldasaurus. 47. VOM 3. Vomer with (0) or without (1) fang pair. STATE 1: derived Amphibamidae (Amphibamus plus Doleserpeton). 48. VOM 4 (YW 77). Vomer with (0) or without (1) denticles. [Repolarised from Ruta et al. (2003).] STATE 1: Dvinosauria, Nigerpeton, Actinodontidae, Superstereospondyli (except Rewana spp. plus Brachyopoidea). 282 K. PAWLEY PHD THESIS

49. VOM 5. Vomer excluded from (0) or contributing to (1) interpterygoid vacuities. STATE 1: derived Dendrerpetontidae (Dendrerpeton plus Balanerpeton), Dvinosauria, Eryopoidomorpha (except Iberospondylus plus Parioxys). 50. VOM 6 (YW 63). Vomers: without a row of teeth between the vomerine tusks (0) with a tooth row running between the vomerine tusks (1). STATE 1: Saharastega, Trimerorhachis, Dvinosauroidea, Konzhukovia plus most Stereospondyli. 51. VOM 7 (YW 64). Vomer not forming (0) or forming (1) suture with maxilla anterior to choana. STATE 1: some Cochleosauridae (Nigerpeton plus Adamanterpeton plus Cochleosaurus floriensis), Dvinosauria, Stereospondylomorpha. 52. VOM 10. Vomer in contact with anterior ramus of pterygoid (0) or not (1). STATE 1: some Dvinosauria (Eobrachyopidae, Tupilakosauridae), Amphibamidae, Konzhukovia plus Stereospondyli. 53. VOM 13 (YW 76). Vomer-palatine suture lateral to choana: absent (0) present (1). STATE 1: some Dvinosauria (Neldasaurus, Eobrachyopidae, Tupilakosauridae); derived Superstereospondyli (Plagiosauroidea plus Rewana spp. plus Brachyopoidea). 54. VOM 14 (YW 79). Tooth row along the medial margin of the choana: absent (0) present (1). STATE 1: derived Dvinosauria (Trimerorhachis, Dvinosauroidea), Stereospondyli (except Laidleria, Brachyopidae). 55. VOM 15. Anterior part of cultriform process of parasphenoid clasped by medial posterior extensions of the vomers: absent (0) present (1). STATE 1: Eryopoidomorpha (except Amphibamidae, Archegosauridae). 56. VOM 16. Vomerine tusks: half the width of palatal tusks or less (0) subequal in diameter to palatal tusks (1). STATE 1: Dvinosauria except Isodectes, Stereospondylomorpha. 57. VOM 17. Vomerine ridges radiating from anterior edge of choanae to snout margins: absent (0) present (1). STATE 1: Edops, derived Cochleosauridae (Adamanterpeton plus Cochleosaurus spp.).

Palatine

58. PAL 1. Palatine with (0) or without (1) tusks. [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: derived Amphibamidae (Amphibamus plus Doleserpeton). 59. PAL 2. Palatine (and ectopterygoid): with (0) or without (1) denticles. [This character was merged with ECT 3 from Ruta et al. (2003) because they are not independent, and repolarised.] STATE 1: some Cochleosauridae (Nigerpeton, Chenoprosopus lewisi); Dvinosauria, Actinodontidae, Superstereospondyli (except Rewana spp. plus Brachyopoidea). 60. PAL 3 (YW 51). Palatine excluded from (0) or contributing to (1) interpterygoid vacuities. STATE 1: some Dvinosauria (Eobrachyopidae, Tupilakosauridae); Amphibamidae; Lydekkerina plus Superstereospondyli. 61. PAL 4 (YW 78). Palatine without (0) or with (1) tooth row (distinct from tusks). [This character was re-written from Ruta et al. (2003) in order to clarify it, and repolarised.] STATE 1: some Dvinosauria (Neldasaurus plus Trimerorhachis); Actinodontidae; Stereospondylomorpha (except Brachyopidae). APPENDIX 8: TEMNOSPONDYL CHARACTER STATE LIST 283

62. PAL 5 (YW 2 & 52). Palatine without (0) or with (1) exposure in lateral orbit margin (LEP). STATE 1: Eobrachyopidae, Dissorophoidea (except Platyrhinops), Xenobrachyops. 63. PAL 8 (YW 69). Posterior process of palatine: absent (0) present, makes point contact with the palatine ramus of the pterygoid and excludes ectopterygoid from border of interpterygoid vacuity (1). STATE 1: Mastodonsauroidea plus Trematosauroidea (except Trematosauridae).

Ectopterygoid

64. ECT 2 (YW 58). Ectopterygoid with (0) or without (1) tusks. [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Stereospondylomorpha (except Trematosauridae, Rewana spp. plus Brachyopoidea). 65. ECT 5 (YW 59). Ectopterygoid without (0) or with (1) tooth row (distinct from tusks). [This character was re-written from Ruta et al. (2003) in order to clarify it, and repolarised.] STATE 1: Dvinosauria (except Eobrachyopidae, Thabanchuia), Actinodontidae, Stereospondylomorpha (except Brachyopidae). 66. ECT 8 (YW 82). The strut of bone that separates the interpterygoid vacuity from the subtemporal fossa consisting entirely of the palatine ramus of the pterygoid (0) invasion of this strut by the ectopterygoid (1). STATE 1: derived Superstereospondyli (Metoposauroidea plus Plagiosauroidea plus Rewana spp. plus Brachyopoidea).

Pterygoid

67. PTE 1 (YW 37). Ascending column of pterygoid: absent (0) present, ascending lamina of the pterygoid with a thickened ascending column on its posterior surface directly anterior to exoccipital (1). STATE 1: some Superstereospondyli (Mastodonsauroidea plus Trematosauroidea, Chigutisauridae). 68. PTE 2 (YW 40). Oblique ridge of pterygoid: absent (0) posterior face of ascending lamina of the pterygoid with a low rounded oblique ridge or large sharp edged crest (1). STATE 1: Iberospondylus, basal Stereospondyli: Rhinesuchidae, Lydekkerina, Mastodonsauroidea plus Trematosauroidea (except Trematosauridae), Metoposauroidea. 69. PTE 5 (YW 45). Quadrate ramus of pterygoid gently concave posteriorly, in horizontal section (0) medial end of the quadrate ramus strongly curved posteriorly in horizontal section (1). STATE 1: derived Chigutisauridae (Siderops plus Keratobrachyops) 70. PTE 6 (YW 47). Quadrate ramus of pterygoid erect (0) dorsal end of quadrate ramus posteriorly curved (1). STATE 1: Brachyopoidea. 71. PTE 7. Quadrate (posterior) ramus of pterygoid laterally oriented in ventral view, rather than posterolaterally oriented: absent (0) or present (1). [This character was re- written from Ruta et al. (2003) in order to clarify it.] STATE 1: Chigutisauridae. 72. PTE 9 (YW 56). Pterygoid with (0) or without (1) postero-lateral flange. [Repolarised from Ruta et al. (2003).] STATE 1: Dvinosauria, Stereospondyli. 284 K. PAWLEY PHD THESIS

73. PTE 13 (YW54). Parasphenoid articulates with depression in the corpus of the pterygoid, behind a short triangular medial process of the pterygoid (0) an elongate cylindrical to hemi-cylindrical process medial process of the pterygoid abuts the parasphenoid (1) corpus of the pterygoid forming a broad contact along the lateral margins of the parasphenoid plate (2). STATE 1: all Temnospondyli more derived than Caerorhachis plus Edops (except Temnospondyli possessing state 2). STATE 2: Tupilakosauridae, Stereospondyli. 74. PTE 14. Absence (0) or presence (1) of condition: quadrate ramus of pterygoid robust, indistinctly merging into basal and palatal processes. STATE 1: Dvinosauroidea, Brachyopoidea. 75. PTE 15. Absence (0) or presence (1) of condition: quadrate ramus of pterygoid straight, rod-like and gently tapering distally. STATE 1: derived Amphibamidae (Amphibamus plus Doleserpeton). 76. PTE 18 (YW 48). Quadrate (posterior) ramus of the pterygoid: twisted dorsally from the plane of the corpus and the palatine ramus, to form a subvertical plate (0) untwisted, forming a near horizontal plate, continuous with the plane of the corpus and the palatal ramus of the pterygoid (1) strongly turned ventrally relative to the palatal ramus, creating a vaulted palate (2). Ordered. STATE 1: Plagiosauroidea. STATE 2: Dvinosauroidea, Rewana spp. plus Brachyopoidea. 77. PTE 19 (YW 49). Substapedial ridge of pterygoid (sensu Warren and Hutchinson, 1983): absent (0) present on the dorsal surface of the quadrate ramus of the pterygoid (1). STATE 1: Chigutisauridae. 78. PTE 20 (YW 53). Palatine ramus of the pterygoid: extends anterior to the anterior most ectopterygoid tooth (0) retracted posterior to the anterior most ectopterygoid tooth (1). STATE 1: Tupilakosauridae, Amphibamidae, Lydekkerina plus Superstereospondyli. 79. PTE 21 (YW 86). Ornament on palatine ramus of pterygoid: absent (0) present (1). STATE 1: Peltobatrachus, Lydekkerina, Mastodonsauroidea (except Mastodonsaurus) plus Trematosauroidea, Dutuitosaurus, Laidleria, Rewana spp., Keratobrachyops. 80. PTE 22 (YW 87). Denticles on palatine ramus of pterygoid: present (0) absent (1). STATE 1: Nigerpeton, derived Dvinosauria (Dvinosaurus plus Tupilakosauridae), Mastodonsaurus, Metoposauroidea, Plagiosauroidea, Brachyopoidea (except Keratobrachyops). 81. PTE 23. Dorsoventrally oriented ridge on medial surface of ascending ramus of pterygoid: absent (0) present (1). STATE 1: Dvinosauria. 82. PTE 24. Lateral pterygoid alae, extended posterior to ectopterygoid: present (0) absent (1). STATE 1: Cochleosaurus bohemicus, derived Dvinosauria (Trimerorhachis plus Dvinosauroidea), Eryopoidomorpha. 83. PTE 25. Denticulate vomerine-pterygoid ridge passes from vomerine tusk to medial edge of interpterygoid vacuity: present (0) absent (1). STATE 1: Dvinosauria plus Eryopoidomorpha.

Epipterygoid

84. EPI 1 (YW 85). Ascending ramus of epipterygoid: slender rod (0) robust (1). STATE 1: some Superstereospondyli (Mastodonsauroidea plus Trematosauroidea, Metoposauroidea). APPENDIX 8: TEMNOSPONDYL CHARACTER STATE LIST 285

Choanae

85. CHO 1. Absence (0) or presence (1) of condition: choanae wider anteriorly than posteriorly. STATE 1: Cochleosauridae more derived than Saharastega.

Anterior palatal fossa

86. ANT FOS 1. Anterior palatal fossa (defined as a depression in the anterior part of the vomers, which may or may not be perforated by an anterior palatal vacuity, see ANT VAC 1): absent (0) present (1). STATE 1: some Cochleosauridae (Chenoprosopus, Cochleosaurus), Eryopoidomorpha (except derived Amphibamidae).

Anterior palatal vacuities

87. ANT VAC 1 (YW 50). Absence (0) or presence (1) of anterior palatal vacuities (defined as perforation/s of the anterior part of the vomers). [Repolarised from Ruta et al. (2003).] STATE 1: Saharastega, Nigerpeton, Dvinosauria, Zatrachydidae, Platyoposaurus, Lydekkerina plus Superstereospondyli (except Plagiosauroidea). 88. ANT VAC 2. Anterior palatal vacuity (where present) single (0) or double (1). [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Saharastega, Nigerpeton, Dvinosauria, Platyoposaurus, Mastodonsaurus, Trematosauroidea, Metoposauroidea, Laidleria.

Interpterygoid vacuities

89. INT VAC 2. Absence (0) or presence (1) of condition: interpterygoid vacuities occupying at least half of palatal width. STATE 1: all Temnospondyli more derived than Caerorhachis plus Edops. 90. INT VAC 3 (YW 75). Absence (0) or presence (1) of condition: interpterygoid vacuities concave along their entire margins. STATE 1: all Temnospondyli more derived than Caerorhachis plus Edops (except Tupilakosauridae).

Parasphenoid

91. PASPHE 2 (YW 39). Pedicels of occipital condyles: not covered by parasphenoid (0) covered in ventral view by a posterior extension of the parasphenoid plate (1). STATE 1: Trematosauroidea. 92. PASPHE 4 (YW 62). Foramen for the internal carotid artery on the ventral or lateral surface of the parasphenoid plate (0) internal carotid and the palatine and intracranial branches pass through the dorsal surface of the parasphenoid plate (1). STATE 1: Stereospondyli. 93. PASPHE 5 (YW 71). Denticles on parasphenoid: confined to medial area of parasphenoid (0) transverse belt extending to parasphenoid-pterygoid articulations (1) absent (2). STATE 1: some Stereospondyli (Rhinesuchidae plus Lydekkerina, Benthosuchus plus Trematosauroidea, Rewana spp). STATE 2: Dvinosauria, Dissorophidae (except Ecolsonia), most Superstereospondyli (Mastodonsauridae, Metoposauroidea, Plagiosauroidea, Chigutisauridae). 286 K. PAWLEY PHD THESIS

94. PASPHE 7 (YW 67). Posterior end of the ventral surface of the parasphenoid plate without sharp rimmed depressions (0) rounded, widely separated depressions at the posterior end of the ventral surface of the ventral surface of the parasphenoid plate, with sharp anterior rims (crista muscularis) (1) depressions transversely widened so that the muscular crests approach each other forming transverse ridges (2). Ordered. [This character was re-written from Ruta et al. (2003) in order to clarify it and extend it to include characters suitable for temnospondyls.] STATE 1: Cochleosauridae, Peltobatrachus plus basal Euskelia (Actinodontidae, Zatrachydidae, Eryops), basal Stereospondylomorpha (Archegosauridae plus Konzhukovia plus Rhinesuchidae plus Lydekkerina, Trematosauroidea except Trematosauridae). STATE 2: Mastodonsauroidea. 95. PASPHE 8 (YW 72). Parasphenoid plate anteriorly expanded so that it is much wider anteriorly than posteriorly: absent (0) present (2). STATE 1: Eobrachyopidae. 96. PASPHE 11. Parasphenoid without (0) or with (1) anterolateral extensions meeting pterygoids and projecting anterior to cultriform process insertion. [This character was re- written from Ruta et al. (2003) in order to clarify it.] STATE 1: Amphibamidae. 97. PASPHE 12. Parasphenoid without (0) or with (1) triangular denticle patch with raised margins at base of cultriform process. STATE 1: Dendrerpetontidae, Cochleosaurus, Amphibamidae, Archegosauridae.

Exoccipital

98. EXOCC 1 (YW 70). Exoccipital-pterygoid suture: absent or contact not visible ventrally (0) visible in ventral view (1). STATE 1: Tupilakosauridae, Lydekkerina plus Superstereospondyli (except Trematosauroidea). 99. EXOCC 3 (YW34). Occipital condyle: single with the basioccipital forming the largest contribution to the articulating surface (0) bilobed with reduced basioccipital contribution (1) double with no basioccipital contribution (2). Ordered. [This character was re-written from Ruta et al. (2003) in order to clarify it and extend it to include characters suitable for temnospondyls.] STATE 1: Eryopoidomorpha (except derived Eryopoidomorpha possessing state 2). STATE 2: Lydekkerina plus Superstereospondyli. 100. EXOCC 5. Absence (0) or presence (1) of condition: exoccipitals expanded and adpressed to each other, so as to obliterate basioccipital posterior surface. STATE 1: Dissorophoidea.

MANDIBLE

101. MAND 1 (YW 89). Retroarticular process: absent (0) present (1). STATE 1: Dvinosauria, Peltobatrachus, Lydekkerina plus Superstereospondyli. 102. MAND 2 (YW 90). Arcadian groove along suture between surangular and angular (sensu Jupp and Warren, 1986): present at the posteroventral end of the mandible (0) absent (1). STATE 1: some derived Superstereospondyli (Dutuitosaurus, Plagiosauridae, Brachyopoidea). 103. MAND 3 (YW 99). Postglenoid area with a transverse trough behind the glenoid socket followed by a dorsally bulging tip: absent (0) present (1). STATE 1: derived Dvinosauria (Dvinosaurus plus Tupilakosauridae). APPENDIX 8: TEMNOSPONDYL CHARACTER STATE LIST 287

Dentary

104. DEN 2 (YW 94). Dentary with (0) or without (1) anterior fang pair. STATE 1: Amphibamidae.

Splenial

105. SPL 1. Splenial: forms part of jaw symphysis (0) excluded from symphysis (1). STATE 1: Edops, Chenoprosopus spp., all Temnospondyli more derived than Cochleosauridae.

Prearticular

106. PREART 1 (YW 91). Prearticular: not extending posterior to the level of the glenoid (0) extending posterior to the level of the glenoid, covering the medial face of the articular (1). STATE 1: derived Superstereospondyli (Trematosauroidea, Metoposauroidea plus Plagiosauroidea plus Rewana spp. plus Brachyopoidea). 107. PREART 5. Prearticular sutured with splenial (0) or not (1). STATE 1: Euskelia (except Parioxys), Lydekkerina plus Superstereospondyli). 108. PREART 6 (YW 102. Hamate process projecting dorsally immediately in front of the antero-medial corner of the glenoid of the mandible: absent (0) present (1). STATE 1: Mastodonsauridae.

Anterior coronoid

109. ANT COR 3 (YW 95). Anterior coronoid with (0) or without (1) denticles. STATE 1: Superstereospondyli (except Plagiosauridae, Rewana spp).

Middle coronoid

110. MID COR 3 (YW 95). Middle coronoid with (0) or without (1) denticles. STATE 1: Superstereospondyli (except Plagiosauridae, Rewana spp., Siderops, Pelorocephalus, Xenobrachyops).

Posterior coronoid

111. POST COR 5. Posterior coronoid with (0) or without (1) a posterodorsal process, which is exposed in lateral view and contributes to the surangular crest. [This character was merged with POST COR 6 and POST COR 7 from Ruta et al. (2003) because they are not independent, and repolarised from Ruta et al. (2003).] STATE 1: Lydekkerina plus Superstereospondyli (except Trematosauridae, Keratobrachyops, Batrachosuchus).

Teeth

112. TEETH 1. Absence (0) or presence (1) of pedicely on marginal teeth. STATE 1: derived Amphibamidae (Amphibamus plus Doleserpeton). 113. TEETH 2. Marginal teeth monocuspid (0) or multicuspid (1). STATE 1: derived Amphibamidae (Platyrhinops plus Amphibamus plus Doleserpeton). 288 K. PAWLEY PHD THESIS

114. TEETH 3. Marginal teeth without (0) or with (1) two cuspules labiolingually arranged. STATE 1: derived Amphibamidae (Platyrhinops plus Amphibamus plus Doleserpeton).

POSTCRANIAL CHARACTERS

Vertebra

Neural spine

115. TRU VER 32. Lateral tubercles on neural spine: absent (0) or present (1). STATE 1: some Euskelia (Parioxys plus Iberospondylus, Ecolsonia).

Centra

116. TRU VER 33. Trunk centra: diplospondylous, two large components per vertebral centra (rhachitomous or embolomerous) (0) essentially monospondylous, pleurocentra dominant, intercentra highly reduced or absent (gastrocentrous) (1) essentially monospondylous, intercentra dominant, pleurocentra highly reduced (stereospondylous) (2). STATE 1: Peltobatrachus, Doleserpeton. STATE 2: some Superstereospondyli (Mastodonsaurus, Dutuitosaurus).

Pleurocentra

117. m TRU VER 8. A majority of trunk pleurocentra not fused mid ventrally (0) or fused (1). [N.B. The derived state of this character develops with morphogenesis; consequently it is indeterminate in morphogenetically immature specimens in which the overall degree of ossification of the postcranial skeleton is low. This character was re- written from Ruta et al. (2003) in order to clarify it.] STATE 1: Caerorhachis, Tupilakosauridae, Peltobatrachus, derived Amphibamidae (Amphibamus plus Doleserpeton). 118. m TRU VER 9. A majority of trunk pleurocentra not fused dorsally (0) or fused (1). [N.B. The derived state of this character develops with morphogenesis; consequently it is indeterminate in morphogenetically immature specimens in which the overall degree of ossification of the postcranial skeleton is low. This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Tupilakosauridae, Peltobatrachus.

Intercentra

119. m TRU VER 14 (YW 104). A majority of trunk anterior intercentra not fused mid dorsally (0) fused (1). [N.B. The derived state of this character develops with morphogenesis; consequently it is indeterminate in morphogenetically immature specimens in which the overall degree of ossification of the postcranial skeleton is low. This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Tupilakosauridae, some Superstereospondyli (Mastodonsaurus, Dutuitosaurus). APPENDIX 8: TEMNOSPONDYL CHARACTER STATE LIST 289

Cervical vertebrae

120. m CER VER 1. Atlantal neural spines not fused (0) or dorsally fused (1). [N.B. The derived state of this character develops with morphogenesis; consequently it is indeterminate in morphogenetically immature specimens in which the overall degree of ossification of the postcranial skeleton is low.] STATE 1: Superstereospondyli. 121. m CER VER 2. Atlantal neural arch and atlantal centrum: separate (0) fused (1). [N.B. The derived state of this character develops with morphogenesis; consequently it is indeterminate in morphogenetically immature specimens in which the overall degree of ossification of the postcranial skeleton is low.] STATE 1: Dissorophoidea where known, Stereospondylomorpha. 122. CER VER 10. Atlantal neural spine with prezygapophyses, which articulate with proatlas: present (0) absent (1). STATE 1: Superstereospondyli. 123. CER VER 11. Atlantal neural spine with postzygapophyses, which articulate with axis: present (0) absent (1). STATE 1: Superstereospondyli. 124. CER VER 12. Axial neural spine with prezygapophyses, which articulate with atlas: present (0) absent (1). STATE 1: Superstereospondyli.

Ribs

125. m RIB 5. Anterior trunk ribs with uncinate processes: absent (0) or present (1). [N.B. The derived state of this character develops with morphogenesis; consequently it is indeterminate in morphogenetically immature specimens in which the overall degree of ossification of the postcranial skeleton is low. This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Peltobatrachus plus Eryopoidomorpha (except Amphibamidae, Plagiosauridae).

Interclavicle

126. INTCLA 1. Posterior border of interclavicle drawn out into parasternal process: absent (0) present (1). STATE 1: derived Dvinosauria (Dvinosaurus plus Tupilakosauridae), some Stereospondyli (Lydekkerina, Mastodonsaurus, Trematosauridae, Chigutisauridae). 127. INTCLA 7. Posterior border of interclavicle broadly hemispherical: absent (0) present (1). STATE 1: derived Euskelia (Zatrachydidae, Eryops, Dissorophoidea), except Platyrhinops plus Amphibamus. 128. INTCLA 8. Parasternal process broadly truncate: absent (0) present (1). STATE 1: Chigutisauridae. 129. INTCLA 9. Location of center of ossification of interclavicle: anterior to or level with line of maximum width (0) posterior to line of maximum width (1). STATE 1: Stereospondylomorpha.

Clavicle

130. CLA 3 (YW 110). Clavicular blades broadly contact anterior to main body of interclavicle: absent (0) present (1). [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Superstereospondyli (except Trematosauridae). 290 K. PAWLEY PHD THESIS

131. CLA 4 (YW 113). Shelf-like, anteroposteriorly-oriented keel along the anteroventral margin of the dorsal clavicular process, separating the ornamented ventral surface from the unornamented dorsal clavicular process: absent (0) present (1). STATE 1: Metoposauroidea. 132. m CLA 5 (YW 112). Anterior clavicular flange present on dorsal clavicular process: absent (0) present (1). [N.B. this character is subject to morphogenetic change; where present, it is observable only in later morphogenetic stages, and is indeterminate in early morphogenetic stages.] STATE 1: some Dvinosauria (Trimerorhachis, Thabanchuia), basal Superstereospondyli (Mastodonsauroidea, Metoposauroidea, Plagiosauroidea). 133. CLA 7. Branchial lamina / clavicular groove (sensu Warren and Turner, 2004): present (0) or absent, ventral anterior border ornamented and not recessed, similar to rest of clavicle (1). STATE 1: Dvinosauria where observable (Trimerorhachis, Thabanchuia), basal Stereospondyli (Lydekkerina plus Superstereospondyli except Plagiosauridae and Brachyopoidea). 134. CLA 8. Clavicular recess (sensu Pawley and Warren, 2006) indenting posterior lamina: absent (0) present (1). STATE 1: derived Euskelia (Eryops, Dissorophus plus Ecolsonia).

Cleithrum

135. CLE 4. Dorsal cleithral process: caps anterodorsal edge of scapulocoracoid (0) does not cap scapulocoracoid (1). STATE 1: Amphibamidae. 136. CLE 5. Dorsal cleithral process: not posteriorly extended (0) posteriorly extended to match length of shaft, covers entire dorsal surface of scapula blade (1). STATE 1: Dissorophidae. 137. CLE 7. Sharp edged cleithral crest (sensu Bystrow and Efremov, 1940) on anterodorsal surface of dorsal cleithral process: present (0) highly reduced or absent (1). STATE 1: Stereospondyli. 138. CLE 8. Ventral junction of dorsal cleithral process and cleithral shaft: smoothly rounded or grooved (0) suprascapula lamina (sensu Bystrow and Efremov, 1940) present on lateroventral surface of dorsal cleithral process, clasps anterodorsal surface of scapular blade (1). STATE 1: Peltobatrachus plus Eryopoidomorpha.

Scapulocoracoid

139. m SCACOR 5 (YW 111). Supraglenoid foramen of scapulocoracoid: enclosed (0) ventrally open notch (1). [N.B. The derived state of this character can only be observed in morphogenetically immature specimens lacking a squared off dorsal scapular blade and/or an ossified glenoid fossa and coracoid plate. The supraglenoid foramen becomes enclosed with morphogenesis, so that the derived state is indeterminate if only morphogenetically mature specimens are available.] STATE 1: Superstereospondyli. 140. m SCACOR 9. Scapular tubercle (sensu Nikitin, 1997): absent (0) present (1). [N.B. The derived state of this character develops with morphogenesis; consequently it is indeterminate the earliest morphogenetic stages in which the scapula is lunate and the glenoid fossa and coracoid plate are unossified.] STATE 1: Dvinosauria where observable (Trimerorhachis, Dvinosaurus). APPENDIX 8: TEMNOSPONDYL CHARACTER STATE LIST 291

Humerus

141. m HUM 2 (YW 114). Distinct supinator process projecting anteriorly: present (0) absent (1). [N.B. The derived state of this character develops with morphogenesis; consequently it is indeterminate in morphogenetically immature specimens lacking an ossified radial condyle. Repolarised from Ruta et al. (2003).] STATE 1: Amphibamidae. 142. HUM 5. Entepicondylar foramen: present (0) or absent (1). STATE 1: Edops, all temnospondyls more derived than Dendrerpetontidae. 143. HUM 12. Humerus slender and elongate, with length less (0) or more (1) than three times the diameter of its distal end. STATE 1: derived Amphibamidae (Platyrhinops plus Amphibamus plus Doleserpeton). 144. HUM 21. Deltopectoral crest of humerus more distal than latissimus dorsi process (0) deltopectoral crest and latissimus dorsi process equidistant from head (1). STATE 1: Euskelia. 145. m HUM 25. Latissimus dorsi process of humerus: large knob on the dorsal surface of humerus (0) small inconspicuous knob or absent (1). [N.B. The latissimus dorsi process may be absent in early morphogenetic stages, due to the low degree of ossification of the proximal humerus. This character develops with morphogenesis; a well-developed latissimus dorsi process may only be present in the largest and more ossified specimens.] STATE 1: Stereospondyli.

Ulna

146. ULNA 2. Posterolateral ulnar crest (sensu Pawley and Warren, 2006) absent (0) present, lateral to posterior ulnar crest (1). STATE 1: Eryopoidomorpha where observable.

Pelvic girdle

Ilium

147. ILI 2. Post-iliac process: projecting distinctly posterior to dorsal iliac process and/or sacral rib facet: present (0) or absent, posterior border of sacral rib facet forms posterior border of dorsal iliac process (1). STATE 1: all Temnospondyli more derived than Dendrerpetontidae. 148. ILI 8. Mesial iliac ridge (sensu Coates, 1996): passes down the midline of the ilium, so that there is a recess present both anterior and posterior to the ridge (0) forms the anterior border of the ilium, with no anterior recess (1). [N.B. the mesial ridge is observable in medial view.] STATE 1: Peltobatrachus plus Eryopoidomorpha.

Femur

149. m FEM 6. Adductor crest of femur approximately as deep as femoral shaft (0) reduced to low ridge (1). [N.B. the adductor crest develops with morphogenesis, and may be low and almost absent in morphogenetically immature specimens. The derived state is relevant where it is present in specimens with an internal trochanter distinctly separated from the proximal articulation surface.] STATE 1: Dvinosauria. 292 K. PAWLEY PHD THESIS

150. FEM 8. In ventral view, adductor blade and crest aligned, pass diagonally across flexor surface of femur towards fibular condyle (0) adductor blade set at an angle to adductor crest, which passes down the midline of the flexor surface of the femur (1). STATE 1: Eryopoidomorpha.

Fibula

151. FIB 5. In lateral view, posterior border of fibular shaft between proximal and distal ends: straight / concave (0) convex (1). STATE 1: Euskelia. 152. FIB 6. Mid shaft area of flexor surface of fibula: convex (0) or concave (1). STATE 1: Eryopoidomorpha.

Dermal ossifications

153. m DERM 1 (YW 120). Enlarged dermal osteoderms: absent (0) present (1). [N.B. Dermal ossifications develop with morphogenesis, they are absent in specimens which exhibit a low overall degree of ossification of the postcranial skeleton.] STATE 1: Edops, Trimerorhachis, Peltobatrachus, Dissorophidae, Plagiosauroidea. 154. m DERM 2. Cycloid dorsal scales: overlapping (0) distinctly separate (1). [N.B. Dorsal scales develop with morphogenesis, they may be absent in morphogenetically immature specimens.] STATE 1: Stereospondylomorpha.

Pawley, K. 2006. 293 The Postcranial Skeleton of Temnospondyls (Tetrapoda: Temnospondyli) PhD Thesis. La Trobe University, Melbourne

APPENDIX 9. DATA MATRIX: THE POSTCRANIAL SKELETON OF TEMNOSPONDYLS

Inapplicable states (-) are not recognised by PAUP 4.0b10 [which treats them as (?)], but they are included here to distinguish from data that is actually missing (?). Note that many of the characters from Ruta et al. (2003) have been recoded.

Acheloma cumminsi 1 000-110101 010?111000 1000110000 0100011000 01?1000010 51 0000100000 010000?000 0010000000 0?00010-10 0?20000011 101 000?10?0?? ?000000000 0?00?0?-?? ????01??00 010???1101 151 ????

Acroplous vorax 1 001-00000? 0000001000 0000110100 0010000110 0000010110 51 1110010011 0100000000 011102?00? 110?001110 0020100000 101 1??01??0?0 000000000? ????000-00 ????0000?? ?10??????? 151 ????

Adamanterpeton ohioensis 1 010-001000 0000000001 000?00000? 000000011? 0????00000 51 1000001000 000000??00 001000?000 ??1?100-10 ??01000??? 101 0?0?000000 ?000?????? ?????????? ?????????? ?????????? 151 ????

Almasaurus habbazi 1 001100000? 1000000001 0000110001 10021001?0 0000000111 51 1101110011 1001110100 0120000101 0101011110 0120000120 101 ???0111011 100000000? ?????00-1? 1?10?????? ?10?1???01 151 ????

Amphibamus grandiceps 1 000-01000? 0011110000 000011?000 0000000000 0111?01010 51 01000-0101 0?0000?000 001010?100 ?100000-10 0000011011 101 0?0????0?? ?111001?0? ?????00-00 ?0??1?0?0? ?11???1??? 151 ??00

Archegosaurus decheni 1 101?000010 1000000001 000011100? 0001000000 00?0000010 51 1000010000 100110??00 001000?000 ?100010-10 0001001010 101 0000110000 000000000? ?????00-00 ?0?0000??0 ?10???11?? 151 0?01 294 K. PAWLEY PHD THESIS

Balanerpeton woodi 1 000-01000? 0000000000 0000000000 0000000000 00?0000010 51 0000000000 000000??00 0010000000 ?010000-10 0000001000 101 00000??00? ?000000000 ?00?000-00 ?0???0???? ?00???000? 151 ??0?

Batrachosuchus spp. 1 001?000000 100000011- 0000110001 0000000100 00?0000?11 51 11101100?1 0?00010001 0121020101 010?0?1010 0120000120 101 1100?11011 00000000?1 111??????? 0000?????? 01001??1?? 151 ????

Benthosuchus sushkini 1 0011000?0? 1?00000001 0000110011 0002000110 0000000111 51 1101110011 1011101100 0120000110 0101011010 0112000120 101 1000101011 100000??0? ????100-11 0110001110 0100111101 151 01??

Broiliellus brevis 1 000-0??0?? 000?110000 100011000? 0000011000 01?1?00010 51 0?00???00? 01??00?000 001000??00 ?10?010-10 0020000??? 101 0?0??????? ?00000??0? ?????????? ????0101?? ?10????10? 151 ??1?

Caerorhachis bairdi 1 000-?????? ?0?0?????? ?00000000? 00?0????00 ?0???00000 51 ?00000?000 0?0000??00 000000?000 ?0??00??00 0?00000000 101 0?00?00000 0000001000 00?00????? ?????????? ??????00?? 151 ????

Capetus palustris 1 000-010000 0000000000 00000000?? 0000000000 00?0?00000 51 ?0?0000000 ?0??????00 001000?000 ??1??00-10 0?0?0??000 101 0?0??????? 0000?????? ?????0000? ?????????? ?????????? 151 ????

Cheliderpeton spp. 1 001?010000 1000000001 00001100?? 00000??000 00?0?00110 51 0000100010 100010??00 00100??000 ?1??010-10 0?0?0000?? 101 0?001110?0 000000???? ????100-00 0???0001?? ?10???1??? 151 ??0?

Chenoprosopus lewisi 1 010-0010?0 1100000001 0000000000 0000000110 0000000000 51 ?0?0000010 0000000000 0010000000 0?1?110-10 0???00??00 101 000?1????? ?000?0??0? ????????0? 00???????? ??????10?? 151 ????

APPENDIX 9: TEMNOSPONDYL DATA MATRIX 295

Chenoprosopus milleri 1 010-001000 1000000001 0010000000 0000000110 0000000000 51 00?0000000 000000?000 0010000000 001?110-10 00010000?0 101 000?1??00? ?000?????? ?????????? ?????????? ?????????? 151 ????

Cochleosaurus bohemicus 1 010-001000 0000000001 0011000000 0000000110 0010000000 51 0000001000 000000?000 001000?000 ?11?110-10 ?001001000 101 0?0?000000 0000?????? ?????????? ?????????? ?????????? 151 ????

Cochleosaurus floriensis 1 010-001000 0000000001 001100000? 0000000110 0010000000 51 1000001000 000000?000 001000?000 00??110-10 0?010010?? 101 0?00000000 0000?????? ?????????? ?????????? ?????????? 151 ????

Dendrerpeton acadianum 1 000-010000 0000000000 0000000000 0000000000 0000000010 51 00?0000000 000000??00 001000?000 0?10000-10 ?000001?00 101 000?0??0?? ?00000000? ????000-00 00??000000 ?00??00000 151 0000

Dissorophus spp. 1 000-0100?? 0000110000 1000110000 01000110?0 0??100???? 51 ?????????? ?1??000000 0010000?00 0??0????10 002000001? 101 0?0????0?? 0000000000 1???101-00 00?1010100 110???1101 151 1?1?

Doleserpeton annectans 1 000-01000? 0001110000 1000110000 0000000000 01?1?01010 51 01000-01?1 010- -0??00 001010?100 ?10?000-10 0000011011 101 ???1?????? ?111011?00 ????0????? ????????00 111???110? 151 1?0?

Dutuitosaurus ouazzoui 1 001100000? 1000000001 0000110001 10021001?0 0000000111 51 1101110011 1001110100 0120000111 0101011110 0120000120 101 1100111011 1000020011 1111100011 11100011?0 0100111101 151 010?

Dvinosaurus spp. 1 0010000010 0000000000 0100010100 0010000010 0000010111 51 1001010010 0000100000 0111020001 110?001110 0020000000 101 10101?0000 0000000000 0000010000 ?0?0000001 0100001010 151 0???

296 K. PAWLEY PHD THESIS

Ecolsonia cutlerensis 1 000-1100?1 0000111000 1000110000 0100011000 01?1000010 51 ?0?0100000 010000???? 001000?000 ?10?01??10 0000000011 101 0?0?101000 ?00010000? ????101-00 00010?0?00 0101011101 151 111?

Edops craigi 1 000-001000 0000000001 0000000000 0000000000 0000000000 51 0000001000 0000000000 0000000000 00?0000-00 0000000000 101 0?0010?000 000000000? ?????????? ?????????? 010?????00 151 ????

Eoscopus lockardi 1 000-01000? 0010110000 1000110??? 0000001000 01?1??0010 51 01?0?000?1 010000??00 00100??000 ?100000-10 ?00?011??? 101 0?0?1???00 ?000000?00 ?00000?-?0 ?0???????? ??0???1101 151 1?00

Eryops megacephalus 1 000-010000 0000000001 0000110000 0000000000 0010000010 51 0000100000 0000000000 0010000000 0100010-10 0001000010 101 0000011000 0000000000 0000101-00 0001000100 0101011101 151 1100

Eryosuchus pronus 1 0011000000 1000000001 1000110011 1002000110 0000000111 51 1101110011 1011101100 0120000110 0101011010 0122000120 101 1000101111 100000000? ????100-11 0110001110 ?1001?1101 151 01??

Iberospondylus schultzei 1 000-010000 0000000000 00001100?? 0000000000 00?0?00000 51 0000?00000 000000?000 0010000000 01000?0-10 ?00?000010 101 000?10?000 0000100000 00001????? ?????????? ?????????? 151 ????

Isodectes obtusus 1 001-010000 0000001000 0000000100 0010000110 00?0010110 51 1110000011 0100?0?000 0111020000 ?10?001110 0020100000 101 10001??0?? 000000000? ????000-00 ????????0? ?10??????? 151 ??0?

Keratobrachyops australis 1 001100000? 100000011- 0000110001 0000000??1 0000000010 51 1111110001 1000111011 1121021110 0100011010 0120000120 101 1?00111011 0000?????? ?????????? ?????????? ?????????? 151 ????

APPENDIX 9: TEMNOSPONDYL DATA MATRIX 297

Konzhukovia vetusta 1 0010000?1? 1000000001 0000111000 0001000110 0000000?11 51 110011?0?0 100110?000 0010000000 01000???10 0?01000010 101 ?????????? ?000?????? ?????????? ?????????? ??????11?? 151 ????

Laidleria gracilis 1 00100000?? 100000011- 1000110001 0010000110 1000200110 51 1110?10011 100?1??000 0120010111 010?011110 0?20000120 101 ???0???0?? ?00000000? ?????????? ?????????? ?????????? 151 ??1?

Lapillopsis nana 1 000-0?000? ?000000010 0011020101 0000000010 001100010 51 111011?001 1000000?10 0001000011 1000?01101 10?210000? 101 1110??0?0? ?00000???? ??????100? 0????????0 ??????????? 151 ????

Luzocephalus spp. 1 001000000? 1000000001 0000110011 0001000110 0000000111 51 1101110011 1010?01100 0120000110 010?011110 1111000020 101 ?????????? ?000?????? ?????????? ?????????? ?????????? 151 ????

Lydekkerina huxleyi 1 0011000000 1000000001 00001100?1 0002000110 0000000011 51 1101110001 1001100100 0120000110 0100011010 0111000120 101 1000101000 1000000000 1000110000 0010001100 01001?1101 151 ??0-

Mastodonsaurus giganteus 1 0011000000 1000000001 1000110011 1002000110 0000000111 51 1101110011 1011101100 0120000101 0101011110 0122000120 101 1000101111 1000020011 1111110011 0110001110 010?1?1101 151 01??

Micropholis stowi 1 000-010001 0?00110000 1000110000 0000000000 01?1000010 51 01000000?1 010000?000 0010000000 010?010-10 0?00011011 101 00011010?? 0000000000 ????001-?0 00??10??0? 110???1101 151 ??0-

Neldasaurus wrightae 1 00100?000? 1000000000 0000000000 0010000000 0000000110 51 1010010010 1000100000 0100000000 ?00?00??10 0?20000000 101 10001??0?? ?00000000? ????0???00 ????????0? ?100??1010 151 ??00

298 K. PAWLEY PHD THESIS

Nigerpeton ricqlesi 1 011?001100 1100000001 0010000000 0000000??0 0010000100 51 1000?00010 000000?000 001000?001 001?101110 0??1000000 101 ???00????? ?????????? ?????????? ?????????? ?????????? 151 ????

Onchiodon labyrinthicus 1 000-010000 0000000001 000011000? 0000000000 00?0?00010 51 0000100000 000000??00 001000??0? ?10?010-10 ?00?000??? 101 0?0011???? 000000??0? ?????00-00 000?000100 ??????1?0? 151 11??

Parioxys spp. 1 000-0?0?00 0?0000000? 00001100?? 0000000000 00?0000000 51 0000?00000 000000??00 001000?000 ?1??010-10 0?0000?010 101 0000100000 0000100000 0000?????? 000?000100 010??11101 151 11??

Pelorocephalus spp. 1 001?00000? 100000011- 0000110001 0000000101 0000000011 51 11?1110001 1000111001 112102110? 010?011010 0120000120 101 11001110?0 1000000001 11??11011? 00?0????10 010?1?110? 151 0???

Peltobatrachus pustulatus 1 ?00-?????? ?????????? 0000110000 0000000??0 00000???1? 51 ?????????? ??????0000 0010000?10 0??0????10 0?01000010 101 100??0???? ????0111-? ????1????? ????00?10? 010????100 151 ??1?

Phonerpeton pricei 1 000-110101 0100111000 1000110000 0000011000 0111000010 51 0000100000 010?000000 0010000000 0?0?010-10 0020000011 101 0000101000 000000000? ?00?1????? ????0?0??? 0101?1???? 151 ????

Plagiosauridae 1 0010000000 1000000000 1000110001 0010000110 1000200110 51 11?1110011 1001110000 0120010101 010?010-10 0120000120 101 1100111000 1000000001 1??1000011 01000011?0 010??????? 151 ??1-

Platyoposaurus stuckenbergi 1 101000001? 1000000001 0000111000 00010000?0 0000000010 51 1000010000 1001100000 0010000000 01000?1110 0001001010 101 0?00110000 0000000000 1000100-?? 00??0001?0 ????0?1?01 151 ????

APPENDIX 9: TEMNOSPONDYL DATA MATRIX 299

Platyrhinops lyelli 1 000-01000? 0010?10000 00001100?? 0000000000 01?1?00?10 51 01000??0?1 000000??00 001000?100 ?????00-10 0000011??? 101 0?0???10?0 001100000? 1000000-0? ?0???0???? ?11???1?0? 151 1?0?

Rewana spp. 1 00100000?? 100000011- 000011000? 0000000100 0000000010 51 ?111?10001 10001?0000 0120010110 010?011010 0110000?20 101 1000111?00 10000?000? ????1????? ????????10 01001?1101 151 01??

Rhinesuchidae 1 0010000000 1000000001 0000111001 0002000110 0000000011 51 1101110000 100110?100 0120000000 010?010-10 0111000010 101 0000100000 000000000? ????100-00 0000001100 0100??1101 151 0?01

Saharastega moradiensis 1 000-0?001? ?000000001 001?00000? 0000000?1? 0??0000?01 51 00000?0000 0000000000 00100000?0 00??001110 0?01000000 101 ?????????? ?????????? ?????????? ?????????? ?????????? 151 ????

Sclerocephalus spp. 1 000-010000 1000000001 000011000? 0000000000 00?0?00110 51 0000100010 100010??00 001000?000 ?10?010-10 ?0010?0??? 101 0?001110?0 0000000000 0000100-00 000?000100 01010?1??1 151 1?0-

Siderops kehli 1 0011????0? 10??00???? 000?11???1 ?????????? 0000000011 51 ?1?1110001 1?00111011 1121021101 0100011010 0120000120 101 1??0?1?010 10000???01 111?110111 0000????10 ?1001????? 151 ????

Tersomius spp. 1 000-010001 0000110010 0011000000 0000000000 0101000011 51 0101110001 0100000000 0010000000 0000010-11 0?00001011 101 01000010?? 0000000000 1100?????? ????????00 110???1101 151 1???

Thabanchuia oomie 1 001?0????? ??00?????? 0100010100 00100000?? 0???010111 51 ?111010011 000000?000 0121020101 110?001111 0?2?000??? 101 1?1010???? ?00000111? ?????10?0? 011??????? ?????????? 151 ????

300 K. PAWLEY PHD THESIS

Thoosuchus yakovlev 1 0011000000 1000000001 0000110011 0001000110 0000000111 51 1101110011 1010101100 0120000110 010?011110 1111000020 101 ?????1???? ?000?????? ?????????? ?????????? ?????????? 151 ????

Trematosauridae 1 001000000? 1000000001 0000110011 0000000110 0000000110 51 1101110011 1000101000 0120000110 0101011110 1110000020 101 1000111011 000000000? ????110010 0???001?1? 010???1??? 151 ??0?

Trimerorhachis insignis 1 0010010000 0000000000 0000000000 0010000000 0000010111 51 1001010010 1000100000 0100000000 110?001110 0020000000 101 1000100000 0000000000 0000000-00 0110????01 0100001010 151 0000

Tupilakosaurus spp. 1 00100????? ?000?00000 010001010? 0010000??? ?00?0?01?? 51 111?010011 0000100000 0121020101 110?001111 0020000100 101 1?1?10?0?? 000000111? ?????100?? ?????????? ?????????? 151 ????

Xenobrachyops allos 1 0011000000 1000000??? 0000110001 0000000?00 0000000?11 51 11101100?1 0100010001 0121020101 0100011010 0120000120 101 ?1?01?1?10 10000????? ?????????? ?????????? ?????????? 151 ????

Zatrachydidae 1 000-010000 0000000000 0000110000 0000000000 0010000010 51 0000100000 0000000000 0010000000 010?01?010 0001000010 101 000?111000 000000000? ?????01-00 ?0???????? ??????1??? 151 ??0?

Pawley, K. 2006. 301 The Postcranial Skeleton of Temnospondyls (Tetrapoda: Temnospondyli) PhD Thesis. La Trobe University, Melbourne

APPENDIX 10. COMPARATIVE ANALYSES: THE POSTCRANIAL SKELETON OF TEMNOSPONDYLS

Results of tests of degrees of character inclusion

Exclusion of postcranial characters from the main analysis resulted in 11 trees (Figure 82), of 288 steps (58 taxa, 114 characters, CI = 0.4167, RI = 0.8437, RC = 0.3516). In this analysis, the Cochleosauridae are removed to a position as the next most derived clade to Edops plus Caerorhachis, forming a stem plesion corresponding to the traditional Edopoidea. The ‘Edopoidea’ are basal to the Dendrerpetontidae (Capetus is excluded from the Dendrerpetontidae, and is a stem plesion immediately basal to the clade containing Balanerpeton plus Dendrerpeton). Iberospondylus plus Parioxys, the Dissorophoidea, Zatrachydidae, and a collapsed clade containing Peltobatrachus, Onchiodon, Eryops and Actinodontidae are included as the most basal taxa within the Eryopoidomorpha. Otherwise the tree based exclusively on cranial characters retains the same tree topology as the main analysis. The analysis removing the historically significant vertebral characters that may reflect only differing degrees of ossification (TRU VER 8, TRU VER 9, TRU VER 14) resulted in one tree of 350 steps (58 taxa, 151 characters, CI = 0.4514, RI = 0.8422, RC = 0.3802), identical in topology to that of the main analysis (Figure 44). Additionally excluding TRU VER 33 did not produce any change in tree topology (one tree, 346 steps, 58 taxa, 150 characters, CI = 0.4509, RI = 0.8436, RC = 0.3804).

TABLE 9. COMPARISON OF DEGREES OF CHARACTER INCLUSION, MAIN ANALYSIS. CHARACTERS NUMBER OF NUMBER OF TREE LENGTH NUMBER OF CONSISTENCY INCLUDED TAXA CHARACTERS (STEPS) TREES INDEX

Main analysis 58 154 359 1 0.4485 (all characters) Main analysis 58 114 288 11 0.4167 (cranial only) Main analysis (vertebral characters 58 150 346 1 0.4509 removed)

Table 9 indicates that exclusion of postcranial characters decreased the consistency index slightly, and slightly decreased resolution. The exclusion of the historically significant vertebral characters resulted in a shorter tree with an improved consistency index, but the tree topology (compared to the main analysis) did not change, suggesting that these postcranial characters are not of major phylogenetic significance.

302 K. PAWLEY PHD THESIS

FIGURE 82. Results of main phylogenetic analysis of the Temnospondyli, postcranial characters omitted. Removal of postcranial characters resulted in a consensus of 12 trees of 288 steps, (58 taxa, 114 characters, CI = 0.4167, RI = 0.8437, RC = 0.3516). APPENDIX 10: TEMNOSPONDYL COMPARATIVE ANALYSES 303

As shown in Table 10, removing postcranial characters from the analysis using the same taxon list as Yates and Warren (2000) decreased phylogenetic resolution somewhat, as shown by the increase in the number of trees, and lowered the consistency index.

TABLE 10. COMPARISON OF DEGREES OF CHARACTER INCLUSION, ANALYSES USING THE SAME TAXA AS YATES AND WARREN (2000). CHARACTERS NUMBER OF NUMBER OF TREE LENGTH NUMBER OF CONSISTENCY INCLUDED TAXA CHARACTERS (STEPS) TREES INDEX

Same taxon list as Y 37 125 303 10 0.4356 & W (2000) Same taxon list as Y & W (2000) 37 91 244 18 0.3975 (cranial only)

Table 11 shows that removing the postcranial characters from the analysis of Yates and Warren (2000) increased the consistency index slightly, but also resulted in an increase in the number of trees. Replacing the postcranial characters and coding with those of the main analysis resulted in an increase in the number of trees, but also an increase in the consistency index.

TABLE 11. COMPARISON OF DEGREES OF CHARACTER INCLUSION, ANALYSES USING THE MATRIX OF YATES AND WARREN (2000). [* from Yates and Warren (2000)]. CHARACTERS NUMBER OF NUMBER OF TREE LENGTH NUMBER OF CONSISTENCY INCLUDED TAXA CHARACTERS (STEPS) TREES INDEX

Y & W (2000) 37* 121* 452* 1* 0.325*

Y & W (2000) 37 102 387 8 0.3282 (cranial only) Y & W (2000) 37 142 457 72 0.3545 (postcranial replaced)

Results of tests of phylogenetic stability

Constraining Edops as a sister taxon to the Cochleosauridae in the traditional definition of the Edopoidea increases tree length by three steps and increases the number of trees to 24, with a polytomy formed of Caerorhachis, Capetus, Balanerpeton plus Dendrerpeton, and Cochleosauridae (total 362 steps, CI = 0.4448). Constraining the Dendrerpetontidae to form sister taxa with Caerorhachis takes two extra steps, creates three trees and places Edops as a stem plesion immediately basal to the Cochleosauridae (total 361 steps, CI = 0.4460). As the tree lengths and consistency indices are not much lower than those of the main analysis, these results indicate that the phylogenetic positions of the most basal taxa within the Temnospondyli are not well supported, also consistent with the bootstrap and Bremer node values (Figure 45). A constraint tree which places Peltobatrachus as sister taxon to the other Upper Permian plesiomorphic temnospondyls Saharastega and Nigerpeton, results in the placement of Peltobatrachus as the most plesiomorphic taxon within the Cochleosauridae, and produces three trees that require an extra six steps (total 365 steps, 304 K. PAWLEY PHD THESIS

CI = 0.4411). Placing Peltobatrachus as sister taxon to the Rhinesuchidae, as in Yates and Warren (2000), produces a single tree, but takes an extra ten steps (total 369 steps, CI = 0.4363). The shortest compatible tree to form the Archegosauroidea as defined by Yates and Warren (2000) is 11 steps longer and produces two trees (total 370 steps, CI = 0.4351). Within the Stereospondyli, constraining Lydekkerina to form a clade with mastodonsaurids requires an extra 11 extra steps (total 370 steps, CI = 0.4351). Constraining the Brachyopidae (Batrachosuchus and Xenobrachyops) as sister taxa to the Dvinosauria takes 18 extra steps and collapses the base of the Dvinosauroidea (three trees, total 387 steps, CI = 0.4160). To place the Brachyopoidea as sister taxa to only the Dvinosauroidea produces a similar result (three trees, total 387 steps, CI = 0.4160). Constraining the whole of the Brachyopoidea as sister taxa to the Dvinosauria takes fewer steps, but results in the mostly Permian Dvinosauria as the most derived taxa within the Triassic Stereospondyli (16 extra steps, two trees, total 385 steps, CI = 0.4182).

FIGURE 83. Results of analysis using the same taxon list as Yates and Warren (2000). Strict consensus of ten trees of 303 steps, (hypothetical ancestor + 37 taxa, 125 characters, CI = 0.4356, RI = 0.7711, RC = 0.3359). APPENDIX 10: TEMNOSPONDYL COMPARATIVE ANALYSES 305

Overall, a sister taxon relationship between any members of the Dvinosauria and Brachyopoidea is not nearly as parsimonious as the hypothesis that they are not sister taxa.

Results of comparative phylogenetic analyses

Taxon sampling effects

The result of a search using the same terminal taxa as Yates and Warren (2000), but the characters and coding of the main analysis, was a consensus of ten trees (Figure 83), of 303 steps (hypothetical ancestor + 37 taxa, 125 characters, CI = 0.4356, RI = 0.7711, RC = 0.3359). All taxa were placed in a topology similar to the main analysis (Figure 44), except Tersomius is a stem taxon to Acheloma plus Dissorophus, and the inclusion of Lapillopsis within the Euskelia, as a as a sister taxon to Peltobatrachus (Figure 83). Omitting postcranial characters from this analysis (37 taxa + hypothetical ancestor,

FIGURE 84. Results of analysis using the same taxon list as Yates and Warren (2000), postcranial characters omitted Removal of postcranial characters resulted in a strict consensus of 17 trees of 244 steps, (hypothetical ancestor + 37 taxa, 91 characters, CI = 0.3975, RI = 0.7678, RC = 0.3052). 306 K. PAWLEY PHD THESIS

91 characters, CI = 0.3975, RI = 07678, RC = 3052) produced a similar but less resolved consensus of 18 trees requiring 244 steps (Figure 84). A constraint tree (including all characters) reproducing the tree topology of Yates and Warren (2000), required 36 extra steps, with reduced consistency and retention indices (339 total steps, CI = 0.3894, RI = 0.7229, RC = 2815), clearly indicating that it is significantly less parsimonious than the unconstrained search. The results of this constraint tree are still more parsimonious than the original analysis of Yates and Warren (2000) which required 452 steps (CI = 0.325, RI = 0.669), despite the inclusion of four extra characters in the analysis performed here.

The effect of changing postcranial characters

Omitting postcranial characters from the original matrix of Yates and Warren (2000) produced a tree topology similar to that of Yates and Warren (2000), with one interesting

FIGURE 85. Results of analysis using the original matrix of Yates and Warren (2000), postcranial characters omitted. Removal of postcranial characters resulted in a strict consensus of eight trees of 387 steps, (hypothetical ancestor + 37 taxa, 102 characters, CI = 0.3282, RI = 0.6845, RC = 0.2246). APPENDIX 10: TEMNOSPONDYL COMPARATIVE ANALYSES 307 exception (Figure 85), the Dvinosauroidea form the most crownward clade in the Stereospondyli, nested within the Brachyopidae. This result did not occur with the removal of postcranial characters from either the main analysis (Figure 82), or with the removal of postcranial characters from the analysis using the same taxon list as Yates and Warren (2000) (Figure 83), suggesting that the characters and coding of the data matrix used here are responsible for the difference. The hypothesis that the phylogenetic positions of the Dvinosauroidea Brachyopidae are better supported by the characters and coding used in the main analysis than by those of Yates and Warren (2000) is also supported by the results of the constraint trees (see above). [The analysis of Yates and Warren (2000) also produced constraint trees and discussed implications for the relationships of the Dvinosauroidea and Brachyopidae.] The result of the analysis using the same matrix as Yates and Warren (2000), but the postcranial characters and coding changed to that of the main analysis, was 72 trees of 457 steps (37 taxa + hypothetical ancestor, 142 characters, CI = 0.3545, RI = 0.6855, RC

FIGURE 86. Results of analysis using the original matrix of Yates and Warren (2000), with the postcranial characters and data replaced with those of the main analysis. Strict consensus of 72 trees of 457 steps, (hypothetical ancestor + 37 taxa, 142 characters, CI = 0.3545, RI = 0.6855, RC = 0.2430). 308 K. PAWLEY PHD THESIS

= 2430). A consensus tree (Figure 86) displays the Dvinosauria as the most basal clade within the Temnospondyli, and as stem taxa to the Eryopoidomorpha, as in the main analysis. The Euskelia form the most basal clade within the Eryopoidomorpha, Peltobatrachus is placed within the basal Stereospondylomorpha, in a collapsed clade with the Actinodontidae and Archegosauridae. The base of the Stereospondyli is also collapsed. A constraint tree reproducing the tree topology of Yates and Warren (2000) required 468 steps, (CI = 0.3462, RI = 0.6738, RC = 2332). This result is again much less parsimonious than that of the unconstrained analysis. The change in postcranial characters therefore has had a significant effect on the original tree topology of Yates and Warren (2000). Overall, the results of these tests indicate that the different tree topology and reduced tree length produced in the main analysis is not due to taxon sampling effects, but rather due to the choice of characters and coding.

Comparison of phylogenetic analyses

Table 12 displays the results of all analyses that contain a full complement of characters, except constraint trees.

TABLE 12. EXPECTED AND OBTAINED CONSISTENCY INDICES FOR ALL FULL ANALYSES EXCEPT CONSTRAINT TREES. [* from Yates and Warren (Y & W) (2000)]. EXPECTED OBTAINED NUMBER OF NUMBER OF TREE LENGTH ANALYSIS CONSISTENCY CONSISTENCY TAXA CHARACTERS (STEPS) INDEX INDEX

Main analysis 58 154 358 0.3405 0.4497

Y & W (2000) 37* 121* 452* 0.3776 0.325*

Same taxon list as Y 37 125 303 0.3776 0.4356 & W (2000) Y & W (2000) postcranial characters 37 142 457 0.3776 0.3545 changed

The obtained consistency index for the new analysis is much higher than the expected consistency index. In comparison, the obtained consistency index of the data set of Yates and Warren (2000) is lower than expected. The obtained consistency index of the test analysis that changed the postcranial characters and data of Yates and Warren (2000) is close to expected. The consistency index of the test analysis that used the same taxa as Yates and Warren (2000) (unconstrained search), has a consistency index much higher than expected. These results indicate that there is considerably less homoplasy and a stronger phylogenetic signal in this analysis, compared to the analysis of Yates and Warren (2000). The improved consistency indexes are due to the changes in both the cranial and postcranial characters, and not due to the choice of taxa. The results of the unconstrained analyses compared to the constraint trees, as laid out in Table 13, indicate that the hypothesis of temnospondyl relationships proposed by Yates and Warren (2000) is less parsimonious than that of the main analysis. This is shown by the lower tree lengths and higher consistency indices of the main analysis and the unconstrained analyses using the same taxon list as Yates and Warren (2000) and the APPENDIX 10: TEMNOSPONDYL COMPARATIVE ANALYSES 309 analysis in which the postcranial characters of the original matrix of Yates and Warren (2000) were replaced.

TABLE 13. EXPECTED AND OBTAINED CONSISTENCY INDICES FOR ALL FULL ANALYSES AND CONSTRAINT TREES. [* from Yates and Warren (Y & W) (2000)]. NUMBER OF TREE LENGTH CONSISTENCY ANALYSIS NUMBER OF TAXA CHARACTERS (STEPS) INDEX

Main analysis 58 154 358 0.4497

Y & W (2000) 37* 121* 452* 0.325*

Same taxon list as Y & W 37 125 303 0.4356 (2000): unconstrained Same taxon list as Y & W 37 125 339 0.3894 (2000): constraint tree Y & W (2000) postcranial characters changed: 37 142 457 0.3545 unconstrained Y & W (2000) postcranial characters changed: 37 142 468 0.3462 constraint tree

Pawley, K. 2006. 311 The Postcranial Skeleton of Temnospondyls (Tetrapoda: Temnospondyli) PhD Thesis. La Trobe University, Melbourne

APPENDIX 11. LIST OF SYNAPOMORPHIES: THE POSTCRANIAL SKELETON OF TEMNOSPONDYLS

The following is a complete list of all synapomorphies for all clades. All character state were optimised under ACCTRAN. Terminal autapomorphies are not listed. The list of synapomorphies is presented thus: character state identifier (e.g. PTE 13), consistency index (e.g. CI = 1.000), and state change (e.g. 0 ==> 1). Unambiguous synapomorphies are denoted ‘==>’, ambiguous synapomorphies are denoted ‘−−>’.

Temnospondyli more derived than Edops Cochleosauridae more derived than PTE 13 CI = 0.400 0 ==> 1 Saharastega INT VAC 2 CI = 1.000 0 ==> 1 L SC SKU 1 CI = 1.000 0 ==> 1 PREMAX 4 CI = 0.500 0 ==> 1 Dendrerpetontidae CHO 1 CI = 1.000 0 ==> 1

PREMAX 1 CI = 0.250 0 ==> 1 Chenoprosopus spp. LAC 2 CI = 0.250 1 ==> 0 ANT FOS 1 CI = 0.250 0 ==> 1 PASPHE 12 CI = 0.250 0 −−> 1 SPL 1 CI = 0.333 0 −−> 1 Nigerpeton plus Adamanterpeton plus HUM 5 CI = 1.000 0 −−> 1 Cochleosaurus spp. QUAJUG 7 CI = 0.500 0 ==> 1 Balanerpeton plus Dendrerpeton VOM 7 CI = 0.250 0 ==> 1 VOM 5 CI = 0.333 0 ==> 1 SPL 1 CI = 0.333 1 ==> 0

Temnospondyli more derived than Adamanterpeton plus Cochleosaurus spp. Dendrerpetontidae MAX 1 CI = 0.250 1 ==> 0 MAX 1 CI = 0.250 0 ==> 1 POSPAR 11 CI = 1.000 0 −−> 1 PASPHE 7 CI = 0.333 0 −−> 1 VOM 17 CI = 0.500 0 ==> 1 ILI 2 CI = 1.000 0 ==> 1 Cochleosaurus spp. Cochleosauridae ANT FOS 1 CI = 0.250 0 ==> 1 PASPHE 12 CI = 0.250 0 ==> 1 PIN FOR 1 CI = 0.333 0 ==> 1 JUG 2 CI = 0.333 0 −−> 1 JUG 3 CI = 0.250 0 ==> 1

312 K. PAWLEY PHD THESIS

Temnospondyli more derived than Eobrachyopidae Cochleosauridae PREFRO 8 CI = 0.500 0 ==> 1 VOM 5 CI = 0.333 0 ==> 1 JUG 2 CI = 0.333 0 ==> 1 VOM 7 CI = 0.250 0 −−> 1 VOM 6 CI = 0.111 0 −−> 1 VOM 16 CI = 0.333 0 −−> 1 VOM 14 CI = 0.200 0 −−> 1 PAL 4 CI = 0.250 0 ==> 1 PAL 5 CI = 0.250 0 ==> 1 ECT 5 CI = 0.200 0 ==> 1 ECT 5 CI = 0.200 0 −−> 1 PTE 24 CI = 0.333 0 −−> 1 PASPHE 8 CI = 1.000 0 ==> 1 PTE 25 CI = 1.000 1 ==> 0 MAND 1 CI = 0.333 0 ==> 1 Dvinosaurus plus Tupilakosauridae PREMAX 6 CI = 0.250 0 −−> 1 Dvinosauria PAR 2 CI = 1.000 0 ==> 1 PTE 22 CI = 0.167 0 ==> 1 SC SK 1 CI = 0.250 0 ==> 1 MAND 3 CI = 1.000 0 ==> 1 LAC 2 CI = 0.250 1 ==> 0 INTCLA 1 CI = 0.200 0 ==> 1 SQU 3 CI = 0.500 0 ==> 1 VOM 4 CI = 0.200 0 ==> 1 Tupilakosauridae VOM 13 CI = 0.250 0 −−> 1 PTE 13 CI = 0.400 1 ==> 2 PAL 2 CI = 0.167 0 ==> 1 PTE 20 CI = 0.333 0 ==> 1 PTE 9 CI = 0.500 0 ==> 1 INT VAC 3 CI = 1.000 0 ==> 1 PTE 13 CI = 0.400 0 −−> 1 EXOCC 1 CI = 0.333 0 −−> 1 PTE 23 CI = 1.000 0 −−> 1 TRU VER 8 CI = 0.250 0 ==> 1 ANT VAC 1 CI = 0.167 0 −−> 1 TRU VER 9 CI = 0.500 0 ==> 1 PASPHE 5 CI = 0.286 0 ==> 2 TRU VER 14 CI = 0.333 0 ==> 1 PASPHE 7 CI = 0.333 0 −−> 1 CLA 5 CI = 0.250 0 −−> 1 Peltobatrachus plus Eryopoidomorpha CLA 7 CI = 0.333 0 −−> 1 POSTFRO 2 CI = 0.500 0 ==> 1 SCACOR 9 CI = 1.000 0 −−> 1 INTEMP 1 CI = 0.333 0 ==> 1 FEM 6 CI = 1.000 0 ==> 1 VOM 15 CI = 0.333 0 −−> 1 ANT FOS 1 CI = 0.250 0 −−> 1 Dvinosauria more derived than EXOCC 3 CI = 1.000 0 ==> 1 Neldasaurus RIB 5 CI = 0.333 0 ==> 1 MAX 1 CI = 0.250 1 ==> 0 CLE 8 CI = 1.000 0 ==> 1 VOM 2 CI = 1.000 0 ==> 1 ULNA 2 CI = 1.000 0 −−> 1 VOM 6 CI = 0.110 0 −−> 1 ILI 8 CI = 1.000 0 ==> 1 VOM 14 CI = 0.200 0 −−> 1 FIB 6 CI = 1.000 0 −−> 1

Dvinosauroidea Eryopoidomorpha INTEMP 1 CI = 0.333 0 −−> 1 TAB 9 CI = 1.000 0 ==> 1 MAND 1 CI = 0.333 0 −−> 1 JUG 3 CI = 0.250 0 ==> 1 FEM 8 CI = 1.000 0 ==> 1 VOM 10 CI = 0.250 0 −−> 1 PAL 3 CI = 0.250 0 −−> 1 PAL 4 CI = 0.250 1 ==> 0 PTE 13 CI = 0.400 0 −−> 1 PTE 14 CI = 0.500 0 ==> 1 PTE 18 CI = 0.500 0 ==> 2 APPENDIX 11: TEMNOSPONDYL SYNAPOMORPHIES 313

Euskelia Dissorophidae PREMAX 1 CI = 0.250 0 ==> 1 SQU 8 CI = 1.000 0 ==> 1 VOM 7 CI = 0.250 0 −−> 1 SQU 9 CI = 0.500 0 ==> 1 VOM 16 CI = 0.333 0 −−> 1 PASPHE 5 CI = 0.286 0 ==> 2 ANT VAC 2 CI = 0.200 0 −−> 1 CLE 5 CI = 1.000 0 ==> 1 PREART 5 CI = 0.333 0 −−> 1 DERM 1 CI = 0.333 0 ==> 1 CLA 8 CI = 1.000 0 −−> 1 HUM 21 CI = 1.000 0 ==> 1 Dissorophidae more derived than FIB 5 CI = 1.000 0 ==> 1 Broiliellus TAB 13 CI = 0.500 0 ==> 1 Actinodontidae VOM 4 CI = 0.200 0 ==> 1 Dissorophidae more derived than PAL 2 CI = 0.167 0 ==> 1 Dissorophus NOS 3 CI = 1.000 0 ==> 1 Euskelia more derived than PREFRO 8 CI = 0.500 0 ==> 1 Actinodontidae CER VER 2 CI = 0.333 0 −−> 1 MAX 1 CI = 0.250 1 ==> 0 HUM 2 CI = 0.500 0 −−> 1 QUAJUG 7 CI = 0.500 0 −−> 1 PAL 4 CI = 0.250 1 ==> 0 Acheloma plus Phonerpeton ECT 5 CI = 0.200 1 ==> 0 PREMAX 5 CI = 0.500 0 ==> 1 MAX 2 CI = 0.333 0 ==> 1 Euskelia more derived than Onchiodon INTCLA 7 CI = 0.500 0 ==> 1 Amphibamidae VOM 10 CI = 0.250 0 ==> 1 Euskelia more derived than Eryops VOM 15 CI = 0.333 1 ==> 0 LAC 2 CI = 0.250 1 ==> 0 PAL 3 CI = 0.250 0 ==> 1 PASPHE 11 CI = 1.000 0 ==> 1 Euskelia more derived than Zatrachydidae PASPHE 12 CI = 0.250 0 ==> 1 PASPHE 7 CI = 0.333 1 ==> 0 DEN 2 CI = 1.000 0 ==> 1 RIB 5 CI = 0.333 1 ==> 0 Iberospondylus plus Parioxys CLE 4 CI = 1.000 0 ==> 1 VOM 5 CI = 0.333 1 ==> 0 PTE 2 CI = 0.250 0 ==> 1 Amphibamidae more derived than PREART 5 CI = 0.333 1 ==> 0 Micropholis TRU VER 32 CI = 0.500 0 ==> 1 MAX 5 CI = 0.500 0 ==> 1 ANT FOS 1 CI = 0.250 1 ==> 0 Dissorophoidea INTCLA 7 CI = 0.500 0 −−> 1 NAS 3 CI = 1.000 0 ==> 1 PREFRO 1 CI = 1.000 0 ==> 1 Amphibamidae more derived than FRO 4 CI = 0.200 0 ==> 1 Eoscopus QUAJUG 6 CI = 1.000 0 ==> 1 FRO 4 CI = 0.200 0 −−> 1 QUA 1 CI = 1.000 0 ==> 1 PTE 20 CI = 0.333 0 ==> 1 PAL 5 CI = 0.250 0 ==> 1 TEETH 2 CI = 1.000 0 ==> 1 EXOCC 5 CI = 1.000 0 ==> 1 TEETH 3 CI = 1.000 0 ==> 1 CER VER 2 CI = 0.333 0 −−> 1 HUM 12 CI = 1.000 0 ==> 1 HUM 2 CI = 0.500 0 −−> 1

314 K. PAWLEY PHD THESIS

Doleserpeton plus Amphibamus Lydekkerina plus Superstereospondyli MAX 8 CI = 1.000 0 ==> 1 SC 3 CI = 0.250 0 ==> 1 VOM 3 CI = 1.000 0 ==> 1 TAB 2 CI = 0.500 0 −−> 1 PAL 1 CI = 1.000 0 ==> 1 PAL 3 CI = 0.250 0 ==> 1 PTE 15 CI = 1.000 0 ==> 1 PTE 20 CI = 0.333 0 ==> 1 TEETH 1 CI = 1.000 0 ==> 1 PTE 21 CI = 0.143 0 ==> 1 TRU VER 8 CI = 0.250 0 ==> 1 ANT VAC 1 CI = 0.167 0 ==> 1 EXOCC 1 CI = 0.333 0 ==> 1 Stereospondylomorpha EXOCC 3 CI = 1.000 1 ==> 2 MAND 1 CI = 0.333 0 ==> 1 SC SK 1 CI = 0.250 0 −−> 1 PREART 5 CI = 0.333 0 ==> 1 PREMAX 6 CI = 0.250 0 −−> 1 POST COR 5 CI = 0.250 0 ==> 1 TAB 2 CI = 0.500 0 −−> 1 CLA 7 CI = 0.333 0 −−> 1 SQU 6 CI = 0.333 0 ==> 1 ECT 2 CI = 0.333 0 ==> 1 Superstereospondyli CER VER 2 CI = 0.333 0 ==> 1 DERM 3 CI = 1.000 0 ==> 1 VOM 4 CI = 0.200 0 ==> 1 PAL 2 CI = 0.167 0 ==> 1 Archegosauridae EPI 1 CI = 0.500 0 −−> 1 SK 1 CI = 1.000 0 ==> 1 PREART 1 CI = 0.500 0 −−> 1 VOM 15 CI = 0.333 0 −−> 1 ANT COR 3 CI = 0.333 0 −−> 1 PASPHE 12 CI = 0.250 0 ==> 1 MID COR 3 CI = 0.250 0 −−> 1 CER VER 1 CI = 1.000 0 ==> 1 Stereospondyli more derived than CER VER 10 CI = 1.000 0 ==> 1 Archegosauridae CER VER 11 CI = 1.000 0 ==> 1 JUG 2 CI = 0.333 0 ==> 1 CER VER 12 CI = 1.000 0 ==> 1 JUG 3 CI = 0.250 0 ==> 1 INTCLA 9 CI = 1.000 0 ==> 1 VOM 6 CI = 0.125 0 ==> 1 CLA 3 CI = 0.500 0 −−> 1 VOM 10 CI = 0.250 0 ==> 1 CLA 5 CI = 0.250 0 ==> 1 PASPHE 4 CI = 1.000 0 −−> 1 SCACOR 5 CI = 1.000 0 ==> 1 CLE 7 CI = 1.000 0 ==> 1 HUM 25 CI = 1.000 0 −−> 1 Mastodonsauroidea plus Trematosauroidea TAB 11 CI = 1.000 0 ==> 1 Stereospondyli PAL 8 CI = 0.500 0 ==> 1 PTE 1 CI = 0.500 0 ==> 1 PREMAX 6 CI = 0.250 0 −−> 1 TAB 11 CI = 1.000 0 ==> 1 Mastodonsauroidea SQU 6 CI = 0.333 1 ==> 2 ANT VAC 2 CI = 0.200 0 −−> 1 VOM 14 CI = 0.200 0 ==> 1 PASPHE 7 CI = 0.333 1 ==> 2 PTE 2 CI = 0.250 0 ==> 1 PREART 1 CI = 0.500 0 −−> 1 PTE 9 CI = 0.500 0 ==> 1 PTE 13 CI = 0.400 1 ==> 2 Mastodonsauridae PASPHE 5 CI = 0.286 0 ==> 1 FRO 4 CI = 0.200 0 ==> 1 TAB 13 CI = 0.500 0 ==> 1 PASPHE 5 CI = 0.286 1 ==> 2 PREART 6 CI = 1.000 0 ==> 1 APPENDIX 11: TEMNOSPONDYL SYNAPOMORPHIES 315

Trematosauroidea Rewana spp. plus Brachyopoidea SQU 6 CI = 0.333 2 ==> 1 JUG 3 CI = 0.250 1 ==> 0 ECT 2 CI = 0.250 0 −−> 1 VOM 4 CI = 0.200 1 ==> 0 PASPHE 2 CI = 1.000 0 ==> 1 PAL 2 CI = 0.167 1 ==> 0 EXOCC 1 CI = 0.333 1 ==> 0 ECT 2 CI = 0.250 0 −−> 1 POST COR 5 CI = 0.250 0 −−> 1 ANT VAC 2 CI = 0.200 1 ==> 0 INTCLA 1 CI = 0.200 0 −−> 1 INTCLA 1 CI = 0.200 0 −−> 1 CLA 3 CI = 0.500 0 −−> 1 INTCLA 8 CI = 1.000 0 −−> 1 CLA 5 CI = 0.250 0 −−> 1 Trematosauridae plus Luzocephalus SC 3 CI = 0.250 1 ==> 0 Brachyopoidea SC 3 CI = 0.250 0 −−> 1 Superstereospondyli more derived than VOM 6 CI = 0.110 1 −−> 0 Mastodonsauroidea plus Trematosauroidea PTE 6 CI = 1.000 0 ==> 1 ECT 8 CI = 1.000 0 ==> 1 PTE 14 CI = 0.500 0 ==> 1 PTE 22 CI = 0.167 0 ==> 1 PTE 18 CI = 0.500 1 ==> 2 PASPHE 5 CI = 0.286 1 ==> 2 PTE 21 CI = 0.143 1 ==> 0 PASPHE 7 CI = 0.333 1 ==> 0 ANT COR 3 CI = 0.333 0 −−> 1 MAND 2 CI = 0.500 0 ==> 1 Chigutisauridae Metoposauroidea QUAJUG 4 CI = 1.000 0 ==> 1 TAB 13 CI = 0.500 0 ==> 1 PTE 1 CI = 0.500 0 ==> 1 SQU 7 CI = 1.000 0 ==> 1 PTE 7 CI = 1.000 0 ==> 1 CLA 4 CI = 1.000 0 ==> 1 PTE 19 CI = 1.000 0 ==> 1

Superstereospondyli more derived than Keratobrachyops plus Siderops Metoposauroidea PTE 5 CI = 1.000 0 ==> 1 SC 3 CI = 0.250 0 −−> 1 PREFRO 9 CI = 0.500 0 −−> 1 Brachyopidae LAC 1 CI = 0.500 0 −−> 1 VOM 14 CI = 0.200 1 ==> 0 LAC 2 CI = 0.250 0 −−> 1 PAL 5 CI = 0.250 0 −−> 1 SQU 6 CI = 0.333 2 ==> 0 VOM 6 CI = 0.111 0 −−> 1 VOM 13 CI = 0.250 0 ==> 1 PTE 2 CI = 0.250 1 ==> 0 EPI 1 CI = 0.500 0 −−> 1 ANT COR 3 CI = 0.333 0 −−> 1 MID COR 3 CI = 0.250 0 −−> 1 CLA 7 CI = 0.333 1 ==> 0 DERM 1 CI = 0.333 0 −−> 1

Plagiosauroidea FRO 4 CI = 0.200 0 ==> 1 SQU 3 CI = 0.500 0 ==> 1 QUAJUG 5 CI = 1.000 0 ==> 1 PTF 1 CI = 1.000 0 ==> 2 RIB 5 CI = 0.333 0 −−> 1

Pawley, K. 2006. 317 The Postcranial Skeleton of Temnospondyls (Tetrapoda: Temnospondyli) PhD Thesis. La Trobe University, Melbourne

APPENDIX 12. TERMINAL TAXA: WALKING WITH EARLY TETRAPODS

(mi) indicates that the endochondral postcranial skeleton of all described specimens is morphogenetically immature (Chapter 4). (la) indicates that the endochondral postcranial skeleton of all described specimens is of larval morphology (Chapter 4).

Acanthostega gunnari (Clack, 1988; Coates and Clack, 1991; Clack, 1992, 1994a, 1994b; Ahlberg et al., 1996; Coates, 1996; Ahlberg and Clack, 1998; Clack, 1998b, 2002c, 2003b). Acheloma cumminsi (Trematops milleri) (Williston, 1909a; Case, 1911a; Olson, 1941; Schaeffer, 1941; Bolt, 1974a, 1974b, 1974c; Dilkes and Reisz, 1987). Specimens examined: AMNH 4205, 7150; MCZ 1413, 2524. Acherontiscus caledoniae (Carroll, 1969b). Acroplous vorax (mi) (Hotton, 1959; Coldiron, 1978; Foreman, 1990). Adamanterpeton ohioensis (Milner and Sequeira, 1998). Adelogyrinus simorhynchus (Andrews and Carroll, 1991). Adelospondylus watsoni (Andrews and Carroll, 1991). Albanerpetontidae (Estes and Hoffstetter, 1976; Estes, 1981; McGowan and Evans, 1995; Gardner, 1999; Venczel and Gardner, 2005). Amphibamus (Miobatrachus) grandiceps (mi) (Watson, 1940; Gregory, 1950; Carroll, 1964a; Bolt, 1979; Daly, 1994). Anthracosaurus russelli (Panchen, 1977b, 1981; Clack, 1987b). Apateon pedestris (la) (Boy, 1972, 1986, 1987; Schoch, 1992). Archegosaurus decheni (mi) (Meyer, 1857; Jaekel, 1896; Whittard, 1928; Gubin, 1997; Witzmann, 2006a, 2006b). Specimens examined: NMV P10836, P24165, P198455, P198457. Archeria crassidisca (Romer, 1957; Holmes, 1984; Clack and Holmes, 1988; Holmes, 1989a). Specimens examined: AMNH 4123, 7117, 23599; MCZ 2045, 2046, 2047, 2049, 2075, 2501, 2503, 2509. Asaphestera intermedia (Carroll and Gaskill, 1978). Balanerpeton woodi (Milner and Sequeira, 1994). Baphetes kirkbyi (mi) (Beaumont, 1977; Milner and Lindsay, 1998). Batrachiderpeton reticulatum (Milner, 1980a; Bossy and Milner, 1998). Batropetes fritschia (B. truncatus, Brachystelechus fritschi) (Carroll and Gaskill, 1971, 1978; Carroll, 1991). Benthosuchus sushkini (mi) (Bystrow and Efremov, 1940). Brachydectes spp. (mi) (Wellstead, 1991). Bruktererpeton fiebigi (Boy and Bandel, 1973). Casineria kiddi (Paton et al., 1999). Caerorhachis bairdi (Holmes and Carroll, 1977; Ruta et al., 2002). 318 K. PAWLEY PHD THESIS

Capetus palustris (Sequeira and Milner, 1993). Captorhinus aguti (Warren, 1961; Fox and Bowman, 1966; Clark and Carroll, 1973; Holmes, 1977; de Ricqlès and Bolt, 1983; Sumida, 1990; Rieppel, 1993; Modesto, 1998; Kissel et al., 2002; Holmes, 2003). Cardiocephalus peabodyi (Carroll and Gaskill, 1978). Chenoprosopus lewisi (mi) (Hook, 1993). Chenoprosopus milleri (Langston, 1953). Cochleosaurus bohemicus (Sequeira, 2004). Cochleosaurus florensis (Rieppel, 1980; Godfrey and Holmes, 1995). Colosteus scutellus (Hook, 1983). Crassigyrinus scoticus (mi) (Panchen, 1985; Panchen and Smithson, 1990; Clack, 1996; Ahlberg and Clack, 1998; Clack, 1998c). Dendrerpeton acadianum (Dawson, 1882; Steen, 1934; Carroll, 1967; Milner, 1980b; Godfrey et al., 1987; Holmes et al., 1998; Robinson et al., 2005). Diadectes absitus (Berman et al., 1998). Diadectes spp. (North American) (Case, 1905, 1910, 1911b; Romer and Byrne, 1931; Welles, 1941; Romer, 1944; Olson, 1947; Lewis and Vaughn, 1965; Sumida, 1990; Sumida and Lombard, 1991; Berman et al., 1992; Berman et al., 1998; Berman, 2000; Sumida and Modesto, 2001; Berman and Henrici, 2003). Specimens examined: AMNH 4357, 4364, 4380, 4381, 4709; MCZ 1035, 1106, 1281, 1335, 1715, 1741.1786, 3002, 3194, 7896, 7938, 7945, 8044, 8051. Diceratosaurus brevirostris (mi) (Bossy and Milner, 1998) Diplocaulus magnicornis (mi) (Williston, 1909b; Case, 1911a; Douthitt, 1917; Bossy and Milner, 1998). Diploceraspis burkei (Beerbower, 1963; Bossy and Milner, 1998). Discosauriscus austriacus (mi) (Klembara, 1996, 1997; Klembara and Bartik, 2000). Specimens examined: AMNH 28453, 28454, 28455, 28456. Dissorophus spp. (Williston, 1910b; Case, 1911a; Carroll, 1964a; de Mar, 1968; Bolt, 1974a, 1974c; Milner, 2003). Specimens examined: AMNH 4343, 4593; MCZ 2122, 4169, 4170, 4172, 4173, 4175, 4176, 4177, 4178, 4180, 4181, 4185, 4186, 4187, 4190, 4194. Doleserpeton annectans (Gregory et al., 1956; Bolt, 1969, 1974a, 1974b; Lombard and Bolt, 1988). Dolichopareias disjectus (Andrews and Carroll, 1991). Dvinosaurus spp. (mi) (Sushkin, 1936; Bystrow, 1938; Shishkin, 1973; Nikitin, 1995, 1997; Shishkin, 2000; Gubin, 2004). Ecolsonia cutlerensis (Berman et al., 1985). Edops craigi (mi) (Romer and Witter, 1942; Clack and Holmes, 1988). Specimens examined: MCZ 1378, 1781, 6489, 6490, 7140. Eocaecilia micropoda (Jenkins Jnr and Walsh, 1993; Carroll, 2000a). Eoherpeton watsoni (Panchen, 1975; Smithson, 1985). Eoscopus lockardi (mi) (Daly, 1994). Eryops megacephalus (Cope, 1884, 1888; Case, 1911a; Cope and Matthew, 1915; Romer, 1922; Miner, 1925; Sawin, 1941; Moulton, 1974; Pawley and Warren, 2006). Specimens examined: AMNH 4086, 4121, 4180, 4183, 4186, 4191, 4198, 4203, 4204, 4208, 4211, 4215, 4255, 4280, 4307, 4661, 4754, 4862, 23449; MCZ 1128, 1137, 1219, 1539, 1858, 1883, 1900, 1984, 2092, 2565, 2604, 2615, 2630, 2631, 2675, 2712, 2716, 4121, 4308, 4763, 6475, 7279, 7551, 7555, 7559, 7645, 7647, 7765, 7766, 7768, 7770, 7773, 7797; MV P202186, P202187. Eryosuchus (Parotosuchus) pronus (mi) (Howie, 1970). APPENDIX 12: EARLY TETRAPOD TERMINAL TAXA 319

Eucritta melanolimnetes (la) (Clack, 1998a, 2001). Euryodus primus (Carroll and Gaskill, 1978). Eusthenopteron foordi (Jarvik, 1944; Andrews and Westoll, 1970; Jarvik, 1980; Smithson and Thomson, 1982; Ahlberg et al., 1996). Gephyrostegus bohemicus (Carroll, 1970, 1972b; Godfrey and Reisz, 1991; Rieppel, 1993; Ahlberg and Clack, 1998). Greererpeton burkemorani (mi) (Smithson, 1982; Godfrey, 1989a; Coates, 1996; Schultze and Bolt, 1996; Ahlberg and Clack, 1998; Bolt and Lombard, 2001; Clack, 2003a). Hapsidopareion lepton (mi) (Daly, 1973; Carroll and Gaskill, 1978). Hyloplesion longicostatum (Carroll and Gaskill, 1978). Ichthyostega stensioei (Coates and Clack, 1990; Coates and Clack, 1995; Jarvik, 1996; Ahlberg and Clack, 1998; Clack et al., 2003b; Clack et al., 2003a; Ahlberg et al., 2005). Isodectes (Eobrachyops, Saurerpeton) obtusus (mi) (Watson, 1956; Milner, 1982a; Sequeira, 1998). Specimens examined: AMNH 6919, 6922, 6923, 6934, 6945, 6937, 6947, 6978; USMN 4471 (cast), 4474. Karaurus sharovi (mi) (Ivachnenko, 1978; Milner, 2000). Keraterpeton galvani (mi) (Jaekel, 1903; Milner, 1980a; Bossy and Milner, 1998). Kotlassia prima (Bystrow, 1944; Bulanov, 2003). Leptorophus tener (la) (Boy, 1986). Lethiscus stocki (Wellstead, 1982; Anderson et al., 2001). Limnoscelis spp. (Williston, 1911a, 1911b, 1912; Romer, 1946; Fracasso, 1983, 1987; Berman and Sumida, 1990; Berman et al., 1992; Sumida et al., 1992; Berman, 2000; Sumida and Modesto, 2001). Specimens examined: MCZ 1947, 1948, 1949. Lydekkerina huxleyi (Watson, 1919; Broom, 1930; Broili and Schröder, 1937; Findlay, 1968; Jupp and Warren, 1986; Shishkin et al., 1996; Jeannot, 2004; Pawley and Warren, 2005). Specimens examined: BP/1/1373, BP/1/5021, BP/1/fn 3900; UMZC T218, T243. Mastodonsaurus giganteus (mi) (Schoch, 1999a, 2000b, 2002a, 2002b). Megalocephalus pachycephalus (Beaumont, 1977; Ahlberg and Clack, 1998). Micraroter erythrogeios (Carroll and Gaskill, 1978). Microbrachis pelikani (mi) (Carroll and Gaskill, 1978; Vallin and Laurin, 2004). Micromelerpeton credneri (la) (Boy, 1971, 1972, 1995; Witzmann and Pfretzschner, 2003). Neldasaurus wrightae (mi) (Chase, 1965). Specimens examined: MCZ 1371, 1463, 2406, 2407, 2516, 2518. Nigerpeton ricqlesi (Sidor et al., 2005; Steyer et al., 2006). Odonterpeton triangulare (mi) (Carroll and Gaskill, 1978) Oestocephalus amphiuminus (Carroll, 1998a; Anderson, 2003). Orobates pabsti (Berman and Henrici, 2003; Berman et al., 2004). Ossinodus pueri (Warren and Ptasznik, 2002; Warren and Turner, 2004). Specimens examined: QM F 34280, 34282, 34284, 34600, 34601, 34607, 34610, 34616, 34621, 34622, 34821, 36903, 36907, 36912, 36916, 36932, 36935, 36937, 36955, 37405, 37415, 37417, 37418, 37420, 37426, 37427, 37429, 37431, 37433, 37434, 37437, 37439, 37440, 37441, 37444, 37449, 37450, 37452, 37453. Paleothyris acadiana (Carroll, 1969a). Panderichthys rhombolepis (Vorobyeva, 1977; Vorobyeva and Schultze, 1991; Vorobyeva and Kuznetsov, 1992; Vorobyeva, 1992, 1995; Ahlberg et al., 1996; Ahlberg and Clack, 1998; Vorobyeva, 1998, 2000; Boisvert, 2005; Brazeau and Ahlberg, 2006). Pantylus cordatus (Carroll, 1968; Carroll and Baird, 1968; Romer, 1969a; Sumida, 1990). 320 K. PAWLEY PHD THESIS

Pederpes finneyae (mi) (Clack and Finney, 2005). Pelodosotis elongatum (Carroll and Gaskill, 1978). Peltobatrachus pustulatus (Panchen, 1959; Jupp and Warren, 1986). Petrolacosaurus kansensis (Lane, 1947; Peabody, 1952; Reisz, 1977, 1981). Philoderpeton (Eogyrinus) attheyi (Panchen, 1966, 1972). Pholiderpeton scutigerum (Clack, 1983, 1987a). Phlegethontia linearis (Carroll et al., 1998; Anderson, 2002a). Phonerpeton pricei (Bolt, 1974a; Dilkes, 1990, 1993). Platyoposaurus (Platyops) stuckenbergi (Konzhukova, 1955a, 1955b; Gubin, 1991; Shishkin, 2000). Platyrhinops (Amphibamus) lyelli (Carroll, 1964a; Milner, 1982a; Hook and Baird, 1984; Clack and Milner, 1993; Daly, 1994; Carroll et al., 1999; Schoch and Milner, 2004). Proterogyrinus scheelei (Holmes, 1980, 1984). Ptyonius marshii (mi) (Milner, 1980a; Bossy and Milner, 1998). Rhinesuchidae (van Hoepen, 1915; Broom, 1930; Watson, 1962; Schoch, 2000a; Shishkin and Rubidge, 2000; Pawley and Warren, 2004). Specimens examined: BP/1/3241; M 460; NMQR 1438; TM 75, 77, 78, 79, 85, 103, 208, 4703; SAM-PK-K100021. Rhynchonkos (Goniorhynchus) stovalli (Olson, 1970; Carroll, 1998b; Berman, 2000). Saharastega moradiensis (Sidor et al., 2005; Damiani et al., in press). Sauropleura pectinata / scalaris (Milner, 1980a; Bossy and Milner, 1998) Saxonerpeton geinitzi (Carroll and Gaskill, 1978). Schoenfelderpeton prescheri (la) (Boy, 1986). Scinosaurus crassus (Bossy and Milner, 1998). Sclerocephalus spp. (Broili, 1926; Boy, 1986; Werneburg, 1992; Meckert, 1993; Lohmann and Sachs, 2001; Schoch, 2002c, 2003). Seymouria spp. (Williston, 1911a; White, 1939; Vaughn, 1966; Berman et al., 1987b; Sumida, 1990; Sumida et al., 1992; Laurin, 1995, 1996; Berman et al., 2000; Klembara et al., 2006). Specimens examined: MCZ 1091, 1092, 1093, 1094, 1096, 1097, 1098, 1100, 1101, 1639, 1640, 1641, 1643, 1644, 1650, 1658, 1661. Solenodosaurus janenschi (Carroll, 1970; Laurin and Reisz, 1999). Stegotretus agyrus (Berman et al., 1988). Thabanchuia oomie (mi) (Warren, 1998b). Tiktaalik roseae (Daeschler et al., 2006; Shubin et al., 2006). Triadobatrachus massinoti (Rage and Rocek, 1989). Trimerorhachis insignis (mi) (Cope, 1884; Case, 1911a; Cope and Matthew, 1915; Case, 1935; Nilsson, 1944; Colbert, 1955; Watson, 1956; Olson, 1979; Lombard and Bolt, 1988; Schoch, 1999b). Specimens examined: AMNH 4578, 4720, 4763, 4765, 23283, 23302, 23319, 23321, 23330; MCZ 840, 8116, 8117, 8128, 8164, 8244, 8245, 8312, 8365, 8413, 8515, 8536, 8577, 8611; TMM 40031:61-110, 40998:25, 40998:39; UMCP 105230, 105232, 105236, 105247, 105249, 105258, 105259, 174885-187001, 24009, 24010; UMMP 15995, 16033, 16055. Tuditanus punctulatus (Carroll and Baird, 1968; Carroll and Gaskill, 1978; Rieppel, 1993). Tulerpeton curtum (Lebedev and Clack, 1993; Lebedev and Coates, 1995). Tupilakosaurus spp. (Nielsen, 1954; Shishkin, 1961; Nielsen, 1967; Shishkin, 1973). Urocordylus wandesfordii (mi) (Milner, 1980a; Bossy and Milner, 1998). Valdotriton gracilis (mi) (Evans and Milner, 1996). Ventastega curonica (Ahlberg et al., 1994; Ahlberg and Clack, 1998; Ahlberg and Luksevics, 1998, 2004). Westlothiana lizziae (Smithson and Rolfe, 1990; Smithson et al., 1994) Whatcheeria deltae (Lombard and Bolt, 1995; Bolt and Lombard, 2000; Clack, 2003a; Clack and Finney, 2005).

Pawley, K. 2006. 321 The Postcranial Skeleton of Temnospondyls (Tetrapoda: Temnospondyli) PhD Thesis. La Trobe University, Melbourne

APPENDIX 13. REINTERPRETATIONS OF PUBLISHED MATERIAL: WALKING WITH EARLY TETRAPODS

Morphologically immature specimens are noted; all other specimens are morphologically mature because they have ossified coracoid, pubis, carpus, and tarsus (Chapter 4).

Acanthostega gunnari: In the scapulocoracoid (Coates, 1996: figure 12a), foramen ‘A’ (Coates, 1996: figure 12a) and ‘C’ (Coates, 1996: figure 12c) are reinterpreted as the supraglenoid foramen, foramen ‘E’ (Coates, 1996: figure 12a) as the coracoid foramen, and foramen ‘D’ (Coates, 1996: figure 12a) as the glenoid foramen. The ‘deltopectoral crest’ is interpreted as the deltoid crest, and the ‘supinator ridge’ (Coates, 1996: figure 16) as the pectoral crest. Acheloma cumminsi (Trematops milleri): The forelimb elements figured by Case (1911a: figure 34) are morphogenetically immature. Acherontiscus caledoniae: The element labelled supertemporal by Carroll (1969b) is interpreted as the tabular following Carroll et al. (1998). Acroplous vorax: The endochondral postcranial skeleton (Hotton, 1959; Coldiron, 1978) is morphogenetically immature. Amphibamus grandiceps: The specimen figured by Daly (1994: figure 18) is morphogenetically immature. Apateon pedestris: The specimens (Boy, 1972, 1986, 1987; Schoch, 1992) are all of larval morphology. Archegosaurus decheni: The endochondral postcranial skeleton (Meyer, 1857) is morphogenetically immature. Archeria crassidisca: Specimens were used as a source of data rather than the descriptions of Romer (1957). Balanerpeton woodi: Some characteristics of the postcranial skeleton (Milner and Sequeira, 1994) are indeterminate die to its state of preservation. Baphetes kirkbyi: The endochondral postcranial skeleton (Milner and Lindsay, 1998) is morphogenetically immature. The pit anterior to the latissimus dorsi process (labelled m.sh Milner and Lindsay, 1998: fig 9B) is interpreted as an accessory foramen. The radius of Ossinodus pueri described by Warren and Ptasznik (2002: figure 2, A-F) is reinterpreted as a left radius, following comparison with Acanthostega gunnari (Coates, 1996). As the Baphetes kirkbyi radius figured by Warren and Ptasznik (2002: figure 2, G-L) is a right radius, all labels of Baphetes kirkbyi transpose accordingly. Benthosuchus sushkini: The endochondral postcranial skeleton (Bystrow and Efremov, 1940) is morphogenetically immature. The tibia (Bystrow and Efremov, 1940: figure 55) is interpreted as a left tibia in A, flexor; and B, extensor views. Brachydectes spp.: The endochondral postcranial skeleton (Wellstead, 1991: figure 19) is morphogenetically immature. 322 K. PAWLEY PHD THESIS

Caerorhachis bairdi: The presence of a parasymphysial plate in Caerorhachis is here regarded as uncertain. The area labelled ‘adductor crest’ (add cr) (Ruta et al., 2002: figure 12) is reinterpreted as the fourth trochanter at the distal end of the adductor blade. Captorhinus aguti: Details of scapulocoracoid were taken from the specimen figured by Fox and Bowman (1966: figure 24), rather than the reconstruction by Holmes (1977). Cardiocephalus peabodyi: In agreement with Carroll and Gaskill (1978) the postcranial elements described by Gregory et al. (1956) do not belong to Cardiocephalus, but to Doleserpeton annectans. The femur and tibia questionably attributed to C. sternbergi (Carroll and Gaskill, 1978: figure 34) were not used as a source of data. Chenoprosopus lewisi: The endochondral postcranial skeleton (Hook, 1993: figure 3) is morphogenetically immature. Colosteus scutellus: Coding for the contact between the supratemporal plus tabular and squamosal was revised following Schultze and Bolt (1996). Crassigyrinus scoticus: The endochondral postcranial skeleton (Panchen, 1985; Panchen and Smithson, 1990) is morphogenetically immature. Dendrerpeton acadianum: The endochondral postcranial skeleton of some specimens (Carroll, 1967) is morphogenetically immature, these specimens are readily determinable by their comparatively small size. Diadectes absitus: The fibula of (Berman et al., 1998: figure 19) is inverted; the proximal end is towards the bottom of the page. Diadectes spp. (North American): Berman et al (1992) considered all the North American species of Diadectes questionably distinguishable, they are treated as synonymous here. Diplocaulus magnicornis: The endochondral postcranial skeleton (Williston, 1909b:plate 5) is morphogenetically immature. Discosauriscus austriacus: The endochondral postcranial skeleton (Klembara and Bartik, 2000) is morphogenetically immature. On the humerus, the ‘tldm’ (Klembara and Bartik, 2000: figure 21d) is the scapulohumeral ridge (sensu Pawley and Warren, 2006) and the ‘pr.scs’ (Klembara and Bartik, 2000: figure 21d) is the broken base of the latissimus dorsi process. Doleserpeton annectans: In accordance with Bolt (1969) the postcranial elements are indistinguishable from those described as Cardiocephalus sternbergi (Gregory et al., 1956). The fibulae (Gregory et al., 1956: figure 30) are inverted. Dvinosaurus spp.: The endochondral postcranial skeleton (Sushkin, 1936; Bystrow, 1938; Nikitin, 1995, 1997) is morphogenetically immature. Ecolsonia cutlerensis: The proximal half of the humerus (Berman et al., 1985: figure 11.D) is a left rather than a right as in the distal half, and the latissimus dorsi is broken off. The ulnae (Berman et al., 1985: figure 11.E, 11.F) are also of the left side, as is the clavicle (Berman et al., 1985: figure 11.B). Edops craigi: The humerus (Romer and Witter, 1942: figure 13) is not completely morphogenetically mature, in that the proximal humeral ridge is unossified and the radial condyle of the humerus is only partially ossified. Eoherpeton watsoni: The coronoids of Eoherpeton watsoni (Smithson, 1985: fig 14) are interpreted as being similar to those of Anthracosaurus (Panchen, 1981), with the tooth row on the posterior coronoid. Eoscopus lockardi: The endochondral postcranial skeleton of some specimens of the hind limb (Daly, 1994: figure 8, 16) are morphogenetically immature. The interclavicle (Daly, 1994: figure 12) is inverted. APPENDIX 13: EARLY TETRAPOD REINTERPRETATIONS 323

Eryops megacephalus: The manus of Eryops has four digits, in agreement with Miner (1925), rather than five as described by Gergaory et al. (1923). The sacral rib of Case (1911a: figure 30) and Moulton (1974: figure 13) is more typical than that of Olson (1936a). Eryosuchus (Parotosuchus) pronus: The endochondral postcranial skeleton (Howie, 1970) is morphogenetically immature. Eucritta melanolimnetes: The endochondral postcranial skeleton (Clack, 2001) is larval in morphology. Eusthenopteron foordi: As the process on the pelvis (Andrews and Westoll, 1970: figure 14, 15) lacks a sacral rib attachment, it is not possible to determine if it is a post- iliac process or a dorsal process, coding for iliac processes in Eusthenopteron is indeterminate. The interpretation of the humerus follows Coates (1996). Gephyrostegus bohemicus: Only the specimen drawings (Carroll, 1970) were used as a source of data. Greererpeton burkemorani: Coding for the contact between the supratemporal plus tabular and squamosal was revised following Schultze and Bolt (1996). The endochondral postcranial skeleton (Godfrey, 1989a) is morphogenetically immature. The iliac process (Godfrey, 1989a: figure 21, 22) is a dorsal process because it has an attachment area for the sacral rib. Hapsidopareion lepton: The endochondral postcranial skeleton (Daly, 1973: figure 18; Carroll and Gaskill, 1978: figure 14 K) is morphogenetically immature. Hyloplesion longicostatum: The ‘intermedium’ and ‘fibulae’ (Carroll and Gaskill, 1978: figure 90 I) are interpreted as an astragalus and calcaneum respectively. Ichthyostega stensioei: The ‘interdorsals’ (Jarvik, 1996: figure 34) are interpreted as pleurocentra. Interpretation of the humerus and femur follows Coates and Clack (1991) and Clack et al. (2003a). Isodectes (Eobrachyops, Saurerpeton) obtusus: The endochondral postcranial skeleton (AMNH 6919, 6922, 6923, 6934, 6945, 6937, 6947, 6978; USMN 4471 (cast), 4474) is morphogenetically immature. Specimens were used as the source of data, rather than the reconstructions of Sequeira (1998). Karaurus sharovi: The endochondral postcranial skeleton (Ivachnenko, 1978) is morphogenetically immature. Kotlassia prima: Cranial coding was revised following Bulanov (2003). The coracoid portion of the scapulocoracoid has been reconstructed (Bystrow, 1944: figure 15, 16), and was not used as a source of data. The left rib on vertebrae number 27 (Bystrow, 1944: figure 14) was used as the representative sacral rib. Leptorophus tener: The specimens (Boy, 1986) are all larval in morphology. Lydekkerina huxleyi: The endochondral postcranial skeleton described by Watson (1919) is morphogenetically immature. Mastodonsaurus giganteus: The ‘radius’ (Schoch, 1999a: figure 44) is reinterpreted as an inverted left fibula in anterior, flexor, posterior and extensor views respectively. The ‘fibula’ (Schoch, 1999a: figure 48) is an inverted subadult right tibia in flexor, posterior, extensor, and anterior views respectively. Most of the specimens of the endochondral postcranial skeleton (Schoch, 1999a) are morphogenetically immature. Microbrachis pelikani: Cranial characters were revised following Vallin and Laurin (2004). The endochondral postcranial skeleton (Carroll, 1998b: figure 74) is morphogenetically immature. Micromelerpeton credneri: The specimens (Boy, 1971, 1972, 1995; Witzmann and Pfretzschner, 2003) are all larval in morphology. 324 K. PAWLEY PHD THESIS

Neldasaurus wrightae: The endochondral postcranial skeleton (Chase, 1965) is morphogenetically immature. The figure of the tibia (Chase, 1965: figure 13 H) is reinterpreted as the extensor surface of the right tibia. Odonterpeton triangulare: The endochondral postcranial skeleton (Carroll and Gaskill, 1978: figure 99 E, F) is morphogenetically immature. Oestocephalus amphiuminus: Coding for Oestocephalus amphiuminus was updated following Anderson (2003). Ossinodus pueri: The radius of Ossinodus described by Warren and Ptasznik (2002: figure 2, A-F) is reinterpreted as a left radius, following comparison with Acanthostega gunnari (Coates, 1996). As the Baphetes kirkbyi radius figured by Warren and Ptasznik (2002: figure 2, G-L) is a right radius, all labels of Baphetes transpose accordingly. Pantylus cordatus: The process on the humerus labelled scs (attachment of subcoracoideus muscle) (Carroll and Baird, 1968: figure 122) is reinterpreted as the latissimus dorsi process. The element labelled supertemporal by Romer (1969a) is interpreted as the tabular following Carroll and Gaskill (1978). Pederpes finneyae: The endochondral postcranial skeleton (Clack and Finney, 2005) is morphogenetically immature. Platyoposaurus (Platyops) stuckenbergi: Platyoposaurus is interpreted as possessing lateral line sulci on the skull roof, these are not obvious in the figures (Konzhukova, 1955a, 1955b; Gubin, 1991). Platyrhinops (Amphibamus) lyelli: The specimens figured by Carroll (1964a: figure 23, 24), Carroll et al. (1999: figure 4 A, B)are morphogenetically immature. Ptyonius marshii: The endochondral postcranial skeleton (Bossy and Milner, 1998) is morphogenetically immature. Rhinesuchidae: Included taxa are Uranocentrodon (Myriodon) senekalensis (Haughton, 1915; van Hoepen, 1915); Broomistega putterilli BP/1/3241; an unnamed rhinesuchid, field no. M460, and SAM-PK-K10021 (Pawley and Warren, 2004). The endochondral postcranial skeletons of Broomistega and SAM-PK-K10021 are morphogenetically immature. Rhynchonkos (Goniorhynchus) stovalli: The ‘intermedium’ and ‘fibulae’ (Carroll and Gaskill, 1978: figure 71 G) are reinterpreted as the astragalus and calcaneum respectively. Saxonerpeton geinitzi: The ‘intermedium’ and ‘fibulae’ (Carroll and Gaskill, 1978: figure 23 C-E) are reinterpreted as the astragalus and calcaneum respectively. Schoenfelderpeton prescheri: The specimens (Boy, 1986) are all larval in morphology. Seymouria spp.: Specimens were used as the source of data, rather than the description of the postcranial skeleton by White (1939). Solenodosaurus janenschi: The photograph of the specimen (Laurin and Reisz, 1999: figure 3) was used for character coding in preference to drawings. See also Materials and Methods. Thabanchuia oomie: The endochondral postcranial skeleton (Warren, 1998b) is morphogenetically immature. Trimerorhachis insignis: The endochondral postcranial skeleton (Pawley, in press) is morphogenetically immature. Tuditanus punctulatus: The element labelled supertemporal by Carroll and Baird (1968) is interpreted as the tabular following Carroll and Gaskill (1978). Tulerpeton curtum: The latissimus dorsi process may be broken off (Lebedev and Coates, 1995: fig. 5) its morphology is therefore indeterminate. APPENDIX 13: EARLY TETRAPOD REINTERPRETATIONS 325

Urocordylus wandesfordii: The endochondral postcranial skeleton (Bossy and Milner, 1998) is morphogenetically immature. Ventastega curonica: The ‘iliac process’ (Ahlberg et al., 1994: figure 13) is interpreted as the left cleithrum following Clack (2002c). Westlothiana lizziae: The flexor surface of the fibula (Smithson et al., 1994: figure 17 M) was reconstructed as damaged, here it is interpreted as complete. The status of the intermedium (Smithson et al., 1994: figure 20 A) is unclear, it may be an astragalus, but has been coded as uncertain.

Pawley, K. 2006. 327 The Postcranial Skeleton of Temnospondyls (Tetrapoda: Temnospondyli) PhD Thesis. La Trobe University, Melbourne

APPENDIX 14. CHARACTER STATE LIST: WALKING WITH EARLY TETRAPODS

The materials and methods section (Chapter 6) outlines the selection criteria used for determining which character states were included or modified from previous analyses, and for formulating new character states. Character states in plain type (e.g. TRU VER 1) were previously published and the relevant character state distributions were discussed in Ruta et al. (2003). Characters used in other previously published analyses (Sequeira and Milner, 1993; Carroll, 1995; Lebedev and Coates, 1995; Coates, 1996; Paton et al., 1999; Yates and Warren, 2000; Anderson, 2001; Damiani, 2001a; Gardner, 2001; Clack, 2002b; Clack and Ahlberg, 2004; Klembara and Ruta, 2004b; Pawley and Warren, 2004; Sequeira, 2004; Clack and Finney, 2005; Pawley and Warren, 2005; Daeschler et al., 2006) have only the numbers in bold (e.g. TRU VER 28). New characters not previously used in phylogenetic analyses of basal tetrapods are entirely in bold (e.g. INTCLA 7.). Many of these characters were also used in the phylogenetic analysis of Chapter 5 (Appendix 8). Postcranial character states subject to morphogenetic development (Chapter 4) are marked ‘m’, which means that the morphogenetic stage of the specimens should be assessed before coding the character states, as the derived state may be absent in a specimen due to its immature morphogenetic stage. Additional notes are provided to clarify character states in cases of potential confusion. Characters marked ‘#’ were not used in the main analysis, but were included in the test analyses. Character state distributions for the cranial characters marked ‘#’ can be found in Ruta et al. (2003). Characters marked ‘¨’ are listed in the evolutionary transformation series in the main analysis. Note that character state distributions refer only to the main analysis (Figure 62).

HIGHER LEVEL TAXA

Included taxa are listed for higher level taxa not denoted in Figure 62.

Archeriamorphs

Anthracosauridae (in this analysis): Anthracosaurus plus Eoherpeton.

Temnospondyli

Dvinosauria Dvinosauroidea: Eobrachyopidae plus (Dvinosaurus plus Tupilakosauridae) Eobrachyopidae: Isodectes plus Acroplous Tupilakosauridae: Tupilakosaurus plus Thabanchuia 328 K. PAWLEY PHD THESIS

Euskelia Trematopidae: Ecolsonia plus (Acheloma plus Phonerpeton) Stereospondylomorpha Archegosauridae: Archegosaurus plus Platyoposaurus Superstereospondyli Mastodonsauroidea: Benthosuchus plus (Eryosuchus plus Mastodonsaurus) Mastodonsauridae: Eryosuchus plus Mastodonsaurus

Microsauria

Tuditanomorpha Tuditanidae: Tuditanus plus Asaphestera Hapsidopareiontidae: Saxonerpeton plus Hapsidopareion Pantylidae: Pantylus plus Stegotretus Ostodolepidae: Micraroter plus Pelodosotis Gymnarthridae: Cardiocephalus plus Euryodus

CRANIAL CHARACTERS

Skull

1. SK 1. Snout margins continually converging towards tip (0) tip of snout expanded so that snout margins run parallel, or are concave before tip (1). STATE 1: some Temnospondyli (Archegosauridae).

Ornament

2. L SC SKU 1. Absence (0) or presence (1) of lightly sculptured area along skull midline. STATE 1: Ossinodus, some Temnospondyli (Cochleosauridae more derived than Saharastega).

Sensory canals

3. ¨ SC SK 1. Sensory canals: present, either enclosed or exposed (0) absent (1). [This character state has been simplified from that of Ruta et al. (2003), and separated into two characters states (see SC SK 2).] STATE 1: Anthracosauridae; Temnospondyli (except Nigerpeton, Dvinosauria, Stereospondyli); Neospondyli (except some Seymouriamorpha: Kotlassia plus Discosauriscus; some Microsauria: Microbrachis, Batropetes; Diplocaulidae). 4. ¨ SC SK 2. Sensory canals: partially or wholly enclosed (0) none enclosed (1). [This character state has been simplified from that of Ruta et al. (2003), and separated into two characters states (see SC SK 1).] STATE 1: Terrapoda. 5. SC 3. Infraorbital sulcus (where present): straight or gently curved (0) with a step- like or 'Z' shaped flexure between the orbit and the naris (1). STATE 1: some derived Temnospondyli (Stereospondyli: Lydekkerina plus Mastodonsauroidea). APPENDIX 14: EARLY TETRAPOD CHARACTER STATE LIST 329

Nares

6. NOS 3. Absence (0) or presence (1) of condition: nostrils elongate and keyhole shaped. STATE 1: some Temnospondyli (Trematopidae).

Median rostral

7. MED ROS 1. Median rostral: present (0) absent (1). STATE 1: Crassigyrinus, Colosteidae, Anthracosauria plus Terrapoda.

Premaxilla

8. PREMAX 1. Absence (0) or presence (1) of alary process. STATE 1: some Temnospondyli (Dendrerpetontidae; some Dvinosauria: Trimerorhachis, Isodectes; Euskelia). 9. PREMAX 4. Premaxillae without (0) or with (1) expanded anteromedial dorsal surface and marginal elongation. STATE 1: some Temnospondyli (Edops, Cochleosauridae more derived than Saharastega). 10. PREMAX 5. Premaxillary tusks: absent (0) present (1). STATE 1: Whatcheeriidae, Colosteidae, some Temnospondyli (Nigerpeton, Acheloma plus Phonerpeton), Seymouria. 11. PREMAX 6. Rugose, medial tubercle on the palatal surface of the premaxillae, posterior to the tooth row: absent (0) present (1). STATE 1: some Temnospondyli (Saharastega, Dvinosaurus, Archegosauridae). 12. PREMAX 8. Mouth subterminal so that anterior most surface of premaxilla faces ventrally: absent (0) present (1). STATE 1: some Microsauria (Batropetes plus clade containing Pantylidae plus Ostodolepidae plus Rhynchonkos plus Gymnarthridae). 13. PREMAX 9. Absence (0) or presence (1) of shelf-like premaxilla-maxilla contact mesial to tooth row on palate. STATE 1: Ichthyostega, Crassigyrinus, Greererpeton. 14. PREMAX 10. Elongate medial dorsal processes of premaxillae: absent (0) present (1). STATE 1: Diadectomorpha plus Captorhinomorpha.

Anterior tectal

15. ¨ TEC 1. Presence (0) or absence (1) of anterior tectal. STATE 1: archeriamorphs plus Terrapoda.

Lateral rostral

16. LAT ROS 1. Presence (0) or absence (1) of lateral rostral. STATE 1: Acanthostega, archeriamorphs plus Terrapoda.

Septomaxilla

17. SPTMAX 1. Absence (0) or presence (1) of septomaxilla. STATE 1: Crassigyrinus, Baphetidae, Terrapoda except Nectridia. 330 K. PAWLEY PHD THESIS

18. SPTMAX 2. Septomaxilla ornamented and part of skull roof (0) or a detached ossification inside nostril (1). [This character was re-written from Ruta et al. (2003) in order to clarify it] STATE 1: some Temnospondyli (Dissorophoidea), Seymouriamorpha plus Cotylosauria.

Maxilla

19. MAX 1. Maxilla-nasal suture: absent (0) present (1). STATE 1: Megalocephalus, Pholiderpeton spp., some Temnospondyli (Nigerpeton plus Chenoprosopus spp., Neldasaurus, Sclerocephalus, Stereospondylomorpha). 20. MAX 2. Maxillary tusks: absent (0) present (1). STATE 1: Whatcheeriidae, Anthracosaurus, some Temnospondyli (Chenoprosopus lewisi, Nigerpeton, Acheloma plus Phonerpeton), Seymouria, Captorhinomorpha, some Microsauria (Hyloplesion, Pantylidae, Euryodus). 21. MAX 5. Maxilla not entering (0) or entering orbit margin (1). STATE 1: some Temnospondyli (Amphibamidae: Eoscopus, Amphibamus), some Nectridia (Batrachiderpeton, Sauropleura). 22. # MAX 6. Maxillary arcade closed (0) or open (1) posteriorly. 23. MAX 7. Dorsal maxillary margin not forming (0) or forming (1) distinct dorsal step (1). STATE 1: some Microsauria (Pantylidae). 24. MAX 8. Absence (0) or presence (1) of condition: maxillary facial process shaped like a rectangular flange. STATE 1: some Temnospondyli (derived Amphibamidae: Amphibamus plus Doleserpeton).

Nasal

25. ¨ NAS 1. Absence (0) or presence (1) of paired dorsal nasals. STATE 1: Tetrapoda (all taxa more derived than Panderichthys) except derived Diplocaulidae (Diplocaulus plus Diploceraspis). 26. NAS 3. Ventral flange of nasal: absent (0) present, forming posterior internal wall of the naris (1). STATE 1: some Temnospondyli (Dissorophoidea).

Prefrontal

27. PREFRO 1. Prefrontal ventral process: absent (0) descending ventral flange (ventral process of the prefrontal- VPP) on the orbit rim (1). STATE 1: some Temnospondyli (Dissorophoidea). 28. PREFRO 6. Prefrontal not sutured with premaxilla (0) or sutured (1). STATE 1: Colosteidae. 29. PREFRO 7. Prefrontal without (0) or with (1) stout lateral outgrowth. STATE 1: Baphetidae. 30. PREFRO 8. Absence (0) or presence (1) of prefrontal entering nostril margin. STATE 1: Crassigyrinus plus Colosteidae, some Temnospondyli (Eobrachyopidae, Trematopidae), Batropetes, Nectridia (except Diplocaulus, Batrachiderpeton). 31. PREFRO 9. Prefrontal not sutured with maxilla (0) or sutured. STATE 1: Colosteidae, derived Diplocaulidae (Diplocaulus plus Diploceraspis). APPENDIX 14: EARLY TETRAPOD CHARACTER STATE LIST 331

Lachrymal

32. # LAC 1. Presence (0) or absence (1) of lachrymal. 33. LAC 2. Lachrymal included in orbital margin, prefrontal-jugal suture absent (0) excluded from orbital margin, prefrontal-jugal suture present (1). [This character was re- written from Ruta et al. (2003) in order to clarify it.] STATE 1: ichthyostegamorphs, some Temnospondyli (Edops, Cochleosauridae, basal Euskelia: Sclerocephalus plus Eryops, Stereospondylomorpha), derived Diplocaulidae (Diplocaulus plus Diploceraspis). 34. LAC 4. Lachrymal without (0) or with (1) dorsomedial digitiform process. STATE 1: some Microsauria (Tuditanomorpha except Rhynchonkos), Urocordylidae. 35. LAC 5. Lachrymal without (0) or with (1) V-shaped emargination along posterior margin. STATE 1: Baphetidae.

Frontal

36. FRO 1. Frontal unpaired (0) or paired (1). STATE 1: all taxa more derived than Eusthenopteron (except Sauropleura, derived Diplocaulidae: Diplocaulus plus Diploceraspis). 37. FRO 4. Frontal excluded from (0) or contributing to (1) margin of orbit. STATE 1: Gephyrostegus, some Temnospondyli (Dissorophoidea except Amphibamus, Mastodonsauridae), Captorhinomorpha, derived Diplocaulidae (Diplocaulus plus Diploceraspis).

Parietal

38. ¨ PAR 1. Absence (0) or presence (1) of parietal-tabular suture. STATE 1: Anthracosauria, Neospondyli. 39. PAR 2. Absence (0) or presence (1) of parietal-postorbital suture. STATE 1: Ichthyostegamorphs (Acanthostega plus Ichthyostega), some Temnospondyli (derived Dvinosauroidea: Dvinosaurus plus Tupilakosauridae), Neospondyli more derived than Seymouriamorpha (except Scinosaurus, derived Microsauria: Tuditanomorpha). 40. PAR 7. Absence (0) or presence (1) of parietal-squamosal suture on skull roof. STATE 1: Captorhinomorpha, Diplocaulidae more derived than Scinosaurus.

Pineal foramen

41. PIN FOR 1. Presence (0) or absence (1) of pineal foramen. STATE 1: some Temnospondyli (Cochleosauridae except Adamanterpeton and Chenoprosopus lewisi), some Microsauria (Pantylus, Ostodolepidae).

Postparietal

42. POSTPAR 1. Presence (0) or absence (1) of postparietals. STATE 1: Batropetes, Scinosaurus. 43. POSTPAR 2. Postparietals paired (0) or unpaired (1). STATE 1: Ichthyostega, Limnoscelis plus Diadectomorpha. 332 K. PAWLEY PHD THESIS

44. ¨ POSTPAR 5. Absence (0) or presence (1) of postparietal-exoccipital suture. STATE 1: Terrapoda except Cotylosauria. 45. POSTPAR 6. Postparietals not entirely on occipital surface (0) or entirely on this surface (1). STATE 1: Cotylosauria, some Microsauria (Hapsidopareiontidae plus Tuditanidae). 46. POSTPAR 7. Postparietals and tabulars without (0) or with (1) posteroventrally sloping occipital exposure. [This character was merged with TAB 8 from Ruta et al. (2003) because they are not independent.] STATE 1: derived tuditanomorph Microsauria (Ostodolepidae plus Rhynchonkos plus Gymnarthridae). 47. POSTPAR 8. Postparietals without (0) or with (1) sinuous posterior ridge. STATE 1: derived tuditanomorph Microsauria (Ostodolepidae plus Rhynchonkos). 48. POSTPAR 9. Postparietals without (0) or with (1) broad, concave posterior emargination. STATE 1: derived Diplocaulidae (Diplocaulus plus Diploceraspis). 49. POSPAR 11. Each postparietal with a medial posterior lappet extending as a continuation of the skull roof: absent (0) present (1). STATE 1: some Temnospondyli (Cochleosauridae: Cochleosaurus spp.).

Postfrontal

50. # POSTFRO 1. Presence (0) or absence (1) of postfrontal. 51. POSTFRO 2. Postfrontal-supratemporal suture: absent (0) present (1). STATE 1: Tiktaalik, Megalocephalus, some Temnospondyli (Acroplous, Peltobatrachus plus Eryopoidomorpha). 52. POSTFRO 3. Postfrontal not contacting tabular (0) or contacting it (1). STATE 1: derived Microsauria (Tuditanomorpha), Scinosaurus.

Intertemporal

53. INTEMP 1. Intertemporal present (0) or absent (1). STATE 1: Tiktaalik plus ichthyostegamorphs (Acanthostega plus Ichthyostega), Colosteidae, Megalocephalus, Temnospondyli (Acroplous, Dvinosaurus plus Tupilakosauridae; Peltobatrachus plus Eryopoidomorpha), Neospondyli more derived than Seymouriamorpha. 54. ¨ INTEMP 2. Intertemporal (where present) not interdigitating with cheek (0) or interdigitating (1). [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Whatcheeriidae, Baphetes, Terrapoda. 55. INTEMP 3. Intertemporal (where present) not contacting squamosal (0) or contacting it (1). [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Whatcheeriidae, Nigerpeton, Seymouriamorpha.

Supratemporal

56. ¨ SUTEMP 1. Presence (0) or absence (1) of supratemporal. STATE 1: Lepospondyli except Urocordylidae. 57. SUTEMP 2. Absence (0) or presence (1) of condition: supratemporal forming anterior edge of otic notch. STATE 1: Baphetidae. APPENDIX 14: EARLY TETRAPOD CHARACTER STATE LIST 333

58. SUTEMP 3. Absence (0) or presence (1) of condition: supratemporal narrow and strap like, at least two or three times longer than broad. STATE 1: Urocordylidae (Ptyonius plus Sauropleura). 59. SUTEMP 4. Supratemporal contact with squamosal smooth (0) or interdigitating (1). STATE 1: Whatcheeriidae, Baphetidae, Temnospondyli.

Tabular

60. # TAB 1. Tabular present (0) or absent (1) as a separate ossification. 61. TAB 2. Absence (0) or presence (1) of subdermal, unornamented blade-like posterolateral portion of tabular. [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Crassigyrinus, Anthracosauria more derived than Gephyrostegus, some Temnospondyli (basal Stereospondylomorpha: Archegosauridae plus Rhinesuchidae). 62. TAB 3. Absence (0) or presence (1) of rounded, subdermal, button-like posterior process of tabular, distinct from and medial to, the posterolateral corner of the tabular. [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Colosteidae, Baphetidae, Edops. 63. TAB 5. Tabular contact with squamosal (where present) smooth (0) or interdigitating (1). [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Temnospondyli, some Microsauria (Ostodolepidae), Diplocaulidae. 64. TAB 6. Absence (0) or presence (1) of condition: tabular elongate posteriorly in the form of massive horn. STATE 1: Diplocaulidae more derived than Scinosaurus. 65. TAB 9. Tabulars: well developed rectangular to triangular bones (0) reduced to thin slivers lying against the posterior margin of the skull roof (1). STATE 1: some Temnospondyli (Dvinosauroidea), derived Captorhinomorpha (Paleothyris plus Petrolacosaurus). 66. TAB 10. Terminal crest of tabular horn: absent (0) present on ventral side of tabular horn, lying lateral and parallel to the paraoccipital process (sensu Bystrow and Efremov, 1940) (1). STATE 1: some Temnospondyli (Mastodonsauroidea). 67. TAB 11. Tabular - exoccipital contact in paraoccipital process, lateral to post- temporal fossa: absent (0) present (1). STATE 1: some Temnospondyli (Stereospondyli). 68. TAB 12. Tabular horns: absent or posteriorly directed (0) distally enlarged and laterally directed towards a process of the squamosal, partly or fully enclosing the otic notch (1). STATE 1: some Temnospondyli (Mastodonsauridae). 69. TAB 13. Tabular sutured with quadrate enclosing otic notch: absent (0) present (1). STATE 1: some Temnospondyli (Dissorophidae except Phonerpeton). 70. TAB 14. Tabular horn distally biramous: absent (0) present (1). STATE 1: some Anthracosauria (Anthracosaurus, Pholiderpeton spp.).

Postorbital

71. # POSORB 1. Postorbital present (0) or absent (1) as a separate ossification. 72. POSORB 2. Postorbital without (0) or with (1) ventrolateral digitiform process fitting into a deep groove. STATE 1: Urocordylidae. 334 K. PAWLEY PHD THESIS

73. POSORB 3. Postorbital contributing to (0) or excluded from (1) margin of orbit. STATE 1: Colosteus, derived Diplocaulidae (Diplocaulus plus Diploceraspis). 74. POSORB 5. Postorbital not contacting tabular (0) or contacting it (1). STATE 1: derived Microsauria (Batropetes plus Tuditanomorpha), Scinosaurus. 75. POSORB 8. Anterior most part of dorsal margin of postorbital with sigmoid profile absent (0) or present (1). STATE 1: derived Microsauria (Batropetes plus Tuditanomorpha except Ostodolepidae)

Squamosal

76. # SQU 2. Absence (0) or presence (1) of condition: posterior margin of squamosal sloping posteriorly. 77. ¨ SQU 3. Squamosal without (0) or with (1) broad, concave semicircular embayment. STATE 1: Temnospondyli (except Dvinosauria) plus Seymouriamorpha, Diadectomorpha. 78. # SQU 4. Absence (0) or presence (1) of single ‘squamoso-tabular’ in the place of squamosal and tabular. 79. SQU 5. Squamosal without (0) or with (1) internal shelf bracing quadrate from behind. STATE 1: Diplocaulidae. 80. SQU 6. Posterior squamosal margin in dorsal view: straight or concave (0) convex, bulging posteromedially narrowing the otic notch (1) convex and overhanging (falciform crest) (2). Ordered. STATE 1: some Temnospondyli (Archegosauridae). STATE 2: some Temnospondyli (Stereospondyli). 81. SQU 8. Supratympanic flange of squamosal: absent (0) present (sensu Bolt, 1974c) (1). STATE 1: some Temnospondyli (Dissorophidae). 82. SQU 9. Semi-lunar flange of squamosal: absent (0) present (sensu Bolt, 1974c) (1). STATE 1: some Temnospondyli (Dissorophidae, Eoscopus). 83. # SUS 1. Absence (0) or presence (1) of anteroposteriorly narrow, bar-like squamosal.

Jugal

84. # JUG 1. Presence (0) or absence (1) of jugal. 85. JUG 2. Jugal not contributing (0) or contributing (1) to ventral margin of skull roof. STATE 1: derived Anthracosauria (Proterogyrinus, Archeria), some Temnospondyli (Cochleosauridae, Eobrachyopidae, Stereospondyli more derived than Archegosauridae), Neospondyli (except derived Urocordylidae: Sauropleura spp., Urocordylus). 86. JUG 3. Jugal not contacting (0) or contacting (1) pterygoid (alar process of jugal). STATE 1: Anthracosauria, some Temnospondyli (Cochleosauridae, Dvinosauroidea, Stereospondyli more derived than Archegosauridae), Captorhinus. 87. JUG 5. Jugal-quadratojugal suture: present (0) absent (1). STATE 1: Eusthenopteron, some Microsauria (Ostodolepidae, Hapsidopareion). 88. JUG 6. Absence (0) or presence (1) of condition: jugal ventrally expanded to form flange overlapping posterior end of maxilla. STATE 1: some Microsauria (Pantylidae). APPENDIX 14: EARLY TETRAPOD CHARACTER STATE LIST 335

89. JUG 7. Jugal without (0) or with (1) V-shaped indentation of dorsal margin. STATE 1: most archeriamorphs (Whatcheeriidae, Crassigyrinus, Baphetidae, some Anthracosauria: Pholiderpeton spp., Anthracosaurus). 90. JUG 9. Jugal-squamosal contact: present (0) absent (1). STATE 1: some Microsauria (Ostodolepidae, Hapsidopareion).

Quadratojugal

91. # QUAJUG 1. Presence (0) or absence (1) of quadratojugal. 92. QUAJUG 3. Absence (0) or presence (1) of condition: quadratojugal an anteroposteriorly elongate and dorsoventrally narrow splinter of bone. STATE 1: derived Microsauria (Rhynchonkos plus Gymnarthridae). 93. QUAJUG 6. Posterior medial process of quadratojugal: absent (0) extending posterior to, and wrapping around, the dorsal process of the quadrate (1). STATE 1: some Temnospondyli (Dissorophoidea). 94. QUAJUG 7. Paraquadrate foramen: absent (0) present on the occipital face of the quadratojugal (1). [Repolarised from Chapter 5 (Appendix 8).] STATE 1: Tetrapoda more derived than Acanthostega, where known: Megalocephalus, Pholiderpeton scutigerum, Temnospondyli (except some Cochleosauridae: Nigerpeton, Cochleosaurus spp., Euskelia), Seymouriamorpha, Cotylosauria, Diploceraspis.

Quadrate

95. QUA 1. Enlarged quadrate dorsal process: absent (0) present (1). STATE 1: some Temnospondyli (Dissorophoidea).

Preopercular

96. PREOPE 1. Presence (0) or absence (1) of preopercular. STATE 1: Tetrapoda more derived than Whatcheeriidae.

Post-temporal fossa

97. PTF 1. Fossa near dorsolateral corner of occiput, roofed over by occipital flanges of tabular and postparietal sometimes bordered laterally and ventrally by ossified dorsolateral extension of opisthotic meeting ventromedial flange of tabular (0) small fossa near ventrolateral corner of occiput bordered laterally by ventromedial flange of tabular, roofed over by dorsal portion of lateral margin of supraoccipital-opisthotic complex and floored by lateral extensions of opisthotic (1) absence of fossa (2). [This character was revised from Ruta et al. (2003) following Clack (2003a)]. STATE 1: Cotylosauria. STATE 2: Anthracosauria.

Opisthotic

98. OPI 1. Absence (0) or presence (1) of condition: opisthotic forming thick plate with supraoccipital, separating exoccipitals from skull table. STATE 1: archeriamorphs, Cotylosauria. 336 K. PAWLEY PHD THESIS

Temporal fenestra

99. # TEM FEN 1. Absence (0) or presence (1) of broad postorbital opening (aïstopod pattern).

Stapes

100. ¨ STAP 1. Hyomandibular / stapes: broad and plate like (0) distally elongated rod (1) rod with broad round footplate (2). STATE 1: basal Terrapoda (Temnospondyli, Seymouriamorpha). STATE 2: Neospondyli more derived than Seymouriamorpha.

Vomer

101. ¨ VOM 1. Absence (0) or presence (1) of condition: ventral, exposed surface of vomers narrow elongate and strip-like, without extensions anterolateral or postero lateral to choana and two and a half to three times longer than wide. STATE 1: Anthracosauria more derived than Gephyrostegus, Neospondyli except Nectridia. 102. VOM 2. Vomers: form a large sheet of bone between the anterior end of the cultriform process and the posterior end of the anterior palatal fossa (0) forming a narrow bar between the anterior end of the cultriform process and the posterior edge of the anterior palatal fossa (1). STATE 1: some Temnospondyli (Dvinosauria more derived than Neldasaurus). 103. VOM 3. Vomer with (0) or without (1) fang pair. STATE 1: Ichthyostega, Anthracosauria more derived than Gephyrostegus, some Temnospondyli (derived Amphibamidae: Amphibamus plus Doleserpeton), Neospondyli more derived than Seymouriamorpha. 104. VOM 4. Vomer without (0) or with (1) denticles. STATE 1: Tulerpeton, Baphetidae, Gephyrostegidae, Temnospondyli (except Nigerpeton, Dvinosauria, Sclerocephalus, Mastodonsauroidea), Neospondyli (except Nectridia). 105. VOM 5. Vomer excluded from (0) or contributing to (1) interpterygoid vacuities. STATE 1: some Temnospondyli (derived Dendrerpetontidae: Dendrerpeton plus Balanerpeton, Dvinosauria plus Eryopoidomorpha), derived Diplocaulidae (Diplocaulus plus Diploceraspis). 106. VOM 6. Vomers: with a tooth row running between the vomerine tusks (0) without a row of teeth between the vomerine tusks (1). [Repolarised from Chapter 5 (Appendix 8).] STATE 1: Tetrapoda more derived than Acanthostega except Whatcheeriidae; Crassigyrinus; some Temnospondyli (Saharastega, Trimerorhachis plus Dvinosauroidea, Lydekkerina plus Mastodonsauroidea), Nectridia. 107. VOM 7. Vomer not forming (0) or forming (1) suture with maxilla anterior to choana. STATE 1: Ichthyostega, Crassigyrinus, some Temnospondyli (some Cochleosauridae: Nigerpeton plus Adamanterpeton plus Cochleosaurus floriensis; Dvinosauria; Stereospondylomorpha). 108. VOM 8. Vomer with (0) or without (1) toothed lateral crest. STATE 1: Tetrapoda more derived than Crassigyrinus. 109. VOM 9. Vomer with (0) or without (1) anterior crest. STATE 1: Ichthyostega, Tulerpeton, Tetrapoda more derived than Crassigyrinus. APPENDIX 14: EARLY TETRAPOD CHARACTER STATE LIST 337

110. VOM 10. Vomer in contact with anterior ramus of pterygoid (0) or not (1). STATE 1: some Temnospondyli (Eobrachyopidae, Tupilakosauridae, Amphibamidae, Stereospondylomorpha more derived than Archegosauridae), derived Diplocaulidae (Diplocaulus plus Diploceraspis). 111. # VOM 11. Vomer without (0) or with (1) transverse patch of small teeth posteromedial to choana. 112. VOM 13. Vomer-palatine suture lateral to choana: absent (0) present (1). STATE 1: some Temnospondyli (some Dvinosauria: Neldasaurus, Eobrachyopidae, Tupilakosauridae), derived Captorhinomorpha (Paleothyris plus Petrolacosaurus). 113. VOM 14. Tooth row along the medial margin of the choana: present (0) absent (1). [Repolarised from Chapter 5 (Appendix 8).] STATE 1: Tetrapoda more derived than Crassigyrinus (except some Temnospondyli: Trimerorhachis, Dvinosauroidea, Stereospondyli; derived Microsauria: Rhynchonkos plus Gymnarthridae). 114. VOM 15. Anterior part of cultriform process of parasphenoid clasped by medial posterior extensions of the vomers: absent (0) present (1). STATE 1: some Temnospondyli (Eryopoidomorpha except Amphibamidae, Archegosauridae). 115. VOM 16. Vomerine tusks: subequal in diameter to palatal tusks (0) half the width of palatal tusks or less (1). [Repolarised from Chapter 5 (Appendix 8).] STATE 1: Colosteidae (Greererpeton), Temnospondyli except Dvinosauria (except Isodectes) and Stereospondylomorpha. 116. VOM 17. Vomerine ridges radiating from anterior edge of choanae to snout margins: absent (0) present (1). STATE 1: some Temnospondyli (Edops, derived Cochleosauridae: Adamanterpeton plus Cochleosaurus spp.).

Palatine

117. PAL 1. Palatine with (0) or without (1) tusks. [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Ichthyostega, some Temnospondyli (derived Amphibamidae: Amphibamus plus Doleserpeton), all Neospondyli (except Seymouria plus Discosauriscus). 118. PAL 2. Palatine (and ectopterygoid): without (0) or with (1) denticles. [This character was merged with ECT 3 from Ruta et al. (2003) because they are not independent.] STATE 1: Pederpes, Baphetidae, Gephyrostegus, Temnospondyli (except some Cochleosauridae: Nigerpeton, Chenoprosopus lewisi; Dvinosauria, Sclerocephalus, Mastodonsauroidea), Neospondyli (except Rhynchonkos, Nectridia). 119. PAL 3. Palatine excluded from (0) or contributing to (1) interpterygoid vacuities. STATE 1: some Temnospondyli (some Dvinosauria: Eobrachyopidae, Tupilakosauridae; Amphibamidae; Lydekkerina plus Mastodonsauroidea), Diplocaulidae. 120. PAL 4. Palatine with (0) or without (1) tooth row (distinct from tusks). [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Baphetidae, Anthracosauria, Temnospondyli (except Neldasaurus plus Trimerorhachis; Sclerocephalus, Stereospondylomorpha), Seymouriamorpha (except Kotlassia) plus Cotylosauria plus Westlothiana, some Microsauria (Microbrachomorpha). 121. PAL 5. Palatine without (0) or with (1) exposure in lateral orbit margin (LEP). STATE 1: some Temnospondyli (Eobrachyopidae, Dissorophoidea). 338 K. PAWLEY PHD THESIS

122. PAL 8. Posterior process of palatine: absent (0) present, makes point contact with the palatine ramus of the pterygoid and excludes ectopterygoid from border of interpterygoid vacuity (1). STATE 1: some Temnospondyli (Mastodonsauroidea).

Ectopterygoid

123. ECT 1. Presence (0) or absence (1) of ectopterygoid. STATE 1: Doleserpeton, Captorhinus, some Microsauria (Pantylidae), Diplocaulidae where known. 124. ECT 2. Ectopterygoid with (0) or without (1) tusks. [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Acanthostega plus Ichthyostega, some Temnospondyli (Stereospondylomorpha), Neospondyli. 125. ECT 5. Ectopterygoid with (0) or without (1) tooth row (distinct from tusks). [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Baphetidae, Temnospondyli (except some Dvinosauria: Neldasaurus plus Trimerorhachis, Dvinosaurus plus Tupilakosaurus; Sclerocephalus, Stereospondylomorpha), Seymouriamorpha (except Kotlassia) plus Cotylosauria. 126. # ECT 7. Absence (0) or presence (1) of condition: ectopterygoid narrowly wedged between palatine and pterygoid.

Pterygoid

127. PTE 1. Ascending column of pterygoid: absent (0) present, ascending lamina of the pterygoid with a thickened ascending column on its posterior surface directly anterior to exoccipital (1). STATE 1: some Temnospondyli (Mastodonsauroidea). 128. PTE 2. Oblique ridge of pterygoid: absent (0) posterior face of ascending lamina of the pterygoid with a low rounded oblique ridge or large sharp edged crest (1). STATE 1: some Temnospondyli (Stereospondyli). 129. ¨ PTE 3. Absence (0) or presence (1) of pterygoid flange oriented transversely. STATE 1: basal Neospondyli (Seymouriamorpha plus Cotylosauria). 130. PTE 4. Pterygoid flange (where present) without (0) or with (1) row of teeth. [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Limnoscelis plus Captorhinomorpha. 131. PTE 7. Quadrate (posterior) ramus of pterygoid laterally oriented in ventral view, rather than posterolaterally oriented: absent (0) or present (1). [This character was re- written from Ruta et al. (2003) in order to clarify it.] STATE 1: Diplocaulidae. 132. PTE 9. Pterygoid without (0) or with (1) postero-lateral flange. STATE 1: Temnospondyli (except Dvinosauria, Stereospondyli), Seymouriamorpha plus Cotylosauria, some Microsauria (Tuditanidae plus Hapsidopareiontidae, Gymnarthridae). 133. PTE 10. Pterygoids not sutured with each other (0) or sutured (1). STATE 1: Tetrapoda (except Edops, some Lepospondyli: Hyloplesion, Pelodosotis, Rhynchonkos, Diplocaulidae except Batrachiderpeton). 134. # PTE 11. Pterygoid not sutured with maxilla (0) or sutured (1). APPENDIX 14: EARLY TETRAPOD CHARACTER STATE LIST 339

135. PTE 13. Parasphenoid articulates with depression in the corpus of the pterygoid, behind a short triangular medial process of the pterygoid (0) an elongate cylindrical to hemi-cylindrical process medial process of the pterygoid abuts the parasphenoid (1) corpus of the pterygoid forming a broad contact along the lateral margins of the parasphenoid plate (2). STATE 1: Temnospondyli (except Caerorhachis plus Edops, Neldasaurus plus Trimerorhachis, Tupilakosauridae, Stereospondyli). STATE 2: some Temnospondyli (Tupilakosauridae, Stereospondyli), derived Diplocaulidae (Diceratosaurus plus Diplocaulus plus Diploceraspis). 136. PTE 14. Absence (0) or presence (1) of condition: quadrate ramus of pterygoid robust, indistinctly merging into basal and palatal processes. STATE 1: some Temnospondyli (Dvinosauroidea). 137. PTE 15. Absence (0) or presence (1) of condition: quadrate ramus of pterygoid straight, rod-like and gently tapering distally. STATE 1: some Temnospondyli (derived Amphibamidae: Amphibamus plus Doleserpeton). 138. PTE 16. Palatal ramus of pterygoid without (0) or with (1) distinct, anterior digitiform process. STATE 1: Colosteidae. 139. PTE 17. Basal region of pterygoid immediately anterior to quadrate ramus without (0) or with (1) sharply defined, elongate longitudinal groove. STATE 1: some Microsauria (Hapsidopareiontidae, Ostodolepidae plus Rhynchonkos plus Gymnarthridae). 140. PTE 18. Quadrate (posterior) ramus of the pterygoid: twisted dorsally from the plane of the corpus and the palatine ramus, to form a subvertical plate (0) untwisted, forming a near horizontal plate, continuous with the plane of the corpus and the palatal ramus of the pterygoid (1) strongly turned ventrally relative to the palatal ramus, creating a vaulted palate (2). Ordered. STATE 1: Diplocaulidae more derived than Scinosaurus, where known (Batrachiderpeton, Diplocaulus plus Diploceraspis). STATE 2: some Temnospondyli (Dvinosauroidea). 141. PTE 20. Palatine ramus of the pterygoid: extends anterior to the anterior most ectopterygoid tooth (0) retracted posterior to the anterior most ectopterygoid tooth (1). STATE 1: some Temnospondyli (Tupilakosauridae, Amphibamidae, Lydekkerina plus Mastodonsauroidea), derived Diplocaulidae (Diplocaulus plus Diploceraspis). 142. PTE 21. Ornament on palatine ramus of pterygoid: absent (0) present (1). STATE 1: some Temnospondyli (Peltobatrachus, Lydekkerina plus Benthosuchus plus Eryosuchus). 143. PTE 22. Denticles on palatine ramus of pterygoid: present (0) absent (1). STATE 1: Ichthyostega, some Temnospondyli (Nigerpeton, Dvinosaurus plus Tupilakosauridae, Mastodonsaurus), Rhynchonkos, derived Diplocaulidae (Diceratosaurus plus Diplocaulus plus Diploceraspis). 144. PTE 23. Dorsoventrally oriented ridge on medial surface of ascending ramus of pterygoid: absent (0) present (1). STATE 1: some Temnospondyli (Dvinosauria). 145. PTE 24. Lateral pterygoid alae, extended posterior to ectopterygoid: absent (0) present (1). [Repolarised from Chapter 5 (Appendix 8).] STATE 1: Anthracosauria, Temnospondyli (except Cochleosaurus bohemicus; Dvinosauria more derived than Neldasaurus; Eryopoidomorpha), Urocordylidae. 146. PTE 25. Denticulate vomerine-pterygoid ridge passes from vomerine tusk to medial edge of interpterygoid vacuity: absent (0) present (1). [Repolarised from Chapter 5 (Appendix 8).] STATE 1: some Temnospondyli (Dendrerpetontidae plus Cochleosauridae). 340 K. PAWLEY PHD THESIS

Epipterygoid

147. EPI 1. Ascending ramus of epipterygoid: unossified or slender rod (0) robust (1). STATE 1: some Temnospondyli (Mastodonsauroidea).

Choanae

148. CHO 1. Absence (0) or presence (1) of condition: choanae wider anteriorly than posteriorly. STATE 1: some Temnospondyli (Cochleosauridae more derived than Saharastega).

Anterior palatal fossa

149. ANT FOS 1. Anterior palatal fossa (defined as a depression in the anterior part of the vomers, which may or may not be perforated by an anterior palatal vacuity, see ANT VAC 1): absent (0) present (1). State 1: Megalocephalus, some Temnospondyli (some Cochleosauridae: Chenoprosopus spp., Cochleosaurus spp.; Euskelia (except derived Amphibamidae); Stereospondylomorpha).

Anterior palatal vacuities

150. ANT VAC 1. Presence (0) or absence (1) of anterior palatal vacuities (defined as perforation/s of the anterior part of the vomers). STATE 1: Baphetes, Anthracosauria plus Temnospondyli (except Saharastega plus Nigerpeton, Dvinosauria, Eryopoidomorpha (except derived Amphibamidae) plus Neospondyli. 151. ANT VAC 2. Anterior palatal vacuity (where present) single (0) or double (1). [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Acanthostega, archeriamorphs, Temnospondyli (except Lydekkerina plus Benthosuchus plus Eryosuchus).

Interpterygoid vacuities

152. INT VAC 1. Absence (0) or presence (1) of interpterygoid vacuities. [Repolarised from Ruta et al. (2003).] STATE 1: Colosteidae, Anthracosauria plus Temnospondyli, Neospondyli more derived than Seymouriamorpha (except Batrachiderpeton). 153. INT VAC 2. Absence (0) or presence (1) of condition: interpterygoid vacuities (where present) occupying at least half of palatal width. [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Temnospondyli more derived than Caerorhachis plus Edops, Diplocaulidae more derived than Scinosaurus, Ptyonius. 154. INT VAC 3. Absence (0) or presence (1) of condition: interpterygoid vacuities (where present) concave along their entire margins. [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Temnospondyli more derived than Caerorhachis plus Edops (except Tupilakosauridae), derived Diplocaulidae (Diceratosaurus plus Diplocaulus plus Diploceraspis).

Parasphenoid

155. PASPHE 3. Parasphenoid without (0) or with (1) posterolaterally directed, ventral thickenings (ridges ending in basal tubera). STATE 1: archeriamorphs, Cotylosauria. APPENDIX 14: EARLY TETRAPOD CHARACTER STATE LIST 341

156. PASPHE 4. Foramen for the internal carotid artery (where observable) on the ventral or lateral surface of the parasphenoid plate (0) internal carotid and the palatine and intracranial branches pass through the dorsal surface of the parasphenoid plate (1). STATE 1: Stereospondyli. 157. PASPHE 5. Denticles on parasphenoid: confined to medial area of parasphenoid (0) transverse belt extending to parasphenoid-pterygoid articulations (1) absent (2). STATE 1: some Temnospondyli (some Stereospondyli: Rhinesuchidae plus Lydekkerina plus Benthosuchus). STATE 2: some Temnospondyli (Dvinosauria, Dissorophidae except Ecolsonia, Mastodonsauridae). 158. PASPHE 7. Posterior end of the ventral surface of the parasphenoid plate without sharp rimmed depressions (0) rounded, widely separated depressions at the posterior end of the ventral surface of the ventral surface of the parasphenoid plate, with sharp anterior rims (crista muscularis) (1) depressions transversely widened so that the muscular crests approach each other forming transverse ridges (2). Ordered. [This character was re- written from Ruta et al. (2003) in order to clarify it and extend it to include characters suitable for temnospondyls.] STATE 1: some Temnospondyli (Cochleosauridae, Peltobatrachus plus basal Euskelia (Sclerocephalus plus Eryops), basal Stereospondylomorpha (Archegosauridae plus Rhinesuchidae plus Lydekkerina). STATE 2: Mastodonsauroidea. 159. PASPHE 8. Parasphenoid plate anteriorly expanded so that it is much wider anteriorly than posteriorly: absent (0) present (2). STATE 1: some Temnospondyli (Eobrachyopidae). 160. PASPHE 9. Ventral cranial fissure not sutured (0) sutured (1) eliminated (2). Ordered. STATE 1: Acanthostega, Crassigyrinus, Whatcheeria. STATE 2: Tetrapoda more derived than Crassigyrinus and Whatcheeria. 161. PASPHE 11. Parasphenoid without (0) or with (1) anterolateral extensions meeting pterygoids and projecting anterior to cultriform process insertion. [This character was re- written from Ruta et al. (2003) in order to clarify it.] STATE 1: some Temnospondyli (Amphibamidae). 162. PASPHE 12. Parasphenoid without (0) or with (1) triangular denticle patch with raised margins at base of cultriform process. STATE 1: some Temnospondyli (Dendrerpetontidae, Cochleosaurus spp., Amphibamidae, Archegosauridae).

Exoccipital

163. EXOCC 1. Exoccipital-pterygoid suture: absent or contact not visible ventrally (0) visible in ventral view (1). STATE 1: some Temnospondyli (Tupilakosauridae, Lydekkerina plus Mastodonsauroidea), derived Diplocaulidae (Diplocaulus plus Diploceraspis). 164. EXOCC 2. Absence (0) or presence (1) of condition: exoccipitals enlarged to form flattened, widely spaced double occipital condyles. STATE 1: Diplocaulidae. 342 K. PAWLEY PHD THESIS

165. EXOCC 3. Occipital condyle: single with the basioccipital forming the largest contribution to the articulating surface (0) bilobed with reduced basioccipital contribution (1) double with no basioccipital contribution (2). Ordered. [This character was re-written from Ruta et al. (2003) in order to clarify it and extend it to include characters suitable for temnospondyls.] STATE 1: some Temnospondyli (Eryopoidomorpha except Lydekkerina plus Mastodonsauroidea). STATE 2: some Temnospondyli (Lydekkerina plus Mastodonsauroidea). 166. EXOCC 4. Absence (0) or presence (1) of condition: exoccipitals forming continuous concave strap-shaped articular surfaces with basioccipital. STATE 1: Microsauria. 167. EXOCC 5. Absence (0) or presence (1) of condition: exoccipitals expanded and adpressed to each other, so as to obliterate basioccipital posterior surface. STATE 1: some Temnospondyli (Dissorophoidea).

Basioccipital

168. BASOCC 1. Basioccipital notochordal (0) or not (1). STATE 1: Tetrapoda more derived than Crassigyrinus. 169. BASOCC 5. Articular surface of basioccipital (where ossified) not convex (0) or convex (1). [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: derived Captorhinomorpha (Paleothyris plus Petrolacosaurus). 170. # BASOCC 6. Absence (0) or presence (1) of condition: basioccipital circular and recessed.

MANDIBLE

171. MAND 1. Retroarticular process: absent (0) present (1). STATE 1: some Temnospondyli (Dvinosauria, Peltobatrachus, Lydekkerina plus Mastodonsauroidea), some Microsauria (Batropetes, Ostodolepidae, Rhynchonkos), Nectridia. 172. MAND 2. Arcadian groove Arcadian groove along suture between surangular and angular (sensu Jupp and Warren, 1986): absent (0) present at the posteroventral end of the mandible (1). [Repolarised from Chapter 5 (Appendix 8).] STATE 1: Temnospondyli. 173. MAND 3. Postglenoid area with a transverse trough behind the glenoid socket followed by a dorsally bulging tip: absent (0) present (1). STATE 1: some Temnospondyli (Dvinosaurus plus Tupilakosauridae).

Parasymphysial plate

174. ¨ PSYM 1. Presence (0) or absence (1) of parasymphysial plate. State 1: Terrapoda. 175. PSYM 2. Parasymphysial plate (where present) without (0) or with (1) paired tusks. [This character was re-written from Ruta et al. (2003) in order to clarify it.] State 1: Acanthostega plus Ichthyostega, Crassigyrinus, Colosteidae, Baphetidae. 176. PSYM 3. Parasymphysial plate (where present) with (0) or without (1) tooth row/s. [This character was re-written and repolarised from Ruta et al. (2003) in order to clarify it.] State 1: archeriamorphs more derived than Whatcheeria. APPENDIX 14: EARLY TETRAPOD CHARACTER STATE LIST 343

177. ¨ PSYM 4. Parasymphysial plate (where present) without (0) or with (1) denticles. [This character was re-written from Ruta et al. (2003) in order to clarify it.] State 1: Tetrapoda.

Dentary

178. ¨ DEN 1. Dentary with (0) or without (1) accessory tooth row. STATE 1: Tetrapoda (except Captorhinus). 179. ¨ DEN 2. Dentary with (0) or without (1) anterior fang pair. STATE 1: some Temnospondyli (Amphibamidae: Doleserpeton), Neospondyli. 180. DEN 3. Dentary without (0) or with (1) chamfered ventral margin. STATE 1: Ventastega plus Acanthostega. 181. DEN 4. Dentary without (0) or with (1) U-shaped notch for premaxillary tusks. STATE 1: Colosteidae.

Splenial

182. SPL 1. Splenial: forms part of jaw symphysis (0) excluded from symphysis (1). STATE 1: some Temnospondyli (Dendrerpetontidae, Edops, Cochleosauridae except Chenoprosopus spp., Eryops), some Microsauria (Microbrachomorpha, Rhynchonkos, Gymnarthridae). 183. ¨ SPL 3. Absence (0) or presence (1) of suture between splenial and anterior coronoid (where present). [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Tetrapoda (except Crassigyrinus). 184. SPL 4. Absence (0) or presence (1) of suture between splenial and middle coronoid (where present). [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Baphetidae plus Anthracosauria, Neospondyli where observable (except Kotlassia).

Postsplenial

185. POSPL 1. Presence (0) or absence (1) of postsplenial. STATE 1: Cotylosauria (except Petrolacosaurus), Westlothiana, some Microsauria (Batropetes, Hapsidopareion), Nectridia. 186. POSPL 2. Postsplenial (where present) without (0) or with (1) mesial lamina. [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Tetrapoda more derived than Colosteidae. 187. POSPL 3. Postsplenial (where present) with (0) or without (1) pit-line. [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Acanthostega, archeriamorphs plus Terrapoda.

Angular

188. # ANG 1. Presence (0) or absence (1) of angular. 189. ANG 2. Angular without (0) or with (1) mesial lamina. STATE 1: Tetrapoda more derived than Tulerpeton. 344 K. PAWLEY PHD THESIS

190. ¨ ANG 4. Angular reaching (0) or not reaching (1) posterior end of lower jaw. STATE 1: Neospondyli.

Surangular

191. # SURANG 1. Presence (0) or absence (1) of surangular. 192. SURANG 3. Surangular with (0) or without (1) pit-line. STATE 1: Tetrapoda more derived than Ventastega. 193. SURANG 5. Absence (0) or presence (1) of condition: lateral exposure of the surangular smaller than that of the angular. STATE 1: some Microsauria (Gymnarthridae), Diplocaulidae.

Prearticular

194. PREART 1. Prearticular: not extending posterior to the level of the glenoid (0) extending posterior to the level of the glenoid, covering the medial face of the articular (1). STATE 1: some Temnospondyli (Sclerocephalus, Archegosauridae). 195. PREART 2. Prearticular-surangular suture: absent (0) present (1). STATE 1: Tetrapoda more derived than Crassigyrinus. 196. PREART 3. Longitudinal dorsal ridge on prearticular: absent (0) present (1). STATE 1: Tiktaalik plus ichthyostegamorphs, Whatcheeriidae, Anthracosauria. 197. PREART 4. Denticle field along at least the whole dorsal edge of prearticular: present (0) absent (1). STATE 1: Tetrapoda more derived than Crassigyrinus (except Caerorhachis). 198. PREART 5. Prearticular sutured with splenial (0) or not (1). STATE 1: some Temnospondyli (Euskelia, Lydekkerina plus Mastodonsauroidea). 199. PREART 6. Hamate process projecting dorsally immediately in front of the anteromedial corner of the glenoid of the mandible: absent (0) present (1). STATE 1: some Temnospondyli (Mastodonsauridae).

Anterior coronoid

200. ANT COR 1. Anterior coronoid present (0) or absent (1). STATE 1: derived Neospondyli: Cotylosauria except Limnoscelis, Lepospondyli). 201. ANT COR 2. Anterior coronoid (where present) with (0) or without (1) tusks. [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Tetrapoda more derived than Ventastega (except Greererpeton, Gephyrostegus, Thabanchuia). 202. ANT COR 3. Anterior coronoid (where present) without (0) or with (1) denticles. [Repolarised from Ruta et al. (2003).] STATE 1: archeriamorphs (except Whatcheeria, Baphetidae) plus Terrapoda (except Mastodonsauroidea). 203. ANT COR 4. Anterior coronoid (where present) with (0) or without (1) tooth row. STATE 1: archeriamorphs (except Whatcheeria) plus Terrapoda. APPENDIX 14: EARLY TETRAPOD CHARACTER STATE LIST 345

Middle coronoid

204. MID COR 1. Middle coronoid present (0) or absent (1). STATE 1: some Neospondyli (Cotylosauria except Limnoscelis), Pantylus, Nectridia where observable. 205. MID COR 2. Middle coronoid (where present) with (0) or without (1) tusks. [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Tetrapoda more derived than Ventastega (except Gephyrostegus, Thabanchuia) 206. MID COR 3. Middle coronoid (where present) without (0) or with (1) denticles. [Repolarised, and re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: archeriamorphs (except Whatcheeria, Baphetidae), Terrapoda (except Mastodonsauroidea) 207. MID COR 4. Middle coronoid (where present) with (0) or without (1) marginal tooth row. [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: archeriamorphs (except Whatcheeria), Terrapoda.

Posterior coronoid

208. POST COR 1. Posterior coronoid present (0) or absent (1). STATE 1: some Microsauria (Gymnarthridae), Nectridia. 209. ¨ POST COR 2. Posterior coronoid (where present) with (0) or without (1) tusks. [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Tetrapoda (except Thabanchuia). 210. POST COR 3. Posterior coronoid (where present) without (0) or with (1) denticles. [Repolarised, and re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Colosteidae, Anthracosauria, Terrapoda (except Captorhinomorpha). 211. POST COR 4. Posterior coronoid (where present) with (0) or without (1) tooth row. [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Archeriamorphs (except Whatcheeria, Anthracosauridae) plus Terrapoda. 212. POST COR 5. Posterior coronoid (where present) without (0) or with (1) posterodorsal process, which is exposed in lateral view and contributes to the surangular crest. [This character was merged with POST COR 6 and POST COR 7 from Ruta et al. (2003) because they are not independent.] STATE 1: Greererpeton, some Anthracosauria (Gephyrostegus, Archeria, Pholiderpeton scutigerum), Temnospondyli (except Lydekkerina, Mastodonsauroidea), Neospondyli.

Adductor fossa

213. ADD FOS 1. Adductor fossa facing dorsally (0) or medially (1). STATE 1: Tetrapoda more derived than Crassigyrinus.

Teeth

214. TEETH 1. Absence (0) or presence (1) of pedicely on marginal teeth. STATE 1: some Temnospondyli (derived Amphibamidae: Amphibamus plus Doleserpeton). 215. TEETH 2. Marginal teeth monocuspid (0) or multicuspid (1). STATE 1: some Temnospondyli (derived Amphibamidae: Amphibamus plus Doleserpeton), Batropetes. 346 K. PAWLEY PHD THESIS

216. TEETH 3. Marginal teeth without (0) or with (1) two cuspules labiolingually arranged. STATE 1: Temnospondyli (derived Amphibamidae: Amphibamus plus Doleserpeton). 217. TEETH 6. Marginal tooth crowns not chisel-tipped (0) or chisel-tipped (1). STATE 1: Anthracosauria more derived than Anthracosauridae. 218. # TEETH 7. Marginal tooth crowns without (0) or with (1) dimple. 219. TEETH 8. Absence (0) or presence (1) of condition: marginal tooth crowns robust, conical structures. STATE 1: some Microsauria (Pantylidae, Gymnarthridae).

POSTCRANIAL CHARACTERS

Vertebra

Neural spine

220. ¨ TRU VER 1. Extra articulations above zygapophyses in at least some trunk and caudal vertebrae: absent (0) or present (1). STATE 1: Nectridia. 221. ¨ TRU VER 2. Neural and haemal spines rectangular to fan-shaped in lateral view: absent (0) or present (1). STATE 1: Nectridia. 222. ¨ TRU VER 10. Neural spines without (0) or with (1) distinct convex “swollen” lateral surfaces. STATE 1: basal Neospondyli: Seymouriamorpha plus Cotylosauria (derived Captorhinomorpha: Paleothyris plus Petrolacosaurus). 223. ¨ m TRU VER 11. Neural spines of trunk vertebrae: not attached to centra (0) closely articulated, suturally attached, or fused (1). [N.B. The derived state of this character develops with morphogenesis; consequently, it is indeterminate in morphogenetically immature specimens in which the overall degree of ossification of the postcranial skeleton is low. This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: some Temnospondyli (Doleserpeton), Neospondyli. 224. ¨ m TRU VER 20. Pre- and postzygapophyses on trunk vertebrae: absent (0) or present (1). [N.B. The derived state of this character develops with morphogenesis; consequently, it is indeterminate in morphogenetically immature specimens in which the overall degree of ossification of the postcranial skeleton is low.] STATE 1: Tetrapoda. 225. ¨ m TRU VER 28. Supraneural canal present in neural arch of at least anterior thoracic vertebrae (0) or does not penetrate neural arch (1). [N.B. The only terrapod known to retain a supraneural canal is Discosauriscus. All specimens of this taxon are morphogenetically immature; therefore, it is plausible that the supraneural canal may be lost with increasing ossification, therefore this character state is indeterminate in morphogenetically immature specimens.] STATE 1: Terrapoda. 226. ¨ m TRU VER 29. Anterior and posterior surfaces of neural spines: parallel in lateral view (0) or sides converge dorsally to form a triangular outline (1). [N.B. The derived state of this character is also characteristic of early morphogenetic stages in which the overall degree of ossification of the postcranial skeleton is low, in which case this derived state is indeterminate. The derived state is relevant in morphogenetically mature specimens with a high overall degree of ossification of the postcranial skeleton.] STATE 1: some Temnospondyli (Doleserpeton), Neospondyli (except Nectridia). APPENDIX 14: EARLY TETRAPOD CHARACTER STATE LIST 347

227. TRU VER 30. Dorsal neural spine: smooth (0) crenulated (1). STATE 1: Nectridia (except derived Diplocaulidae: Diceratosaurus plus Diplocaulus plus Diploceraspis). 228. TRU VER 31. Height of neural spines: even throughout (0) or alternating (1). [N.B. The derived state of this character develops with morphogenesis; consequently, it is indeterminate in morphogenetically immature specimens in which the overall degree of ossification of the postcranial skeleton is low.] STATE 1: Limnoscelis plus Captorhinomorpha, Microsauria.

Centra

229. ¨ TRU VER 33. Trunk centra: diplospondylous, two large components per vertebral centra (rhachitomous or embolomerous) (0) essentially monospondylous, pleurocentra dominant, intercentra highly reduced or absent (gastrocentrous) (1) essentially monospondylous, intercentra dominant, pleurocentra highly reduced (stereospondylous) (2). STATE 1: some Temnospondyli (Peltobatrachus, Doleserpeton), Neospondyli. STATE 2: derived Temnospondyli (some Mastodonsauroidea: Mastodonsaurus).

Pleurocentra

230. TRU VER 6. Long, distally bifurcated transverse processes on trunk pleurocentra: absent (0) or present (1). [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: derived Diplocaulidae (Diplocaulus plus Diploceraspis). 231. ¨ m TRU VER 8. A majority of trunk pleurocentra not fused ventrally (0) or fused. [N.B. The derived state of this character develops with morphogenesis and varies along the vertebral column; examination of a large number of trunk vertebrae from the same specimen is necessary for correct coding. This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Anthracosauria, some Temnospondyli (Caerorhachis, Tupilakosauridae, Peltobatrachus, derived Amphibamidae: Amphibamus plus Doleserpeton), Neospondyli. 232. m TRU VER 9. A majority of trunk pleurocentra not fused dorsally (0) or fused (1). [N.B. The derived state of this character develops with morphogenesis and varies along the vertebral column; examination of a large number of trunk vertebrae from the same specimen is necessary for correct coding. This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Derived Anthracosauria (Archeria plus Pholiderpeton spp.), some Temnospondyli (Tupilakosauridae, Peltobatrachus), Neospondyli. 233. TRU VER 12. Bicipital rib bearers on trunk pleurocentra: absent (0) or present (1). [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: derived Diplocaulidae (Diplocaulus plus Diploceraspis). 234. # TRU VER 15. Enlarged ventral carinae, forming pronounced ridges on trunk pleurocentra: absent (0) or present (1). [This character was re-written from Ruta et al. (2003) in order to clarify it.] 235. TRU VER 18+. Striate ornament on vertebral pleurocentra absent (0) or present (1). STATE 1: derived Diplocaulidae (Diceratosaurus plus Diplocaulus plus Diploceraspis) 236. # TRU VER 34. Vertebrarterial canals pierce bases of trunk diapophyses: absent (0) present (1). 348 K. PAWLEY PHD THESIS

Intercentra

237. TRU VER 13. Trunk intercentra: present (0) or absent (1). STATE 1: some Temnospondyli (Peltobatrachus), Lepospondyli except derived Microsauria (Ostodolepidae plus Rhynchonkos plus Gymnarthridae). 238. m TRU VER 14. A majority of anterior trunk intercentra (where present): not fused mid dorsally (0) fused (1). [N.B. The derived state of this character develops with morphogenesis and varies along the vertebral column; examination of a large number of trunk vertebrae from the same specimen is necessary for correct coding. This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: derived Anthracosauria (Archeria plus Pholiderpeton spp.), some Temnospondyli (Tupilakosauridae, Mastodonsaurus). 239. TRU VER 35. Posterolaterally directed tubercles on cervical intercentra (where present) absent (0) or present (1). [N.B. the distribution of this characteristic can be difficult to determine, due to lack of detailed descriptions of cervical vertebrae.] STATE 1: Greererpeton, Anthracosauria, Trimerorhachis.

Cervical vertebrae

240. ¨ m CER VER 1. Atlantal neural spines not fused (0) or dorsally fused (1). [N.B. The derived state of this character develops with morphogenesis; consequently it is indeterminate in morphogenetically immature specimens in which the overall degree of ossification of the postcranial skeleton is low.] STATE 1: derived Temnospondyli (Mastodonsauroidea), Westlothiana, Lepospondyli. 241. ¨ m CER VER 2. Atlantal neural arch and atlantal centrum: separate (0) fused (1). [N.B. The derived state of this character develops with morphogenesis; consequently it is indeterminate in morphogenetically immature specimens in which the overall degree of ossification of the postcranial skeleton is low.] STATE 1: derived Temnospondyli (Dissorophoidea where known, Stereospondylomorpha), Lepospondyli. 242. CER VER 4. Axial centrum orientation: in plane of axial skeleton (0) or sloping anterodorsally (1). STATE 1: derived Captorhinomorpha (Paleothyris plus Petrolacosaurus). 243. ¨ CER VER 5. Atlantal neural spine: similar to but smaller than axial spine (0) reduced to cylindrical spinous process (1). STATE 1: Terrapoda except Microsauria. 244. ¨ CER VER 6. Axial centrum: with rounded anterodorsal edge (0) or with strong anterior (odontoid) process (1). STATE 1: Lepospondyli. 245. ¨ CER VER 7. Anterior surface of atlas centrum: equal in diameter to posterior surface (0) or laterally expanded (1). STATE 1: Lepospondyli, except Sauropleura spp. 246. CER VER 8. Atlantal pleurocentrum and axial intercentrum discrete elements (0) or fused (1). STATE 1: Cotylosauria, indeterminate in the Lepospondyli. 247. # CER VER 9. Foramen for exit of first spinal nerve in atlas absent (0) or present (1). 248. CER VER 10. Atlantal neural spine with prezygapophyses, which articulate with proatlas: present (0) absent (1). STATE 1: derived Temnospondyli (Mastodonsauroidea), Nectridia. APPENDIX 14: EARLY TETRAPOD CHARACTER STATE LIST 349

Sacral vertebrae

249. ¨ SAC VER 1. Sacral vertebrae and/or ribs: absent (0) one pair (1) two or more pairs (2). STATE 1: Tetrapoda. STATE 2: Diadectomorpha plus Captorhinomorpha, tuditanomorph Microsauria (except Pelodosotis, Saxonerpeton, Pantylus).

Caudal vertebrae

250. ¨ CAU VER 1 (TRU VER 3). Neural and haemal spines facing each other dorsoventrally: absent (0) or present (1). STATE 1: Nectridia. 251. CAU VER 2 (TRU VER 5). Extra articulations on haemal spines absent (0) or present (1). STATE 1: some Nectridia (Urocordylidae). 252. CAU VER 3 (TRU VER 16). Strong proximal emargination in haemal spines of posterior tail vertebrae absent (0) or present (1). STATE 1: Diplocaulidae more derived than Scinosaurus. 253. CAU VER 4 (TRU VER 17). Strong proximal emargination in haemal spines of anterior tail vertebrae absent (0) or present (1). STATE 1: derived Diplocaulidae (Diceratosaurus plus Diplocaulus plus Diploceraspis). 254. ¨ CAU VER 5 (TRU VER 22). Pre- and postzygapophyses on proximal tail vertebrae: absent (0) or present (1). STATE 1: Tetrapoda. 255. ¨ CAU VER 7 (TRU VER 24). Pre- and postzygapophyses on distal tail vertebrae: absent (0) or present (1). STATE 1: archeriamorphs plus Terrapoda. 256. ¨ CAU VER 9. Caudal intercentra present (haemal arches attached to intercentra) (0) or absent, haemal arches attached to pleurocentra (1). STATE 1: Nectridia. 257. ¨ CAU VER 10. Supraneural spines: present (0) absent (1). STATE 1: archeriamorphs plus Terrapoda. 258. # CAU VER 11. Spinal nerve foramina in centra of caudal vertebra: absent (0) present (1).

Ribs

259. ¨ RIB 1. Atlantal rib: present / diapophyses on atlas neural arch present (0) absent (1). STATE 1: basal Terrapoda (Temnospondyli plus Seymouriamorpha), Nectridia. 260. RIB 2. Cervical ribs with (0) or without (1) expanded distal ends. STATE 1: some Nectridia (Urocordylidae). 261. ¨ m RIB 3. Ribs mostly straight (0) or ventrally curved (along ventral margin) (1) in at least part of the trunk. [N.B. The derived state of this character develops with morphogenesis; the curvature of the ribs increases with increasing morphogenetic stage. The derived state is therefore indeterminate in morphogenetically immature specimens. This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Tetrapoda more derived than Acanthostega. 350 K. PAWLEY PHD THESIS

262. ¨ m RIB 5. Anterior trunk ribs with uncinate processes: absent (0) or present (1). [N.B. The derived state of this character develops with morphogenesis; consequently it is indeterminate in morphogenetically immature specimens in which the overall degree of ossification of the postcranial skeleton is low. This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: archeriamorphs, derived Temnospondyli (Peltobatrachus plus Eryopoidomorpha, except Amphibamidae). 263. # RIB 6. Distally expanded, elongate posterodorsal flange (alar process) in midtrunk ribs absent (0) or present (1). [This character was re-written from Ruta et al. (2003) in order to clarify it.] 264. # m RIB 7. Absence (0) or presence (1) of condition: longest trunk ribs poorly ossified slender rods, the length of which is smaller than the length of three trunk vertebrae. [N.B. the derived state is present in all larval specimens; in this instance, the derived state only pertains to taxa that are otherwise morphogenetically mature. This character was re-written from Ruta et al. (2003) in order to clarify it.] 265. RIB 8. Trunk ribs: tapered distally or parallel sided (0) distally expanded (1). STATE 1: Acanthostega, archeriamorphs (except Anthracosauria), Temnospondyli. 266. # RIB 9. Rib heads: (obscurely) bicapitular (0) unicapitular (1). 267. m RIB 10. Anterior trunk ribs with proximally expanded flange: absent (0) present (1). [N.B. The derived state of this character develops with morphogenesis; consequently, it may be indeterminate in morphogenetically immature specimens in which the overall degree of ossification of the postcranial skeleton is low.] STATE 1: Tiktaalik, Ichthyostega, Whatcheeriidae. 268. ¨ m RIB 11. Posteroventral edge of (first) sacral rib not significantly thickened (0) or posteroventral border thickened and distinct from main body of rib (1). [N.B. The derived state of this character develops with morphogenesis; consequently, it is indeterminate in morphogenetically immature specimens in which the overall degree of ossification of the postcranial skeleton is low.] STATE 1: Terrapoda where known. 269. ¨ m RIB 12. Distal expansion of sacral rib symmetrical relative to proximal head (0) or strongly recurved (1). [N.B. The derived state of this character develops with morphogenesis; consequently, it is indeterminate in morphogenetically immature specimens in which the overall degree of ossification of the postcranial skeleton is low.] STATE 1: Terrapoda where known.

Interclavicle

270. INTCLA 1. Posterior border of interclavicle drawn out into parasternal process: absent (0) present (1). STATE 1: archeriamorphs (except Colosteidae), some Temnospondyli (Dvinosaurus plus Tupilakosauridae, some Stereospondyli (Lydekkerina, Mastodonsaurus), Seymouriamorpha plus Cotylosauria, Microsauria, derived Diplocaulidae (Diplocaulus plus Diploceraspis). 271. INTCLA 2. Parasternal process (where present): tapered (0) elongate and parallel sided for most of its length (1). STATE 1: Ichthyostega, Whatcheeria plus Ossinodus, Dvinosaurus, Seymouriamorpha plus Cotylosauria, Microsauria. 272. INTCLA 5. Interclavicle external ornament: absent (0) present (1). STATE 1: Tetrapoda except Paleothyris, Microsauria. 273. # INTCLA 6. Interclavicle: present (0) absent (1). APPENDIX 14: EARLY TETRAPOD CHARACTER STATE LIST 351

274. INTCLA 7. Posterior border of interclavicle broadly hemispherical: absent (0) present (1). STATE 1: some Temnospondyli (derived Euskelia: Eryops plus Dissorophoidea, except Amphibamus). 275. INTCLA 8. Parasternal process broadly truncate: absent (0) present (1). STATE 1: STATE 1: derived Diplocaulidae (Diplocaulus plus Diploceraspis). 276. INTCLA 9. Location of center of ossification of interclavicle: anterior to or level with line of maximum width (0) posterior to line of maximum width (1). STATE 1: some Temnospondyli (Mastodonsauroidea).

Clavicle

277. # CLA 1. Clavicles: present (0) or absent (1). 278. CLA 3. Clavicular blades broadly contact anterior to main body of interclavicle: absent (0) present (1). [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Colosteidae, some Temnospondyli (Mastodonsauroidea), Neospondyli except Urocordylidae. 279. m CLA 5. Anterior clavicular flange present on dorsal clavicular process: absent (0) present (1). [N.B. this character is subject to morphogenetic change; where present, it is observable only in later morphogenetic stages, and is indeterminate in early morphogenetic stages.] STATE 1: some Temnospondyli (some Dvinosauria: Trimerorhachis, Thabanchuia; basal Mastodonsauroidea: Mastodonsauroidea). 280. ¨ CLA 6. Posterior border of dorsal clavicular process (posterior lamina): concave in lateral view (0) posteriorly expanded dorsal to center of ossification at the junction with the ventral blade, so that it is posteriorly convex between the ventral blade and the dorsal tip (1). STATE 1: Terrapoda except Nectridia. 281. CLA 7. Branchial lamina / clavicular groove (sensu Warren and Turner, 2004): present (0) or absent, ventral anterior border ornamented and not recessed, similar to rest of clavicle (1). STATE 1: Anthracosauria where known, some Temnospondyli (Dvinosauria where observable: Trimerorhachis, Thabanchuia, Lydekkerina plus Mastodonsauroidea), derived Nectridia (Diplocaulus plus Diploceraspis). 282. CLA 8. Clavicular recess (sensu Pawley and Warren, 2006) indenting posterior lamina: absent (0) present (1). STATE 1: some Temnospondyli (derived Euskelia: Eryops, Dissorophus, Ecolsonia). 283. CLA 9. Clavicular rod (sensu Pawley and Warren, 2006) connects smoothly into anterior border of clavicle blade (0) connects medially into blade, so that an anterolateral flange is present on the clavicular blade (1). STATE 1: derived Anthracosauria (Proterogyrinus plus Archeria plus Pholiderpeton), Seymouria.

Anocleithrum

284. ANOCLE 1. Anocleithrum present (0) absent (1). State 1: Anocleithra are typically absent within the Tetrapoda (except Acanthostega, Tulerpeton, Pholiderpeton, and Discosauriscus). 285. ¨ ANOCLE 2. Anocleithrum (where present) external ornament: present (0) absent (1). STATE 1: Tetrapoda. 352 K. PAWLEY PHD THESIS

Cleithrum

286. CLE 1. T-shaped dorsal expansion of cleithrum: absent (0) or present (1). STATE 1: Diplocaulidae more derived than Scinosaurus. 287. ¨ CLE 2. Medially directed flange (sometimes enlarged into postbranchial lamina) along medial edge of clethrum: present (0) or absent (1). [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Terrapoda. 288. ¨ m CLE 3. Cleithrum a large flattened plate, or co-ossified with scapulocoracoid, cleithral shaft absent (0) or long ventral cleithral shaft present with expanded dorsal process (1). [N.B. the initially separate, long stemmed cleithrum sometimes becomes co- ossified with the scapulocoracoid in old specimens of some taxa within the Terrapoda. This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: archeriamorphs plus Terrapoda. 289. CLE 4. Dorsal cleithral process (where present): does not cap scapulocoracoid (0) caps anterodorsal edge of scapulocoracoid (1). [Repolarised from Chapter 5 (Appendix 8).] STATE 1: Whatcheeriidae, Temnospondyli (except Amphibamidae), Diadectes, Microsauria, Diplocaulidae more derived than Scinosaurus. 290. CLE 5. Dorsal cleithral process: not posteriorly extended (0) posteriorly extended to match length of shaft, covers entire dorsal surface of scapula blade (1). STATE 1: some Temnospondyli (Dissorophidae), Diplocaulidae more derived than Scinosaurus. 291. # CLE 6. Cleithrum: present (0) absent (1). 292. CLE 7. Sharp edged cleithral crest (sensu Bystrow and Efremov, 1940) on anterodorsal surface of dorsal cleithral process: present (0) highly reduced or absent (1). STATE 1: derived Temnospondyli (Stereospondyli), Neospondyli. 293. CLE 8. Ventral junction of dorsal cleithral process and cleithral shaft: smoothly rounded or grooved (0) suprascapula lamina (sensu Bystrow and Efremov, 1940) present on lateroventral surface of dorsal cleithral process, clasps anterodorsal surface of scapular blade (1). STATE 1: derived Whatcheeriidae (Ossinodus plus Pederpes) Pholiderpeton scutigerum, derived Temnospondyli (Peltobatrachus plus Eryopoidomorpha), Cotylosauria. 294. ¨ CLE 9. Anterior clavicular recess of the cleithral shaft (sensu Warren and Turner, 2004): present (0) absent (1). STATE 1: Terrapoda. 295. ¨ CLE 10. Rounded, thickened ridge (lateral cleithral ridge, new term) passes down external surface of cleithral shaft: absent (0) present (1). STATE 1: archeriamorphs. 296. ¨ CLE 11. Anteroventral process of cleithrum (sensu Coates and Clack, 1991) clasping dorsal clavicular process of clavicle: present (0) absent (1). STATE 1: archeriamorphs plus Terrapoda. 297. ¨ CLE 12. Cleithrum external ornament: present (0) absent (1). STATE 1: Tetrapoda.

Scapulocoracoid

298. SCACOR 1. Supraglenoid foramen: present (0) absent (1). STATE 1: Greererpeton, Microsauria (except Pantylus). APPENDIX 14: EARLY TETRAPOD CHARACTER STATE LIST 353

299. m SCACOR 2. Glenoid subterminal (0) or not (coracoid extending ventral to posteroventral margin of glenoid) (1). [N.B. This character is indeterminate in morphogenetically immature specimens, in which the glenoid fossa and coracoid plate are unossified.] STATE 1: Tetrapoda except derived Anthracosauria (Archeria plus Proterogyrinus plus Pholiderpeton). 300. m SCACOR 3. Glenoid foramen ventral to midpoint of glenoid: present (0) absent (1). [N.B. This character is indeterminate in morphogenetically immature specimens, in which the glenoid fossa and coracoid plate are unossified. This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Captorhinomorpha, some Microsauria (Pantylidae). 301. ¨ m SCACOR 4. Ventromedially extended infraglenoid buttress: absent (0) present (1). [N.B. The derived state of this character develops with morphogenesis; consequently it is indeterminate in morphogenetically immature specimens, in which the glenoid fossa and coracoid plate are unossified.] STATE 1: Tetrapoda more derived than Acanthostega. 302. m SCACOR 5. Supraglenoid foramen of scapulocoracoid (where present): enclosed (0) ventrally open notch (1). [N.B. The derived state of this character can only be observed in morphogenetically immature specimens lacking a squared off dorsal scapular blade and/or an ossified glenoid fossa and coracoid plate. The supraglenoid foramen becomes enclosed with morphogenesis, so that the derived state is indeterminate if only morphogenetically mature specimens are available.] STATE 1: some Temnospondyli (Mastodonsauroidea). 303. ¨ m SCACOR 6. Scapular blade: narrows dorsally (0) dorsally flared (1). [N.B. The derived state of this character develops with morphogenesis; consequently it is indeterminate in morphogenetically immature specimens, identifiable by the semicircular anterodorsal edge of the scapular blade and unossified glenoid fossa and coracoid plate.] STATE 1: Terrapoda (except derived Captorhinomorpha: Paleothyris plus Petrolacosaurus, and Lepospondyli where known). 304. ¨ SCACOR 7. Canal of supraglenoid foramen oriented: mediolaterally (0) anteroposteriorly (1). [N.B. The derived state of this character develops with morphogenesis; consequently it is indeterminate the earliest morphogenetic stages in which the scapula is lunate and the glenoid fossa and coracoid plate are unossified.] STATE 1: Terrapoda. 305. ¨ m SCACOR 8. Supraglenoid fossa: absent (0) present (1). [N.B. The derived state of this character develops with morphogenesis; consequently it is indeterminate in the earliest morphogenetic stages in which the scapula is lunate and the glenoid fossa and coracoid plate are unossified.] STATE 1: Ichthyostega, Terrapoda. 306. m SCACOR 9. Scapular tubercle (sensu Nikitin, 1997): absent (0) present (1). [N.B. The derived state of this character develops with morphogenesis; consequently it is indeterminate the earliest morphogenetic stages in which the scapula is lunate and the glenoid fossa and coracoid plate are unossified.] STATE 1: some Temnospondyli (Dvinosauria where observable: Trimerorhachis, Dvinosaurus). 307. m SCACOR 10. Glenoid dorsoventrally constricted below area of supraglenoid fossa (0) glenoid width constant along entire length (1). [N.B. This character is indeterminate in morphogenetically immature specimens, in which the glenoid fossa and coracoid plate are unossified.] STATE 1: Anthracosauria. 354 K. PAWLEY PHD THESIS

308. ¨ m SCACOR 11. Posterior end of glenoid essentially laterally oriented (0) glenoid twisted so that the posterior most surface is oriented almost dorsally (1). [N.B. This character is indeterminate in morphogenetically immature specimens, in which the glenoid fossa and coracoid plate are unossified.] STATE 1: Terrapoda. 309. ¨ m SCACOR 12. Infraglenoid ridge (sensu Pawley and Warren, 2005): absent (0) present (1). [N.B. This character is indeterminate in morphogenetically immature specimens, in which the coracoid plate is unossified.] STATE 1: Terrapoda where observable (except Pantylidae). 310. ¨ m SCACOR 13. Infraglenoid recess (sensu Pawley and Warren, 2005): absent (0) present (1). [N.B. This character is indeterminate in morphogenetically immature specimens, in which the coracoid plate is unossified.] STATE 1: Terrapoda where observable (except Pantylidae). 311. ¨ SCACOR 14. Scapular ridge (new term) on the posteromedial surface of the supraglenoid buttress: present (0) absent (1). STATE 1: Terrapoda. 312. ¨ m SCACOR 15. In medial view, anterior edge of the supra/infraglenoid buttress (with glenoid horizontally positioned): anterodorsally oriented (0) dorsally oriented, hiding the supraglenoid foramen within the subscapular fossa (1). [N.B. The derived state of this character develops with morphogenesis; consequently it is indeterminate in morphogenetically immature specimens, in which the glenoid fossa and coracoid plate are unossified.] STATE 1: Ichthyostega, Terrapoda. 313. m SCACOR 16. Mesocoracoid arch: absent (0) or present (1). [N.B. This character is indeterminate in morphogenetically immature specimens, in which the coracoid plate is unossified.] STATE 1: Captorhinomorpha. 314. ¨ m SCACOR 17. Anterior border of glenoid: posterior to the posterior edge of scapular blade (0) or aligned with the posterior edge of the scapular blade (1). [N.B. This character is indeterminate in morphogenetically immature specimens, in which the glenoid fossa is unossified.] STATE 1: Terrapoda ( except tuditanomorph Microsauria). 315. m SCACOR 18. Enlarged coracoid foramen (at least three times the size of the glenoid foramen): absent (0) or present (1). [N.B. This character is indeterminate in morphogenetically immature specimens, in which the coracoid plate is unossified.] STATE 1: Panderichthys plus Tiktaalik. 316. ¨ m SCACOR 19. Presence (0) or absence (1) of accessory foramina in coracoid. [N.B. This character is indeterminate in morphogenetically immature specimens, in which the coracoid plate is unossified.] STATE 1: archeriamorphs plus Terrapoda. 317. m SCACOR 20. Scapulocoracoid: not fused to cleithrum (0) indistinguishably fused (1). [N.B. the initially separate, long stemmed cleithrum sometimes becomes fused to the scapulocoracoid in very mature specimens of some taxa within the Terrapoda.] STATE 1: ichthyostegamorphs.

Humerus

318. ¨ HUM 1. Latissimus dorsi process: offset anteriorly (0) aligned with ectepicondyle (1). STATE 1: Anthracosauria plus Terrapoda. APPENDIX 14: EARLY TETRAPOD CHARACTER STATE LIST 355

319. ¨ m HUM 2. Distinct supinator process projecting anteriorly: absent (0) or present (1). [N.B. The derived state of this character develops with morphogenesis; consequently it is indeterminate in morphogenetically immature specimens lacking an ossified radial condyle, and ichthyostegamorphs and archeriamorphs in which the anterior humeral keel is unossified.] STATE 1: Terrapoda (except some Temnospondyli: Amphibamidae, Captorhinus, Microsauria). 320. m HUM 3. Ventral humeral ridge: present (0) or absent (1). [N.B. The derived state of this character develops with morphogenesis; consequently it is indeterminate in morphogenetically immature specimens lacking an ossified radial condyle.] STATE 1: The distribution of this characteristic is difficult to determine, within the Terrapoda, torsion of the humerus often obscures the presence of the ventral humeral ridge. Conservatively, the derived state is present in the Captorhinomorpha and Lepospondyli. 321. HUM 5. Entepicondylar foramen: present (0) or absent (1). STATE 1: derived Temnospondyli (Edops, temnospondyls more derived than Dendrerpetontidae); some Microsauria, (Stegotretus, Rhynchonkos plus Gymnarthridae); some Nectridia (Keraterpeton plus Diceratosaurus, Urocordylidae). 322. ¨ HUM 6. Ectepicondylar foramen: present (0) or absent (1). STATE 1: archeriamorphs (except Crassigyrinus) plus Terrapoda. 323. HUM 7. Distinct ectepicondyle: present (0) or absent (1). STATE 1: Microsauria. 324. ¨ m HUM 9. Distal extremity of ectepicondylar ridge: aligned with ulnar condyle (0) aligned between ulnar and radial condyles (1) aligned with radial condyle (2). [N.B. This character is indeterminate in morphogenetically immature specimens lacking an ossified radial condyle.] Ordered. STATE 1: archeriamorphs. STATE 2: Terrapoda. 325. ¨ HUM 10. Humerus not constricted below deltopectoral crest (0) or constricted (waisted) (1). [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Terrapoda. 326. ¨ m HUM 11. Radial condyle: visible in both extensor and flexor views (terminal) (0) visible in flexor view only (ventral) (1). [N.B. This character is indeterminate in morphogenetically immature specimens lacking an ossified radial condyle. This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Anthracosauria plus Terrapoda. 327. HUM 12. Humerus slender and elongate, with length less (0) or more (1) than three times the diameter of its distal end. STATE 1: some Temnospondyli (derived Amphibamidae: Amphibamus plus Doleserpeton), derived Captorhinomorpha (Paleothyris plus Petrolacosaurus). 328. ¨ HUM 17. Presence (0) or absence (1) of accessory foramina in humerus. STATE 1: Anthracosauria plus Terrapoda. 329. ¨ HUM 19. Proximal and distal articulation surfaces twisted at approximately 45 degrees or less (0) set at approximately right angles to each other (1). [N.B. this character is subject to phenotypic variation in some taxa.] STATE 1: Terrapoda. 330. ¨ HUM 20. Latissimus dorsi process of humerus: an elongate ridge connected to deltoid crest (0) knob on extensor surface of humerus (1). [N.B. The latissimus dorsi process may be absent in early morphogenetic stages, due to the low degree of ossification of the proximal humerus.] STATE 1: archeriamorphs plus Terrapoda. 356 K. PAWLEY PHD THESIS

331. HUM 21. Deltopectoral crest of humerus more distal than latissimus dorsi process (0) deltopectoral crest and latissimus dorsi process equidistant from head (1). [N.B. The latissimus dorsi process may be absent in early morphogenetic stages, due to the low degree of ossification of the proximal humerus.] STATE 1: some Temnospondyli (Euskelia). 332. ¨ HUM 22. Anterior humeral keel originates: on the extensor side of the deltopectoral crest (0) distal to deltopectoral crest (1). STATE 1: Terrapoda. 333. ¨ m HUM 23. Humeral process 2 (sensu Jarvik, 1996): absent (0) or present (1). [N.B. The derived state of this character develops with morphogenesis (increasing ossification of the proximal humerus); consequently it may be indeterminate in morphogenetically immature specimens lacking an ossified radial condyle.] STATE 1: basal Tetrapoda (ichthyostegamorphs plus archeriamorphs). 334. ¨ m HUM 24. Latissimus dorsi process of humerus set at right angles to extensor surface of humerus (0) flattened towards the surface of the humerus and proximally directed (1). [N.B. The latissimus dorsi process may be absent in early morphogenetic stages, due to the low degree of ossification of the proximal humerus.] STATE 1: Terrapoda. 335. m HUM 25. Latissimus dorsi process of humerus: large knob on the dorsal surface of humerus (0) small inconspicuous knob or absent (1). [N.B. The latissimus dorsi process may be absent (unossified) in early morphogenetic stages, due to the low degree of ossification of the proximal humerus.] STATE 1: some Temnospondyli (Stereospondyli), Captorhinomorpha. 336. ¨ m HUM 26. Ectepicondyle: does not project anterior to anterodistal corner of humerus (0) or projects anterior to radial condyle (1). [N.B. The derived state of this character develops with morphogenesis, consequently it is indeterminate in morphogenetically immature specimens lacking an ossified radial condyle.] STATE 1: Terrapoda. 337. HUM 27. Attachment are for pectoralis muscle (on posterior surface of deltopectoral crest of humerus) bulbous with a deep central pit: absent (0) present (1). STATE 1: Anthracosauria where observable.

Radius

338. ¨ RAD 1. Radius longer (0) subequal to (1) or shorter (2) than humerus. Ordered. [N.B. as the radius and humerus both increase in length with morphogenesis, it is necessary for the specimens of the humerus and femur to be from the same specimen for this comparison to be valid.] STATE 1: Tiktaalik plus Panderichthys. STATE 2: Tetrapoda. 339. ¨ RAD 2. Radius longer than (0) as long as (1) ulna without olecranon process. [N.B. The olecranon process of the ulna develops with morphogenesis; consequently this character is indeterminate in morphogenetically immature specimens. This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: all Tetrapoda more derived than Tulerpeton. 340. ¨ RAD 3. Dorsomedial radial ridge (sensu Warren and Ptasznik, 2002): present (0) absent (1). [N.B. the radius of Ossinodus pueri described by Warren and Ptasznik (2002) is here reinterpreted as a left radius, following comparison with Acanthostega (Coates, 1996).] STATE 1: Terrapoda. APPENDIX 14: EARLY TETRAPOD CHARACTER STATE LIST 357

341. ¨ RAD 4. Proximoventral radial ridge (sensu Warren and Ptasznik, 2002): present (0) absent (1). [N.B. the radius of Ossinodus pueri described by Warren and Ptasznik (2002) is here reinterpreted as a left radius, following comparison with Acanthostega (Coates, 1996).] STATE 1: Terrapoda.

Ulna

342. ¨ m ULNA 1. Absence (0) or presence (1) of olecranon process, here defined as an extension of the lateral surface of the ulna above the medial edge of the articulation surface. [N.B. The derived state develops with morphogenesis; consequently this character is indeterminate in morphogenetically immature specimens. This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Tetrapoda more derived than Acanthostega. 343. ULNA 2. Posterolateral ulnar crest (sensu Pawley and Warren, 2006) absent (0) present, lateral to posterior ulnar crest (1). STATE 1: some Temnospondyli (Eryopoidomorpha where observable).

Manus

344. ¨ DIG 2. Number of digits in manus: more than five (0) five (1) four (2) three (3). [This character was merged with DIG 3 and DIG 4 from Ruta et al. (2003) because they are not independent.] STATE 1: Tetrapoda more derived than Tulerpeton. STATE 2: Temnospondyli more derived than Casineria, Westlothiana, Lepospondyli. State 3: some Microsauria (Microbrachomorpha).

Digits

345. ¨ DIG 1. Digits: absent (lepidotrichia present) (0) present (1). STATE 1: Tetrapoda. 346. ¨ m DIG 5. Proximal ends of metacarpals/tarsals: symmetrical (0) asymmetrical [N.B. The derived state develops with morphogenesis; therefore this character may be indeterminate in morphogenetically immature specimens.] (1). STATE 1: archeriamorphs more derived than Tulerpeton, plus Terrapoda.

Pelvic girdle

Ilium

347. ILI 1. External iliac shelf: absent (0) present (1). STATE 1: Seymouriamorpha plus Cotylosauria (except Captorhinomorpha). 348. ILI 2. Post-iliac process: projecting distinctly posterior to dorsal iliac process and/or sacral rib facet: present (0) or absent, posterior border of sacral rib facet forms posterior border of dorsal iliac process (1). STATE 1: Greererpeton, Temnospondyli more derived than Dendrerpetontidae, Lepospondyli (except some Microsauria: Tuditanidae, Hapsidopareiontidae). 349. ILI 3. Dorsal iliac process: present (0) absent (not extended above junction with post-iliac process) (1). [N.B. This character is indeterminate for taxa that lack a post-iliac process. [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Temnospondyli, Captorhinomorpha, Westlothiana. 358 K. PAWLEY PHD THESIS

350. ¨ ILI 4. Dorsal iliac ridge (new term) passes across dorsal iliac process (where present) from anterior edge to the dorsal surface of post-iliac process: present (0) or absent (1). [N.B. This character is indeterminate for taxa that lack a post-iliac process.] STATE 1: Anthracosauria plus Terrapoda. 351. ¨ ILI 5. In medial view, mesial iliac ridge (sensu Coates, 1996) offset anteriorly to iliac neck (0) aligned with iliac neck (1). STATE 1: Terrapoda. 352. ¨ ILI 6. Supracetabular iliac buttress less (0) or more (1) prominent than post- acetabular buttress. STATE 1: Tetrapoda more derived than Acanthostega. 353. ¨ ILI 7. Transverse pelvic ridge: absent (0) or present, passes from the border of the ilium anterior to the supracetabular buttress towards the ventral border of the post- iliac process (1). [N.B. This character was separated into three parts in order to clarify it; the two additional characters are ILI 1. and ILI 4.] STATE 1: Anthracosauria plus Temnospondyli plus basal Neospondyli (Seymouriamorpha plus Cotylosauria). 354. ILI 8. Mesial iliac ridge (sensu Coates, 1996): passes down the midline of the ilium, so that there is a recess present both anterior and posterior to the ridge (0) forms the anterior border of the ilium, with no anterior recess (1). [N.B. the mesial ridge is observable in medial view.] STATE 1: derived Temnospondyli (Peltobatrachus plus Eryopoidomorpha). 355. ILI 11. Post-iliac ridge (new term): absent (0) present on medial side of post-iliac process, passes from junction with dorsal iliac process along medial surface of post-iliac process (1). STATE 1: Whatcheeriidae plus Crassigyrinus.

Pubis

356. ¨ m PUB 1. Accessory pelvic foramina (0) absent, obturator foramen only (1). [N.B. This character is indeterminate in morphogenetically immature specimens lacking an ossified pubis.] STATE 1: Terrapoda.

Femur

357. ¨ FEM 5. Femur shorter than (0) subequal to (1) or longer than humerus (2). Ordered. [N.B. as the humerus and femur both increase in length with morphogenesis, it is necessary for the specimens of the humerus and femur to be from the same specimen for this comparison to be valid.] STATE 1: ichthyostegamorphs. STATE 2: archeriamorphs plus Terrapoda. 358. m FEM 6. Adductor crest of femur approximately as deep as femoral shaft (0) reduced to low ridge (1). [N.B. the adductor crest develops with morphogenesis, and may be low and almost absent in morphogenetically immature specimens. The derived state is relevant where it is present in specimens with an internal trochanter distinctly separated from the proximal articulation surface.] STATE 1: some Temnospondyli (Dvinosauria). 359. ¨ FEM 7. Posterior intertrochanteric ridge (sensu Pawley and Warren, 2006) joins the adductor blade proximal to adductor crest (0) joins at the junction of the adductor blade and adductor crest (1). [N.B. The posterior intertrochanteric ridge increases in height with morphogenesis.] STATE 1: Terrapoda. APPENDIX 14: EARLY TETRAPOD CHARACTER STATE LIST 359

360. FEM 8. In ventral view, adductor blade and crest aligned, pass diagonally across flexor surface of femur towards fibular condyle (0) adductor blade set at an angle to adductor crest, which passes down the midline of the flexor surface of the femur (1). STATE 1: derived Temnospondyli (Eryopoidomorpha). 361. ¨ FEM 9. In distal view, intercondylar fossa aligned with the distal end of the adductor crest (0) aligned with the midline of the popliteal area (1). STATE 1: Terrapoda.

Tibia

362. ¨ m TIB 1. Tibial distal articulation surface: L shaped with projection on anterior side of flexor surface (0) ovoid (1). [N.B. The derived state is affected by the degree of ossification of the distal end of the tibia, it is indeterminate in specimens that exhibit a low overall degree of ossification of the postcranial skeleton.] STATE 1: Anthracosauria plus Terrapoda. 363. ¨ TIB 2. Posterior tibial keel (new term): present (0) absent (1). STATE 1: Terrapoda. 364. ¨ TIB 3. Cnemial extensor flange (new term): absent (0) present (1). STATE 1: Ichthyostega plus archeriamorphs. 365. ¨ TIB 4. Junction of anterior and posterior tibial flexor crests with distal tibial flexor crests (sensu Pawley and Warren, 2006) on the anterior side of the flexor surface (0) junction on ventral midline of shaft, distal tibial flexor crest located down midline of the shaft (1). STATE 1: Terrapoda. 366. TIB 7. Posterior tibial flange (new term) absent (0) present, distinct from main body of tibia (1). [This character was re-written from Ruta et al. (2003) in order to clarify it.] STATE 1: Westlothiana, Captorhinomorpha.

Fibula

367. ¨ FIB 1. Fibula not waisted (0) or waisted, interepipodial space present (1). STATE 1: archeriamorphs plus Terrapoda. 368. ¨ FIB 2. Flexor surface of fibula medially bulbous below femoral articulation surface (0) concave (1). STATE 1: Terrapoda. 369. FIB 5. In lateral view, posterior border of fibular shaft between proximal and distal ends: straight / concave (0) convex (1). STATE 1: some Temnospondyli (Euskelia). 370. FIB 6. Mid shaft area of flexor surface of fibula: convex (0) or concave (1). STATE 1: some Temnospondyli (Eryopoidomorpha). 371. FIB 7. Fibular sulcus across anterodistal surface: present (0) absent (1). STATE 1: Anthracosauria, Captorhinomorpha. 360 K. PAWLEY PHD THESIS

Pes

372. ¨ m TAR 3. Ossified proximal edge of separate tarsal intermedium: concave / convex (0) asymmetrical notch, with the fibulare side longest (1). STATE 1: [N.B. This character does not pertain to the absence or presence of an astragalus (TAR 6). Taxa that possess an astragalus can be coded, as the intermedium forms the proximal edge of the astragalus (Berman and Henrici, 2003; O'Keefe et al., 2006). This character was re- written from Ruta et al. (2003) in order to clarify it.] STATE 1: archeriamorphs, Caerorhachis, Cotylosauria plus Westlothiana. 373. ¨ m TAR 4. Distal tarsals between fibulare and digits: absent (0) or present (1). [N.B. This character is indeterminate in morphogenetically immature specimens lacking ossified tarsals.] STATE 1: archeriamorphs plus Terrapoda. 374. ¨ m TAR 5. Distal tarsals between tibiale and digits: absent (0) or present (1). [N.B. This character is indeterminate in morphogenetically immature specimens lacking ossified tarsals.] STATE 1: archeriamorphs plus Terrapoda. 375. m TAR 6. Astragalus, composed of fused tibiale, intermedium and proximal centrale (sensu Kissel et al., 2002; Berman and Henrici, 2003): absent (0) or present (1). [N.B. The astragalus develops with morphogenesis (Kissel et al., 2002; Berman and Henrici, 2003).] STATE 1: Neospondyli more derived than Seymouriamorpha.

Dermal ossifications

376. m DERM 1. Dermal osteoderms: absent (0) present (1). [N.B. Dermal ossifications develop with morphogenesis, they are absent due to immaturity in specimens which exhibit a low overall degree of ossification of the postcranial skeleton.] STATE 1: some Temnospondyli (Peltobatrachus, Dissorophidae). 377. m DERM 2. Cycloid dorsal scutes: overlapping (0) distinctly separate (1). [N.B. Dorsal scales develop with morphogenesis, they may be absent in morphogenetically immature specimens.] STATE 1: some Temnospondyli (Stereospondylomorpha).

Pawley, K. 2006. 361 The Postcranial Skeleton of Temnospondyls (Tetrapoda: Temnospondyli) PhD Thesis. La Trobe University, Melbourne

APPENDIX 15. DATA MATRIX: WALKING WITH EARLY TETRAPODS

Inapplicable states (-) are not recognised by PAUP 4.0b10 [which treats them as (?)], but they are included here to distinguish from data that is actually missing (?).Note that many of the characters from Ruta et al. (2003) have been recoded.

Acanthostega gunnari 1 0000000000 0000010-00 0000100000 0010010010 0000000000 51 001- -00000 0000000000 0000000000 0000000000 0000000000 101 0000000000 0000000000 000100000- 0010000000 0000000000 151 10- -0-0??1 0000-000- - 0000101101 0010001000 0100010000 201 1000100010 0000000000 0001000000 0000000000 0000000010 251 0001000000 0000100001 0100000000 00001000- - 0- -0-01010 301 00-0000000 0- -0001000 0000000000 001- -00100 0000100000 351 0000001000 0000000000 0?0000-

Acheloma cumminsi 1 001- -11101 0000111101 000?111001 0000011000 0001000000 51 101- -00010 00-0000010 0000001000 1100000000 001?110001 101 0001110110 0011100101 100010?00- 0100100000 0000?00011 151 -1110?2002 0000101100 0101- - -1?0 00?????0?0 0100101?0? 201 ?????????? ??10000000 0001?00000 0000000000 0010000?10 251 0001?01010 1?00100?10 -???-?0??? ???1-01111 0?????1010 301 1011100111 1101010?10 110211011- ?10??10211 11??1101- - 351 ?111-12011 1110?01??? ?0110??

Acherontiscus caledoniae 1 000??0???? ????11??0? 00001?0101 ?000010110 ?????????0 51 -????1- - -0 0??000??00 001?1000?0 00001?0000 00???1??0? 101 ?????????? ?????????? 0????????? ?????????? ?????????? 151 ?????????? ?????????? 0????????0 0??????0?1 ?????????? 201 ?????????? ???0000000 000??0??00 11010?01?? 0????????0 251 000??????? 1??0?????0 -?0?-?0??? ?????????? ?????????? 301 ?????????? ?????????? ?????????? ?????????? ?????????? 351 ?????????? ?????????? ???????

362 K. PAWLEY PHD THESIS

Acroplous vorax 1 000?-01000 000011??00 0000100001 0000010000 0001000000 51 101- -00010 0??0100000 0000000000 0000110000 000101000? 101 0100111111 0110000011 100010000- 0000110002 00?100?000 151 1111002012 0000000100 1??????100 00??011010 01??????00 201 1?10111011 1110000000 0001?00000 00000?00?? ?????????0 251 ?????????? ?00010???0 -100-0?0?? ?????0?110 000?01???? 301 ?????????? ?????????0 1???1?0??1 ??0??????? ?????????? 351 ?????????? ?????????? ???????

Adamanterpeton ohioensis 1 011- -01010 0000111000 0000100000 0010010000 000?0????0 51 0001000010 0??000?000 0000001000 0000110000 00??????0? 101 0001011110 0010110101 000010??0- 0100100000 000??1?101 151 -111??0102 00???????? 0?0????1?0 0110011010 ???01??000 201 1110111011 ??1000000? ?????????? ?????????? ?????????? 251 ?????????? ?????????? ?????????? ?????????? ?????????? 301 ?????????? ?????????? ?????????? ?????????? ?????????? 351 ?????????? ?????????? ???????

Adelogyrinus simorhynchus 1 0?0????00? ?0?011??00 10001??00? 10?0010?1? 000?000000 51 -?1- -1- - -1 ------001?0001?0 0000??0000 00??01??0? 101 ?????????? ?????????? 0????????? ?????????? ?????????? 151 ?????????? ???0000101 1?0?????10 01??0010?1 010?????0? 201 ?????????? ???0001100 ?001??0?10 1101001- -0 ?????-???? 251 ???????10? 01110?00?? 0-?00-?0?0 ????1-0?10 00?0??11?? 301 ?????????? ?????????? ?????????? ?????????? ?????????? 351 ?????????? ?????????? ?????0?

Adelospondylus watsoni 1 0?0??????? ?????????? ?0001?000? 0000010??? 000?000000 51 -?1- -1- - -1 ------1- - - -0?100 0000100000 00??010?0? 101 ?????????? ???????10? 000110??0- 00?0000?0? 000?10???? 151 ?100??2?02 ?????????? 100?????10 0???0??0?1 0100????0? 201 ?????????? ???0001100 0001100?10 1101001- -? 0????-???? 251 ??????10?? 1110?????0 -???-?0?01 ?????????? ?????????? 301 ?????????? ?????????? ?????????? ?????????? ?????????? 351 ?????????? ?????????? ???????

Albanerpetontidae 1 001- -01100 00?111??00 1101100101 0000001- -0 11------01 51 - -1- -1- - -1 ------1- - - -100?0 00001?0000 ??0?01??0? 101 ?????????? ?????????? ?????????? ?????????? ?????????? 151 ?????????? ???0101100 0001- - -110 0- - -1- -01- 1- -0-??-0 201 - - -1- - -1- - - -10100000 0011110?10 1100011- -1 1??11-1110 251 00010?1?11 0001110??- - -1- - -1------1------1------??? 301 ?????????? ????????01 111-11111? ??0??-?211 11?21??1-? 351 ???0-?2??? ??1??01??? ??110??

APPENDIX 15: EARLY TETRAPOD DATA MATRIX 363

Amphibamus grandiceps 1 001- -01100 00?011??00 1001111000 0000010000 000?000000 51 101- -00010 ???0000000 0000001000 0000000000 001011??01 101 0011110111 0010-01111 ?00010?00- 0100101000 100?000001 151 -111000002 1100101100 0?0??????0 0???0??0?? 010?10??00 201 ???0???0?? ??11110000 00011?0000 1?000000?? ?01?????10 251 00011010?0 ??0??00??0 -?00-0000? ??01-0110? 00????1010 301 10?1?????? ???1010??? 11??1?111? ???????21? ???21?01- - 351 ?1??-?2??? ??10?01??? ?????00

Anthracosaurus russelli 1 001- -01000 000011??01 0000100000 0000010100 0000000000 51 0000000000 10-0000001 0000000000 0000??0010 000?01210? 101 1010?10110 0?10?00001 000000000- 00?0000000 0000?0?001 151 -1001??002 00?0000100 ???????1?0 001?01101? 010??110?0 201 1110111011 0??000000? ?????????? ?????????? ?????????? 251 ?????????? ?????????? ?????????? ?????????? ?????????? 301 ?????????? ?????????? ?????????? ?????????? ?????????? 351 ?????????? ?????????? ???????

Apateon pedestris 1 001- -0100? 000011??0? 1001100000 0000011000 000?000000 51 101- -00010 0?-00??000 0000001000 0000000000 000??10?0? 101 0000110111 0000000010 000?00??0- 0000111000 000??0?0?1 151 -111002002 10???????? ?????????0 0???0??0?? 010??????? 201 ?????????? ??1?000000 00?1?????0 ??00??0??? ?????????0 251 00011????0 ??0??????0 -??????1?? ???1-011?? ?????????? 301 ?????????? ?????????? ????1??1?? ???????21? ???21????? 351 ??????2??? ?????01??? ???????

Archegosaurus decheni 1 101- -01000 100011??10 0000100000 0010010000 000?000000 51 101- -00010 10-000?000 0000001001 0000??0000 000?01??01 101 0001111110 0010?00100 000100??0- 0100100000 000?00?0?1 151 -111000102 0100100100 0101- - -100 0010011010 0101101000 201 1110111011 1110000000 0001100000 00000000?? ????????10 251 0001?010?0 1?00100??0 -100-00001 ?001-01110 00?1011??? 301 ?????0???? ??????0??? 1???1?0??? ???????211 1???1?01- - 351 1??1-?2??? ??????1?0? ?????01

Archeria crassidisca 1 000?001000 0000110-00 0000100000 0000010100 0000000000 51 0000000000 10-0000000 0000000000 0000110000 000?01210? 101 ????0????? 0??00???0? 0?0?00000- 0010000000 000010??0? 151 ?100102002 0000000100 0000011100 0011011010 0100111000 201 1110111011 1110001000 0001000000 1100000110 0000000010 251 0001101000 11000000?1 0100000000 1011-00100 000011100? 301 1000001000 0000010100 0101010101 0010001110 0101110001 351 0110002000 0101001000 11110??

364 K. PAWLEY PHD THESIS

Asaphestera intermedia 1 001- -01000 000011??00 0000100000 000?010100 0001100000 51 -11- -1- - -0 000000-000 0001100000 0000100000 000??1000? 101 ????0?0??0 ???0-?1100 000100000- 0110000000 000010???? 151 ?100?????? ???0010100 ????????10 0??????0?? 010?????0? 201 ?????????? ???0000000 0011110?10 1100001- -? ?????-???? 251 ?????????? 1000000??1 ?000?001?? ?????????? ???????110 301 1-0-100111 ???0010??? 011211011? ??0??1???? ????1??00? 351 1??00????? ?????????? ?????00

Balanerpeton woodi 1 001- -01100 000011??00 0000100000 0000010000 000?000000 51 0001000010 0?-0000000 0000001000 0000000000 000?010?01 101 0001110110 0010100101 000010??0- 0100100000 000?110001 151 -111000002 0100000100 0101- - -100 0110011010 010?101?00 201 1110???011 1?10000000 0001?00000 00000000?0 ?????00010 251 0001?0???0 1000100??0 -100-0000? ???1-011?0 0?????1??? 301 ?????????? ??????0??? 01??1?01?? ??0????21? ?1?21??01- 351 11?00?20?? ???01011?? ?01100?

Baphetes kirkbyi 1 000?000000 0000111000 0000100010 0000110000 0000000000 51 0001001010 01-0000000 0000000000 0000000010 000?01010? 101 0001010110 0010000101 000010??0- 0010000000 000?00?001 151 -0- -100002 0000000?00 ???0111100 001?01101? ?????????0 201 101??0???? ???0000000 00???00??0 ??000000?? ?????????0 251 000??0???? ?1?0?????? ??????0000 00????01?? 00?0?11??? 301 ?????????? ???????0?0 010?0?0001 00?00001?0 0?????000? 351 ?10??????? ?001001?00 0??????

Batrachiderpeton reticulatum 1 000??01000 0000110-00 1000100000 0000010111 0001000000 51 -01- -1- - -0 001100?000 0000000010 0000100000 00??010?0? 101 001000?110 0010-010?0 001- - -000- 1010000001 -00?-0?001 151 -0- -0??0?2 00?1000100 1001- - -110 00- -1- -011 0110101001 201 - - -1- - -1- - - -?0000001 1011?01010 1100001- -? ?????-???1 251 00011110?? ?????????0 -???-????? ?????1?1?? ?????????? 301 ?????????? ?????????? ?????????? ?????????? ???21????? 351 ?????????? ?????????? ???????

Batropetes fritschia 1 000??01000 010011??00 0000100001 000?010110 01------0 51 -01- -1- - -0 000000-000 0001100000 0010100000 0?0?010002 101 ?????????? ???0-0???? ?0????000- 00???0???0 ?00?10??0? 151 -???000002 0000010100 100????110 00??1- -0?? 010?????0? 201 ?????????? ??10100000 0011?10110 1100001- -1 1??11-0?1? 251 ???1??1000 1000000??1 10000?0101 ??01-0?110 01??011110 301 1-0-100111 ???0010?01 01??11011? ?10???0211 11?21101-1 351 110?-120?0 ?11??0110? ?011???

APPENDIX 15: EARLY TETRAPOD DATA MATRIX 365

Benthosuchus sushkini 1 000110100? 000011??1? 0000100000 0010010000 0001000000 51 101- -00010 0010011000 0000001002 0000110000 0001010001 101 0000101111 0001000010 010100110- 0000200000 1100001010 151 0111011202 00102001- - 1101- - -100 0010011010 0100101100 201 1010101011 1010000000 0001100000 ??00?000?? ?????????0 251 ?????????? 1100100110 -100-10111 1001-01110 01110110?? 301 11?110???? 1?????0110 110?110111 01011?0211 111?1?01- - 351 1111-?2011 1110101101 0??????

Brachydectes spp. 1 001- -01000 0000110-00 1000100001 00010101-0 1001010001 51 - -1- -1- - -0 0000000000 1- - - -00000 0011------1-0?010002 101 1010010110 0000-01001 001- - -000- 000000000? -0?0-0?001 151 -0- -002002 0000010100 0001- - -110 01- -1- -011 01??????01 201 - - -1- - -1- - - -?0000000 0011?1??10 1101001- -? 10?11-?010 251 0001?01011 1010000??0 -?0?-?0?0? ???1-011?? 0????11??? 301 ?????????? ?????????? ?1??1?011? ???????21? ????1????? 351 ??????2??? ?????01??? ?????0-

Broiliellus brevis 1 001- -01??0 ?0?011??00 000?111000 0000011000 000?000000 51 101- -00010 00-000?000 0000001000 1100000000 001?11??01 101 000111011? 001???01?1 1?0?10?00- 0100100000 ?00?00?011 151 -111002002 00???????? 0?0??????0 0??????0?0 ?????????? 201 ?????????? ??10000000 0001?00000 ??000?00?? ????????10 251 000??????? ??00?0011? ?????????? ???1-01111 0011011??? 301 1?1??????? ?????????? 11??1?011? ?1???????? ?????????? 351 11?1?1?01? ?????????? ?????1?

Bruktererpeton fiebigi 1 00???0100? ???011??0? ?00010?0?0 00???11100 000?00000? 51 0?00?00000 0?-00??000 ???????00? ?????????? ?????????? 101 0001?1?110 0?10000101 ??0??0???? ?????????? ?00?1???0? 151 ?????????? ?????????? ?????????? ?????????? ?????????? 201 ?????????? ???0000000 0001?0??00 10000000?? ?????????0 251 000110?0?0 1100000??? ?10??0???? ???????1?? ???????0?? 301 ?0???0???? 00????0100 010?0?01?1 ???????21? ?1??11???? 351 ??????20?0 ??????1?0? ?1??00?

Caerorhachis bairdi 1 001- -?1??? ???0?????0 ?0001????? ?????1?000 000?000000 51 0001?00010 0?-000?000 000000?000 ??0??0???? 000??1??0? 101 000101?110 00101?0101 ?00010??0- 01?0000000 000?1??00? 151 ?1000?0002 0000000??? 0?0????100 0?1001?010 0100100000 201 1110111011 1110000000 0001100000 10000000?0 0?1????0?0 251 000110?0?0 1000?00??? ?????????? ?????????? ?????????? 301 ?????????? ?????????? ?????????? ?????????? ?1??11001- 351 ?110?1???? ??1?101??? 01110??

366 K. PAWLEY PHD THESIS

Capetus palustris 1 001- -01100 0000111000 0000100000 0000010000 000?000000 51 0001000010 0?-00??000 0000001000 0000000000 000?01??0? 101 000101?110 0?1010010? 0???????0- 0100100000 000??1??01 151 -1110?0?02 ??00000100 0?0????1?? ????01?0?0 010??????? 201 ???????0?? ?11000000? ?????????? ?????????? ?????????? 251 ?????????? ?????????0 -100000??? ?????????? ?????????? 301 ?????????? ?????????? ?????????? ?????????? ?????????? 351 ?????????? ?????????? ???????

Captorhinus aguti 1 001- -01000 0001111101 0000100000 0000011-11 0000100000 51 0?1- -00001 - - - - -00000 0000000000 0000110000 000?011102 101 1011010110 0010-01101 001- - -0011 0110000000 -000100001 151 -1001?0002 0000000100 0001- - -010 00- -1- -011 0100101001 201 - - -1- - -010 1110000000 0111110110 11000000?0 0010010020 251 0001101000 1000000?11 1100000101 0001-????? ???????011 301 1011100111 1111010101 0102110111 010?110211 110111001- 351 1110012010 1110111100 11111??

Cardiocephalus peabodyi 1 001- -01000 010?111000 0000100000 0001010100 0001010000 51 -11- -1- - -0 000000-000 0001110000 0000100000 010?010002 101 ?011010110 0000-01?0? 0001?0000- 0110000010 000010?001 151 -1000?0002 0000010100 0001- - -110 0?-1011011 0110101001 201 - - -01111- - - -10000010 0011?10?10 11000000?1 10011?0020 251 ???1??1000 1000000??? ??????0?01 ?00??????? ?????????? 301 ?????????? ????????0? 11??11011? ??0????21? ?1??11???? 351 ??????2010 1110?01?0? 0?11???

Casineria kiddi 1 ?????????? ?????????? ?????????? ?????????? ?????????? 51 ?????????? ?????????? ?????????? ?????????? ?????????? 101 ?????????? ?????????? ?????????? ?????????? ?????????? 151 ?????????? ?????????? ?????????? ?????????? ?????????? 201 ?????????? ?????????0 0001?00000 100?0000?? ????????1? 251 ?????????0 1000100??? ??????0?0? ???1-0?110 001??1??1? 301 ?????????? ?????????? 01??1?0?1? ??0????11? ?1?1??001- 351 ?1????2??? ?????????? ???????

Chenoprosopus lewisi 1 011- -01010 ?000111011 0000100000 0010010000 0001000000 51 0001000010 00-0000000 0000001000 0000110000 000101000? 101 000101?110 0?10100001 000010000- 0100100000 0000?1?111 151 -1110???02 0??0000100 010??????0 00??0?101? 010??????0 201 ???0???0?? ???0000000 0001????00 ??00?000?? ?????????? 251 ?????????? ?????????? ?10??00?0? ??0??????? ?????????? 301 ?????????? ?????????? ?????????? ?????????? ??????01- - 351 11?0-???1? ?????????? ???????

APPENDIX 15: EARLY TETRAPOD DATA MATRIX 367

Chenoprosopus milleri 1 011- -01010 0000111010 0000100000 0010010000 1001000000 51 0001000010 00-0000000 0000001000 0000110000 0001010?0? 101 0001010110 0?10100101 000010?00- 0100100000 000011?111 151 -111000102 0000?00100 010??????? ?0?????010 010?101?0? 201 ?1???????? ???000000? ?????????? ?????????? ?????????? 251 ?????????? ?????????? ?????????? ?????????? ?????????? 301 ?????????? ?????????? ?????????? ?????????? ?????????? 351 ?????????? ?????????? ???????

Cochleosaurus bohemicus 1 011- -01010 0000111000 0000100000 0010010000 100?000010 51 0001000010 00-0000000 0000001000 0000110000 000001000? 101 0001010110 0010110101 000010?00- 0100100000 000?01?111 151 -111000102 0100000100 0?0????1?0 0110011010 0100101000 201 1110111011 111000000? ?????????? ?????????? ?????????? 251 ?????????? ?????????? ?????????? ?????????? ?????????? 301 ?????????? ?????????? ?????????? ?????????? ?????????? 351 ?????????? ?????????? ???????

Cochleosaurus floriensis 1 011- -01010 0000111000 0000100000 0010010000 100?000010 51 0001000010 00-000?000 0000001000 0000110000 0000010?0? 101 0001011110 0010110101 000010?00- 0100100000 00001??111 151 -1110?0102 010??????? 0?01- - -100 0110011010 0100101000 201 1110111011 111000000? ?????????? ?????????? ?????????? 251 ?????????? ?????????? ?????????? ?????????? ?????????? 301 ?????????? ?????????? ?????????? ?????????? ?????????? 351 ?????????? ?????????? ???????

Colosteus scutellus 1 0000001001 00?0110-00 0000100101 1000010010 000?000000 51 001- -00000 0?0000?000 0010000000 0000??0000 000??1??0? 101 00000?0110 0010?00000 0?0000???? 0010000100 000?00?000 151 1100????02 ?????????? 000????100 10?0?????? ???0101?00 201 111????011 10?0000000 0001?0??00 00000?00?? ?????????0 251 ?????????? 1100100??0 -100-001?? ?????????? ?????????? 301 ?????????? ?????????? ?????????? ???????1?? ????1????? 351 ?????????? ?????????? ?????0?

Crassigyrinus scoticus 1 00???01000 0010111000 0000100001 0000010000 0000000000 51 0000?00000 10-0000000 0000000000 0000000010 000?01??0? 101 0000001000 0000000000 000000??0- 00??000000 0?00000000 151 1???1??001 000????0- - 00001?1100 00000?1010 01000?0000 201 111011101? ??00000000 00010000?0 ??00?000?0 ?????????0 251 000??????? 1?00100001 0100000000 000??0?100 00??111??? 301 ?????????? ???????0?0 000?0?000? 00???0011? ?1??1?0000 351 ?10?1?2?00 0?0??01000 0??????

368 K. PAWLEY PHD THESIS

Dendrerpeton acadianum 1 001- -01100 0000111000 0000100000 0000010000 000?000000 51 0001000010 0?-0000000 0000001000 000000?00? 0001010001 101 0001110110 0?10100101 000010??0- 0100100000 0000?10001 151 -111000002 01?0000100 010??????0 01??011010 010?1???00 201 ???0???0?? ??10000000 0001?00000 00000000?? ?????????0 251 000??????0 1000100??0 -100-0000? ??01-01110 000101101? 301 1011?00??? ???1010??? 010211011? ?10???0211 11021?001- 351 1110?1?010 1110?01?00 ?011?00

Diadectes absitus 1 001- -01000 0001111100 0000100000 0000010110 0010100000 51 001- -00000 00-0000000 0000001000 0000100000 0001011102 101 101?010110 0010-01?01 0001100010 0110000000 000010?001 151 -10-1??002 0000000100 0001- - -110 00- -1- -011 010?101001 201 - - -1- - -01? 1110000000 0111?10010 11000000?0 0010010020 251 0001101000 1000000??1 1?????0101 ?001-01110 011101101? 301 1011100111 ???1010?10 010211011? 01???1?211 11?1111001 351 ?110?12010 ??10?01100 ??1?10-

Diadectes spp. (Nth American) 1 001- -01000 0001111100 0000100000 0000010110 0010100000 51 001- -00000 00-0000000 0000001000 0000100000 0001011102 101 101?010110 0010-01?01 0001100010 0110000000 00?0100001 151 -10-1??002 0000000100 0001- - -110 00- -1- -01? 0100101001 201 - - -1- - -01? 1110000000 0111110010 11000000?0 0010010020 251 0001101000 1000000??1 1100000101 ?001-01110 0111011010 301 1011100111 1101010110 0102110111 0101010211 1101111001 351 1110012010 1110101100 011110-

Diceratosaurus brevirostris 1 0001?0?00? ?0??110-0? 0000100001 0000010111 000?0000?0 51 -01- -1- - -0 0?110??000 0000000010 ??001??000 00???1??0? 101 ?????????? ?????????? 0???????0- 10?020000? ??1??????? 151 ?111??2??? ?????????? ????????10 0????????? ?????????? 201 ?????????? ???0000001 1011?00?10 1100101- -? ?????????1 251 01111?10?? 1000?????0 -100-00100 ?001-1?111 01??011??? 301 ?????????? ??????0??? 11????01?? ????????1? ?1?21?01-1 351 ????-?2??? ?????01??? ???????

Diplocaulus magnicornis 1 0001?01000 0000110- -0 0000000000 1010001111 0001000100 51 -01- -1- - -0 0011000000 0010000010 0000100000 000?01??0? 101 0010100111 0010-01010 00????000- 1000200001 1010-0?001 151 -1110??0?2 0011000100 1001- - -110 00- -1- -011 0110101001 201 - - -1- - -1- - - -10000001 1011100011 1110101- -1 10111-0111 251 0111111010 1000?00??1 01001?01?? 1??1-1?1?? ?????????? 301 ?????????? ??????0??? 010?1?01?? ?1?????21? ???21????? 351 ??????20?0 ?????01??? ???????

APPENDIX 15: EARLY TETRAPOD DATA MATRIX 369

Diploceraspis burkei 1 0001001000 0000110- -0 0000000001 101- -01111 0001000100 51 -01- -1- - -0 0011000000 0010000010 0000100000 000101000? 101 0010100111 0010-01010 00????000- 1000200001 1010-0?001 151 -1110??0?2 0011000100 1001- - -110 00- -1- -011 0110101001 201 - - -1- - -1- - - -10000001 1011100011 1110101- -1 10111-01?1 251 011111101? ?????0???1 010010010? 1001-????? ?????????? 301 ?????????? ?????????? ?????????? ?????????? ???21????? 351 ?????????? ?????????? ???????

Discosauriscus austriacus 1 0001001000 0000111100 0000100000 0000010100 000?000000 51 0001100000 00-0000000 0000001000 0000100000 000?010?0? 101 1001010110 0010000101 0001100010 0110000000 000010?001 151 -0- -0?0002 0000000?00 0001- - -110 0011011011 0100101000 201 1110111011 1110000000 01?1?10010 1100000000 0010000010 251 0001101010 1000000?11 1100000101 0000101100 01?10110?? 301 10?110???? 110???0110 01021?01?1 010????211 1?011?1001 351 ??1???20?0 11?0?0110? ??????0

Dissorophus spp. 1 001- -01100 ?0?011??00 0000111000 0000011000 000?000000 51 101- -00010 00-0000010 0000001000 11000?0000 00??110001 101 ?0???????? ?????????? 1?0?10000- 01?0100000 ?000??0??? 151 ?111002002 000010?000 0?0??????? 0??????010 010?101?0? 201 ???????011 1110000000 0001100000 00000000?0 1010000?10 251 ???1???0?0 1100100110 -?01-0000? ?101-0?111 0011011010 301 ?011100111 1101010?00 110211011- ?10??10211 11??1?01- - 351 11?1-12011 111010111? 0????1?

Doleserpeton annectans 1 001- -01100 000011??00 0001111000 0000011000 000?000000 51 101- -00010 00-0000000 0000001000 0000000000 001?11??01 101 0011110111 1010-01?11 101- - -??0- 0101101000 100?00?001 151 -111000002 1100101100 ???1- - -110 0???0??0?0 010??????0 201 ???0???0?? ???1110000 0011110?10 1?000000?0 ?0?00?0??0 251 00011???1? ?001?????? ?????????? ???1-??1?? ???????010 301 1011100111 1101010?00 110211111? ?10??10211 11?21?01- - 351 11?1-1201? 111010111? 0????0?

Dolichopareias disjectus 1 000??01000 ?0?0110-00 ?0001??001 1000010?1? 000?0?00?0 51 -01- -1- - -1 ------001?00?1?? ??00???00? ?????1??0? 101 ?????????? ?????????? ?????????? ?????????? ?????????? 151 ?????????? ?????????? ?????????? ?????????? ?????????? 201 ?????????? ???000110? ?????????? ?????????? ?????????? 251 ?????????? ?????????? ?????????? ?????????? ?????????? 301 ?????????? ?????????? ?????????? ?????????? ?????????? 351 ?????????? ?????????? ???????

370 K. PAWLEY PHD THESIS

Dvinosaurus spp. 1 0001001000 1000111000 0000100000 0000010010 0001000000 51 001- -00010 0010100000 0000000000 0000010000 000101000? 101 0100101110 0000000001 000000000- 0000110002 001100?000 151 1111002002 0000000100 1111- - -100 0010011010 010?101000 201 1110111011 1110000000 0001100000 00000000?0 00100000?0 251 000??0??1? 1000?00??1 1100000001 ?001-01110 00010110?? 301 1011?1???? 11????0110 1102110111 0101010211 1102?101- - 351 1?10-?2110 1110?0110? 0??????

Ecolsonia cutlerensis 1 001- -11100 ?0?0111100 0000111001 0000011000 000?000000 51 101- -00010 00-0000010 0000001000 1100000000 001?110001 101 000111?110 0?11100101 100010??0- 0100100000 000?00?01? 151 ?111000002 0000101100 0?01- - -1?0 001001?010 0100101100 201 1110111011 1?10000000 0001100000 00000000?? ????????10 251 000110?0?? 1100?0???0 -101-00001 0101-0111? 00??01101? 301 10111001?1 ???????110 110211011- 110?010??? ?11?1101- - 351 11?1-12011 1110?01?11 ?0??01?

Edops craigi 1 001- -0?010 000?111000 0000100000 0010010000 0001000000 51 0001000010 01-0000000 0000001000 0000000000 0001010001 101 0001010110 0010110101 000010000- 0110000000 00001?0001 151 -100000002 0000000100 0?01- - -100 001?011010 0100101?00 201 1110111011 1110000000 0001???000 00000000?? ?????????? 251 0????0???? ?????0???? ?????????? ?????????? ?????????? 301 ?????????? ????????10 110211011? ?10??10??? ?????????? 351 ??1????0?0 1?????11?? 0??????

Eocaecilia micropoda 1 001- -01100 0001111?00 10011??00? 0000010-?1 11------00 51 -01- -1- - -1 ------1- - - -00000 00001?0000 00???1??01 101 0010111111 0110-01010 001- - -??0- 0100011000 -00?-0?001 151 -1110?2002 00-0101100 1001- - -110 0- - -1- -1- - 1- -1-??-01 201 - - -1- - -1- - - -?1110000 00-1?- - -10 11000?00?1 10?1??11?0 251 00011?1?11 0001000??- - -1- - -1------1- - -1- - 1------??? 301 1?1??????1 ??????0??? 11??1?11?? ???????21? ????1????? 351 ??????1??? ?????01??? ???????

Eoherpeton watsoni 1 001- -0?000 ?000110-00 0000100000 0000010100 0000000000 51 0000000000 10-0000000 0000000000 0000??0000 000?01210? 101 1????????? ??????0001 000000000- 00??000000 0000?0???? 151 ????1??002 00???????? 000????1?0 0?11011010 0100111000 201 11?011?011 0010000000 0001000?00 10000000?0 ??0??????0 251 0001?????? 1?00000??? ??????0?0? ?????????? ???????01? 301 10??001000 ?0?0010100 0101010101 0010001??? ??????0001 351 01100??000 01??001?00 1??????

APPENDIX 15: EARLY TETRAPOD DATA MATRIX 371

Eoscopus lockardi 1 001- -01100 000011??00 1000111000 0000011000 000?000000 51 101- -00010 00-0???000 0000001000 0100000000 001?11??01 101 0?01110111 0?1?100?11 100010??0- 010010000? 000?000001 151 -111?00?0? 11???????? 0?01- - -??0 00??01101? ????101??0 201 ?1?0?1?0?1 ???0000000 0001?00000 0?000000?0 ?01??00010 251 0001101010 1000100110 -?0?-?0001 ??0????1?? ????????1? 301 1?1??????1 ?1???????0 ????1?0?1? ??0????21? ?1?21101- - 351 11?101?011 ?110?0111? 0011000

Eryops megacephalus 1 001- -01100 0000111100 0000100000 0010010000 0001000000 51 101- -00010 00-0000000 0000001000 0000000000 0000010001 101 0001110110 0011100101 000010000- 0100100000 0000000011 151 -111000102 0000100100 0101- - -100 0110011010 0100101100 201 1110111011 1110000000 0001100000 0000000000 0010000010 251 0001101010 1100100110 -101-00001 0101-01110 0011011010 301 1011100111 1101010110 1102110111 1101010211 11121101- - 351 1111-12011 1110101111 0011000

Eryosuchus pronus 1 0001101000 0000111010 0000100000 0010011000 0001000000 51 101- -00010 0010011100 0000001002 0000110000 0001010001 101 0000101111 0001000010 010100110- 0000200000 1100001010 151 0111012202 00102001- - 1101- - -100 0010011010 0100101110 201 1010101011 1010000000 0001100000 00000000?? ????????10 251 ?????????? 1100100110 -100-10111 1001-01110 01110110?0 301 11?110???? 1?????01?0 110?1?0111 010?1?0211 11??1?01- - 351 1111-?2011 1110101101 0??????

Eucritta melanolimnetes 1 00???0100? ?0?011??0? 0000100000 0000110000 000?000000 51 0001001010 0?-00??000 0000000000 0000??0010 000??1??0? 101 ??010?0110 ?0?0??0101 0?0000??0- 0?1?00000? 0?0??????? 151 ?0- -1??002 00???????? 0?0??????0 0????????? ????????0? 201 ?????????? ???000000? ?????????? ?????????? ?????????? 251 ?????????0 ??000????1 0100000000 0?01-0?100 00???????? 301 ?????????? ?????????? 0?0?0?0??? ?????0?21? ?1??11?00? 351 ??????2??? ?????01??? ??????-

Euryodus primus 1 001- -01000 ?100111001 0000100000 0001010100 0001010000 51 -11- -1- - -0 000000-000 000110?000 0000100000 010?010002 101 1011010110 0000-01100 000100000- 0110000010 -00010??01 151 -1000??002 0000010100 000????110 01??011011 0110101?0? 201 ???????1- - - -10000010 0011110110 11000000?1 10?11?1?20 251 ???????000 1000000??? ?????????? ?????????? ???????11? 301 1-0??????? ???????101 111-110111 01010-0211 1??????1-? 351 ?????1???? ?????????? ?????0-

372 K. PAWLEY PHD THESIS

Eusthenopteron foordi 1 0000000000 0000000-00 0000000000 0000000000 0000000000 51 0000000000 0000000000 0000000000 0000001000 0000000000 101 0000000000 0000000000 000000000- 000000000? 000000?000 151 00- -000000 0000-000- - 0000000000 0-00000000 0000000000 201 0000000000 0000000000 0000000000 0000000000 0000000000 251 00000000?? 0000000??0 -000-00000 00000000- - 0- -0-00000 301 00-0------0000 0000000000 ??0- -0000? ?00-0-???? 351 0000??0??? ?????00?0? ??- -000

Gephyrostegus bohemicus 1 000??01000 000011??00 0000100000 0000011100 0000000000 51 0000?00000 0?-000?000 0000000000 00000?0000 000?01??0? 101 000101011? 00100?0101 0000?0000- 0010000000 000010?001 151 -100????02 0?00000?00 000????100 0011011010 0100111000 201 0110011011 1110000000 0001?00000 1000000010 0000000010 251 000?????00 1100000001 0100000000 ??01-0?100 ?????110?? 301 100?00?000 000001010? 010?010101 0?1?00?11? ??01110001 351 0110??2000 ?????0100? ?11100-

Greererpeton burkemorani 1 0000001001 0010110-00 0000100101 1000010010 0000000000 51 001- -00000 0100000000 0000000000 0000000000 000?010100 101 0000010110 0010100000 000000000- 0010000100 000000?000 151 1100100002 0000000100 0000111100 1010001010 0100101000 201 011????011 1110000000 0001000000 0000000010 0000000010 251 0001?01000 1100100000 -100-00100 0001-00100 0000?11110 301 1-0?00?000 ?0000100?0 010?0?0001 0010000110 01011101- - 351 0100-02000 0001001000 0?11000

Hapsidopareion lepton 1 001- -01000 ?000111000 0000100000 0001010100 0001100000 51 -11- -1- - -0 000000-000 000?100000 0000101001 000?010002 101 1011010110 0010-01100 0001?0000- 0110000010 -00010?001 151 -1000?0002 0000010100 000?????10 00-11- -011 010010100? 201 ?????????? ??10000000 0011?10110 1100001- -? 1??11-??20 251 ??????100? ?????????? ?00???0??? ???1-????? ?????????? 301 ?????????? ?????????? ?????????? ?????????? ?1??1????? 351 ?????1???? ?????0???? ???????

Hyloplesion longicostatum 1 001- -01000 000?111001 0000100000 0000010110 0001000000 51 -01- -1- - -0 000000-000 0000000000 0000100000 000?01?002 101 1011?10110 0010-01101 000101??0- 0000000000 000??0??01 151 -1000?0002 0000010100 000????110 01???- -011 01?0???00? 201 ?????????? ??10000000 0011110110 11000010?? 10011?0010 251 ????????0? ?000000??1 10000?01?? ???????1?? ???????11? 301 1-0-??0111 ???101010? 0112110111 010?010??? ?1?31101-1 351 ?10?-?2010 1110?01??? ?????0?

APPENDIX 15: EARLY TETRAPOD DATA MATRIX 373

Ichthyostega stensioei 1 0000000000 0010000-00 0000100000 0010010010 0010000000 51 001- -00000 0000000000 0000000000 0000000000 000?000000 101 0010011010 0000-01000 000100000- 0010000000 001000?000 151 00- -0-0000 0000-000- - 0000101100 0010000000 0100010000 201 1000100010 0000000000 0001?00000 00000000?? ?????????0 251 00010000?? 1000001??1 1100000000 ?00??000- - 0- -0-01010 301 10-0100000 0100001000 0000000000 001- -00100 010?100000 351 0100001000 0001000000 ??00000

Isodectes obtusus 1 0001-01100 0000111000 0000100001 0000010000 0001000000 51 0001000010 0?10100000 0000000000 0000110000 000?01000? 101 0100111111 0110100011 1000?0?00- 0000110002 000?00?000 151 1111002012 0000000100 1101- - -100 00??0?10?? 010???1?0? 201 ???????011 1110000000 0001?00?00 00000?00?? ?????????0 251 ?????????0 1000100??0 -100-000?1 ???????1?? ??????10?? 301 ?0???????? ?????????0 11??1?01?? ??0????21? ?1?21????? 351 ?????????? ?????????? ?????0?

Karaurus sharovi 1 001- -0110? ?00011??0? 1101100001 10?0011-?1 11------01 51 - -1- -1- - -1 ------1- - - -00000 0001------000??1??02 101 001010?111 1??0-0???? ??1- - -??0- 1000110000 -01?-??011 151 -111002012 10?0101100 ????????1? ?- - -1- -01? 1- -?-??-?1 201 - - -1- - -1- - - -???????0 0011????10 11100????1 1??11??110 251 00011?1??0 0001100??- - -1- - -1------1- - -1- - 1------??? 301 1?1??????? ??????0?0? 111-1?111? ?????-?211 11?211???? 351 ??????2??? ??????1??? ?????0-

Keraterpeton galvani 1 0001?0100? ?0?0110-0? 0000100001 0000010111 000?000000 51 -01- -1- - -0 0?110??000 0000000010 00001?0000 00???1??0? 101 ?????????? ?????????? 0???????0- ?0?000000? ??0??????? 151 ?????????? ?????????? 1?0?????10 0??????0?1 01???????? 201 ?????????? ???0000001 1011?01010 1100001- -? ?????-??11 251 010111101? 1000000??0 -100-00100 ?001-1?111 01??011??? 301 ?????????? ??????0??? 11????01?? ????????1? ?1??1?01-1 351 ????-?2??? ?????01??? ???????

Kotlassia prima 1 000??01000 000011??00 0000100000 0000010100 0001000000 51 00011000?0 00-0000000 0000001000 0000?00000 0001010001 101 10??010110 0??0??1100 0001000010 0110000000 000010?001 151 -0- -0?0002 0000000100 0001- - -110 0010011011 0100101000 201 1110111011 1110000000 0111110?10 110000000? ?01???0?10 251 0001101010 1000000111 1100000101 ?001-0?100 01??0110?? 301 101110???? ??????0??? ??0211011? ?????1???? ?1??111001 351 ?110?12010 1?10?0110? 0?110??

374 K. PAWLEY PHD THESIS

Leptorophus tener 1 001- -0??0? ?00?11??0? 1001100000 0000011000 000?000000 51 101- -00010 ??-00??000 0000001000 0000000000 000??1??0? 101 0010110111 0001-01010 ?00?00??0- 0000111000 000??0?0?1 151 -111002002 00???????? 0?01- - -1?? ?01001101? 0101?0?100 201 1110111011 1110110000 00?1?????? ??00?0???? ?????????0 251 000??????? ?????????0 -10?-????? ?????0?1?0 00????1??? 301 ?????????? ?????????? 11??1??1?? ???????21? ????1????? 351 ?????????? ?????????? ??????0

Lethiscus stocki 1 0?1- -0?00? ????11??0? 10001?0000 000001??10 00??000000 51 00???00100 0??0???000 00000000?0 ??1???0?01 0????1??1? 101 ????0????? ????????00 ??1- - -???? ?????????? -????????? 151 ?????????? ?????????? 000????11? ???????0?? 010?????0? 201 ?????????? ?110000000 00111?0?10 1100001- -? ?????-???? 251 ???11?11?? 0???00???? ?????????? ?????????? ?????????? 301 ?????????? ?????????? ?????????? ?????????? ????- -???? 351 ?????????? ?????????? ??- -?0-

Limnoscelis spp. 1 001- -01000 0000111100 0000100000 0000010110 0010100000 51 001- -00000 00-0000000 0000000000 0000100000 000101110? 101 1011010110 0010-01101 0001100011 0110000000 0000100001 151 -1001?0002 0000000100 0001- - -110 00111- -011 0100101000 201 1110111011 1110000000 0111110110 11000000?0 0010010010 251 0001101000 1000000??1 1?000?0?01 ?001-01100 01?????010 301 1011100111 ???1010110 0102110111 0101010211 11011?1001 351 ?11??12010 1110101100 01??1??

Lydekkerina huxleyi 1 0001101000 0000111010 0000100000 0010010000 0001000000 51 101- -00010 00100?1000 0000001002 0000110000 0001010001 101 0001101111 0001000110 000000010- 0000200000 1100000010 151 0111011102 00102001- - 1101- - -100 0010011010 0100101100 201 1110111011 1010000000 0001100000 00000000?0 1010000010 251 000110??1? 1100100111 0100000001 1001-0?110 0111011010 301 1011100111 1101010110 1102110111 0101110211 11??1101- - 351 1111-12011 11?0?01??? ?????0-

Mastodonsaurus giganteus 1 0001101000 0000111010 0000100000 0010011000 0001000000 51 101- -00010 0010011100 0000001002 0000110000 0001010001 101 0000101111 0001000010 010100110- 0000200000 1010001010 151 1111012202 00102001- - 1101- - -100 0010011010 0100101110 201 1010101011 1010000000 0001100020 0000000101 1010000110 251 0001101010 1100100111 0100010111 1001-01110 0111011010 301 1111100111 ??01010?10 110?11011? ?10?1?0??? ?1??1?01- - 351 1111-?2011 1110101101 0??????

APPENDIX 15: EARLY TETRAPOD DATA MATRIX 375

Megalocephalus pachycephalus 1 000?000000 0000111010 0000100010 0000110000 0000000000 51 101- -01010 01-0000000 0000000000 0000000010 000101010? 101 0001010110 0010000101 000010000- 0010000000 000000?010 151 10- -100002 0000000100 0000111100 0011011010 0100101000 201 1010101010 101000000? ?????????? ?????????? ?????????? 251 ?????????? ?????????? ?????????? ?????????? ?????????? 301 ?????????? ?????????? ?????????? ?????????? ?????????? 351 ?????????? ?????????? ???????

Micraroter erythrogeios 1 001- -01000 ?100111000 0000100000 0001010100 1001011000 51 -11- -1- - -0 001000-000 0001000000 0000101001 000?010?02 101 1?110?0110 0??0?01100 000100??0- 0010000?10 00001???01 151 -1000?0002 0000010100 000????110 01??011011 0100101001 201 - - -0111011 1110000000 0011110110 1100001- -? 10011-0010 251 0001101000 1000000?11 10000?01?? 0?01-01110 0??????11? 301 ?-?-?????? ????????01 01??1?011? ?????????? ???31?0??1 351 ?10??1???? ?????????? ?????0?

Microbrachis pelikani 1 0001-01000 0000111000 0000100000 0000010110 0001000000 51 -01- -1- - -0 000000-000 0000000000 0000100000 000?010002 101 1011010110 0010-01101 000100000- 0010000000 0000100001 151 ?1000??002 00?0010100 10???????0 0?-????0?1 010?101001 201 - - -01?101? 1110000000 0011?10110 1100000- -? 10?11-??20 251 0001??1000 1000000??1 10000?010? ???1-01110 0??1011??? 301 ?????????? ????????0? 011?1?011? ?10????21? ???2110??1 351 ?10??1???? ??1?101?0? ?01110?

Micromelerpeton credneri 1 000100110? 000011110? 0000101000 0000011?00 000?000000 51 101- -00010 0?-00??000 0000001000 0000000000 0001?1??01 101 000?110111 00?0000010 100?00??0- 0100101000 000?00?0?0 151 01110?0?02 0????????? 1101- - -100 0110011010 0101101100 201 ???0111011 1110000000 00?1?????0 ??00??0??? ?????????? 251 ???11????0 ??0??????0 -101-00001 0?01-??11? 0?1?011??? 301 ???1?????? ?????????? 11??1?01?? ???????21? ???21?01- - 351 ????-?2??? ??????1??? ???????

Neldasaurus wrightae 1 0001001?00 000011??10 0000100000 0000010000 000?000000 51 0001000010 0010000000 0000000000 0000000000 000101000? 101 0000111110 0110000000 000000000- 0000000000 000?10?00? 151 ?1110?2002 0000000100 1101- - -100 00??011010 010?1???0? 201 ?????????? ??10000000 0001?00000 00000?00?? ?????????0 251 000??????? ?000?00??? ?10??000?? ???????1?? ???????0?? 301 10???????? 11????01?? 110?1?01?1 0101???211 11?21?01- - 351 11?0-??110 1?10?0???? ?????00

376 K. PAWLEY PHD THESIS

Nigerpeton ricqlesi 1 0101?01011 0000111011 0000100000 0010010000 1001000000 51 0001100010 00-0000000 0000001000 0000??0000 000001000? 101 0000011110 001?100001 000010?00- 0100100000 001011?100 151 11110??102 0000000100 ???1- - -10? ?1???????? ?????????? 201 ?????????? ?????????? ?????????? ?????????? ?????????? 251 ?????????? ?????????? ?????????? ?????????? ?????????? 301 ?????????? ?????????? ?????????? ?????????? ?????????? 351 ?????????? ?????????? ???????

Odonterpeton triangulare 1 001- -01000 ?00011??00 00001?0000 0000010110 00??000000 51 -01- -1- - -1 ------000 0000010000 00001?0000 000?01??02 101 ?011010110 0010-?1?00 00???1000- 0010000000 000010?001 151 -1000?0002 0000010100 ?00??????0 01?????0?1 0100????0? 201 ?????????? ???0000000 0011?10110 1100?010?? ???????0?? 251 ????????00 10000001?1 10000?0101 ??01-01110 0????????? 301 ?????????? ?????????? ????1?01?? ???????21? ?1?31????? 351 1????12?10 ?110?0110? ???????

Oestocephalus amphiuminus 1 001- -0?000 ?00?11??00 1000100001 00000100?0 00010000?0 51 -01- -00100 0000000000 1- - - -00000 0010001001 000?01001? 101 ?0???????0 ???0??1000 00????000- 00?000000? 0??010??0? 151 ?1000?00?2 00?0000101 0001- - -110 01??1- -011 010010100? 201 ?????????? ???000?000 0011??0010 1100001- -? 1??1?-11?? 251 0001111110 0000010??1 0?001????? ?????????? ?????????? 301 ?????????? ?????????? ?????????? ?????????? ????- -???? 351 ?????????? ?????????? ??- -?0?

Orobates pabsti 1 001- -01000 ?0?1111100 0000100000 0000010110 0010100000 51 001- -00000 00-0000000 0000001000 00001?0000 000?011102 101 10??01???0 0?10-?1?01 0001100010 0110000000 00?010??0? 151 ?10?1???0? 0000000??? 000????1?0 ?0??1- -01? 010010100? 201 ???1- - -01? 1110000000 0111?10010 110000???0 0010010020 251 0??11?1000 1000000??1 ?100000?01 ?001-0?100 01110110?? 301 101110???? ???101011? 0102110111 0101010211 11?1111001 351 ?110?12010 ??10?01?0? ?11110-

Ossinodus pueri 1 ?100?0???1 0??0?????1 0000?????? ?00001?000 000?000000 51 000??00010 0?-000?000 000000000? ??0000?01? 000?00??0? 101 ???00??0?0 0?????0000 0?0000??0- 001000000? ?????????? 151 ?????????? ?????????? 000??????? ?????????? 01???????? 201 ?????????? ???0000000 0001000??? ?????????? ?????????? 251 ?????????? 1??010???1 1100000000 000??00110 00101110?? 301 1000000000 00?00?0??? ?????????? ?????????0 0???110000 351 01001??000 000100???? ???????

APPENDIX 15: EARLY TETRAPOD DATA MATRIX 377

Paleothyris acadiana 1 001- -01?00 000?111?01 0000100000 0000011111 0000100000 51 001- -00000 00-0100000 0000000000 0000100000 000101110? 101 1011010110 0110-01101 00????0011 0110000000 0000100001 151 -100??0002 0000000110 000????110 0???1- -011 010?1??00? 201 ???????010 1110000000 0011?10?10 11000000?0 0110010020 251 0001???000 1000000111 10000?010? ??01-011?0 ?????1101? 301 100?10???? 11?1010111 010211111? ?10??1?211 11?111001- 351 11?0012010 ??10?11?0? ?1111??

Panderichthys rhombolepis 1 0000000000 0000000-00 0000000000 0000010000 0000000000 51 00???00000 0000000000 0000000000 0000000000 000?000?00 101 0000000000 0000000000 000000000- 0000000000 000000?000 151 00- -000000 0000-000- - 0000000000 0-00000000 0000000000 201 0000000010 0000000000 00?0?0?0?0 - -00-00000 00000000?0 251 00000?0??? 00000?0??0 -0??-?0000 00000000- - 0- -0-00?00 301 00-0------100000 0000000000 000- -0000? ?00-0-???? 351 ?????????? ?????0???? ??- -???

Pantylus cordatus 1 001- -01000 0100111001 0010100000 0001010100 1001000000 51 -11- -1- - -0 000000-000 0001100000 0000100100 000?010002 101 1011010110 0010-01?0? 001- - -000- 0010000000 000?1?0001 151 -1000?0002 0000010100 ???1- - -110 00- -011011 010?1010?1 201 - - -1- - -011 1110000010 0011?10110 1100001- -1 10011-0010 251 0001101000 1000000??1 10000?0101 ?001-01110 0101?11011 301 1001100100 ??00010101 0112110111 010101021? ?1?21?01-1 351 ?10?-?2010 1110?01?0? ?0111??

Pederpes finneyae 1 0000?0?0?? ?0?011??01 0000100000 00000?00?0 000?0???00 51 0001100010 0?-0000000 0000000000 0000???01? 000?000?00 101 0?0?00???0 0?000?0100 0000000?0- 0?1000000? 000??????? 151 ????1?000? 000??????? 000??????? ????0??01? 010?0????0 201 ???0???0?? ???0000000 0001?0?000 00000000?0 000????0?0 251 ???????0?0 ?100101??1 ?100?00000 000??001?0 00101110?? 301 100000???? ???????0?0 010?0?0001 00?0000110 010?110000 351 01001?2?00 000?0?1000 0????0?

Pelodosotis elongatum 1 001- -01000 ?10011??00 0000100000 0001010100 100?011000 51 -11- -1- - -0 001000-000 0001000000 0000101001 000?010002 101 1011010110 0010-01100 000100??0- 0000000010 000?10?001 151 -1000?0002 00?0010100 100?????10 0??????0?1 ????????0? 201 ?????????? ???0000000 0011?10110 1100000- -? ???11-??10 251 0001??1000 1000000??1 10000?0101 ??01-01110 0??????11? 301 1-?-?????? ???0010?01 011?11011? ?10???0211 11?211?1-? 351 ?10?-12010 ?110?01100 0-11100

378 K. PAWLEY PHD THESIS

Peltobatrachus pustulatus 1 ?01- -????? ?????????? ????1????? ?????10000 0001000000 51 101- -0?0?0 00-0000000 0000001000 0000??0?00 00010?000? 101 0???1????? ?????????? ??????000- 010?100000 ?100??0??? 151 ?1110?0102 0000100100 110??????? ????011010 01?01????? 201 ?????????? ??1??????0 0001100?10 1100001- -? ????????10 251 0001?0?0?? 110010011? ?????????? ??????1110 0?11??1010 301 10111?0111 ??01010?10 11??1?011? ?1???????? ??????0??- 351 1111?1?0?0 ?????????? ?????1?

Petrolacosaurus kansensis 1 001- -01000 ?001111?01 0000100000 0000011111 0000100000 51 001- -00000 00-0100000 0000000000 0000100000 000101110? 101 1011010110 0110-01101 0001100011 0110000000 000010?001 151 -1001?0002 0000000110 0001- - -110 01- -011011 0100???001 201 - - -1- - -010 1110000000 0011110110 11000000?0 0110010020 251 0001101000 1000000111 1100000?01 ?001-01100 0111011011 301 100?100111 1111010?11 010211111? ?10?110211 11?111001- 351 1110012010 1110?11100 11111??

Phlegethontia linearis 1 001- -01000 ?00?110-?0 1000100001 ?1- - -0???? 01------?0 51 -01- -1- - -? ?????????- 1- - - -0000? ??1010-000 ????01??1? 101 ?????????? ?????????? 0??????0?? 1?00000?0? ????????0? 151 ?10??0???? ???0000101 0??1- - -110 0- - -1- -1- - 1- -?-???01 201 - - -1- - -1- - - -?0000000 0011?0?010 11000?1- -0 10?10-11?? 251 0??11?1111 0000010??- - -1- - -???? ?????????? ?????????? 301 ?????????? ?????????? ?????????? ?????????? ????-????? 351 ?????????? ?????????? ?????0-

Pholiderpeton attheyi 1 0000001000 000011??10 0000100000 0000010100 000?000000 51 0000000000 10-0000001 0000000000 0000?10010 000?01??0? 101 ?010010110 0010-00001 000000??0- 0010000000 000?10?001 151 -10010?002 000??????? 000????1?0 0011011010 0100111000 201 1110111011 1010001000 0001?00?00 11000001?? ????????10 251 0001?0?0?? 110000000? ?????????? ?????????? ?????????? 301 ?????????? ?????????? ?????????? ?????????? ?????????? 351 ????????00 ?????????? ???????

Pholiderpeton scutigerum 1 0?0?00?000 0000110-10 0?001???00 ?000010100 0000000000 51 0000000000 10-0000001 0000000000 0000?10010 0001012100 101 1010010110 0010?0000? ??0000000- 001?000000 000010?001 151 -1001?2002 0000000100 0??0011100 0?11011010 010?111000 201 1110111011 1110001000 0001000?00 1100?00110 000??000?0 251 ????????00 110000???1 ?100?0000? ??10100100 0010111000 301 10000010?0 00?0010??0 0?0?0?0??1 0??0001110 01??1????? 351 ?????????? ?????????? ???????

APPENDIX 15: EARLY TETRAPOD DATA MATRIX 379

Phonerpeton pricei 1 001- -11101 0000111101 0000111001 0000011000 0001000000 51 101- -00010 00-0000000 0000001000 1100000000 0010110001 101 0001110110 0011100101 100?10000- 0100100000 0000?0?011 151 -111002002 0000101100 0101- - -100 0010011010 0100101100 201 1110111011 1110000000 0001?00000 00000000?? ?????000?0 251 00011010?? 1100100??? ?????????? ?????0111? 00?1011??? 301 1?1?1????? 1???????1? 110211011- 110??102?1 111??????? 351 ?????????? ?????????? ???????

Platyoposaurus stuckenbergi 1 101- -01000 100011??10 0000100000 0010010000 0001000000 51 101- -00010 10-0000000 0000001001 0000?10000 000101000? 101 0001111110 0010000100 000?00000- 0100100000 00000000?0 151 1111000102 0100100100 0?01- - -100 0010011010 0101101000 201 1110111011 1110000000 0001100000 00000000?0 10100000?0 251 0001?01010 1100100??0 -?00-?0?01 ?????01110 00110110?? 301 1?1110???? 1?????01?? ?????????1 ??0?0??211 11??1101- - 351 111?-?2011 1????????? ??11???

Platyrhinops lyelli 1 001- -01100 000011??00 10001?1000 0000010000 000?000000 51 101- -00010 0?-00??000 0000001000 0000000000 001?11??0? 101 000?110111 0010??0?11 000010??0- 0100100000 100?????01 151 -111000002 11???????? 0?01- - -1?0 0?1001101? 010?101100 201 ???0111011 1110110000 0001?00000 00000?00?? 1010?00010 251 0001101010 ?00??00110 -?00-00?01 ???1-011?0 ?????11??? 301 1????????? ?????????? 110?11111? ???????21? ?1?21101- - 351 111?-?20?? ??????1?1? ??1100?

Proterogyrinus scheelei 1 000??01000 0000110-00 0000100000 00?0010100 0000000000 51 0000000000 10-0000000 0000000000 00001100?0 000?01210? 101 1???0????0 ??10??0001 000000000- 0010000000 000010??0? 151 ?1001?0002 0000000100 000????100 001?0?1010 0100?11?00 201 ?1?0111011 1?10001000 000100000? 1000000010 0000000010 251 0001101000 1?00000001 0100000000 1011-00100 0000111000 301 1000001000 0??0010100 0101010101 001000?110 0101110001 351 0110002000 0101001000 111100-

Ptyonius marshii 1 001- -01000 0000110-00 0000100001 0001010110 0001000000 51 001- -00100 0??000?000 0100000000 0000100000 00???1??0? 101 001000?110 0010-01000 000100??0- 0010000000 000?00?001 151 -1100??0?2 00?00?0100 1?0?????10 0??????0?1 010?????0? 201 ?????????? ???0000001 1011?01010 1100001- -1 ?0111-?111 251 100111101? 1000000??0 -100-00000 ??01-0?1?0 0????11??? 301 ?????????? ??????0??? 11??1101?? ??0????21? ?1?21101-1 351 ???0-?2??? ?????01??? ?????0-

380 K. PAWLEY PHD THESIS

Rhinesuchidae 1 0001001000 0000111010 0000100000 0010010000 0001000000 51 101- -00010 10-0001000 0000001002 0000110000 0001010001 101 0001101111 0001000100 000000?10- 0000200000 000000?011 151 -111011102 0000100100 0101- - -100 0010011010 0100101000 201 1110111011 1110000000 0001100000 00000000?? ????????10 251 00011010?0 1100100110 -100-00001 0001-01110 011101101? 301 101110???? 1??1010?10 110211011? 01???10211 11?21101- - 351 11?1-12011 1110?01?0? ??11001

Rhynchonkos stovalli 1 001- -01000 0100111000 0000100000 0000010100 0001011000 51 -11- -1- - -0 000000-000 0001100000 000010?000 010?010002 101 1010010110 0000-01000 000100000- 0000000010 001010??01 151 ?1000?0002 0000010100 100????110 01-1011011 0100101001 201 - - -01?101? 1110000000 0011?10?10 1100000- -1 10011-002? 251 ????????00 ?????????? ?????????1 ?????????? ?????????? 301 ?????????? ????????01 1???11011? ?1???????? ??????01-1 351 ?10?-1???? ?????????? ???????

Saharastega moradiensis 1 001- -0??00 100?11???0 0000100000 0010010?00 10010000?0 51 0001?000?0 00-000?000 0000001000 0000?1?000 00??010?0? 101 000?000110 0010?00101 000010000- 01?0100000 0?001??000 151 11110?0102 0000000100 ?????????? ?????????? ?????????? 201 ?????????? ?????????? ?????????? ?????????? ?????????? 251 ?????????? ?????????? ?????????? ?????????? ?????????? 301 ?????????? ?????????? ?????????? ?????????? ?????????? 351 ?????????? ?????????? ???????

Sauropleura spp. 1 001- -01000 0000110-00 10001?0001 0001000110 000?000000 51 001- -00100 000000?000 0100000000 0000000000 000?010?0? 101 ?010?00110 0010-01000 00?????00- 00?0000000 000?00?001 151 -1000?20?2 00?0000100 1?01- - -110 00??1- -0?1 0100??100? 201 ???????1- - - -10000001 1011?01010 1100001- -1 101?0-?111 251 1001111111 1000000??0 -100-00000 ??01-0?100 0????11??? 301 ?????????? ??????0??? 110211011? ??0??1?21? ?1?21101-1 351 ???0-?20?0 ?????01?0? ?01110-

Saxonerpeton geinitzi 1 001- -01000 0000111000 0000100000 0001010100 0001100000 51 -11- -1- - -0 000000-000 0001100000 0000100000 000?01000? 101 1011010110 0010-0110? 0001?0000- 0110000010 -000-0?001 151 -100000002 0000010100 0001- - -110 00-????011 0100???001 201 - - -????011 1110000000 0011?10110 1100001- -1 10011-0010 251 0001101000 1000000??1 1000000101 ?001-01110 0101011?1? 301 1?0??????? ???????101 0112110111 010101021? ?1?2110??1 351 ?10?????10 ?110?01??? ?01110?

APPENDIX 15: EARLY TETRAPOD DATA MATRIX 381

Schoenfelderpeton prescheri 1 001- -0??0? ?00?11??0? 1101100000 0000010000 000?000000 51 101- -000?0 ??-00???0? 0000001000 1?00100000 000??1??0? 101 0010110111 001?-01010 ??0?00??0- 0000111000 001??0?0?1 151 -111002002 00???????? ???????1?0 00??01101? 010??01?0? 201 ???????0?? ?11?000000 0??1?????? ???0?????? ?????????? 251 ?????????? ?????????0 -????????? ?????????? ?????????? 301 ?????????? ?????????? 11??1??1?? ???????21? ???21????? 351 ?????????? ?????????? ???????

Scinosaurus crassus 1 0001?01000 0000110-00 0000100001 0000010100 01------00 51 -11- -1- - -0 0??00??000 0001000010 0000100000 000?01??0? 101 001????11? 0010-01010 001- - -??0- 1000000000 -0??-0?001 151 -1000??002 0001000100 ?????????? ?????????? ?????????? 201 ?????????? ???0000001 1011?01?10 1100001- -? 1????-??11 251 00011?10?? 1000?????0 -100-00100 ?001-0?100 0????11??? 301 1????????? ????????11 0???1101?? ???????21? ?1??1?01-1 351 ?1??-?2??? ?????01??? ???????

Sclerocephalus spp. 1 001- -01100 0000111010 0000100000 0010010000 000?000000 51 101- -00010 00-000?000 0000001000 0000000000 000?01??01 101 0000110110 0011100000 000000??0- 0100100000 000?00?011 151 -111000102 ?0???????? 0?01- - -100 0010011010 0101101100 201 1?10111011 1110000000 0001?00000 00000000?0 001???0010 251 0001101010 1100100??0 -100-00001 0?01-0?110 0011011010 301 10111001?? 110101011? 1102110111 110101?21? ?1?21101-? 351 ?1??-?2?11 ??10?01?1? 001100-

Seymouria spp. 1 001- -01001 0000111101 0000100000 0000010100 0001000000 51 0001100000 00-0000000 0000001000 0000100000 000?010001 101 1001010110 0010000101 0001100010 0110000000 000010?001 151 -0- -0?0002 0000000100 0001- - -110 0011011011 010?101000 201 1110111011 1110000000 0111110010 11000000?0 0010000010 251 0001101010 1000000111 1100000101 0011-01100 0??1011010 301 1011100111 1101010110 0102110111 0101010211 11?1111001 351 1110012010 1110101100 0?110??

Solenodosaurus janenschi 1 001- -???0? ????11??0? 00001?0000 0000010110 0000100000 51 001- -00000 00-0000000 00000010?0 00001?0000 000?01?10? 101 ?????????? ?????????? ??????00?? ?????????? ??????0??? 151 ?????????? ?????????? 000??????? ???????0?? 010?101??? 201 ?????????? ???0000000 0011?00010 11000000?? ????????10 251 0001?010?0 1000000?11 1100000?01 ??0??011?? ?????????? 301 1?1?10???? ???????1?? 0?02110111 ??0101?21? ????11???? 351 ?????????? ?????????? ???????

382 K. PAWLEY PHD THESIS

Stegotretus agyrus 1 001- -01000 010011??01 0010100000 0001010100 0001000000 51 -11- -1- - -0 000000-000 0001100000 0000100100 000?010002 101 10?1010110 ?010-01?0? 001- - -?00- 0010000000 000010?001 151 -1000?0002 0000010100 000????110 00??011011 01001??00? 201 ?????????? ??10000010 0011110?10 1100001- -? 10?11-00?? 251 ????????0? ?????????? ?0????01?? ?????????? ???????111 301 1-?-??0100 ???0010?0? 111?11011? ??0????21? ?1??11???? 351 ?????1???? ?????????? ???????

Thabanchuia oomie 1 0001?01??? ???0?????? 00001??00? ?????10010 000?000000 51 001- -0?010 0010100000 0000000000 00000?0000 ?????1000? 101 010010?111 0100000011 000010?00- 0000210002 101100?000 151 1110??2?02 00?????1?? 1?11- - -100 00?????0?0 0?00?????0 201 0??00??00? ??10000000 0001?0?000 11000001?? ?????????0 251 ?????????? 1?0???0??1 ?100?00?11 1?0??0?1?? 0??????0?? 301 1????????? ?????????? ?????????? ?????????? ?????????? 351 ?????????? ?????????? ???????

Tiktaalik roseae 1 0000?00??? ?0??00??0? 0000??0?00 00000?0000 000?00?000 51 101- -00000 0?-00??000 00000000?0 00000?0000 000?00??0? 101 00??0????0 ???0????0? 0?00?0??0- 0000000000 00???????? 151 ????0?000? 0000-00??? 0000000000 0000000000 0000010000 201 0000000010 000000000? ?????????? ?????????? ?????????? 251 ?????????? 0000001??? ?????????? ???00000- - 0- -0-00?10 301 0?0?000000 0000100000 0000?00000 0?0- -00?00 000-0-???? 351 ?????????? ?????????? ?????00

Triadobatrachus massinoti 1 001- -????? ?????????? 1?01100?0? ??????1-?1 01------01 51 - -1- -1- - -1 ------1- - - -0100? ?001------?????1??0? 101 ?????????1 ???0??1011 ??1- - -??0- 000111000? -????????? 151 ?1110?20?2 10?0101100 ?????????? ??- -1- -1- - 1- -?-??-?1 201 - - -1- - -1- - - -???????0 0011??0?10 11000?1- -1 10??1???10 251 0001111?00 0001010?1- - -1- - -0?0? ??01-0?100 0????110?? 301 ??1??????? ??????0?0? 111-1?11?? ??0??-?21? ?1??1?01- - 351 1?0?-?2??? ?110?01??? ?0???0-

Trimerorhachis insignis 1 0001001100 0000111000 0000100000 0000010000 000?000000 51 0001000010 0010000000 0000000000 0000000000 0001010001 101 0100101110 0000000000 000000000- 0000000000 000100?000 151 1111002002 0000000100 1101- - -100 0010011010 0100101000 201 1110111011 1110000000 0001100000 0000000010 0010000010 251 0001101010 1000100110 -100-00011 1001-??1?? ??????10?? 301 10?1?1???? 11????0110 110?110111 0101010211 110?1101- - 351 1110-?2110 1110101100 0????00

APPENDIX 15: EARLY TETRAPOD DATA MATRIX 383

Tuditanus punctulatus 1 001- -01000 ?000111000 0000100000 0001010100 000?100000 51 -11- -1- - -0 000000-000 0001100000 00001?0000 000?01?002 101 1011010110 0010??110? 0??1?0??0- 0?10000000 00??1??001 151 -1000?0002 0000010100 000????110 00?????0?? 01?????00? 201 ?????????? ???0000000 0011110?10 1100001- -? 10?11-0??0 251 0001101000 1000000??1 10000?0101 0?01-0?110 0101011??? 301 ?????????? ?????????1 01??11011? ?10???021? ???2110001 351 ?100?1201? ?110?01100 ?01110-

Tulerpeton curtum 1 ??00????00 0????????? ?????????? ???????0?? ?????????? 51 ?????????? ????00?000 ?????????? 00???????? ?????????? 101 ?001?1?01? 0?0??0???? ?????????? ?????????? ????????0? 151 ?????????? ?????????? ???????100 0??????000 ?????????? 201 ?????????? ???000000? ????????00 00000000?? ?????????? 251 ?????????? 1??0?0???1 ?100000000 000010?100 000?111010 301 1000000000 00000?0000 010100000? 001- -00100 01?0100000 351 0?0?0?2000 0001001000 01110?0

Tupilakosaurus spp. 1 000100???? ?00??????0 00001?0000 0000010010 0001000000 51 001- -00010 0?1010?000 0000000000 0000??0000 0001?1000? 101 ??00??1111 01?0000011 000000000- 0000210002 101100?000 151 1110002002 0010000100 1?1??????? ?0???????? ??001???0? 201 ???????0?? ?11000000? 0001?00000 11000001?? ?????????0 251 ?????????? 1?00100??1 0??00?0??? ?????????? ?????????? 301 ?????????? ?????????? ?????????? ?????????? ?????????? 351 ?????????? ?????????? ???????

Urocordylus wandesfordii 1 0?1- -??000 ?0??1???00 0000??0??? ?00101???? ?????????? 51 0????????0 0?00???000 010?000000 ??000000?0 000??1??0? 101 ?????????? ??????1?00 ?????????? 0????00??? ?00?0???0? 151 ?100?????? ?????????? 1?0????110 0??????0?1 010?????0? 201 ?????????? ???0000001 1011?01010 1100001- -1 ?01??-??11 251 1001111011 1000000??0 -100-00000 ??01-0?1?0 0????11??? 301 ?????????? ??????0??? 110??101?? ???????21? ?1?21101-1 351 ???0-?2??? ?????01??? ?????0-

Valdotriton gracilis 1 001- -01100 ?00011??00 11011??001 11- - -11-?? 11------?1 51 ?-1- -1- - -1 ------1- - - -0000? ??01------1-???1??02 101 0010101111 1??0-0???? ??1- - -??0- 100101000? -00?-0??01 151 -1110?2002 00?0101100 ???1- - -110 0- - -1- -1- - 1- -?-??-?1 201 - - -1- - -1- - - -?1??0000 0011?10?11 1110011- -? 1??11-1?10 251 00011?1110 0001000??- - -1- - -1------1------1------?1? 301 ??1??????? ??????0?0? 111-11111? ?????-?21? ?1?21????? 351 ???????2?? ???1??01?? ???????

384 K. PAWLEY PHD THESIS

Ventastega curonica 1 0?000?0000 00??????00 0000??0??? ?01001?0?? ?????????? 51 ?????????? ?????????? ?????????? ??000?0000 00???0???? 101 ?000?0?00? ??0?000000 0???????0- 00?0000?00 000?0???00 151 0????????? ?????????? 0000001101 0010000000 0000010000 201 0000000010 000000000? ?????????? ?????????? ?????????? 251 ?????????? 0????????1 01000?0000 000??0?0- - 0- -0-?1??? 301 ??-??????? ??????1??? ?????????? ?????????? ?????????? 351 ?????????? ?????????? ???????

Westlothiana lizziae 1 001- -01000 0000111000 000010000? 00??010110 000?000000 51 001- -00000 0?-00??000 0000000000 00001?0000 000?01??0? 101 ????0?0??0 ?0?0??1101 ??01?0??0- 0010000000 000?1????? 151 ?100??0002 00???????? 000?????10 0???1- -0?1 010?????0? 201 ?????????? ???0000000 0011110?10 11010000?? ?????????0 251 00011010?0 1000000??? ?????????? ???????1?? ????????1? 301 1????????? ?10????110 0102110111 ?1?10?0211 11??11001- 351 11?0012010 1110?11100 011?100

Whatcheeria deltae 1 0000001001 ?0?011??01 0000100000 00?0010000 0000000000 51 0001100010 00-0000000 00000000?0 0000??0010 000?000100 101 ??000????? ??????00?0 0?0000??0- 0?1??????0 0?0??????? 151 ???????0?1 ???0000??? 0000001100 00??0?10?0 ???001??00 201 1000100010 00?0000000 000100?000 0?000?00?? ????????10 251 000??0???0 1100101??1 1100000000 ??01-00110 000011101? 301 ?00??????? ???001000? 0101000?01 0?1000?110 01??110000 351 ?10??02000 00??001000 ?????0?

Pawley, K. 2006. 385 The Postcranial Skeleton of Temnospondyls (Tetrapoda: Temnospondyli) PhD Thesis. La Trobe University, Melbourne

APPENDIX 16. COMPARATIVE ANALYSES: WALKING WITH EARLY TETRAPODS

Results of tests of degrees of character inclusion

An analysis using only the cranial characters of the main analysis (245 characters, 92 taxa with Casineria deleted) resulted in 6487 trees of 549 steps (CI = 0.3825, RI = 0.8289, RC = 0.3171) (Figure 87).

TABLE 14. COMPARISON OF DEGREES OF CHARACTER INCLUSION. CHARACTERS NUMBER OF NUMBER OF TREE LENGTH NUMBER OF CONSISTENCY INCLUDED TAXA CHARACTERS (STEPS) TREES INDEX

Main analysis 93 349 832 9 0.4387 (all characters) Main analysis (cranial 92 210 611 6487 0.3825 only) Main analysis (historic vertebral 93 345 820 49 0.4412 removed)

Table 14 clearly indicates that in the main analysis, the inclusion of postcranial characters increases resolution and decreases homoplasy. The results of the analysis removing historically significant vertebral characters (TRU VER 8, TRU VER 8, TRU VER 14) resulted in 47 trees of 820 steps (93 taxa, 346 characters, CI = 0.4415, RI = 0.8559, RC = 0.3778). The tree topology is identical in topology to that of the main analysis (Figure 62), except that Edops is collapsed into a clade with Casineria and Caerorhachis. Additionally excluding TRU VER 33 did not produce any further change in tree topology (49 trees of 816 steps, 93 taxa, 345 characters, CI = 0.4412, RI = 0.8549, RC = 0.3772).

TABLE 15. COMPARISON OF DEGREES OF CHARACTER INCLUSION, ANALYSES USING THE SAME TAXA AS RUTA ET AL. (2003). CHARACTERS NUMBER OF NUMBER OF TREE LENGTH NUMBER OF CONSISTENCY INCLUDED TAXA CHARACTERS (STEPS) TREES INDEX

Same taxon list as 90 352 892 1130 0.4063 Ruta et al. (2003) Same taxon list as Ruta et al. (2003) 90 201 589 2477 0.3497 (cranial only)

386 K. PAWLEY PHD THESIS

FIGURE 87. Results of the main phylogenetic analysis of early tetrapods, postcranial characters and Casineria omitted. Removal of postcranial characters and Casineria resulted in a strict consensus of 6487 trees of 549 steps (92 taxa, 201 characters, CI = 0.3825, RI = 0.8289, RC = 0.3171). APPENDIX 16: EARLY TETRAPOD COMPARATIVE ANALYSES 387

As shown in Table 15, removing postcranial characters from the analysis using the same taxon list as Ruta et al. (2003) decreased phylogenetic resolution, as shown by the increase in the number of trees, and lowered the consistency index, as in the main analysis. Table 16 shows that removing the postcranial characters from the analysis of Ruta et al. (2003) increased the consistency index slightly but also resulted in an increase in the number of trees [data taken from Ruta et al. (2003)]. Replacing the postcranial characters and coding of Ruta et al. (2003) with those of the main analysis resulted in an huge increase in the number of trees, which interestingly is not associated with a corresponding lack of resolution (Figure 91), and also an improvement in the consistency index.

TABLE 16. COMPARISON OF DEGREES OF CHARACTER INCLUSION, ANALYSES USING THE MATRIX OF RUTA ET AL. (2003). [*taken from Ruta et al. (2003)]. CHARACTERS NUMBER OF NUMBER OF TREE LENGTH NUMBER OF CONSISTENCY INCLUDED TAXA CHARACTERS (STEPS) TREES INDEX

Ruta et al. (2003) 90* 319* 1375* 64* 0.2392*

Ruta et al. (2003) 90* 224* 1022* 1188* 0.2485* (cranial only) Ruta et al. (2003) 90 371 1326 47952 0.2949 (postcranial replaced)

Results of tests of phylogenetic stability

Constraining the Anthracosauria and Gephyrostegidae to fall within the Neospondyli (forming the traditional ‘reptiliomorph’ or stem amniote lineage) takes 24 extra steps (61 trees, total 856 steps, CI = 0.4206, RI = 0.8422, RC = 0.3542), indicating that this hypothesis of phylogenetic relationships of early tetrapods is significantly less parsimonious than that of the main analysis. Constraining Caerorhachis and Casineria as sister taxa to the Anthracosauria + Neospondyli (stem amniote lineage) takes an extra 29 extra steps (18 trees, total 861 steps, CI = 0.4181, RI = 0.8406, RC = 0.3515), and is less parsimonious than the previous analysis. The Temnospondyli are the stem group to the Anthracosauria + Neospondyli in both these analyses.

Results of comparative phylogenetic analyses

Taxon sampling effects

This comparative analysis used the same taxon list as Ruta et al. (2003), but changed both the cranial (201 included) and postcranial (151 included) characters (90 taxa, total 352 characters). The result was 1130 trees of 891 steps (CI = 0.4063, RI = 0.8194, RC = 0.3329), with a consensus tree (Figure 88) of essentially the same tree topology as the main analysis. Re-weighting the characters by their consistency indices (Figure 89) resolved the polytomies within the Aïstopoda and Lissamphibia. 388 K. PAWLEY PHD THESIS

FIGURE 88. Results of the analysis using the same taxon list as Ruta et al. (2003). Strict consensus of 1130 trees of 892 steps using the same taxon list as Ruta et al. (2003), but the characters and data of the main analysis (90 taxa, 352 characters, CI = 0.4053, RI = 0.8194, RC = 0.3329). APPENDIX 16: EARLY TETRAPOD COMPARATIVE ANALYSES 389

The effect of changing postcranial characters

This comparative analysis used the original matrix of Ruta et al. (2003), but changed the postcranial characters (90 taxa, total 376 characters, including 158 postcranial). The result was 47952 trees of 1326 steps (CI = 0.2949, RI = 0.7341, RC = 0.2165), a consensus tree is shown in (Figure 91). Generally the tree topology is similar to that of the main analysis, the Temnospondyli are sister taxa to the Neospondyli, as in the main analysis. Despite the high number of trees, the tree is reasonably well resolved, with collapses within the archeriamorphs, basal Temnospondyli, tuditanomorph Microsauria, and basal Nectridia. Re-weighting the characters by their consistency indices resulted in a single tree (Figure 92). The re-weighted tree resolves all collapsed clades, again the Temnospondyli are sister taxa to the Neospondyli. The results of these analyses are summarised in Table 16. The results of the two comparative analyses clearly indicate that the novel phylogenetic position of the Temnospondyli presented here (as sister taxa to the Neospondyli, forming the Terrapoda), remains unchanged in all analyses except constraint trees, and is due to the change in postcranial characters and coding, rather than a taxon sampling effect.

Comparison of phylogenetic analyses

Table 17 contains the results of all analyses that contain a full complement of characters, except constraint trees.The analyses presented in Table 17 are similar in the number of taxa, but differ somewhat in the number of characters. Tree length varies considerably, the shortest tree is that of the main analysis, which is only 61% of the length of the analysis of Ruta et al. (2003), despite the increased number of taxa (3% more) and characters (9% more).

TABLE 17. COMPARISON OF PHYLOGENETIC ANALYSES USING ALL CHARACTERS. [*taken from Ruta et al. (2003)]. NUMBER OF NUMBER OF TREE LENGTH NUMBER OF CONSISTENCY ANALYSIS TAXA CHARACTERS (STEPS) TREES INDEX

Main analysis 93 349 832 9 0.4387

Ruta et al. (2003) 90* 319* 1375* 64* 0.2392*

Same taxon list as 90 352 892 1130 0.4063 Ruta et al. (2003) Ruta et al. (2003) postcranial characters 90 371 1326 47952 0.2949 changed

The tree of Ruta et al. (2003) is the longest, and also has the lowest consistency index. The combination of the cranial characters of Ruta et al. (2003) and the postcranial characters of the main analysis resulted in a dramatic increase in the number of trees, but not in tree length. Interestingly, the high number of trees did not result in a reduction in the consistency index, which is midway between that of the main analysis and that of Ruta et al. (2003). 390 K. PAWLEY PHD THESIS

FIGURE 89. Results of the analysis using the same taxon list as Ruta et al. (2003), and all characters re- weighted. Strict consensus of three trees obtained using the results of analysis using the same taxon list as Ruta et al. (2003), but the characters and data of the main analysis (Figure 88), with all characters re- weighted by their consistency indices. APPENDIX 16: EARLY TETRAPOD COMPARATIVE ANALYSES 391

The comparative analysis using the same taxa as Ruta et al. (2003) shows little evidence of sampling effect, the tree is somewhat longer and the consistency index slightly lower than that of the main analysis, which is expected given the more diverse taxon sample.

TABLE 18. COMPARISON OF PHYLOGENETIC ANALYSES USING ONLY CRANIAL CHARACTERS. [*taken from Ruta et al. (2003)]. NUMBER OF NUMBER OF TREE LENGTH NUMBER OF CONSISTENCY ANALYSIS TAXA CHARACTERS (STEPS) TREES INDEX

Main analysis 93 201 549 6487 0.3825 cranial characters only Ruta et al. (2003) 90* 224* 1022* 1188* 0.2485* cranial characters only Same taxon list as Ruta et al. (2003) 90 201 589 2427 0.3497 cranial characters only

The results of the analyses using only cranial characters (Table 18) all have much higher numbers of trees (and thus are less resolved) and possess lower consistency indices than the analyses that include all postcranial characters, listed in Table 17. 392 K. PAWLEY PHD THESIS

FIGURE 90. Results of the analysis using the same taxon list as Ruta et al. (2003), postcranial characters omitted. Removal of postcranial characters resulted in a strict consensus of 2427 trees of 589 steps using the same taxon list as Ruta et al. (2003), but the cranial characters and data of the main analysis (90 taxa, 201 characters, CI = 0.3497, RI = 0.7860, RC = 0.2749), with all postcranial characters omitted. APPENDIX 16: EARLY TETRAPOD COMPARATIVE ANALYSES 393

FIGURE 91. Results of the analysis using the original matrix of Ruta et al. (2003), with the postcranial characters and data replaced with those of the main analysis. Strict consensus of 47952 trees of 1326 steps obtained using the original cranial characters and data of Ruta et al. (2003), with the postcranial characters and coding replaced with those of the main analysis (90 taxa, 371 characters, CI = 0.2949, RI = 0.7341, RC = 0.2165). 394 K. PAWLEY PHD THESIS

FIGURE 92. Results of the analysis using the original matrix of Ruta et al. (2003), with the postcranial characters and data replaced, and all characters re-weighted. Single tree obtained using the original cranial characters and matrix of Ruta et al. (2003), with the postcranial characters replaced with those of the main analysis (Figure 91), with all characters re-weighted by their consistency indices. Pawley, K. 2006. 395 The Postcranial Skeleton of Temnospondyls (Tetrapoda: Temnospondyli) PhD Thesis. La Trobe University, Melbourne

APPENDIX 17. LIST OF SYNAPOMORPHIES: WALKING WITH EARLY TETRAPODS

Taxa more derived than Panderichthys Tetrapoda NAS 1 CI = 0.500 0 −−> 1 INTEMP 1 CI = 0.125 0 −−> 1 Ichthyostegamorphs more derived than INTEMP 2 CI = 0.333 0 −−> 1 Tiktaalik PASPHE 9 CI = 0.667 0 −−> 1 LAT ROS 1 CI = 0.500 0 −−> 1 PREART 3 CI = 0.333 0 ==> 1 LAC 2 CI = 0.143 0 ==> 1 TRU VER 20 CI = 1.000 0 −−> 1 PAR 2 CI = 0.143 0 −−> 1 SAC VER 1 CI = 0.400 0 −−> 1 ECT 2 CI = 0.200 0 −−> 1 CAU VER 5 CI = 1.000 0 −−> 1 PTE 10 CI = 0.143 0 −−> 1 RIB 10 CI = 0.333 0 −−> 1 PSYM 4 CI = 1.000 0 ==> 1 INTCLA 1 CI = 0.125 0 −−> 1 DEN 1 CI = 0.500 0 ==> 1 INTCLA 5 CI = 0.333 0 −−> 1 DEN 3 CI = 0.500 0 −−> 1 SCACOR 2 CI = 0.500 0 ==> 1 SPL 3 CI = 0.500 0 ==> 1 RAD 1 CI = 1.000 0 −−> 1 RIB 8 CI = 0.250 0 −−> 1 FEM 5 CI = 1.000 0 −−> 1 ANOCLE 2 CI = 1.000 0 −−> 1 CLE 12 CI = 1.000 0 ==> 1 SCACOR 18 CI = 1.000 1 −−> 0 SCACOR 20 CI = 0.500 0 ==> 1 HUM 23 CI = 0.500 0 −−> 1 DIG 1 CI = 1.000 0 −−> 1

Ichthyostegamorphs more derived than Ventastega ANT VAC 2 CI = 0.250 0 −−> 1 PSYM 2 CI = 0.333 0 ==> 1 POSPL 3 CI = 0.500 0 −−> 1 SURANG 3 CI = 1.000 0 ==> 1 ANT COR 2 CI = 0.250 0 ==> 1 MID COR 2 CI = 0.333 0 ==> 1

396 K. PAWLEY PHD THESIS

Ichthyostegamorphs more derived than Archeriamorphs more derived than Acanthostega Tulerpeton PREMAX 9 CI = 0.500 0 ==> 1 VOM 6 CI = 0.125 1 −−> 0 QUAJUG 7 CI = 0.333 0 −−> 1 ANG 2 CI = 1.000 0 ==> 1 VOM 6 CI = 0.125 0 −−> 1 ANOCLE 1 CI = 0.333 0 ==> 1 VOM 7 CI = 0.167 0 −−> 1 RAD 2 CI = 1.000 0 ==> 1 VOM 9 CI = 0.500 0 ==> 1 DIG 2 CI = 0.750 0 −−> 1 DEN 3 CI = 0.500 1 −−> 0 DIG 5 CI = 1.000 0 ==> 1 TRU VER 35 CI = 0.333 0 −−> 1 ILI 11 CI = 0.500 0 −−> 1 RIB 3 CI = 1.000 0 ==> 1 INTCLA 2 CI = 0.200 0 −−> 1 Whatcheeriidae SCACOR 4 CI = 1.000 0 ==> 1 PREMAX 5 CI = 0.200 0 ==> 1 ULNA 1 CI = 1.000 0 ==> 1 MAX 2 CI = 0.100 0 ==> 1 ILI 6 CI = 1.000 0 ==> 1 INTEMP 3 CI = 0.333 0 ==> 1 TIB 3 CI = 0.500 0 −−> 1 SUTEMP 4 CI = 0.333 0 ==> 1 PSYM 2 CI = 0.333 1 −−> 0 Archeriamorphs CLE 4 CI = 0.167 0 ==> 1

MED ROS 1 CI = 0.500 0 −−> 1 Pederpes plus Ossinodus TEC 1 CI = 1.000 0 −−> 1 CLE 8 CI = 0.200 0 ==> 1 SPTMAX 1 CI = 0.250 0 −−> 1 Archeriamorphs more derived than LAC 2 CI = 0.143 1 −−> 0 Whatcheeriidae PAR 2 CI = 0.143 1 −−> 0 PREFRO 8 CI = 0.125 0 −−> 1 INTEMP 1 CI = 0.125 1 −−> 0 PREOPE 1 CI = 1.000 0 ==> 1 JUG 7 CI = 0.250 0 −−> 1 PSYM 3 CI = 1.000 0 −−> 1 OPI 1 CI = 0.333 0 −−> 1 PREART 3 CI = 0.333 1 −−> 0 ECT 2 CI = 0.200 1 −−> 0 ANT COR 3 CI = 0.333 0 ==> 1 INT VAC 1 CI = 0.250 0 −−> 1 ANT COR 4 CI = 1.000 0 ==> 1 PASPHE 3 CI = 0.333 0 −−> 1 MID COR 3 CI = 0.333 0 −−> 1 CAU VER 7 CI = 1.000 0 −−> 1 MID COR 4 CI = 1.000 0 ==> 1 CAU VER 10 CI = 1.000 0 −−> 1 POST COR 3 CI = 0.333 0 −−> 1 RIB 5 CI = 0.250 0 −−> 1 POST COR 4 CI = 0.333 0 −−> 1 CLE 3 CI = 1.000 0 ==> 1 RIB 10 CI = 0.333 1 −−> 0 CLE 11 CI = 1.000 0 ==> 1 INTCLA 2 CI = 0.200 1 −−> 0 SCACOR 19 CI = 1.000 0 −−> 1 HUM 11 CI = 1.000 0 −−> 1 SCACOR 20 CI = 0.500 1 ==> 0 HUM 6 CI = 0.500 0 ==> 1 HUM 9 CI = 1.000 0 ==> 1 HUM 20 CI = 1.000 0 −−> 1 FEM 5 CI = 1.000 1 ==> 2 FIB 1 CI = 1.000 0 ==> 1 TAR 4 CI = 1.000 0 ==> 1 TAR 5 CI = 1.000 0 ==> 1

APPENDIX 15: EARLY TETRAPOD SYNAPOMORPHIES 397

Archeriamorphs more derived than Baphetidae Crassigyrinus MED ROS 1 CI = 0.500 1 ==> 0 TAB 3 CI = 0.250 0 −−> 1 PREFRO 7 CI = 1.000 0 ==> 1 JUG 7 CI = 0.250 1 −−> 0 LAC 5 CI = 1.000 0 ==> 1 VOM 6 CI = 0.125 0 −−> 1 SUTEMP 2 CI = 1.000 0 ==> 1 VOM 7 CI = 0.167 1 −−> 0 SUTEMP 4 CI = 0.333 0 ==> 1 VOM 8 CI = 1.000 0 ==> 1 JUG 7 CI = 0.250 0 −−> 1 VOM 14 CI = 0.200 0 ==> 1 INT VAC 1 CI = 0.250 1 −−> 0 PASPHE 9 CI = 0.667 1 ==> 2 ANT COR 3 CI = 0.333 1 ==> 0 BASOCC 1 CI = 0.500 0 ==> 1 MID COR 3 CI = 0.333 1 −−> 0 PREART 2 CI = 1.000 0 ==> 1 POST COR 3 CI = 0.333 1 −−> 0 PREART 4 CI = 0.500 0 ==> 1 ADD FOS 1 CI = 1.000 0 ==> 1 Archeriamorphs more derived than ILI 4 CI = 1.000 0 −−> 1 Baphetidae ILI 11 CI = 0.500 1 −−> 0 PAR 1 CI = 0.500 0 −−> 1 TAB 3 CI = 0.333 1 −−> 0 Colosteidae PTE 24 CI = 0.200 0 ==> 1 PREMAX 5 CI = 0.200 0 ==> 1 PSYM 2 CI = 0.333 1 −−> 0 SPTMAX 1 CI = 0.250 1 −−> 0 POST COR 5 CI = 0.200 0 ==> 1 PREFRO 6 CI = 1.000 0 ==> 1 HUM 1 CI = 1.000 0 ==> 1 PREFRO 9 CI = 0.500 0 ==> 1 HUM 17 CI = 1.000 0 ==> 1 PAR 2 CI = 0.143 0 ==> 1 ILI 7 CI = 0.500 0 ==> 1 INTEMP 1 CI = 0.125 0 ==> 1 TIB 1 CI = 1.000 0 ==> 1 VOM 16 CI = 0.200 0 −−> 1 PTE 16 CI = 1.000 0 ==> 1 Anthracosauria DEN 4 CI = 1.000 0 ==> 1 SPTMAX 1 CI = 0.250 1 −−> 0 INTCLA 1 CI = 0.125 1 ==> 0 INTEMP 2 CI = 0.333 1 −−> 0 CLA 3 CI = 0.250 0 ==> 1 JUG 3 CI = 0.200 1 −−> 0 SCACOR 1 CI = 0.333 0 −−> 1 PTF 1 CI = 1.000 0 −−> 2 ILI 2 CI = 0.250 0 −−> 1 ECT 5 CI = 0.125 1 −−> 0 PREART 3 CI = 0.333 0 ==> 1 Archeriamorphs more derived than CLA 7 CI = 0.250 0 −−> 1 Colosteidae SCACOR 10 CI = 1.000 0 −−> 1 PREMAX 9 CI = 0.500 1 ==> 0 HUM 27 CI = 1.000 0 −−> 1 PREFRO 8 CI = 0.125 1 −−> 0 FIB 7 CI = 0.500 0 −−> 1 VOM 4 CI = 0.111 0 ==> 1 PAL 2 CI = 0.100 0 ==> 1 Anthracosauria more derived than PAL 4 CI = 0.143 0 ==> 1 Gephyrostegus ECT 5 CI = 0.125 0 −−> 1 TAB 2 CI = 0.250 0 ==> 1 ANT VAC 1 CI = 0.143 0 −−> 1 JUG 2 CI = 0.167 0 −−> 1 SPL 4 CI = 0.333 0 ==> 1 VOM 1 CI = 0.333 0 ==> 1 POSPL 2 CI = 1.000 0 ==> 1 VOM 3 CI = 0.250 0 ==> 1 TRU VER 8 CI = 0.200 0 −−> 1 VOM 4 CI = 0.111 1 ==> 0 RIB 8 CI = 0.250 1 −−> 0 PAL 2 CI = 0.100 1 ==> 0 CLA 9 CI = 0.500 0 −−> 1

398 K. PAWLEY PHD THESIS

Anthracosauridae Terrapoda SC SK 1 CI = 0.111 0 ==> 1 POST COR 4 CI = 0.500 1 ==> 0 SC SK 1 CI = 0.111 0 ==> 1 POST COR 5 CI = 0.200 1 −−> 0 SC SK 2 CI = 1.000 0 ==> 1 POSTPAR 5 CI = 0.500 0 ==> 1 Anthracosauria more derived than SQU 3 CI = 0.250 0 −−> 1 Anthracosauridae OPI 1 CI = 0.333 1 ==> 0 TEETH 6 CI = 1.000 0 ==> 1 STAP 1 CI = 1.000 0 ==> 1 SCACOR 2 CI = 0.500 1 ==> 0 PTE 9 CI = 0.167 0 ==> 1 PASPHE 3 CI = 0.333 1 ==> 0 Embolomeri PSYM 1 CI = 1.000 0 ==> 1 PASPHE 5 CI = 0.286 0 ==> 2 TRU VER 28 CI = 1.000 0 ==> 1 TRU VER 9 CI = 0.250 0 ==> 1 TRU VER 35 CI = 0.333 1 −−> 0 TRU VER 14 CI = 0.333 0 ==> 1 CER VER 5 CI = 0.500 0 ==> 1 RIB 1 CI = 0.333 0 −−> 1 Pholiderpeton spp. RIB 5 CI = 0.250 1 ==> 0 MAX 1 CI = 0.167 0 ==> 1 RIB 11 CI = 1.000 0 ==> 1 TAB 15 CI = 0.500 0 ==> 1 RIB 12 CI = 1.000 0 ==> 1 JUG 7 CI = 0.200 0 ==> 1 CLA 6 CI = 0.500 0 ==> 1 ANOCLE 1 CI = 0.333 1 −−> 0 CLE 2 CI = 1.000 0 ==> 1 CLE 8 CI = 0.200 0 −−> 1 CLE 9 CI = 1.000 0 ==> 1 CLE 10 CI = 1.000 1 ==> 0 SCACOR 6 CI = 0.333 0 ==> 1 SCACOR 7 CI = 1.000 0 ==> 1 SCACOR 8 CI = 0.500 0 ==> 1 SCACOR 11 CI = 1.000 0 ==> 1 SCACOR 12 CI = 0.500 0 ==> 1 SCACOR 13 CI = 0.500 0 ==> 1 SCACOR 14 CI = 1.000 0 ==> 1 SCACOR 15 CI = 0.500 0 ==> 1 SCACOR 17 CI = 0.500 0 ==> 1 HUM 2 CI = 0.200 0 ==> 1 HUM 9 CI = 1.000 1 ==> 2 HUM 10 CI = 1.000 0 ==> 1 HUM 19 CI = 1.000 0 ==> 1 HUM 22 CI = 1.000 0 ==> 1 HUM 23 CI = 0.500 1 ==> 0 HUM 24 CI = 1.000 0 ==> 1 HUM 26 CI = 1.000 0 ==> 1 RAD 3 CI = 1.000 0 ==> 1 RAD 4 CI = 1.000 0 ==> 1 ILI 5 CI = 1.000 0 ==> 1 PUB 1 CI = 1.000 0 ==> 1 FEM 7 CI = 1.000 0 ==> 1 FEM 10 CI = 1.000 0 ==> 1 TIB 2 CI = 1.000 0 ==> 1 TIB 3 CI = 0.500 1 ==> 0 TIB 4 CI = 1.000 0 ==> 1 FIB 2 CI = 1.000 0 ==> 1 APPENDIX 15: EARLY TETRAPOD SYNAPOMORPHIES 399

Temnospondyli Cochleosauridae more derived than Saharastega LAC 2 CI = 0.143 0 −−> 1 L SC SKU 1 CI = 0.500 0 ==> 1 PAR 1 CI = 0.500 1 −−> 0 PREMAX 4 CI = 0.500 0 ==> 1 SUTEMP 4 CI = 0.333 0 ==> 1 CHO 1 CI = 1.000 0 ==> 1 TAB 5 CI = 0.333 0 −−> 1 VOM 16 CI = 0.200 0 ==> 1 Chenoprosopus spp. PTE 25 CI = 0.500 0 −−> 1 ANT FOS 1 CI = 0.200 0 ==> 1 MAND 2 CI = 1.000 0 −−> 1 SPL 4 CI = 0.333 1 ==> 0 Nigerpeton plus Adamanterpeton plus RIB 8 CI = 0.250 0 −−> 1 Cochleosaurus QUAJUG 7 CI = 0.333 1 ==> 0 INTCLA 1 CI = 0.125 1 −−> 0 VOM 7 CI = 0.167 0 ==> 1 CLE 4 CI = 0.167 0 ==> 1 SPL 1 CI = 0.167 0 ==> 1 ILI 3 CI = 0.333 0 ==> 1

Adamanterpeton plus Cochleosaurus Temnospondyli more derived than MAX 1 CI = 0.167 1 ==> 0 Caerorhachis plus Casineria TRU VER 8 CI = 0.200 1 ==> 0 POSPAR 11 CI = 1.000 0 −−> 1 VOM 17 CI = 0.500 0 ==> 1 HUM 5 CI = 0.143 0 −−> 1

DIG 2 CI = 0.750 1 −−> 2 Cochleosaurus spp. TAR 3 CI = 0.500 1 −−> 0 ANT FOS 1 CI = 0.200 0 ==> 1 PASPHE 12 CI = 0.250 0 ==> 1 Temnospondyli more derived than Edops PTE 10 CI = 0.143 1 ==> 0 Temnospondyli more derived than PTE 13 CI = 0.333 0 ==> 1 Cochleosauridae INT VAC 2 CI = 0.333 0 ==> 1 VOM 5 CI = 0.333 0 ==> 1 INT VAC 3 CI = 0.333 0 ==> 1 VOM 7 CI = 0.167 0 −−> 1

VOM 16 CI = 0.200 1 −−> 0 Dendrerpetontidae PAL 4 CI = 0.143 1 ==> 0 PREMAX 1 CI = 0.250 0 ==> 1 ECT 5 CI = 0.125 1 ==> 0 LAC 2 CI = 0.143 1 −−> 0 PTE 24 CI = 0.200 1 −−> 0 PASPHE 12 CI = 0.250 0 −−> 1 PTE 25 CI = 0.500 1 −−> 0 SPL 1 CI = 0.167 0 −−> 1 MAND 1 CI = 0.143 0 ==> 1 HUM 5 CI = 0.143 1 −−> 0

Balanerpeton plus Dendrerpeton VOM 5 CI = 0.333 0 ==> 1

Temnospondyli more derived than Dendrerpetontidae MAX 1 CI = 0.167 0 ==> 1 PASPHE 7 CI = 0.500 0 −−> 1 ILI 2 CI = 0.250 0 ==> 1

Cochleosauridae PIN FOR 1 CI = 0.200 0 ==> 1 JUG 2 CI = 0.167 0 −−> 1 JUG 3 CI = 0.200 0 ==> 1

400 K. PAWLEY PHD THESIS

Dvinosauria Dvinosaurus plus Tupilakosauridae SC SK 1 CI = 0.111 1 ==> 0 PREMAX 6 CI = 0.333 0 −−> 1 LAC 2 CI = 0.143 1 −−> 0 PAR 2 CI = 0.143 0 ==> 1 SQU 3 CI = 0.250 1 ==> 0 PTE 22 CI = 0.167 0 ==> 1 VOM 4 CI = 0.111 1 ==> 0 MAND 3 CI = 1.000 0 ==> 1 VOM 13 CI = 0.250 0 −−> 1 INTCLA 1 CI = 0.125 0 ==> 1 PAL 2 CI = 0.100 1 ==> 0 PTE 9 CI = 0.167 1 ==> 0 Tupilakosauridae PTE 13 CI = 0.333 1 −−> 0 PTE 13 CI = 0.333 1 ==> 2 PTE 23 CI = 1.000 0 −−> 1 PTE 20 CI = 0.250 0 ==> 1 ANT VAC 1 CI = 0.143 1 −−> 0 INT VAC 3 CI = 0.333 1 ==> 0 PASPHE 5 CI = 0.286 0 ==> 2 EXOCC 1 CI = 0.333 0 −−> 1 PASPHE 7 CI = 0.500 1 −−> 0 ANT COR 2 CI = 0.250 1 −−> 0 TRU VER 35 CI = 0.333 0 −−> 1 MID COR 2 CI = 0.333 1 −−> 0 CLA 5 CI = 0.333 0 −−> 1 POST COR 2 CI = 0.500 1 −−> 0 CLA 7 CI = 0.250 0 −−> 1 TRU VER 8 CI = 0.200 0 ==> 1 SCACOR 9 CI = 1.000 0 −−> 1 TRU VER 9 CI = 0.250 0 ==> 1 FEM 6 CI = 1.000 0 ==> 1 TRU VER 14 CI = 0.333 0 ==> 1

Dvinosauria more derived than Peltobatrachus plus Eryopoidomorpha Neldasaurus POSTFRO 2 CI = 0.250 0 ==> 1 MAX 1 CI = 0.167 1 ==> 0 INTEMP 1 CI = 0.125 0 ==> 1 VOM 2 CI = 1.000 0 ==> 1 VOM 15 CI = 0.333 0 −−> 1 VOM 6 CI = 0.125 1 −−> 0 ANT FOS 1 CI = 0.200 0 −−> 1 VOM 14 CI = 0.200 1 −−> 0 EXOCC 3 CI = 1.000 0 ==> 1 PREART 5 CI = 0.500 0 −−> 1 Dvinosauroidea RIB 5 CI = 0.250 0 ==> 1 INTEMP 1 CI = 0.125 0 −−> 1 CLE 8 CI = 0.200 0 ==> 1 TAB 9 CI = 0.500 0 ==> 1 ULNA 2 CI = 1.000 0 −−> 1 JUG 3 CI = 0.200 0 ==> 1 ILI 8 CI = 1.000 0 ==> 1 VOM 10 CI = 0.200 0 −−> 1 FIB 6 CI = 1.000 0 −−> 1 PAL 3 CI = 0.200 0 −−> 1 PAL 4 CI = 0.143 0 ==> 1 Eryopoidomorpha PTE 13 CI = 0.333 0 −−> 1 MAND 1 CI = 0.143 1 −−> 0 PTE 14 CI = 1.000 0 ==> 1 FEM 9 CI = 1.000 0 ==> 1 PTE 18 CI = 0.667 0 ==> 2 Euskelia PREMAX 1 CI = 0.250 0 ==> 1 Eobrachyopidae QUAJUG 7 CI = 0.333 1 −−> 0 PREFRO 8 CI = 0.125 0 ==> 1 VOM 7 CI = 0.167 1 −−> 0 JUG 2 CI = 0.167 0 ==> 1 VOM 16 CI = 0.200 0 −−> 1 VOM 6 CI = 0.125 0 −−> 1 PREART 5 CI = 0.500 0 ==> 1 VOM 14 CI = 0.200 0 −−> 1 CLA 8 CI = 1.000 0 −−> 1 PAL 5 CI = 0.500 0 ==> 1 HUM 21 CI = 1.000 0 ==> 1 ECT 5 CI = 0.125 0 −−> 1 FIB 5 CI = 1.000 0 ==> 1 PASPHE 8 CI = 1.000 0 ==> 1

APPENDIX 15: EARLY TETRAPOD SYNAPOMORPHIES 401

Euskelia more derived than Amphibamidae Sclerocephalus MAX 5 CI = 0.250 0 −−> 1 SPTMAX 2 CI = 0.333 0 ==> 1 VOM 10 CI = 0.200 0 ==> 1 MAX 1 CI = 0.167 1 ==> 0 VOM 15 CI = 0.333 1 −−> 0 PAL 4 CI = 0.143 0 ==> 1 PAL 3 CI = 0.200 0 ==> 1 ECT 5 CI = 0.125 0 ==> 1 ANT FOS 1 CI = 0.200 1 ==> 0 INTCLA 7 CI = 0.500 0 −−> 1 PASPHE 11 CI = 1.000 0 ==> 1 PASPHE 12 CI = 0.250 0 ==> 1 Dissorophoidea DEN 2 CI = 0.500 0 −−> 1 NAS 3 CI = 1.000 0 ==> 1 RIB 5 CI = 0.250 1 ==> 0 PREFRO 1 CI = 1.000 0 ==> 1 INTCLA 7 CI = 0.500 1 −−> 0 LAC 2 CI = 0.143 1 ==> 0 CLE 4 CI = 0.167 1 −−> 0 FRO 4 CI = 0.167 0 ==> 1 SQU 9 CI = 0.500 0 −−> 1 Doleserpeton plus Amphibamus QUAJUG 6 CI = 1.000 0 ==> 1 MAX 8 CI = 1.000 0 ==> 1 QUA 1 CI = 1.000 0 ==> 1 SQU 9 CI = 0.500 1 −−> 0 PAL 5 CI = 0.500 0 ==> 1 VOM 3 CI = 0.250 0 ==> 1 PASPHE 7 CI = 0.500 1 ==> 0 PAL 1 CI = 0.250 0 ==> 1 EXOCC 5 CI = 1.000 0 ==> 1 PTE 15 CI = 1.000 0 ==> 1 CLE 5 CI = 0.500 0 −−> 1 PTE 20 CI = 0.250 0 ==> 1 HUM 2 CI = 0.200 1 −−> 0 TEETH 1 CI = 1.000 0 ==> 1 TEETH 2 CI = 0.500 0 ==> 1 Dissorophidae TEETH 3 CI = 1.000 0 ==> 1 TAB 14 CI = 0.500 0 ==> 1 TRU VER 29 CI = 0.333 0 −−> 1 SQU 8 CI = 1.000 0 ==> 1 TRU VER 8 CI = 0.200 0 ==> 1 PASPHE 5 CI = 0.286 0 −−> 2 HUM 12 CI = 0.500 0 ==> 1 DERM 1 CI = 0.500 0 ==> 1 Stereospondylomorpha Dissorophidae more derived than SC SK 1 CI = 0.111 1 ==> 0 Dissorophus TAB 2 CI = 0.250 0 −−> 1 NOS 3 CI = 1.000 0 ==> 1 SQU 6 CI = 1.000 0 ==> 1 PREFRO 8 CI = 0.125 0 ==> 1 ECT 2 CI = 0.200 0 −−> 1 HUM 2 CI = 0.200 0 −−> 1 CER VER 2 CI = 0.333 0 ==> 1 DERM 3 CI = 1.000 0 ==> 1 Acheloma plus Phonerpeton PREMAX 5 CI = 0.200 0 ==> 1 Archegosauridae MAX 2 CI = 0.100 0 ==> 1 SKU 1 CI = 1.000 0 ==> 1 PREMAX 6 CI = 0.333 0 ==> 1 VOM 15 CI = 0.333 1 −−> 0 PASPHE 12 CI = 0.250 0 ==> 1 PREART 1 CI = 0.500 0 ==> 1

402 K. PAWLEY PHD THESIS

Stereospondyli Mastodonsauroidea TAB 12 CI = 1.000 0 ==> 1 FRO 4 CI = 0.167 0 ==> 1 SQU 6 CI = 1.000 1 ==> 2 TAB 13 CI = 1.000 0 ==> 1 JUG 2 CI = 0.167 0 −−> 1 PASPHE 5 CI = 0.286 1 ==> 2 JUG 3 CI = 0.200 0 ==> 1 PREART 6 CI = 1.000 0 ==> 1 VOM 6 CI = 0.125 1 ==> 0 VOM 10 CI = 0.200 0 ==> 1 Neospondyli VOM 14 CI = 0.200 1 ==> 0 PTE 2 CI = 1.000 0 ==> 1 SPTMAX 2 CI = 0.333 0 −−> 1 PTE 9 CI = 0.167 1 ==> 0 INTEMP 3 CI = 0.333 0 −−> 1 PTE 13 CI = 0.333 1 ==> 2 JUG 2 CI = 0.167 0 ==> 1 ANT VAC 2 CI = 0.250 1 −−> 0 VOM 1 CI = 0.333 0 ==> 1 PASPHE 4 CI = 1.000 0 ==> 1 ECT 2 CI = 0.200 0 ==> 1 PASPHE 5 CI = 0.286 0 ==> 1 PTE 3 CI = 0.500 0 −−> 1 CLE 7 CI = 0.500 0 ==> 1 DEN 2 CI = 0.500 0 ==> 1 HUM 25 CI = 0.500 0 −−> 1 ANG 4 CI = 1.000 0 ==> 1 TRU VER 10 CI = 0.333 0 −−> 1 Lydekkerina plus Superstereospondyli TRU VER 11 CI = 0.500 0 ==> 1 SC 3 CI = 1.000 0 ==> 1 TRU VER 29 CI = 0.333 0 ==> 1 TAB 2 CI = 0.250 1 −−> 0 TRU VER 33 CI = 0.500 0 ==> 1 TAB 11 CI = 1.000 0 −−> 1 TRU VER 9 CI = 0.250 0 ==> 1 PAL 3 CI = 0.200 0 ==> 1 INTCLA 2 CI = 0.200 0 ==> 1 PTE 20 CI = 0.250 0 ==> 1 CLA 3 CI = 0.250 0 ==> 1 PTE 21 CI = 0.333 0 ==> 1 CLE 7 CI = 0.500 0 ==> 1 ANT VAC 1 CI = 0.143 1 ==> 0 ILI 1 CI = 0.333 0 −−> 1 EXOCC 1 CI = 0.333 0 ==> 1 EXOCC 3 CI = 1.000 1 ==> 2 Seymouriamorpha MAND 1 CI = 0.143 1 ==> 0 INT VAC 1 CI = 0.250 1 ==> 0 PREART 5 CI = 0.500 1 ==> 0 POST COR 5 CI = 0.200 1 ==> 0 Kotlassia plus Discosauriscus CLA 7 CI = 0.250 0 ==> 1 SC SK 1 CI = 0.111 1 ==> 0

Superstereospondyli Neospondyli more derived than VOM 4 CI = 0.111 1 ==> 0 Seymouriamorpha PAL 2 CI = 0.100 1 ==> 0 PAR 2 CI = 0.143 0 ==> 1 PAL 8 CI = 1.000 0 ==> 1 INTEMP 1 CI = 0.125 0 ==> 1 PTE 1 CI = 1.000 0 ==> 1 SQU 3 CI = 0.250 1 −−> 0 EPI 1 CI = 1.000 0 ==> 1 STAP 1 CI = 1.000 1 ==> 2 PASPHE 7 CI = 0.500 1 ==> 2 VOM 3 CI = 0.250 0 ==> 1 ANT COR 3 CI = 0.333 1 ==> 0 PAL 1 CI = 0.250 0 ==> 1 MID COR 3 CI = 0.333 1 ==> 0 PTE 4 CI = 0.500 0 −−> 1 CER VER 1 CI = 0.500 0 −−> 1 POSPL 1 CI = 0.250 0 ==> 1 CER VER 10 CI = 0.500 0 −−> 1 ANT COR 1 CI = 0.500 0 −−> 1 INTCLA 9 CI = 1.000 0 ==> 1 TRU VER 31 CI = 0.333 0 −−> 1 CLA 3 CI = 0.250 0 ==> 1 CER VER 8 CI = 1.000 0 −−> 1 CLA 5 CI = 0.333 0 ==> 1 RIB 1 CI = 0.333 1 −−> 0 SCACOR 5 CI = 1.000 0 ==> 1 TAR 6 CI = 1.000 0 ==> 1

APPENDIX 15: EARLY TETRAPOD SYNAPOMORPHIES 403

Cotylosauria Neospondyli more derived than POSTPAR 2 CI = 0.333 0 −−> 1 Cotylosauria POSTPAR 5 CI = 0.500 1 ==> 0 SPTMAX 2 CI = 0.333 1 −−> 0 POSTPAR 6 CI = 0.500 0 ==> 1 ECT 5 CI = 0.125 1 −−> 0 PTF 1 CI = 1.000 0 ==> 1 PTE 3 CI = 0.500 1 −−> 0 OPI 1 CI = 0.333 0 ==> 1 PTE 9 CI = 0.167 1 ==> 0 PASPHE 3 CI = 0.333 0 ==> 1 TRU VER 10 CI = 0.333 1 −−> 0 CLE 8 CI = 0.200 0 −−> 1 CER VER 1 CI = 0.500 0 −−> 1 CER VER 2 CI = 0.333 0 −−> 1 Diadectomorpha plus Captorhinomorpha CER VER 6 CI = 1.000 0 −−> 1 PREMAX 10 CI = 1.000 0 ==> 1 CER VER 7 CI = 0.500 0 −−> 1 MID COR 1 CI = 0.333 0 ==> 1 SCACOR 1 CI = 0.333 0 −−> 1 POST COR 3 CI = 0.333 1 −−> 0 SCACOR 6 CI = 0.333 1 −−> 0 SAC VER 1 CI = 0.400 1 ==> 2 DIG 2 CI = 0.750 1 −−> 2

Diadectomorpha ILI 1 CI = 0.333 1 −−> 0 SQU 3 CI = 0.250 0 ==> 1 ILI 7 CI = 0.500 1 −−> 0 PTE 4 CI = 0.500 1 −−> 0 Lepospondyli TRU VER 31 CI = 0.333 1 −−> 0

Diadectes spp. SUTEMP 1 CI = 0.500 0 −−> 1 CLE 4 CI = 0.167 0 ==> 1 SUTEMP 3 CI = 1.000 0 −−> 1 PAL 4 CI = 0.143 1 −−> 0 Captorhinomorpha TRU VER 13 CI = 0.333 0 ==> 1 MAX 2 CI = 0.100 0 ==> 1 HUM 3 CI = 0.500 0 ==> 1 FRO 4 CI = 0.167 0 ==> 1 ILI 2 CI = 0.250 0 ==> 1 PAR 7 CI = 0.500 0 ==> 1 TAR 3 CI = 0.500 1 ==> 0 POSTPAR 2 CI = 0.333 1 −−> 0 TAB 9 CI = 0.500 0 −−> 1 Microsauria SCACOR 3 CI = 0.500 0 ==> 1 EXOCC 4 CI = 1.000 0 ==> 1 SCACOR 16 CI = 1.000 0 ==> 1 CER VER 5 CI = 0.500 1 ==> 0 HUM 3 CI = 0.500 0 ==> 1 INTCLA 5 CI = 0.333 1 ==> 0 HUM 25 CI = 0.500 0 ==> 1 CLE 4 CI = 0.167 0 ==> 1 HUM 2 CI = 0.200 1 ==> 0 ILI 1 CI = 0.333 1 −−> 0 HUM 7 CI = 1.000 0 ==> 1 ILI 3 CI = 0.333 0 ==> 1

TIB 7 CI = 0.500 0 ==> 1 Microbrachomorpha FIB 7 CI = 0.500 0 ==> 1 PAL 4 CI = 0.143 0 −−> 1 Paleothyris plus Petrolacosaurus SPL 1 CI = 0.167 0 ==> 1 VOM 13 CI = 0.250 0 ==> 1 POSPL 1 CI = 0.250 1 −−> 0 BASOCC 5 CI = 1.000 0 ==> 1 DIG 2 CI = 0.750 2 ==> 3 SPL 1 CI = 0.167 0 −−> 1 TRU VER 10 CI = 0.333 1 ==> 0 Tuditanomorpha CER VER 4 CI = 1.000 0 ==> 1 PREMAX 8 CI = 0.500 0 −−> 1 SCACOR 6 CI = 0.333 1 ==> 0 LAC 4 CI = 0.333 0 −−> 1 HUM 12 CI = 0.500 0 ==> 1 POSORB 5 CI = 0.500 0 ==> 1 POSORB 8 CI = 0.500 0 ==> 1 SCACOR 17 CI = 0.500 1 ==> 0

404 K. PAWLEY PHD THESIS

Tuditanomorpha more derived than Tuditanidae plus Hapsidopareiontidae Batropetes PREMAX 8 CI = 0.500 1 −−> 0 PAR 2 CI = 0.143 1 ==> 0 POSTPAR 6 CI = 0.500 0 ==> 1 POSTFRO 3 CI = 0.500 0 ==> 1 PTE 9 CI = 0.167 0 ==> 1 ILI 2 CI = 0.250 1 −−> 0 Pantylidae plus Ostodolepidae plus Rhynchonkos plus Gymnarthridae Hapsidopareiontidae POSPL 1 CI = 0.250 1 ==> 0 PTE 17 CI = 0.500 0 ==> 1 SCACOR 3 CI = 0.500 0 −−> 1 SCACOR 12 CI = 0.500 1 −−> 0 Nectridia SCACOR 13 CI = 0.500 1 −−> 0 SPTMAX 1 CI = 0.250 1 ==> 0 PREFRO 8 CI = 0.125 0 ==> 1 Pantylidae VOM 1 CI = 0.333 1 ==> 0 MAX 2 CI = 0.100 0 ==> 1 VOM 4 CI = 0.111 1 ==> 0 MAX 7 CI = 1.000 0 ==> 1 VOM 6 CI = 0.125 1 ==> 0 JUG 6 CI = 1.000 0 ==> 1 PAL 2 CI = 0.100 1 ==> 0 ECT 1 CI = 0.250 0 ==> 1 PTE 24 CI = 0.200 1 −−> 0 MID COR 1 CI = 0.333 0 −−> 1 PASPHE 5 CI = 0.286 0 ==> 2 TEETH 8 CI = 0.500 0 ==> 1 MAND 1 CI = 0.143 0 ==> 1 MID COR 1 CI = 0.333 0 −−> 1 Ostodolepidae plus Rhynchonkos plus POST COR 1 CI = 0.500 0 ==> 1 Gymnarthridae TRU VER 1 CI = 1.000 0 ==> 1 POSTPAR 7 CI = 1.000 0 ==> 1 TRU VER 2 CI = 1.000 0 ==> 1 POSTPAR 8 CI = 0.500 0 −−> 1 TRU VER 29 CI = 0.333 1 ==> 0 PTE 17 CI = 0.500 0 ==> 1 TRU VER 30 CI = 0.500 0 ==> 1 MAND 1 CI = 0.143 0 −−> 1 TRU VER 31 CI = 0.333 1 −−> 0 SPL 1 CI = 0.167 0 −−> 1 CER VER 10 CI = 0.500 0 ==> 1 TRU VER 13 CI = 0.333 1 ==> 0 CAU VER 1 CI = 1.000 0 ==> 1 SAC VER 1 CI = 0.400 1 −−> 2 CAU VER 9 CI = 1.000 0 ==> 1 RIB 1 CI = 0.333 0 −−> 1 Ostodolepidae INTCLA 1 CI = 0.125 1 ==> 0 PIN FOR 1 CI = 0.200 0 ==> 1 INTCLA 2 CI = 0.200 1 −−> 0 TAB 5 CI = 0.333 0 ==> 1 INTCLA 8 CI = 1.000 0 −−> 1 POSORB 8 CI = 0.500 1 ==> 0 CLA 6 CI = 0.500 1 ==> 0 JUG 5 CI = 0.333 0 ==> 1 CLA 7 CI = 0.250 0 −−> 1 JUG 9 CI = 0.500 0 ==> 1 HUM 5 CI = 0.143 0 −−> 1

Rhynchonkos plus Gymnarthridae Diplocaulidae QUAJUG 3 CI = 1.000 0 ==> 1 SC SK 1 CI = 0.111 1 ==> 0 VOM 14 CI = 0.200 1 ==> 0 TAB 5 CI = 0.333 0 −−> 1 HUM 5 CI = 0.143 0 ==> 1 SQU 5 CI = 1.000 0 ==> 1 PAL 3 CI = 0.200 0 ==> 1 Gymnarthridae ECT 1 CI = 0.250 0 ==> 1 POSTPAR 8 CI = 0.500 1 −−> 0 PTE 7 CI = 1.000 0 ==> 1 PTE 9 CI = 0.167 0 ==> 1 PTE 10 CI = 0.143 1 −−> 0 MAND 1 CI = 0.143 1 −−> 0 PTE 20 CI = 0.250 0 −−> 1 SURANG 5 CI = 0.500 0 ==> 1 EXOCC 2 CI = 1.000 0 ==> 1 POST COR 1 CI = 0.500 0 ==> 1 SURANG 5 CI = 0.500 0 −−> 1 TEETH 8 CI = 0.500 0 ==> 1 APPENDIX 15: EARLY TETRAPOD SYNAPOMORPHIES 405

Diplocaulidae more derived than Sauropleura plus Urocordylus Scinosaurus JUG 2 CI = 0.167 1 ==> 0 PAR 7 CI = 0.500 0 ==> 1 CER VER 7 CI = 0.500 1 −−> 0 TAB 6 CI = 1.000 0 ==> 1 PTE 18 CI = 0.667 0 ==> 1 EXOCC 1 CI = 0.333 0 −−> 1 CLE 1 CI = 1.000 0 ==> 1 CLE 4 CI = 0.167 0 −−> 1 CLE 5 CI = 0.500 0 −−> 1

Diplocaulidae more derived than Batrachiderpeton VOM 5 CI = 0.333 0 −−> 1 VOM 10 CI = 0.200 0 −−> 1 CAU VER 3 CI = 1.000 0 ==> 1

Diplocaulidae more derived than Keraterpeton PTE 13 CI = 0.333 0 ==> 2 PTE 22 CI = 0.167 0 ==> 1 INT VAC 2 CI = 0.333 0 ==> 1 INT VAC 3 CI = 0.333 0 ==> 1 TRU VER 30 CI = 0.500 1 ==> 0 TRU VER 18 CI = 1.000 0 ==> 1 CAU VER 4 CI = 1.000 0 ==> 1

Diploceraspis plus Diplocaulus NAS 1 CI = 0.500 1 ==> 0 PREFRO 9 CI = 0.500 0 ==> 1 LAC 2 CI = 0.143 0 ==> 1 FRO 1 CI = 0.333 1 ==> 0 FRO 4 CI = 0.167 0 ==> 1 POSTPAR 9 CI = 1.000 0 ==> 1 POSORB 3 CI = 0.500 0 ==> 1 TRU VER 6 CI = 1.000 0 ==> 1 TRU VER 12 CI = 1.000 0 ==> 1 INTCLA 1 CI = 0.125 0 ==> 1 HUM 5 CI = 0.143 1 −−> 0

Urocordylidae LAC 4 CI = 0.333 0 ==> 1 SUTEMP 1 CI = 0.500 1 −−> 0 POSORB 2 CI = 1.000 0 ==> 1 CAU VER 2 CI = 1.000 0 ==> 1 RIB 2 CI = 1.000 0 −−> 1 CLA 3 CI = 0.250 1 ==> 0