Geological Society, London, Special Publications

The Triassic timescale based on nonmarine tetrapod biostratigraphy and biochronology

Spencer G. Lucas

Geological Society, London, Special Publications 2010; v. 334; p. 447-500 doi:10.1144/SP334.15

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© 2010 Geological Society of London The Triassic timescale based on nonmarine tetrapod biostratigraphy and biochronology

SPENCER G. LUCAS New Mexico Museum of Natural History and Science, 1801 Mountain Road NW, Albuquerque, NM 87104-1375 USA (e-mail: [email protected])

Abstract: The Triassic timescale based on nonmarine tetrapod biostratigraphy and biochronology divides Triassic time into eight land-vertebrate faunachrons (LVFs) with boundaries defined by the first appearance datums (FADs) of tetrapod genera or, in two cases, the FADs of a tetrapod species. Definition and characterization of these LVFs is updated here as follows: the beginning of the Lootsbergian LVF ¼ FAD of Lystrosaurus; the beginning of the Nonesian ¼ FAD Cynognathus; the beginning of the Perovkan LVF ¼ FAD Eocyclotosaurus; the beginning of the Berdyankian LVF ¼ FAD Mastodonsaurus giganteus; the beginning of the Otischalkian LVF ¼ FAD Parasu- chus; the beginning of the Adamanian LVF ¼ FAD Rutiodon; the beginning of the Revueltian LVF ¼ FAD Typothorax coccinarum; and the beginning of the Apachean LVF ¼ FAD Redonda- saurus. The end of the Apachean (¼ beginning of the Wasonian LVF, near the beginning of the Jurassic) is the FAD of the crocodylomorph Protosuchus. The Early Triassic tetrapod LVFs, Loots- bergian and Nonesian, have characteristic tetrapod assemblages in the Karoo basin of South Africa, the Lystrosaurus assemblage zone and the lower two-thirds of the Cynognathus assemblage zone, respectively. The Middle Triassic LVFs, Perovkan and Berdyankian, have characteristic assem- blages from the Russian Ural foreland basin, the tetrapod assemblages of the Donguz and the Bukobay svitas, respectively. The Late Triassic LVFs, Otischalkian, Adamanian, Revueltian and Apachean, have characteristic assemblages in the Chinle basin of the western USA, the tetrapod assemblages of the Colorado City Formation of Texas, Blue Mesa Member of the Petrified Forest Formation in Arizona, and Bull Canyon and Redonda formations in New Mexico. Since the Triassic LVFs were introduced, several subdivisions have been proposed: Lootsbergian can be divided into three sub-LVFs, Nonesian into two, Adamanian into two and Revueltian into three. However, successful inter-regional correlation of most of these sub-LVFs remains to be demonstrated. Occasional records of nonmarine Triassic tetrapods in marine strata, palynostrati- graphy, conchostracan biostratigraphy, magnetostratigraphy and radioisotopic ages provide some basis for correlation of the LVFs to the standard global chronostratigraphic scale. These data indicate that Lootsbergian ¼ uppermost Changshingian, Induan and possibly earliest Olene- kian; Nonesian ¼ much of the Olenekian; Perovkan ¼ most of the Anisian; Berdyankian ¼ latest Anisian? and Ladinian; Otischalkian ¼ early to late Carnian; Adamanian ¼ most of the late Carnian; Revueltian ¼ early–middle Norian; and Apachean ¼ late Norian–Rhaetian. The Trias- sic timescale based on tetrapod biostratigraphy and biochronology remains a robust tool for the correlation of nonmarine Triassic tetrapod assemblages independent of the marine timescale.

Triassic tetrapod (amphibian and reptile) fossils In this paper: FAD ¼ first appearance datum; have long been used in biostratigraphy, a tradition HO ¼ highest occurrence; LO ¼ lowest occur- extending back to at least the 1870s. Lucas (1990) rence; LMA ¼ land–mammal ‘age’; LVA ¼ advocated developing a global Triassic timescale land-vertebrate ‘age’; LVF ¼ land–vertebrate fau- based on tetrapod evolution, and subsequently nachron; and SGCS ¼ standard global chronostrati- Lucas (1998a) presented a comprehensive global graphic scale (the marine timescale). Triassic tetrapod biochronology (Fig. 1). This bio- chronological timescale divides the Triassic into eight time intervals (land-vertebrate faunachrons, Previous studies LVFs) based on successive changes in faunal com- position driven by tetrapod evolution. This model Although tetrapods have been used to correlate non- has been tested and refined for more than a marine Triassic strata since the 1800s, before the decade. Here, I present the current status of the 1990s few attempts were made to establish a Triassic tetrapod-based timescale, incorporating global tetrapod biostratigraphy or biochronology new data, analyses and modifications published of the Triassic (Fig. 2). In the late 1800s, some since 1998. workers did use tetrapod fossils to correlate

From:LUCAS, S. G. (ed.) The Triassic Timescale. Geological Society, London, Special Publications, 334, 447–500. DOI: 10.1144/SP334.15 0305-8719/10/$15.00 # The Geological Society of London 2010. 448 S. G. LUCAS

J Hettangian Wassonian 201 Protosuchus Rhaetian Apachean Redondasaurus

210 Norian Revueltian

Typothorax coccinarum 220 Adamanian Late

Carnian

Rutiodon 230 Otischalkian Parasuchus TRIASSIC

Berdyankian

Ladinian Mastodonsaurus 240 giganteus

Perovkan Anisian Middle

Eocyclotosaurus 250 Olenekian Nonesian Cynognathus Early Induan Lootsbergian 252 Lystrosaurus Permian Changxingian Platbergian

Fig. 1. The Triassic timescale based on tetrapod biostratigraphy and biochronology. Restoration of Typothorax by Matt Celeskey. nonmarine Triassic strata on a broad scale, for tetrapod biostratigraphy, for the Lower Triassic of example, Cope (1875) who correlated part of the the Karoo basin in South Africa. He identified German Keuper to the Upper Triassic strata of the three successive biostratigraphic intervals, the American Southwest based on shared taxa of fossil Lystrosaurus, Procolophon and Cynognathus reptiles such as the phytosaur ‘Belodon’. ‘beds’. Watson (1914a, b) later termed these Broom (1906, 1907, 1909) introduced the ear- ‘zones’ and, since Kitching (1970), the Lystrosaurus liest, and perhaps the most influential, Triassic and Procolophon zones have been combined into a i.2. Fig.

TRIASSIC Period rvosttao-ae udvsoso rasctime. Triassic of subdivisions tetrapod-based Previous EARLY MIDDLE LATE (1966) (Argentina) Ischigualastian Puestoviejan Coloradian Chanarian Bonaparte Romer (1975) C A B Dinodontosaurus Kannemeyeria Cooper (1982) Plateosaurus Stahleckeria Tetragonias Placerias Placerias zone zone zone zone zone zone Ochev & Shishkin protero- (1989) kannemeyeroidean dinosaurian suchian epoch epoch epoch Lucas (1993a) Ningwuan lvf Jimsarian lvf Jimsarian Ordosian lvf Fuguan lvf China Lucas &Hunt(1993a) Otischalkian lvf Adamanian lvf Revueltian lvf Apachean lvf (western USA) (western (eastern North America) North (eastern Huber etal. (1993b) Conewagian lvf Neshanician lvf Economian lvf Sanfordian lvf Cliftonian lvf Lootsbergian lvf Berdyankian lvf Otischalkian lvf Adamanian lvf Revuletian lvf Apachean lvf Nonesian lvf Perovkan lvf

(1998a) Lucas

RASCTETRAPODS TRIASSIC 449 450 S. G. LUCAS single, Lystrosaurus zone (e.g. Rubidge et al. 1995; and made some necessary modifications that are Botha & Smith 2007). Recognition elsewhere of the incorporated and elaborated upon here. Lystrosaurus and/or Cynognathus ‘beds’ or ‘zones’ has long been possible in Antarctica, South America, India, China and/or Russia because Vertebrate biostratigraphy and some Early Triassic tetrapod taxa are virtually cos- biochronology mopolitan, especially the genera Lystrosaurus and Cynognathus (Lucas 1998a). The term LMA has long referred to intervals of geo- Romer (1975; also see Cox 1973) presented the logical (mostly Cenozoic) time characterized by dis- first global Triassic tetrapod biochronology, by tinctive mammalian fossil assemblages. LMAs have identifying three successive Triassic land– been defined to encompass Cenozoic time intervals vertebrate ‘faunas’: A, Early Triassic; B, Middle on most of the world’s continents (Savage & Russell Triassic; and C, Late Triassic (Fig. 2). Cosgriff 1983), and for the Late Cretaceous of western North (1984) divided Romer’s division A into A1 (¼ America (Cifelli et al. 2004). However, more Lystrosaurus biochron) and A2 (¼ Cynognathus broadly-based LVA or LVF have been introduced biochron). Ochev & Shishkin (1989; also see for parts of the Mesozoic record of Asia, South Anderson & Cruickshank 1978) recognized the America and North America (Lucas 1997b, 2008). same intervals as Romer, but chose to name them Thus, LVAs or LVFs have been proposed for the the: A, proterosuchian epoch; B, kannemeyerioi- Triassic and Jurassic of China (Lucas 1993a, 1996); dean epoch; C, and dinosaurian epoch. the Triassic of Argentina (Bonaparte 1966); the Late Cooper (1982) proposed a more detailed global Triassic of western North America (Lucas & Hunt tetrapod biostratigraphy of the Triassic than did 1993a); the Middle Triassic–Early Jurassic of east- Romer and other workers of the 1970s and 1980s ern North America (Huber et al. 1993a; Lucas & (Fig. 2). In this, he recognized a succession of six Huber 2003; Lucas & Tanner 2007a, b); the Late Triassic zones based largely on a perceived strati- Jurassic–Early Cretaceous of western North graphic succession of dicynodonts (Lucas & Wild America (Lucas 1993e); the Late Cretaceous of 1995 later presented a revized Triassic dicyno- western North America (Russell 1964, 1975; dont biozonation). Subsequent workers have not Sullivan & Lucas 2003, 2006); the Late Jurassic- adopted Cooper’s zonation. Indeed, prior to Lucas Cretaceous of Mongolia and China (Jerzykiewicz & (1998a), the concept of a global Triassic tetrapod Russell 1991; Lucas & Estep 1998; Lucas 2006a); biostratigraphy and biochronology had not pro- and the Cretaceous of Argentina (Leanza et al. gressed beyond Romer (1975). 2004). Russell (1993) proposed marine vertebrate Tetrapod-based subdivisions of Triassic time ages for the Cretaceous of western North America. have been proposed as local, provincial biochronol- Mammals are not the only tetrapods that can ogies for Argentina, North America and China. be used to recognize intervals of geologic time. In Bonaparte (1966, 1967, 1982) introduced a set of the Mesozoic, especially prior to the Late Cretac- ‘provincial ages’ for the Triassic of Argentina, but eous, when mammal fossils are very rare, non- he never defined these terms (Fig. 2). However, mammalian tetrapods can be biochronologically since then Lucas & Harris (1996) have defined the useful. For this reason, some workers use the term Chanarian as a LVF, and Langer (2005b) has LVA. Because LMAs and LVAs are not formal defined the Ischigualastian as a LVF. Lucas ages in stratigraphy, Lucas (1993a) introduced the (1993a) proposed a succession of LVFs for the term faunachron (essentially the same concept as Chinese Early–Middle Triassic tetrapod record. At Dunbar & Rodgers’ [1957] ‘faunichron’) to refer about the same time, Lucas & Hunt (1993a) pro- to the time interval that is equivalent to the duration posed Late Triassic LVFs based on the Chinle of a ‘fauna’. I, thus, use the more precise term LVF Group tetrapod record from the western United instead of LMA or LVA. States, and Huber et al. (1993b) proposed LVFs are biochronological units, and I define Middle–Late Triassic LVFs based on the Newark their beginnings by biochronological events. Each Supergroup record of eastern North America LVF begins with the FAD of a tetrapod index (Fig. 2). Lucas et al. (1997a) since then have pre- taxon, usually a genus, though species are used if sented revized definitions of some of the Late Trias- they provide greater biostratigraphic resolution. In sic LVFs proposed by Lucas & Hunt (1993a). so doing, the end of an LVF is defined by the begin- Lucas & Huber (2003) reviewed global Late ning of the succeeding LVF, which is the FAD of Triassic tetrapod biochronology and demonstrated another tetrapod index taxon. This is a precise way the broad applicability of the LVFs proposed by to define LVF boundaries. LVFs thus are interval Lucas and Hunt (1993a; also see Lucas 1997a). biochrons. Lucas et al. (2007e) reviewed the status of the Trias- A distinctive assemblage of vertebrate fossils sic timescale based on patterns of tetrapod evolution characterizes each LVF. The name of the LVF is a TRIASSIC TETRAPODS 451 geographical name taken from the place where (or correlatives of the characteristic tetrapod assem- very close to where) the characteristic example of blage of each LVF are listed in this article. These the vertebrate fossil assemblage was collected. are tetrapod assemblages that are reasonably well Many LMA and LVA names have been taken studied, diverse and unambiguously correlated. from the rock formation in which the fossils are Although I make a strong effort here to correlate found, and the rock formation name is based on a the LVFs to the SGCS, the tetrapod biochronology place name. However, using the rock formation of the Triassic is a timescale independent of the name may cause confusion because it can imply SGCS. It is also important to keep in mind that, that the LMA or LVA refers to the entire duration although global LVF’s could not be defined today of deposition of the formation and not just to the due to the wide separation of most of the continents, duration of interval in which the vertebrate fossil in the Triassic Pangaean world it was possible for at assemblage is found, which is often much shorter. least some of the land vertebrates to spread across It is less confusing to choose another place name most of the world’s land area. Some degree of for the LMA or LVA. For example, the Late Triassic endemism is apparent, but it was not so great as Ischigualastian LVA of Argentina (Bonaparte 1966) to prevent definition of global or near-global was named for the Ischigualasto Formation, but the faunachrons. Ischigualastian LVF vertebrates do not occur throughout the Ischigualasto Formation, which is potentially confusing. In contrast, the Late Triassic Triassic land–vertebrate faunachrons AdamanianLVF of western North America (Lucas & Introduction Hunt 1993a) is named after Adamana, where the fossils occur, not after the Blue Mesa Member of The Triassic tetrapod timescale is based on tetrapod the Petrified Forest Formation, which contains the assemblages from the Karoo basin in South Africa characteristic fossil assemblage. This prevents con- (Early Triassic: Lootsbergian–Nonesian), the Ural fusion between the concept of a formation and the forelandbasininRussia(MiddleTriassic:Perovkan– concept of a LVF. Berdyankian) and the Chinle basin of the western The characteristic tetrapod assemblage is the USA (Upper Triassic: Otsichalkian–Apachean) primary basis for characterization of the LVF. (Fig. 3). The Karoo basin contains the tetrapod Index fossils identified here meet the criteria of assemblages characteristic of the Lootsbergian and true index fossils (temporally restricted, common, Nonesian LVFs. These assemblages are stratigra- widespread, easily identified) and do not include phically superposed and are thus demonstrably endemic or rare taxa that happen to be restricted to time successive; they are the classic Lystrosaurus a LVF, usually as single records. Principal assemblage zone and most of the Cynognathus

Ural basin

Chinle basin

Karoo basin

Fig. 3. Map of Triassic Pangaea showing the three areas that provide the fossils and strata that form the standards for the Triassic tetrapod timescale: Karoo basin, South Africa (Lootsbergian and Nonesian), Russian Urals basin (Perovkan and Berdyankian) and Chinle basin (Otischalkian, Adamanian, Revueltian and Apachean). Base map drawn by Matt Celeskey. 452 S. G. LUCAS assemblage zone (e.g. Rubidge et al. 1995; the cynodont Cynognathus (Fig. 1). Its character- Groenewald & Kitching 1995; Kitching 1995; istic tetrapod assemblage is the Lystrosaurus Hancox & Rubidge 1997; Hancox 2000; Smith & Assemblage Zone found in the Balfour (Palingkloof Botha 2005; and Botha & Smith 2007 provide an Member), Katberg and Burgersdorp (lower part) overview). These assemblages include amphibians, formations of the Karoo basin of South Africa parareptiles, dicynodonts and cynodonts particu- (e.g. Groenewald & Kitching 1995; Damiani et al. larly useful for broad correlation. 2003; Smith & Botha 2005; Botha & Smith 2006, The South African Triassic tetrapod record con- 2007). This assemblage zone has a type locality tains a long hiatus between the uppermost strata of designated by Groenewald & Kitching (1995) the Lower Triassic Cynognathus assemblage zone around Lootsberg Pass. Lootsbergian time begins and southern African rocks that contain tetrapods with the FAD of Lystrosaurus, which is the end of of certain Late Triassic age (notably the lower the Late Permian Platbergian LVF of Lucas (2005, Elliot Formation: Lucas & Hancox 2001). This 2006b). The end of the Lootsbergian is equivalent forces the tetrapod biochronological standards for to the beginning of the Nonesian LVF, which is Middle Triassic time to be moved elsewhere. For defined by the FAD of Cynognathus. this part of the standards, Lucas (1998a) used two Broom (1906) introduced two successive zones – superposed tetrapod assemblages from the Russian Lystrosaurus and Procolophon – that Kitching Ural foreland basin (e.g. Shishkin et al. 1995b, (1970, 1977) later combined into a single, Lystro- 2000a, b; Ivakhnenko et al. 1997; Novikov et al. saurus Zone. Keyser (1979) referred to this same 2000; Battail & Surkov 2000; Gower & Sennikov zone as the Lystrosaurus-Thrinaxodon Assemblage 2000; Spencer & Benton 2000; Ivakhnenko 2008a, Zone. The original name Lystrosaurus Zone (or b, c; Sennikov 2008; Tatarinov 2008) as the basis Assemblage Zone) continues to be used (e.g. for the Middle Triassic Perovkan and Berdyankian Groenewald & Kitching 1995; Lucas 1998a; LVFs. The presence of some temporal overlap Damiani et al. 2001; Botha & Smith 2006, 2007; between the top of the South African section Smith & Botha 2005). (upper Cynognathus Zone) and the Urals foreland basin section makes correlation between these sec- Characteristic tetrapod fossil assemblage. The tions considerably easier. The Russian assemblages characteristic tetrapod fossil assemblage of the yield amphibians, archosaurs and dicynodonts of Lootsbergian LVF is the Lystrosaurus Assemblage value for broad correlation. No Upper Triassic Zone of the Karoo basin, South Africa. It consists tetrapod assemblages are known from the Russian of amphibians, parareptiles, prolacertiforms, archo- Ural foreland basin (e.g. Shishkin et al. 2000b), saurs, dicynodonts, therocephalians and cynodonts. so the tetrapod biochronology standard for Late Kitching (1977) reviewed the Lystrosaurus Triassic time again must be moved elsewhere. Assemblage Zone localities, Groenewald & The Chinle Group strata of the American South- Kitching (1995) provided a synopsis of the strati- west provide the best studied and most complete graphic ranges of the genera, and Botha & Smith record for defining the Late Triassic LVFs: Otis- (2006, fig. 7) have presented the most recent data. chalkian, Adamanian, Revueltian and Apachean. The Lystrosaurus Assemblage Zone has long pro- Of great importance, tetrapod assemblages from vided a standard for correlation of the oldest Triassic Texas (Otischalkian characteristic assemblage), tetrapod assemblages, so it logically serves as the Arizona (Adamanian characteristic assemblage) basis for the oldest Triassic LVF (though it encom- and New Mexico (Revueltian and Apachean charac- passes the Permo-Triassic boundary and includes teristic assemblages) are stratigraphically super- some uppermost Permian strata, see below). posed and thus are time successive (e.g. Lucas 1993c, 1997a; Lucas et al. 2001; Heckert & Lucas Index fossils. The following tetrapod genera are 2002a, b, 2003; Heckert 2004; Heckert et al. restricted to Lootsbergian time and are widespread 2005a, b; Parker et al. 2006). The Chinle assem- and/or common enough to be useful as index blages yield phytosaurs, aetosaurs and metoposaurs fossils (Fig. 4): the amphibians Wetlugasaurus, useful for broad correlation, and a burgeoning Tupilakosaurus, Luzocephalus, and Lydekkerina; microvertebrate biostratigraphy also supports ths the parareptile Procolophon; the prolacertiform macrovertebrate-based correlation (Heckert 2004; Prolacerta; the archosaur Proterosuchus (¼ Heckert & Lucas 2006). Chasmatosaurus); the dicynodont Lystrosaurus; and the cynodonts Scaloposaurus and Thrinaxodon. Lootsbergian LVF Principal correlatives. Recognition of and corre- Definition. Lucas (1998a) introduced the term lation within the Lootsbergian is one of the most Lootsbergian LVF for the time between the FAD stable parts of the Triassic tetrapod timescale. of the dicynodont Lystrosaurus and the FAD of Thus, the terms Lystrosaurus zone, beds or fauna TRIASSIC TETRAPODS 453

taxa Lootsbergian Nonesian Perovkan Berdyankian amphibians: Eocyclotosaurus Eryosuchus Luzocephalus Lydekkerina Mastodonsaurus Odenwaldia Paracyclotosaurus Parotosuchus Trematosaurus Trematosuchus Tupilakosaurus Wetlugasaurus parareptile: Procolophon prolacertiform: Prolacerta archosaurs: Arizonasaurus Erythrosuchus dicynodonts: Kannemeyeria Lystrosaurus Parakannemeyeria Shansiodon Sinokannemeyeria Stahleckeria cynodonts: Cynognathus Diademodon Massetognathus Scalenodon Scaloposaurus Thrinaxodon Trirachodon

Fig. 4. Temporal ranges of selected genera of Early and Middle Triassic tetrapods. have long been applied to a wide geographical range formations, Junggur basin, China; Heshanggou For- of strata/fossils of Lootsbergian age. mation, Ordos basin, China; Panchet Formation, Most significant correlatives are the vertebrate India; Sanga do Cabral Formation, Parana´ basin, fossil assemblages of the: Wordy Creek Formation, Brazil; Rewan Formation, SE Galilee basin, eastern Greenland; Vokhmian, Sludkian and Ust- Australia; Arcadia Formation, SW Bowen basin, mylian horizons of the Vetluga Series, Russian Australia; and lower part of Fremouw Formation, Urals; upper Guodikeng and lower Jiucaiyuan Antarctica. Note that the alleged Lystrosaurus 454 S. G. LUCAS record from Laos (Repelin 1923; Piveteau 1938) has In southern Brazil, the Sanga do Cabral For- been re-identified as the Late Permian dicynodont mation in the Parana´ basin yields a rhytidosteid Dicynodon (Battail et al. 1995; Battail 1997). amphibian, indeterminate temnospondyls, Procolo- The Wordy Creek Formation in eastern Green- phon, ?thrinaxodontids and ?Lystrosaurus (e.g. land yields the amphibians Luzocephalus, Wetluga- Barbarena et al. 1985; Lucas 2002; Abdala et al. saurus and Tupilakosaurus (Sa¨ve-So¨derbergh 1935; 2002; Cisneros 2008a, b; Cisneros & Schultz 2002; Nielsen 1954) and thus is of Lootsbergian age. Dias-da-Silva et al. 2005, 2006a, b; Dias-da-Silva These strata also yield Induan ammonites, and are & Marsicano 2006; Dias-da-Silva & Schultz 2008). key to correlation of the Lootsbergian to part of A putative Permian tetrapod record from the Buena the Induan (see below). Vista Formation of Uruguay (Pin˜eiro et al. 2003, In the Russian Urals, the Lootsbergian interval is 2004, 2007) is more likely correlative to the Loots- equivalent to Zone V of Efremov (1937, 1952), bergian Sanga do Cabral assemblage (Dias-da-Silva which has most recently been called the Vokhmian, et al. 2006b). Sludkian and Ustmylian horizons of the Vetlugan In eastern Australia, the Arcadia Formation (SW Series (Superhorizon) (Ivakhnenko et al. 1997; Bowen basin) and the Rewan Formation (SE Galilee Shishkin et al. 2000b). Tetrapod taxa include basin) yield small assemblages of tetrapods of anthracosaurs, the temnospondyls Luzocephalus, Lootsbergian age. The Arcadia Formation assem- Benthosuchus, Wetlugasaurus and Tupilakosaurus, blage encompasses a diversity of mostly endemic procolophonids, a prolacertiform, the proterosuchid amphibians, including fragmentary lydekkerinids, Chasmatosuchus and other (mostly fragmentary) a primitive procolophonid, a possible Prolacerta, archosaurs and the dicynodont Lystrosaurus an archosaur similar to Proterosuchus and ?Lystro- (Shishkin et al. 1995b; Ivakhnenko et al. 1997; saurus (e.g. Bartholomai 1979; King 1983; Battail & Surkov 2000; Gower & Sennikov 2000; Thulborn 1983; Warren 1991; Damiani 2001; Novikov et al. 2000; Shishkin et al. 2000a, b; Warren et al. 2006). In the SE Galilee basin, the Spencer & Benton 2000). occurrence of Lydekkerina in the Rewan Formation In northwestern China, land-vertebrates of supports a Lootsbergian age assignment (Warren Lootsbergian age come from the upper part of et al. 2006). the Guodikeng Formation and the lowermost Southwest of the Transantarctic Mountains in Jiucaiyuan Formation (both in the Cangfanggou southern Antarctica, the lower part of the Fremouw Group) near Jimsar NE of Urumqi in western Formation yields a vertebrate fossil assemblage of Xinjiang (e.g. Cheng 1981; Metcalfe et al. 2009). Lootsbergian age that includes temnospondyls, These vertebrates are the ‘Lystrosaurus fauna’ of a rhytidosteid, the procolophonid Procolophon, northwestern China of some workers (e.g. Sun the prolacertiform Prolacerta, a proterosuchid or 1972), and they provided the basis for the Jimsarian erythrosuchid, a rauisuchian, the dicynodonts Myo- LVF of Lucas (1993a). Taxa present are a pro- saurus and Lystrosaurus, the cynodont Thrinaxodon lacertid, a ?procolophonid, the proterosuchian and scaloposaurs (e.g. Colbert 1972, 1991; Hammer Proterosuchus (¼ Chasmatosaurus), a regisaurid 1990; Collinson et al. 2006). This Lootsbergian therocephalian and the dicynodont Lystrosaurus, assemblage has been referred to as the lower of which seven species have been named, most of Fremouw fauna or lower tetrapod fauna of the which are invalid (Colbert 1974; Colbert & Kitching Fremouw Formation (Colbert 1972, 1991). 1977; Lucas 2001). In the Ordos basin of north–central China, near Comments. Most Lootsbergian vertebrate fossil Fugu, Shanxi, the upper part of the Heshanggou assemblages are readily recognized by the presence Formation yields a vertebrate fauna that was the of Lystrosaurus. Procolophon and Proterosuchus basis of the Fuguan LVF of Lucas (1993a). Taxa are also important to the correlation of Lootsbergian present are indeterminate capitosauroids, procolo- tetrapod assemblages. However, temnospondyl- phonids, an erythrosuchid and an ordosiid theroce- dominated assemblages occur that lack Lystrosaurus phalian; based primarily on the procolophonids, and thus are more difficult to correlate. I have used these are of likely Lootsbergian age. the temporal overlap of Lystrosaurus and the amphi- In India, the Panchet Formation along the bians Tupilakosaurus and Luzocephalus in Russian Damodar River northwest of Calcutta has produced strata as the primary basis for equating Lootsbergian a Lootsbergian vertebrate assemblage that includes dicynodont-dominated assemblages with temnos- a lydekkerinid, ?benthosuchid, ?capitosaurids, an pondyl-dominated assemblages. indobrachyopid, trematosaurids, a procolophonid, Cosgriff (1984) assigned several temnospondyl- the proterosuchian Proterosuchus and Lystrosaurus dominated assemblages to his A1 ‘horizon’ (Lydekker 1882; Sahni & Huene 1958; Tripathi (¼ Lootsbergian), even though these lack any 1961, 1969; Tripathi & Satsangi 1963; Hughes index taxa of the Lootsbergian: the Knocklofty 1963; Ray 2005). Sandstone/Shale in SE Tasmania (Cosgriff 1974), TRIASSIC TETRAPODS 455 the Sticky Keep Formation in Svalbard (Wiman P 1910, 1915; Nilsson 1942, 1943; Cox & Smith TRIASSIC 1973), the upper Andavakoera Formation (Middle

Sakamena Group or Formation) in NW Madagascar Lootsbergian Nonesian (Lehman 1961, 1966; Steyer 2002; Maganuco & Perovkan Pasini 2009) and the Arcadia Formation of southern Queensland (Warren 1991). Except for the Arcadia Formation, I assign these assemblages a Nonesian age (see below). Lootsbergian time encompasses both the ‘Lystro- saurus zone’ and ‘Procolophon zone’ of classic usage (e.g. Broom 1906). Thus, two distinct tetrapod assemblages (at least in the Karoo basin) can be C C recognized within the Lootsbergian, simply based A B A B on the stratigraphic distribution of Procolophon. According to Botha & Smith (2007), all records Lystrosaurus with Procolophon Lystrosaurus with Dicynodon Lystrosaurus without Dicynodon and Procolophon Cynognathus without Kannemeyeria Cynognathus with Kannemeyeria of Lystrosaurus maccaigi in the Karoo basin are and Shansiodon Angonisaurus Permian (they co-occur with the Permian dicynodont Dicynodon), whereas L. curvatus straddles the Permo-Triassic boundary, and records of L. murrayi and L. declivus are Triassic. This provides a basis for a threefold subdivision of the Lootsbergian (Fig. 5): (1) Lootsbergian A is the time of overlap of Dicyno- don and Lystrosaurus; (2) Lootsbergian B is the succeeding interval with Lystrosaurus without Pro- colophon; and (3) Lootsbergian C is the temporal overlap of Lystrosaurus and Procolophon. These subdivisions have some value outside of the Karoo basin. For example, in the Guodikeng Formation in the Junggur basin of northwestern China, there is a stratigraphic overlap of Lystrosaurus and Dicynodon (Lootsbergian A) followed by an interval of Lystro- saurus without Procolophon (Lootsbergian B) (Cheng 1981; Metcalfe et al. 2009). Indeed, in north- western China, the co-occurrence of Lystrosaurus and Dicynodon at Dalongkou was first assigned to the upper Changhsingian Falsisca postera conchos- tracan zone and uppermost part of the F. eotriassica conchostracan zone by Kozur (1998a, b) (see also Kozur & Weems 2010). Therefore, a formal subdivi- sion of the Lootsbergian into sub-LVFs has merit and should provide more precise correlation within the Lootsbergian interval.

Nonesian LVF

Definition. The term Nonesian LVF refers to the time Fig. 5. Subdivisions of the Lootsbergian and Nonesian between the FAD of the cynodont Cynognathus and LVFs (based primarily on Hancox 2000). Restoration of the FAD of the amphibian Eocyclotosaurus. The Lystrosaurus by Matt Celeskey. characteristic tetrapod assemblage is found in the lower two-thirds of the Cynognathus Assemblage Zone, which is from the upper two-thirds of the Bur- Cynognathus, which is the end of the Lootsbergian gersdorp Formation in the Karoo basin of South LVF. The end of the Nonesian is the beginning of Africa (e.g. Kitching 1995; Hancox et al. 1995; the Perovkan LVF, which is defined by the FAD of Hancox 2000). The type section of the Cynognathus Eocyclotosaurus. Assemblage Zone encompasses Nonesi’s Nek, from Broom (1906, 1907) coined the name Cynog- which the name Nonesian is derived (Kitching nathus ‘beds’, which was later transmuted to ‘zone’ 1995). Nonesian time begins with the FAD of by other workers (Watson 1914a, b; Kitching 1970, 456 S. G. LUCAS

1977). Keyser & Smith (1978) renamed it the Kan- Arizona (Damiani 2001; Lucas & Schoch 2002; nemeyeria Assemblage Zone, and Keyser (1979) Heckert et al. 2005a; Nesbitt 2005). These records termed it the Kannemeyeria–Diademodon Assem- of Nonesian index taxa are of late Olenekian age blage Zone. Kitching (1984) called it the Cynog- (see below). nathus–Diademodon Assemblage Zone. The term In the Germanic basin, the Middle Buntsandstein Cynognathus Assemblage Zone has been used (upper Volpriehausen, Hardegsen and Solling for- most recently (e.g. Kitching 1995; Rubidge et al. mations) yields fossils of Parotosuchus, Oldenwal- 1995; Lucas 1998a; Hancox 2000). dia and Trematosaurus, indicative of a Nonesian age (e.g. Schroeder 1913; Werneburg 1993; Lucas Characteristic tetrapod assemblage. The character- 1999; Schoch & Werneburg 1999; Lucas & istic assemblage of the Nonesian LVF occurs in sub- Schoch 2002; Schoch 2008). Specifically, Odenwal- zones A and B of the Cynognathus Assemblage dia occurs only in the Solling Formation, and Zone of the Karoo basin (Hancox 2000) (Fig. 5). Trematosaurus is common in the Hardegsen For- The tetrapod taxa present are amphibians, including mation and present in the Solling Formation. Paro- Parotosuchus, Wellesaurus and Trematosuchus, tosuchus is known from the Hardegsen and the captorhinids, a ?sphenodontid (or ?procolophonid), Solling formations. One specimen of Parotosuchus rhynchosaurs, the archosaurs Erythrosuchus and (the holotype of P. helgolandicus) is known from Euparkeria, the dicynodonts Kannemeyeria and the uppermost Volpriehausen Formation, from the Kombuisia, therocephalians and cynodonts, includ- upper Gerviellia beds assigned by Kozur & ing Cynognathus, Diademodon and Trirachodon Bachmann (2008), based on conchostracans, to the (e.g. Kitching 1977, 1995; Hancox & Rubidge Spathian. 1994; Hancox et al. 1995; Shishkin et al. 1995a; Temnospondyls of the Sticky Keep Formaton in Damiani 2001; Damiani & Rubidge 2003; Abdala Svalbard co-occur with early Olenekian (Smithian) et al. 2005). ammonites (Buchanen et al. 1965; Tozer 1967). The temnospondyls are: Sasenisaurus, Peltostega, Index fossils. The following tetrapod genera are Aphanerama (¼ Lonchorhynchus), Lyrocephalis- restricted to Nonesian time and are widespread cus, Teretrema and Boreaosaurus (Wiman 1910, and/or common enough to be considered index 1915, 1916; Nilsson 1942, 1943; Cox & Smith fossils (Fig. 4): the amphibians Parotosuchus, 1973). Such an acme in trematosaur diversity may Odenwaldia, Wellesaurus, Trematosaurus and characterize the Nonesian. I assign a Nonesian age Trematosuchus and the cynodont Trirachodon. to the Sticky Keep tetrapods based mostly on the The LOs of the the archosaur Erythrosuchus, the marine evidence that they are Olenekian and that cynodonts Cynognathus and Diademodon and of the Nonesian is equivalent to at least part of the Ole- the dicynodont Kannemeyeria are in the Nonesian. nekian (see below). The species K. simocephalus is restricted to Nones- The Petropavlovsk svita in the Russian Urals ian time, but the species K. cristarhynchus is (Yarenskiy horizon) yields anthracosaurs, temnos- younger, of Perovkan age. pondyls (including Parotosuchus), procolophonids, a prolacertid, and various archosaurs, including ery- Principal correlatives. Principal correlatives of the throsuchids and rauisuchids (Shishkin et al. 1995b, type Cynognathus Assemblage Zone are: Wupatki 2000a, b ; Ivakhnenko et al. 1997; Battail & and Torrey formations of the Moenkopi Group/ Surkov 2000; Gower & Sennikov 2000; Novikov Formation, Utah/Arizona, USA; Sticky Keep et al. 2000; Spencer & Benton 2000). The Parotosu- Formation of Svalbard, Arctic Norway; Middle chus record is the primary basis for a Nonesian Buntsandstein (upper Volpriehausen, Hardegsen age assignment. and Solling formations), Germany; Petropavlovsk In China, the lower Ermaying Formation in the Formation (Yarenskiy horizon) in the Russian Ordos basin produces a vertebrate fauna upon Urals; lower part of Ermaying Formation, Ordos which Lucas (1993a) based the Ordosian LVF. basin, China; Puesto Viejo and Rio Mendoza for- Taxa present are a procolophonid, a proterosuchian, mations, Argentina; base of the Lower Sandstone euparkeriids, a therocephalian and the dicynodonts of the Zarzaitine Series in Algeria; lower N’tawere Parakannemeyeria and Kannemeyeria(¼ Shaan- Formation, Zambia; K7 horizon of the Kingori beikannemeyeria) (Lucas 2001). The Kannemeyeria Sandstone, Tanzania; and upper Fremouw For- record as well as the overall composition of the mation, Antarctica. assemblage suggest a Nonesian age. The Torrey Formation of the Moenkopi Group in In Argentina, the upper part of the Puesto Viejo Utah, USA, has yielded a skull of Parotosuchus Formation produces the dicynodont Kannemeyeria, (Lucas & Schoch 2002). Specimens of Wellesaurus a traversodontid and Cynognathus (Bonaparte 1970, as well as an Odenwaldia-like form are from the 1978, 1982). The co-occurrence of Cynognathus Wupatki Member of the Moenkopi Formation in and Kannemeyeria supports a Nonesian age TRIASSIC TETRAPODS 457 assignment. The correlative fauna from the middle and Hancox (2000) provided a synopsis of the strati- part of the Rio Mendoza Formation (but see Zavat- graphic ranges of the genera. Watson (1942) and tieri & Arcucci 2007 for a different correlation) Kitching(1977)subdividedtheCynognathusAssem- includes Kannemeyeria (Vinceria andina Bonaparte blage Zone into two subzones. Hancox & Rubidge is not Shansiodon, as Lucas [1993e] suggested, but (1994), Hancox et al. (1995), Shishkin et al. instead is Kannemeyeria), traversodontids and (1995a), Hancox (2000; Hancox et al. 1995, 2000) a galeosaurid. and Abdala et al. (2005) divided the Cynognathus Bonaparte (1981) described dicynodonts and Assemblage Zone into three stratigraphically dis- proterosuchian postcrania from the lower part of crete assemblages (Fig. 5). These assemblages have the Puesto Viejo Formation. He referred to them been called subzones A, B and C by Hancox et al. as the Agua de los Burros local fauna. He assigned (1995), and the upper one is now assigned a Perovkan the dicynodonts to ‘Vinceria’(¼ Kannemeyeria) age (Hancox 2000; Abdala et al. 2005; Lucas et al. and claimed correlation to the Lystrosaurus Assem- 2007e). This means that the South African Nonesian blage Zone based on a mean value of K/Ar ages of (which encompasses subzones A and B) is divisible 232 + 4 Ma from basalts and tuffs that bracket the into two biochronological units (Hancox 2000). A fossils (Valencio et al. 1975, fig. 2). Given that we more important point is that recognizing subzone now know that the Induan is approximately 251– C as Perovkan means that not all of the classically- 252 Ma (Bachmann & Kozur 2004; Kozur & recognized ‘Cynognathus zone’ is Nonesian. Weems 2010; Mundil et al. 2010), the Argentinian Following Hancox (2000), the Nonesian can be dates (which are Carnian by current Triassic subdivided into older (Nonesian A) and younger timescale calibration) do not support Bonaparte’s (Nonesian B) sub-LVFs (Fig. 5). Nonesian A correlation, nor do the fossils, which instead begins with the FAD of Cynognathus, and Nonesian suggest a Nonesian age. B begins with the FAD of Kannemeyeria. The FAD The baseof the Lower Sandstone of the Zarzaitine of Eocyclotosaurus (beginning of the Perovkan Series in southeastern Algeria yields the amphibians LVF) is the end of Nonesian B. In the Karoo basin, Odenwaldia and ‘Wellesaurus’ (an indeterminate where Eocyclotosaurus is so far unknown, the LO heylerosaurid according to Damiani 2001) as well of shansiodont dicynodonts approximates the begin- as a ?brachyopid, trematosaurid and the prolacerti- ning of Perovkan time (Fig. 5). In Nonesian A time form Jesairosaurus (Lehman 1957, 1971; Welles in the Karoo basin, the amphibian Kestrosaurus is 1993; Jalil 1990, 1993, 1994, 1997, 1999). The common and associated with Trematosuchus as record of Odenwaldia supports a Nonesian age well as theriodonts, Cynognathus, Diademodon, assignment. Trirachodon and Bauria. During Nonesian B time, The lower part of the N’tawere Formation in characteristic taxa are Parotosuchus, Kannemeyeria, Zambia produces Diademodon and Kannemeyeria Cynognathus, Diademodon, Trirachodon, Bauria, (Crozier 1970). In the Ruhuhu Valley of Tanzania, Erythrosuchus and Euparkeria. Most of the Nones- the K7 horizon of the Kingori Sandstone Formation ian correlative tetrapod assemblages (see above) of Stockley (1932) yields Kannemeyeria (Cruick- include Kannemeyeria, so they are of Nonesian shank 1986). These are likely (though not defini- B age. tively) Nonesian records. In Antarctica, the upper part of the Fremouw Perovkan LVF Formation yields capitosaurid temnospondyls inc- luding Parotosuchus, Cynognathus, a diademodon- Definition. The term Perovkan LVF refers to the tid and a kannemeyeriid (Colbert 1991; Hammer time interval between the FADs of the amphibians 1988, 1990, 1995; Damiani 2001; Collinson et al. Eocyclotosaurus and Mastodonsaurus giganteus 2006). This has long been regarded as an assemblage (Fig. 1). The characteristic tetrapod assemblage is of the ‘Cynognathus zone’, and is of Nonesian age. the vertebrate fossil assemblage of the Donguz svita (Eryosuchus fauna) in the Russian Urals Comments. Most Nonesian vertebrate assemblages (Shishkin et al. 1995b, 2000b; Ivahknenko et al. have long been recognized by the presence of 1997). Lucas (1998a) termed this the Shansiodon Cynognathus and/or Diademodon, but these taxa Assemblage Zone, after the distinctive dicynodont have temporal ranges that extend into the Perovkan. Shansiodon (¼ Rhinodicynodon). These fossils are Parotosuchus is a key temnospondyl taxon to corre- from an approximately 175-m-thick section late many Nonesian assemblages (Damiani 2001). exposed in the Donguz River drainage near the The temporal succession of Kannemeyeria species city of Perovka, from which the name of the LVF is important, with K. simocephalus restricted to the is taken (Lucas 1998a, fig. 8). The beginning of Nonesian and K. cristarhynchus a Perovkan taxon. the Perovkan is defined by the FAD of the amphi- Kitching (1977) reviewed the Cynognathus bian Eocyclotosaurus. The end of the Perovkan Assemblage Zone localities, and Kitching (1995) LVF is the beginning of the Berdyankian LVF, 458 S. G. LUCAS which is defined by the FAD of the amphibian index taxa of the Perovkan (Fig. 4): the amphibians Mastodonsaurus giganteus. Eryosuchus, Eocyclotosaurus and Paracycloto- Lucas (1998a) originally defined the beginning saurus, the archosaur Arizonasaurus, the cynodont of Perovkan time as the FAD of the dicynodont Scalenodon and the dicynodonts Shansiodon, Para- Shansiodon. However, Shishkin (2000) has argued kannemeyeria and Sinokannemeyeria. Kannemeyria that the type assemblage of the Perovkan LVF is christarhynchus is a Perovkan index fossil, and the late Anisian, so it is younger than the Eocycloto- HOs of Kannemeyeria, Cynognathus and Diademo- saurus assemblage that typically represents the Per- don are Perovkan. ovkan in western Europe and North America and is of unambiguous early Anisian age (Lucas & Schoch Principal correlatives. Principal correlatives of the 2002). A more circumspect reading of the same data type Perovkan assemblage are from the Holbrook (e.g. Ivakhenko et al. 1997) simply regards the and Anton Chico members of the Moenkopi For- Donguz assemblage as Anisian, with no more mation, Arizona–New Mexico USA; lower part of precise correlation to the SGCS. Wolfville Formation at Lower Economy, Fundy Lucas (1993d) argued that the LO of the dicyno- basin, Nova Scotia, Canada; Otter Sandstone of dont Shansiodon is Anisian, and this is why Lucas the United Kingdom; Upper Buntsandstein (Ro¨t (1998a) used it to define the beginning of the Perov- Formation), Germany-France; lower Kelamayi kan. However, if the LO of Shansiodon is actually Formation, Junggur basin, Xinjiang, China; upper younger than the LO of Eocyclotosaurus, then Ermaying Formation, Ordos Basin, China; Yerra- records of Eocyclotosaurus (Upper Buntsandstein palli Formation, India; Lower Zarzaitine Formation, in Germany and France, upper Moenkopi Group in Algeria; upper part of the Burgersdorp Formation in USA) are of Nonesian age. The temporal succession the Karoo basin of South Africa; Omingonde For- of Eocyclotosaurus and Shansiodon is not easily mation, Namibia; and lower Manda Formation, resolved, but Lucas et al. (2007e) noted that the Tanzania. LO of Kannemeyeria in China predates the LO of The Holbrook and Anton Chico members of the Shansiodon, and in South Africa the LO of Kanne- Moenkopi Formation, in Arizona–New Mexico, meyria predates the LO of shansiodonts (Fig. 5), USA, yield the characteristic Perovkan capitosaur- and there is no conclusive evidence that the young- oid amphibian Eocyclotosaurus, very similar to est Nonesian assemblage in South Africa (subzone E. lehmanni from the Upper Buntsandstein (Ro¨t B of Hancox et al. 1995) is equivalent to the Eocy- Formation), as well as other capitosaurs, brachyo- clotosaurus zone. Lucas et al. (2007e) therefore pids, and the ctenosauriscid Arizonasaurus (Lucas recognized problems in establishing the temporal & Morales 1985; Lucas & Hunt 1987; Morales succession of Perovkan assemblages, but believe 1987; Schoch 2000b; Boy et al. 2001; Lucas & all are broadly Anisian, and some (part of American Schoch 2002; Heckert et al. 2005a; Nesbitt 2005). Moenkopi Group, German Ro¨t Formation) are A Shansisuchus-like erythrosuchian from the Anton clearly early Anisian. The easiest way to remove Chico Member in New Mexico (Lucas et al. 1998b; ambiguity here is to redefine the beginning of the Nesbitt et al. 2006) is consistent with a Perovkan Perovkan as the FAD of Eocyclotosaurus, as did age assignment. Lucas et al. (2007e). In the Fundy basin of Nova Scotia, Canada, the lower part of the Wolfville Formation (also referred Characteristic tetrapod assemblage. Three princi- to as the ‘Lower Economy Beds’) yields a small tet- pal sites in the Donguz svita produce the following rapod assemblage that was the basis of the Econo- taxa: various amphibians, including Eryosuchus, mian LVF of Huber et al. (1993b). The presence Bukobaja, Plagiosternum and Plagioscutum,a of a trematosaur (cf. Cosgriffius) and the lepidosaur procolophonid, a prolacertid, a proterosuchid, the cf. Tanystropheus suggests a possible Perovkan age erythrosuchid Erythrosuchus, rauisuchids, a eupar- (Lucas & Huber 2003). keriid, the dicynodonts Kannemeyeria (¼ Rhadio- The Otter Sandstone in Devon, United Kingdom, dromus, Rabidosaurus, Edaxosaurus, Calleonassus yields the temnospondyl Eocyclotosaurus, the and Rhinocerocephalus) and Shansiodon (¼ Rhino- rhynchosaur Fodonyx, the prolacertiform Tanystro- dicynodon), therocephalians, the cynodonts Scale- pheus, a procolophonid, a rauisuchian and a ?cteno- nodon, Antecosuchus and a traversodontid (Shishkin sauriscid archosaur (Benton et al. 1994; Hone & et al. 1995b, 2000a, b; Ivakhnenko et al. 1997; Benton 2008). As Milner et al. (1990) stressed, Surkov 1999; Battail & Surkov 2000; Gower & ‘Mastodonsaurus’ lavisi Seeley from the Otter Sennikov 2000; Spencer & Benton 2000; Tverdokh- Sandstone is a nomen dubium upon which it is lebov et al. 2002). risky to draw stratigraphic conclusions, so I do not consider it a Perovkan record of Mastodonsaurus. Index fossils. The following tetrapod genera are Indeed, Damiani (2001) considered the type common and/or widespread enough to be useful material of ‘M.’ lavisi to be indeterminate. TRIASSIC TETRAPODS 459

In Germany and France, the Upper Buntsandstein The lower Manda Formation in Tanzania pro- (Ro¨t Formation) yields Eocyclotosaurus (Heyler duces the amphibian Eryosuchus, the rhynchosaur 1969, 1976; Ortlam 1970; Kamphausen & Morales Stenaulorhynchus, the archosaur ‘Mandasuchus’, 1981; Lucas & Schoch 2002) and is of Perovkan age. the dicynodonts Shansiodon (¼ Tetragonius) and In the Junggur basin of Xinjiang, China, the Angonisaurus and the cynodont Scalenodon lower part of the Kelamayi (¼ Karamay) Formation (Huene 1938a, b; Crompton 1955; Cruickshank produces a vertebrate fauna that consists of indeter- 1965, 1967; Cox & Li 1983; Damiani 2001). This minate labyrinthodonts (including the holotype of is a Perovkan assemblage. the nomen dubium ‘Parotosaurus’[¼ Parotosu- chus] turfanensis Young: Lucas & Hunt 1993b), a Comments. Lucas (1998a) defined the Perovkan euparkeriid, an erythrosuchid and the dicynodonts LVF as the time between the FAD of the dicynodont Parakannemeyeria and Xiyukannemeyeria (Liu & Shansiodon and the FAD of the temnospondyl Mas- Li 2003; Liu 2004). The bauriid therapsid Traver- todonsaurus. Its characteristic assemblage is the sodontoides from Jiyuan, Henan may also be of tetrapod fauna from the Russian Donguz svita, so Perovkan age (Cheng 1981; Sun 1989). the land-vertebrate biochronology shifts here from The upper part of the Ermaying Formation in the superposed South African assemblages (the charac- Ordos basin in northern China produces what has teristic assemblages of the Lootsbergian and Nones- been called the Perovkan-age ‘Sinokannemeyeria ian LVFs) to the superposed Russian assemblages fauna’ or ‘kannemeyeriid fauna’ of China (e.g. Sun (the characteristic assemblages of the Perovkan 1972; Cheng 1981; Lucas 2001). Lucas (1993a) and Berdyankian LVFs). This geographical shift based the Ningwuan LVF on this assemblage. The poses problems for the biochronology, particularly vertebrate fossil assemblage includes indeterminate in demonstrating the temporal succession (and not labyrinthodonts, a procolophonid, erythrosuchids, overlap) of Nonesian and Perovkan assemblages. an ?ornithosuchid, a ?euparkeriid, a cynodont, and Indeed, the reassignment of the upper ‘Cynognathus the dicynodonts Shansiodon, Sinokannemeyeria zone’ to the Perovkan LVF discussed above directly and Parakannemeyeria (Lucas 2001). reflects such problems (Hancox 2000; Abdala et al. In the Pranhita–Godavari Valley of India, the 2005; Lucas et al. 2007e). The easiest way to reduce Yerrapalli Formation yields an assemblage of Per- ambiguity here was to redefine the beginning of the ovkan age. It includes the amphibian Eryosuchus, Perovkan as the FAD of Eocyclotosaurus (Lucas the rhynchosaur Mesodapedon, a prolacertid, the et al. 2007e). archosaur Erythrosuchus, a raisuchid, the dicyno- Perovkan tetrapod assemblages are best known donts Wadiasaurus and Kannmeyeria (¼ Rechni- in Russia and China where they contain numerous saurus), and a trirachodontid, (e.g. Roychowdhury dicynodonts. Correlatives are either dicynodont 1970a, b; Chatterjee 1980b; Damiani 2001; Sen dominated (Manda Formation, upper Burgersdorp 2003, 2005; Bandyopadhyay & Sengupta 2006). Formation) or amphibian dominated (upper Moen- The Omingonde Formation in Namibia produced kopi, upper Buntsandstein). a Perovkan-age assemblage that includes an eryo- poid temnospondyl, the dicynodonts Kannemeyeria Berdyankian LVF cristarhynchus, Dolichuranus, and Rhopalorhinus, a bauriamorph, and cynodonts, including ?Cynog- Definition. The term Berdyankian LVF is the time nathus, Diademodon and Trirachodon (Keyser interval between the FAD of the amphibian Masto- 1973a, b, 1978; Pickford 1995; Smith & Swartt donsaurus giganteus and the FAD of the phytosaur 2002). Parasuchus (¼ Paleorhinus) (Fig. 1). The charac- In the Karoo basin of South Africa, the upper teristic tetrapod assemblage is the vertebrate fossil part of the Burgersdorp Formation yields the upper assemblage of the Bukobay svita in the Russian part of the Cynognathus Assemblage Zone (sub- Urals (e.g. Ivakhnenko et al. 1997; Shishkin et al. zone C of Hancox 2000, see discussion above and 2000b). Relevant vertebrate-fossil localities are Fig. 5). Characteristic taxa are the amphibian near the Berdyank River, from which the LVF Paracylotosaurus, the dicynodonts Cynognathus, takes its name. The characteristic Berdyankian tet- Diademodon and Cricodon, and the dicynodonts rapod assemblage is directly superposed on the Angonisaurus and Kannemeyeria, which support a characteristic Perovkan assemblage. The beginning Perovkan age assignment (e.g. Hancox & Rubidge of the Berdyankian is defined by the FAD of Masto- 1994, 1996; Damiani 2001; Damiani & Hancox donsaurus giganteus, whereas the end of the Ber- 2003; Abdala et al. 2005). Paracylotosaurus is dyankian is the beginning of the Otischalkian, also known from the Denwa Formation in the which is defined by the FAD of Parasuchus. Satpura basin, India and the Wianamatta Group of the Sydney basin, Australia (Damiani & Hancox Characteristic tetrapod assemblage. The assem- 2003), so these may also be Perovkan records. blage from the Bukobay Formation includes an 460 S. G. LUCAS anthracosaur, the amphibians Mastodonsaurus, Probainognathus (Bonaparte 1970; Romer 1973; Bukobaja,?Cyclotosaurus, Plagioscutum and Pla- Sereno & Arcucci 1993, 1994; Lucas & Harris giosternum, an erythrosuchid, a rauisuchid, and 1996; Bonaparte 1997; Arcucci & Marsicano the dicynodonts ‘Elephantosaurus jachimovitschi’ 1998; Hsiou et al. 2002). Bonaparte (1966, 1967, Vyushkov (a Stahleckeria-like form) and a generic- 1982) based the Chanarian ‘provincial age’ on ally indeterminate kannemeyeriid (Shishkin et al. this assemblage. 1995b, 2000a, b; Ivakhnenko et al. 1997; Battail & The lower part of the Santa Maria Formation in Surkov 2000; Gower & Sennikov 2000). the Parana´ basin of Rio Grande do Sul, Brazil yields vertebrate fossil assemblages from Cande- Index fossils. The following tetrapod genera are laria and Chiniqua´ considered by Barberena common and/or widespread enough to be index (1977) and Barberena et al. (1985) to be two differ- fossils of the Berdyankian (Fig. 4): the cynodont ent local faunas of different ages. Lucas (2002) Massetognathus and the dicynodonts Dinodonto- regarded them as a single biostratigraphic assem- saurus and Stahleckeria. The LO of the amphibian blage that includes a procolophonid, archosaurs, Mastodonsaurus giganteus is Berdyankian. An the dicynodonts Dinodontosaurus and Stahleckeria, acme in plagiosaur diversity and abundance charac- chiniquodontids, and the traversodontids Massetog- terizes Berdyankian time. No procolophonids are nathus, Belesodon, Traversodon, Exaeretodon, known from Berdyankian strata (Cisneros 2008a), Santacruzodo, Protuberum and Probelesodon (e.g. but this must be due to a lack of discovery, not a Abdala & Ribeiro 2003; Cisneros et al. 2004; real absence, as both pre- and post-Berdyankian Langer et al. 2007; Reichel et al. 2009). This ass- procolophonids are known. emblage and the Chanarian type assemblage in Argentina are assigned a Berdyankian age based Principal correlatives. The Lettenkohle (Lettenkeu- largely on their dicynodonts and traversodontids per, Lower Keuper, Erfurt Formation) in Germany and their stratigraphic position, which places them and the Chanarian LVF localities in Argentina and between the Nonesian and the Adamanian. Brazil are the principal correlatives of the Berdyan- kian type assemblage. The Lettenkohle record is Comments. Previously, I used the FAD of the genus important because it establishes the Ladinian age Mastodonsaurus to define the beginning of the Ber- of at least part of the Berdyankian (see below). dyankian. This was based on a taxonomy in which The Lettenkohle fossils are from the Grenze Mastodonsaurus (typified by the species M. gigan- bonebed, the laterally equivalent/overlying teus) was distinguished from the older (Perovkan) Vitriolschiefer and the Kupferzell locality, so they Heptasaurus (e.g. Schoch 1999; Schoch & Milner are above the unconformity that separates the 2000). However, taxonomists who study these Keuper from the underlying Muschelkalk. Letten- amphibians have suggested that Mastodonsaurus kohle tetrapods include a chroniosuchian, the and Heptasaurus be combined into a single genus, amphibians Mastodonsaurus giganteus, Calli- Mastodonsaurus (Rayfield et al. 2009). Thus, I stomordax, Plagiosternum, Plagiosuchus and now use the FAD of the species M. giganteus to Kupferzella, the rauisuchian Batrachotomus, the define the beginning of the Berdyankian so as not prolacertiform Tanystropheus and small cynodonts to be subject to the shifting opinions of taxonomists (e.g. Wild 1978, 1980; Schoch 1997, 2000a; Lucas revising the genus-level taxonomy of stereospondyl 1999; Schoch & Werneburg 1999; Witzmann amphibians. This preserves the original intent of the et al. 2008; Damiani et al. 2009; Gower & Schoch Berdyankian, as no temnospondyl worker has advo- 2009). A Dinodontosaurus-like humerus from the cated the synonymy of Heptasaurus cappelensis and Vitriolschiefer (Lucas & Wild 1995) may link the Mastodonsaurus giganteus at the species-level. Lettenkohle to the South American Chanarian. As noted by Lucas (1998a), global correlations However, a Dinodontosaurus-like radius is also within the Berdyankian interval are confounded by known from the upper Anisian interval of the the near endemism of South American tetrapod Muschelkalk in Germany, so this may indicate that assemblages that are apparently of this age (the the Berdyankian also encompasses part of late Dinodontosaurus faunas of Argentina and Brazil, Anisian time (Lucas 2007b). classically assigned to the Chanarian LVA of The Chan˜ares local fauna from the Ischichuca Bonaparte 1966, 1967). Recognition of Berdyankian- (formerlyChan˜ares)FormationoftheIschigualasto– age assemblages in Russia and Germany is rendered Villa Unio´n basin of northwestern Argentina easy by the presence of the key taxon Mastodon- includes various archosaurs such as Tarjadia, saurus giganteus (Lucas 1999). Lagerpeton, Marasuchus and Chanaresuchus, the The Berdyankian is difficult to correlate glob- dicynodont Dinodontosaurus, the traversodontids ally, largely because of a paucity of tetrapod assem- Massetognathus and Megagomphodon, the chini- blages of this age. Two clusters of localities quodontid Probelesodon and the probainognathid (European and South American) are equated, TRIASSIC TETRAPODS 461 largely on the basis of the Lettenkohle dicynodont has both Otischalkian and earliest Adamanian and the conclusion that ‘Elephantosaurus’ is a ‘stah- records. The LOs of the widespread temnospondyl leckeriid’, possibly a synonym of Stahleckeria Metoposaurus and of the rhynchosaur Hyperodape- (Lucas & Wild 1995). The South American Chanar- don are Otischalkian, and these taxa are also known ian LVF thus is the provincial secondary standard in Adamanian strata (Lucas et al. 2002a, 2007e). correlative to the Berdyankian. The Berdyankian may be relatively long, at least Principal correlatives. Besides Chinle Group corre- correlative to the latest Anisian and Ladinian (see latives, principal Otischalkian vertebrate assem- below). Nevertheless, Berdyankian tetrapod fossil blages are from the Sanfordian interval of the assemblages probably only represent the earlier Newark Supergroup basins of eastern North part of this time interval. Indeed, the paucity of tetra- America; Schilfsandstein (Stuttgart Formation) of pod assemblages of Berdyankian age represents one the German Keuper; the Irohalene Member (T4) of of the most substantial deficits in the global record of the Timesgadiouine Formation, Argana Group, Triassic tetrapods. This is an important deficit Morocco; and the basal part of the Maleri For- because many characteristic Late Triassic tetrapod mation, Pranhita–Godavari Valley, India. taxa, such as metoposaurs, phytosaurs, aetosaurs Otischalkian principal correlatives and the and dinosaurs, so far lack evolutionary antecedents characteristic tetrapod assemblage encompass a that should occur in Berdyankian-age strata. broad geographical range of Chinle Group outcrops in Wyoming, New Mexico and Texas. They occur Otischalkian LVF in units of the lower part of the Chinle Group that have been correlated with each other on a lithostrati- Definition. The Otischalkian LVF is the time inter- graphic basis (Lucas 1993b). The most well-known val between the FADs of the phytosaurs Parasuchus principal correlative of the type Otischalkian fauna (¼ Paleorhinus) and Rutiodon (Fig. 1). Lucas & in the Chinle Group is the vertebrate-fossil assem- Hunt (1993a) proposed the Otischalkian LVF blage from the Popo Agie Formation of Wyoming, based on the vertebrate fossil assemblage of the Col- principally Fremont County (Branson & Mehl orado City Formation of the Chinle Group near the 1928; Mehl 1928; Colbert 1957; Lucas 1994; Lucas defunct town of Otis Chalk, Howard County, et al. 2002a) that includes the amphibian Buettneria, Texas, USA (Lucas & Anderson 1993a, b, 1994, the phytosaurs Parasuchus and Angistorhinus, the 1995; Lucas et al. 1993, 1994, 1997a). The begin- aetosaur Desmatosuchus, the archosaurs Popo- ning of the Otischalkian is the FAD of Parasuchus. saurus and Heptasuchus, the rhynchosaur Hypero- The end of the Otischalkian is the beginning of the dapedon, and the dicynodont Placerias. A less Adamanian, which is defined by the FAD of the well-known principal correlative is the small assem- phytosaur Rutiodon. blage from the Salitral Formation in Rio Arriba County, New Mexico that consists of a metoposaur, Characteristic tetrapod assemblage. The character- Longosuchus, a phytosaur, and an indeterminate istic tetrapod assemblage of the Otischalkian is the dinosaur (Lucas & Hunt 1992). Heckert (2004; assemblage of vertebrate fossils from just north of Heckert & Lucas 2006) provided some microverte- the defunct town of Otis Chalk in Howard County, brate basis for recognition of the Otischalkian in Texas. Lucas et al. (1993) reviewed the fauna, Chinle Group strata, such as the LO of the ‘dino- which is from the Colorado City Formation of the saur’ Protecovasaurus and the archosaur Trilopho- Chinle Group. The following taxa are present: the saurus buettneri (also see Spielmann et al. 2008). amphibians Latiscopus, Buettneria and Apache- In the Newark Supergroup of eastern North saurus, a procolophonid, the rhynchosaur Otis- America, the stratigraphically lower formations of chalkia, the archosaurs Doswellia, Trilophosaurus the Deep River, Gettysburg, Newark and Fundy (¼ Malerisaurus) and Poposaurus, the aetosaurs basins contain two distinct vertebrate fossil assem- Longosuchus (¼ Lucasuchus) and Coahomasuchus, blages. The older of these was used by Huber and the phytosaurs Parasuchus and Angistorhinus et al. (1993b) as the basis of the Sanfordian LVF, (Lucas et al. 1993; Long & Murry 1995; Heckert & after the characteristic assemblage from the Lucas 1999; Spielmann et al. 2006c). middle Pekin Formation in the Sanford sub-basin of the Deep River basin complex. An age-equivalent Index fossils. The following tetrapod genera are assemblage from the middle Wolfville Formation restricted to Otischalkian time and are widespread (Fundy basin) is also assigned to this LVF. The col- and/or common enough to be useful as index lective Newark tetrapod fauna of this Sanfordian fossils (Fig. 6): the aetosaur Longosuchus, and the LVF includes the amphibian Metoposaurus, proco- archosaur Doswellia. Parasuchus and Angistorhi- lophonids, the traversodontids Arctotraversodon nus are mostly of Otischalkian age, but also have and Plinthogomphodon, the dicynodont Placerias, early Adamanian records. The dicynodont Placerias the rhynchosaur Hyperodapedon, the archosaur 462 S. G. LUCAS

taxa Otischalkian Adamanian Revueltian Apachean amphibians: Apachesaurus Buettneria Metoposaurus phytosaurs: Angistorhinus Mystriosuchus Nicrosaurus Parasuchus Pseudopalatus Redondasaurus Rutiodon aetosaurs: Aetosaurus Desmatosuchus Longosuchus Paratypothorax Redondasuchus Rioarribasuchus Stagonolepis Typothorax others: Doswellia Eudimorphodon Hyperodapedon Placerias Revueltosaurus Riojasaurus

Fig. 6. Temporal ranges of selected genera of Late Triassic tetrapods.

Doswellia, the aetosaurs Desmatosuchus and Long- Lucas 1991; Lucas 1999; Schoch & Werneburg osuchus, indeterminate rauisuchians (‘Zamotus’), 1999; Hungerbu¨hler 2001b). the rauisuchian Postosuchus, the ‘sphenosuchian’ The500-m-thickIrohaleneMemberoftheTimes- Dromicosuchus, indeterminate phytosaur fragments gadiouine Formation (interval T-5 of Dutuit 1966; and fragmentary dinosaur remains (e.g. Cope 1871; Tixeront 1971) has produced most of the Late Trias- Olsen et al. 1989; Hunt & Lucas 1990; Huber et al. sic vertebrate fauna from Morocco. It contains the 1993a; Hunt 1993; Sues et al. 1994, 1999, 2003; majority of vertebrate fossil localities described by Langer et al. 2000b; Lucas et al. 2002a; Peyer Dutuit (1972, 1976, 1977, 1988, 1989a, b). Most of et al. 2008; Dilkes & Sues 2009). The Sanfordian these occur in the lower part of the member and correlates with the Chinle Group Otischalkian have produced a moderately diverse fauna that LVF based on the shared presence of Buettneria, includes the amphibians Almasaurus and Dutuito- Hyperodapedon, Desmatosuchus, Longosuchus, saurus, the phytosaur Parasuchus, the aetosaur Doswellia, and Placerias. Longosuchus, the dicynodont Placerias (¼ Mog- In Germany, the Schilfsandstein produces Meto- hreberia, ¼ Azarifeneria: Cox 1991; Lucas & Wild posaurus and Parasuchus but lacks Stagonolepis,so 1995), the dinosauriform Azendohsaurus (Gauffre it can be assigned an Otischalkian age (Hunt & 1993; Lucas 1998b; Jalil 1999) and at least one TRIASSIC TETRAPODS 463 ornithischian dinosaur. Several of Dutuit’s (1976) are regarded as Adamanian (as they were by Lucas localities occur in the upper part of the Irohalene 1998a), and the conchostracan-based correlations Member, which is a distinct faunal horizon that of the Adamanian are accepted, then records of includes the amphibian Arganasaurus, the phytosaur Parasuchus from the German Kieselsandstein and Angistorhinus, and the dicynodont Placerias. The Blasensandstein and the Polish Krasiejo´w locality presence of Parasuchus, Angistorhinus, Longosu- are Adamanian. This is also consistent with the chus and Placerias supports assigning the Irohalene Chinle Group record of Parasuchus at the Placer- Membertetrapod assemblage(s) an Otischalkian age. ias/Downs quarries in the Bluewater Creek For- In the Pranhita–Godavari Valley of India, the mation of the Chinle Group in Arizona, in what I basal Maleri Formation produces a tetrapod assem- have regarded as oldest Adamanian strata (Lucas blage that includes the amphibian Metoposaurus, et al. 1997a). Thus, recognizing that Parasuchus the rhynchosaur Paradapedon, the phytosaur Para- records are not strictly Otischalkian (some are suchus, the archosaur ‘Malerisaurus’, an aetosaur, early Adamanian: Fig. 6), and that Stagonolepis the theropod dinosaur Alwalkeria, a prosauropod records are strictly Adamanian, clarifies correlation (‘cf. Massospondylus’ of Kutty & Sengupta 1989), in the Otischalkian–Adamanian interval. a large dicynodont, and the cynodont Exeraetodon The Otischalkian index taxa Longosuchus (¼ (e.g. Huene 1940; Jain et al. 1964; Roychowdhury Lucasuchus) and Doswellia still stand. Metopo- 1965; Chatterjee 1967, 1974, 1978, 1980a, 1982, saurus also has only Otischalkian and early Adama- 1987; Chatterjee & Roychowdhury 1974; Jain & nian records, though Milner & Schoch (2004) Roychowdhury 1987; Bandyopadhyay & Sengupta recently claimed its presence in the Revueltian 2006; Spielmann et al. 2006c). This is the only well- Stubensandstein of Germany, a claim that met a described Upper Triassic tetrapod assemblage from detailed refutation from Lucas et al. (2007e). The the Pranhita–Godavari Valley. It includes Parasu- last Otischalkian index fossil listed by Lucas chus and Metoposaurus, taxa indicative of a likely (1998a) is the phytosaur Angistorhinus. Its records Otischalkian age. are Otischalkian (Long & Murry 1995) except one, near Lamy, New Mexico, USA, where it Comments. The Otischalkian LVF was originally co-occurs with Rutiodon in the earliest Adamanian defined as the time between the FADs of the phyto- (Hunt et al. 1993, 2005) (Fig. 7). saurs Parasuchus (¼ Paleorhinus) and Rutiodon (Lucas & Hunt 1993a; Lucas et al. 1997a; Lucas 1998a). It is important to note that a little advertised Adamanian LVF petition to the International Commission on Zoolo- gical Nomenclature by Chatterjee (2001) resulted in Definition. The Adamanian is the time interval establishing a diagnostic lectotype for Parasuchus between the FAD of the phytosaur Rutiodon and (long a nomen dubium: Hunt & Lucas 1991), so the FAD of the aetosaur Typothorax coccinarum that this name should be regarded as the senior (Fig. 1). Lucas & Hunt (1993a) based the Adama- synonym of Paleorhinus (Lucas et al. 2007c). Fur- nian LVF on the vertebrate fauna of the Blue thermore, even though Hunt & Lucas (1991) pro- Mesa Member of the Petrified Forest Formation in vided a careful taxonomic revision of Parasuchus, the Petrified Forest National Park, Arizona, USA and provided a clear diagnosis of the genus that (Lucas 1993b; Lucas & Hunt 1993a; Lucas et al. has never been contested, some taxonomists have 1997a). Lucas (1998a) termed this the Rutiodon relegated all primitive phytosaurs to a metataxon Assemblage Zone. The beginning of the Adamanian (grade) and then claimed these phytosaurs (long is defined as the FAD of the phytosaur Rutiodon. and widely known as Paleorhinus/Parasuchus) The end of the Adamanian is the beginning of the are of no value to biostratigraphy (e.g. Fara & Revueltian, which is defined by the FAD of the Hungerbu¨hler 2000; Rayfield et al. 2005, 2009). I aetosaur T. coccinarum. reject such an approach to primitive phytosaur tax- onomy and recognize Parasuchus as a diagnosable Characteristic tetrapod assemblage. The character- genus (Lucas et al. 2007c). istic tetrapod assemblage of the Adamanian is the I have long regarded Parasuchus as a robust assemblage of vertebrate fossils found in the Blue index taxon of the Otischalkian (Hunt & Lucas Mesa Member of the Petrified Forest Formation in 1991; Lucas et al. 2007c, d). However, recently the Petrified Forest National Park, near the defunct developed Upper Triassic conchostracan biostrati- railroad siding of Adamana, Arizona. Recent graphy (Kozur & Weems 2005, 2007) and European faunal lists have been provided by Murry & Long records of the characteristic Adamanian aetosaur (1989), Long & Murry (1995), Heckert et al. Stagonolepis suggest that some Parasuchus records (2005a) and Parker et al. (2006). The fauna includes should be considered early Adamanian in age (Kozur the following tetrapods: the amphibians Apache- & Weems 2005). Thus, if all Stagonolepis records saurus and Buettneria, the aetosaurs Desmatosuchus 464 S. G. LUCAS Petrified Forest National Park Chinle Group Bluewater Shinarump Creek Petrified Forest Formation Formation Blue Mesa Formation Sonsela Member Painted Member Desert Rainbow Agate Bridge Member Forest Bed Jim Camp Wash Bed Bed

Chinle Group east – central NM Formation Canyon

Santa Rosa Formation Garita Creek Trujillo Bull Tecolotito Los Esteros Tres Lagunas Formation Formation Member Member Member htsusatsusothers aetosaurs phytosaurs Rutiodon Paleorhinus Pseudopalatus Angistorhinus Stagonolepis Rioarribasuchus Desmatosuchus haplocerus chamaensis Paratypothorax Typothorax Placerias Typothorax antiquum coccinarum Poposaurus Revuletosaurus hunti R. callenderi Trilophosaurus buettneri Trilophosaurus jacobsi

ADAMANIAN REVUELTIAN LVF Barrancan sub- St. Johnsian Lamyan LVF

Fig. 7. Lithostratigraphy and tetrapod biostratigraphy of the Adamanian and Revueltian intervals in east–central New Mexico and in the Petrified Forest National Park, Arizona. The Lamyan interval is shaded (after Hunt et al. 2005).

(¼ Acaenasuchus), Stagonolepis, Adamanasuchus and the dicynodont Placerias, as well as many and Paratypothorax, Rutiodon-grade phytosaurs microvertebrate taxa. (including Leptosuchus and Smilosuchus), the raui- suchian Postosuchus, the archosaurs Hesperosu- Index fossils. The following tetrapod genera are chus, Acallosuchus, Parrishea and Vancleavea, restricted to Adamanian time and are widespread TRIASSIC TETRAPODS 465 and/or common enough to be useful as index fossils The following tetrapod taxa are known from the (Fig. 6): Rutiodon-grade phytosaurs, including Lep- Los Esteros Member, Santa Rosa Formation, near tosuchus and Smilosuchus, the trilophosaurid archo- Lamy, New Mexico: the amphibian Apachesaurus, saur Spinosuchus and the aetosaur Stagonolepis. The the phytosaurs Rutiodon and Angistorhinus, the HO of dicynodonts was long thought to be Adama- aetosaurs Desmatosuchus, Tecovasuchus and nian. However, there is a putative Cretaceous Stagonolepis and the dicynodont cf. Ischigualastia record from Australia (Thulborn & Turner 2003), (Hunt & Lucas 1993a, 1994; Hunt et al. 2005; and Dzik et al. (2008) recently reported a Triassic Heckert et al. 2007b). The overlying Garita Creek dicynodont from Poland in strata they deemed Rhae- Formation contains the following taxa: the amp- tian based on palaeobotany. The HO of the wide- hibian Buettneria, phytosaurs, rauisuchians, and spread rhynchosaur Hyperodapedon is Adamanian the aetosaurs Desmatosuchus, Stagonolepis and (Lucas & Heckert 2001; Lucas et al. 2002a) Paratypothorax (Hunt et al. 2005). (Fig. 8). Within the Chinle Group, various microver- The Tecovas Formation of West Texas yields the tebrate taxa, including Colognathus, Tecovasurus, following tetrapod taxa: the amphibians Buettneria and Crosbysaurus, are index taxa of the Adamanian and Apachesaurus, the probable tetrapod Colog- (Heckert 2004; Heckert & Lucas 2006). nathus, the archosauromorphss Trilophosaurus, Parrishea, Tecovasaurus, and Crosbysaurus, the Principal Correlatives. Besides the Chinle Group phytosaurs Rutiodon, Leptosuchus and Smilosuchus, correlatives, major Adamanian faunas are those the aetosaurs Desmatosuchus and Stagonolepis, the of the Conewagian interval of the Newark Super- rauisuchian Postosuchus, and the oldest known group basins of eastern North America; Lossie- mammal, Adelobasileus (Lucas & Luo 1993; mouth Sandstone Formation, Scotland; Lehrberg Lucas et al. 1994; Long & Murry 1995; Spielmann Schichten interval of the German Keuper; the Kra- et al. 2008). siejo´w locality in Poland; Ischigualasto Formation, In the Deep River basin of North Carolina, an Argentina; and upper Santa Maria Formation, assemblage of the Conewagian LVF from the Brazil. Cumnock Formation is superposed on the character- In the Chinle Group, Adamanian vertebrates are istic Sanfordian assemblage. Conewagian assem- widespread and include the vertebrate fossil assem- blages are characterized by the tetrapod assemblage blages of the Placerias and Downs’ quarries, Blue- in the basal Gettysburg Formation (Kozur & water Creek Formation, Arizona (Camp & Welles Weems 2010) along Little Conewago Creek in 1956; Kaye & Padian 1994; Long & Murry 1995; south-central Pennsylvania (Gettysburg basin: Lucas et al. 1997a; Heckert 2004; Heckert et al. Huber et al. 1993b; Sullivan et al. 1995; Lucas & 2005a); the Bluewater Creek Formation and Blue Sullivan 1997) and also are known from the Cow Mesa Member of the Petrified Foreset Formation Branch Formation (Dan River basin), and upper in the Blue Hills, Arizona; the Bluewater Creek For- Stockton and Lockatong formations (Newark mation and Blue Mesa Member of the Petrified basin). The most widespread and characteristic Con- Forest Formation, McKinley and Cibola counties, ewagian tetrapod is the phytosaur Rutiodon, which New Mexico (Heckert 1997); the Los Esteros and co-occurs with the amphibian Buettneria, archosaurs Tres Lagunas members, Santa Rosa Formation, of uncertain affinity, an aetosaur (Desmatosuchus), vicinity of Lamy, Santa Fe County, New Mexico one or more ‘ornithischian dinosaurs’ (e.g. Pekino- (Hunt et al. 2005) (Fig. 7); Garita Creek Formation, saurus and Galtonia), the archosaur Tanytrachelos Santa Rosa and vicinity, Guadalupe County, New (¼ ?Gwyneddosaurus) and the lepidosauromorph Mexico (Hunt & Lucas 1993a); and Tecovas For- Icarosaurus (e.g. Emmons 1856; Olsen 1980, mation, West Texas (Murry 1986, 1989; Long & 1988; Olsen et al. 1989; Sues 1992; Huber et al. Murry 1995). 1993a; Hunt 1993; Hunt & Lucas 1994; Doyle & The fauna at the Placerias and Downs’ quarries Sues 1995; Lucas & Huber 2003). Conewagian has most recently been discussed by Kaye & assemblages correlate with the Adamanan LVF of Padian (1994), Long & Murry (1995), Lucas et al. the Chinle Group, based on the shared presence of (1997a) and Heckert (2004). It includes the Buettneria, Rutiodon and other Rutiodon-grade amphibians Buettneria and Apachesaurus, the pro- phytosaurs (Smilosuchus of Long & Murry 1995), lacertiform Tanytrachelos, the phytosaurs Parasu- Desmatosuchus and broadly similar ‘ornithischian chus and Rutiodon/Leptosuchus, the aetosaurs dinosaurs’ (e.g. Murry & Long 1989; Lucas et al. Stagonolepis and Desmatosuchus (¼ Acaenasu- 1992, 1997a; Huber et al. 1993b; Hunt 1993; Hunt chus), the rauisuchid Postosuchus, the archosaurs & Lucas 1994; Heckert 2004). Trilophosaurus, Acallosaurus, Poposaurus, Chat- The tetrapod assemblage of the Lossiemouth terjeea, Hesperosuchus, Tecovasaurus and cf. Sandstone Formation of Grampian (Elgin) Scotland Uatchitodon, an indeterminate ceratosaur and the comes from small quarries and the coastal section at dicynodont Placerias. Lossiemouth. Benton & Spencer (1995) provided a Nova Mada- Stage lvf Wyoming Scotland India Zimbabwe Argentina Brazil Scotia gascar biochron of Otischalkian– Caturrita Formation Hyperodapedon Hyperodapedon assemblage zone assemblage Formation Sandstone Beds Lossiemouth Adamanian Isalo II Ischigualasto upper assemblage Formation S. G. LUCAS Pebbly Arkose Pebbly Carnian localities, which identify a Hyperodapedon biochron Maleri Formation lower (Alemoa Member) assemblage Santa Maria Formation Wolfville Formation Formation Popo Agie Popo Otischalkian Hyperodapedon Chinle Group

Hyperodapedon record Global correlation of Adamanian age. Fig. 8. 466 TRIASSIC TETRAPODS 467 detailed summary and indicate that all sites come the aetosaur Stagonolepis (¼ Aetosauroides); tra- from a narrow stratigraphic range, so I treat the ver- versodontids, proterochampsids; the archetypal tebrates as a single biostratigraphic assemblage. rauisuchian Rauisuchus and the primitive dinosaur It includes the procolophonid Leptopleuron, the Staurikosaurus (Barberena et al. 1985; Lucas 2002; sphenodontid Brachyrhinodon, the rhynchosaur Lucas & Heckert 2001; Langer et al. 2007). Clearly, Hyperodapedon, the aetosaur Stagonolepis, the the presence of Scaphonyx and Stagonolepis (‘Aeto- ornithosuchid Ornithosuchus, the crocodylomorph sauroides’) supports correlation with the vertebrates Erpetosuchus, the probable ornithodiran Scleromo- of the Ischigualasto Formation in Argentina, and chlus and the ‘dinosaur’ Saltopus. The presence of therefore an Adamanian (¼ Ischigualastian) age Hyperodapedon and Stagonolepis supports corre- (Lucas & Heckert 2001; Heckert & Lucas 2002c; lation of this assemblage to the Chinle Group Lucas 2002). Adamanian. The tetrapod assemblage of the Caturrita For- In Germany, the stratigraphic interval between mation, which overlies the Santa Maria Formation, the Schilfsandstein and the Stubensandstein (Lehr- includes a mastodonsauroid amphibian, the spheno- berg Schichten, Blasensandstein and Kieselsan- dont Clevosaurus, the rhynchosaur Hyperodapedon, dandstein) produces Stagonolepis, Parasuchus and the proterochampsid Proterochampsa, the dinosaurs Metoposaurus (e.g. Lucas 1999), and is assigned Guabisaurus and Saccasaurus, a phytosaur, the an Adamanian age (Kozur & Weems 2005). cynodonts Exaeretodon and Riograndia, the dicy- In Poland, the Krasiejo´w tetrapod assemblage nodont Ischigualastia (¼ Jachaleria) and cyno- includes the amphibians Cyclotosaurus and Meto- donts (Arau´jo & Gonzaga 1980; Barbarena et al. posaurus, the phytosaur Parasuchus, the aetosaur 1985; Dornelles 1990; Bonaparte et al. 1999, Stagonolepis, the rauisuchian Teratosaurus and the 2001; Kischlat & Lucas 2003; Ferigolo & Langer dinosauriform Silesaurus (Dzik 2001, 2003; Sulej 2006; Bonaparte & Sues 2006; Bonaparte et al. 2002, 2005, 2007; Sulej & Majer 2005; Dzik & 2007; Langer et al. 2007; Dias-da-Silva et al. 2009). Sulej 2007; Lucas et al. 2007d). This assemblage Most South American workers (e.g. Bonaparte is from strata c. 80 m above the Reed Sandstone 1982; Barberena et al. 1985; Langer 2005a; Rubert (a Schilfsandstein equivalent) that are homotaxial & Schultz 2004; Dias-da-Silva et al. 2007; Langer to the German Lehrberg Schichten and is of et al. 2007) advocate dividing the Brazilian Upper Adamanian age. Triassic tetrapod succession into two biostratigra- In Argentina, the Ischigualasto Formation is phically distinct assemblages largely based on their 500–900 m thick and consists of drab mudstones, judgment that the dicynodonts Jachaleria and Ischi- tuffs and sandstones that produce an extensive gualastia are not the same taxon. They, therefore, tetrapod assemblage including: the amphibian correlate the Brazilian Caturrita Formation to the Promastodonosaurus, the archosaurs Saurosuchus, Argentinian Los Colorados Formation. Langer Sillosuchus, and Proterochampsa, the aetosaur (2005b) also claimed that the Ischigualastian ¼ Stagonolepis (¼ Aetosauroides), the rhynchosaur Otischalkian þ Adamanian, largely based on not Hyperodapedon, the dinosaurs Herrerasaurus (¼ recognizing the temporal range of Hyperodapedon ?Ischisaurus ¼ Frenguellisaurus), Eoraptor and as longer than the temporal range of the Ischigua- Pisanosaurus, the chiniquodontid cynodont Chini- lastian. I do not accept either evaluation of the quodon, the gomphodont cynodonts Exeraetodon, Brazilian Upper Triassic tetrapod biostratigraphy Proexaraetodon, and Ischignathus and the dicyno- (Lucas 2002). dont Ischigualastia (e.g. Cabrera 1944; Reig 1959, In the Pranhita–Godavari Valley of India, the 1961, 1963; Casamiquela 1960, 1962; Cox 1965; upper vertebrate fossil assemblage from the Maleri Bonaparte 1976; Rogers et al. 1993; Sereno et al. Formation is stratigraphically above the lower 1993; Bonaparte 1997; Alcober & Parrish 1997; assemblage, but its stratigraphic range is not clear. Heckert & Lucas 2002c). The assemblage slightly This upper assemblage includes an aetosaur, pro- overlaps and mostly overlies the Herr Toba bentonite sauropods and a large dicynodont. Chigutisaurid that yielded a 40Ar/39Ar age of 227.8 + 0.3 Ma amphibians (Compsocerops and Kuttycephalus: (Rogers et al. 1993), which was ‘recalculated’ to Sengupta 1995) and a ‘Rutiodon-like’ phytosaur 231.4 Ma by Irmis and Mundil (2008). are also present (Bandyopadhyay & Sengupta In Brazil, the principal Upper Triassic vertebrate 2006). Therefore, this assemblage may be Adama- assemblage from the Santa Maria Formation is nian, but needs further documentation. from the vicinity of Santa Maria City. This is the In western Madagascar, the Isalo group (‘Groupe Rhynchocephalia assemblage zone of Barberena d l’Isalo’ of Besarie 1930; also see Besarie & (1977) or the Scaphonyx assemblage of Barberena Collignon 1960, 1971) has long been divided into et al. (1985), from the upper part of the Santa Isalo I, Isalo II and Isalo III based on perceived geo- Maria Formation. The assemblage consists of abun- logical age. The Isalo II strata yield Late Triassic dant fossils of the rhynchosaur Hyperodapedon and tetrapods, including metoposaurs, sphenodontids, 468 S. G. LUCAS phytosaurs, the rhynchosaur Hyperodapedon, the pattern of its dorsal paramedian osteoderms’, con- aetosaur Desmatosuchus, the archosaur Azendoh- trary to the published work of Lucas & Heckert, as saurus, cynodonts and dicynodonts (Guth 1963; well as those of Long & Ballew (1985), Parrish Westphal 1970; Dutuit 1978; Buffetaut 1983; Flynn (1994), Long & Murry (1995) and Parker (2007), et al. 1999, 2000, 2008; Langer et al. 2000a; among others. Lucas et al. 2002a; Burmeister et al. 2006). The Langer (2005b) also used the conclusions of stratigraphic range of the Isalo II tetrapods is Sulej (2002) regarding the taxonomy of Metopo- c. 1200 m, but the rhynchosaur Hyperodapedon is saurus and Buettneria to question using amphibians one of the stratigraphically lowest taxa in the assem- to distinguish the Otischalkian and Adamanian. blage. This means the Isalo assemblage is no older However, a review of the metoposaur specimens than Otischalkian and, based on the Desmatosuchus described by Sulej (2002) does not support some record, likely to be Adamanian. of his basic anatomical observations or his taxon- omy (Lucas et al. 2007d). Rayfield et al. (2005, Comments. Lucas (1998a) listed as Adamanian 2009) also argued for merging of the Otischalkian index fossils the rhynchosaur Scaphonyx, the aeto- and Adamanian based largely on the same argu- saur Stagonolepis and Rutiodon-grade phytosaurs ments as Langer (2005b), but Lucas et al. (2007e) (including Leptosuchus and Smilosuchus). The have presented a detailed refutation of their dicynodont Ischigualastia (¼ Jachaleria) was also arguments. considered an Adamanian index taxon. Taxonomic What these workers have failed to recognize is revisions and range extensions have necessitated a that: (1) Otischalkian and Adamanian tetrapod reconsideration of some of these index taxa. assemblages are stratigraphically superposed and Stagonolepis now co-occurs with Parasuchus at readily distinguished in the Chinle Group of the Krasiejo´w in southern Poland (Dzik 2001; Lucas American Southwest; (2) there is no evidence that et al. 2007d). This lends support to Heckert & the ‘Ischigualastian’ of South America is Otischalk- Lucas’ (2000) conclusion that Ebrachosaurus sin- ian and much more evidence that it is Adamanian, gularis Kuhn 1936, from the Adamanian German so Ischigualastian should not be redefined to encom- Blasensandstein (type destroyed in World War II), pass both Otischalkian and Adamanian time; and (3) was based on specimens of Stagonolepis. These recognition of distinct Otischalkian and/or Adama- European Adamanian records of Stagonolepis are nian assemblages has been achieved in North consistent with regarding its stratigraphically lowest America, South America, Europe, India and North records in North America, such as at the Placerias/ Africa (e.g. Fig. 8). The fact that Langer (2005b) Downs quarries in Arizona, as early Adamanian and Rayfield et al. (2005, 2009) do not accept a (Lucas et al. 1997a). well-documented alpha taxonomy of Otischalkian An extensive revision of Late Triassic rhyncho- and Adamanian index fossils is not a valid reason saurs (Langer & Schultz 2000; Langer et al. to merge the Otischalkian and Adamanian LVFs. 2000a, b) indicates that specimens previously Recent work in the Chinle Group of the western assigned to Scaphonyx are mostly of Hyperodape- USA has refined the stratigraphic ranges of known don. Lucas et al. (2002a) reviewed these records tetrapod taxa and has produced new records in in detail and demonstrated that a Hyperodapedon strata of Adamanian age. These new data are princi- biochron is of Otischalkian and Adamanian age pally from the Petrified Forest National Park in (Fig. 8). Thus, the rhynchosaur Hyperodapedon Arizona (Heckert & Lucas 2002a; Hunt et al. cannot be used to discriminate the Otischalkian 2002; Woody 2003, 2006; Heckert 2004; Woody & and Adamanian. Parker 2004; Heckert et al. 2005a) and the extensive Largely based on this, Langer (2005a, b; also see exposures of the Chinle Group in east–central Schultz 2005) claimed that the Otischalkian and New Mexico (Lucas et al. 2001, 2002b), though Adamanian cannot be distinguished and they should there are also other new records from the Tecovas be abandoned and replaced by a single LVF, the and Trujillo formations in Texas (Heckert 2004; Ischigualastian. To do so, Langer (2005b) dismissed Heckert et al. 2006; Martz & Small 2006). phytosaur-based distinctions of the Otischalkian and Clearly, there is a ‘transitional’ fauna between the Adamanian, basing his rejection largely on the cla- Adamanian and Revueltian LVFs (Woody & dotaxonomy of primitive phytosaurs advocated in Parker 2004), and this prompted Hunt et al. (2005) published abstracts by Hungerbu¨hler (2001a; Hun- to subdivide the Adamanian into two sub- gerbu¨hler & Chatterjee 2002). Langer (2005b) also faunachrons, St. Johnsian (older) and Lamyan rejected aetosaur-based correlations based on the (younger), of regional biochronological significance taxonomy of South American aetosaurs published (Fig. 7). The aetosaur Tecovasuchus is a St. Johnsian by Heckert & Lucas (2000) and Lucas & Heckert index taxon (Heckert et al. 2007b), whereas the (2001). Thus, Langer (2005b, p. 228) states that aetosaur Typothorax antiquum is a Lamyan index ‘Stagonolepis wellesi lacks a unique ornamentation taxon (Hunt et al. 2005). TRIASSIC TETRAPODS 469

Heckert & Lucas (2006) built upon the micro- see Dalla Vecchia 2009). The stratigraphic co- vertebrate collections documented by Heckert occurrence of dinosaurs and dinosauromorphs (2001, 2004) to demonstrate that there are multiple (Sullivan & Lucas 1999; Ezcurra 2006; Irmis et al. microvertebrate index taxa of Adamanian 2007; Nesbitt et al. 2007, 2009; Spielmann et al. (St. Johnsian) time, including the xenacanth ‘Xena- 2007b; Nesbitt & Chatterjee 2008) also aids in canthus’ moorei, the enigmatic tetrapod Colog- recognition of Revueltian time. nathus obscurus and the archosaurs (possibly ornithischian dinosaurs) Tecovasaurus murryi, Principal correlatives. Besides Chinle Group Crosbysaurus harrisae, and Krzyzanowskisaurus assemblages, which are primarily from Texas, New hunti. So far, these taxa are presently known only Mexico and Arizona (e.g. Zeigler et al. 2003; from the Chinle Group of the American Southwest, Heckert et al. 2005a, b; Parker et al. 2006; so they may not be of broad biostratigraphic utility. Spielmann et al. 2007a, b; Nesbitt & Stocker 2008), the principal Revueltian tetrapod assem- Revueltian blages are those of the Newark Supergroup of eastern North America of Neshanician and Clifto- Definition. The Revueltian is the time interval nian (part) age; Ørsted Dal Member of the Fleming between the FAD of the aetosaur Typothorax cocci- Fjord Formation, Greenland; Stubensandstein (Lo¨w- narum and the FAD of the phytosaur Redonda- enstein Formation) of the German Keuper; Calcare saurus (Fig. 1). Lucas & Hunt (1993a) introduced di Zorzino (Zorzino Limestone) and Dolomia di the term Revueltian LVF to refer to the time equiv- Forni (Forni Dolomite), northern Italy; and lower alent to the vertebrate fossil assemblage of the Bull part of Dharmaran Formation, India. Canyon Formation in east–central, New Mexico, In West Texas-eastern New Mexico, the Bull USA (Lucas et al. 1985; Hunt 1994, 2001; Hunt & Canyon Formation of the Chinle Group yields exten- Lucas 1997). Lucas (1998a) termed this the Pseudo- sive assemblages of Revueltian tetrapods, including palatus Assemblage Zone. The name of the LVF is the characteristic tetrapod assemblage (e.g. Hunt for Revuelto Creek, one of the key collecting areas 2001; Lehman & Chatterjee 2005). In the Chama in eastern New Mexico. Revueltian time begins basin of north–central New Mexico, the Petrified with the FAD of the aetosaur T. coccinarum. The Forest Formation of the Chinle Group also yields end of the Revueltian is the beginning of the Apa- Revueltian tetrapods, especially from the Snyder chean, which is defined by the FAD of the phytosaur and Canjilon phytosaur-dominated bonebeds Redondasaurus. (Zeigler et al. 2003; Heckert et al. 2005b; Nesbitt & Stocker 2008). In northern Arizona, two Chinle Characteristic tetrapod assemblage. The character- Group units, the Painted Desert Member of the Pet- istic tetrapod assemblage of the Revueltian is that of rified Forest Formation and the overlying Owl the Bull Canyon Formation in east–central New Rock Formation, have produced numerous Revuel- Mexico (Quay and Guadalupe counties), and the tian fossils, especially from the Petrified Forest following taxa are present: the amphibian Apache- National Park and from localities on Ward’s saurus, the turtle Chinlechelys, the phytosaur Terrace north of Flagstaff (e.g. Kirby 1989, 1991, Pseudopalatus and other Pseudopalatus-grade 1993; Heckert et al. 2005a; Spielmann et al. 2007a). phytosaurs, the aetosaurs Rioarribasuchus, Paraty- In eastern North America, the provincial Nesha- pothorax, Typothorax coccinarum, and Aetosaurus, nician LVF is based on a limited fossil assemblage the suchian Revueltosaurus, the ‘dinosaur’ Luciano- typified by the aetosaur Aetosaurus arcuatus saurus, the rauisuchian Postosuchus, the chatter- (Lucas et al.1998a; Lucas & Huber 2003). This jeeids Shuvosaurus (¼ Effigia) and Chatterjeea, taxon is present in ‘Lithofacies Association II’ of the sphenosuchian Hesperosuchus; and the cynodont the Chatham Group (Durham sub-basin of the Pseudotriconodon (e.g. Hunt 1994, 2001; Lucas Deep River basin), the Newark Basin (range zone: et al. 2001; Joyce et al. 2009). Warford through Neshanic Members of the lower Passaic Formation), and the middle New Haven Index fossils. The following tetrapod taxa are Arkose of central Connecticut. Other vertebrates restricted to Revueltian time and are widespread from the Neshanician LVF include indeterminate and/or common enough to be useful as index metoposaurid and phytosaur teeth, skull and scute fossils: the crurotarsan Revueltosaurus, the aeto- fragments (e.g. ‘Belodon validus’), a rauisuchian, saurs Aetosaurus, Rioarribasuchus and Typothorax crocodylomorph, traversodontid and a sphenodontid coccinarum, and Pseudopalatus-grade phytosaurs. (lower New Haven Arkose) as well as a dominance The pterosaur Eudimorphodon is present in Revuel- of the primitive neopterygian Semionotus sp. over tian assemblages in Italy and Greenland (e.g. Jenkins other fish taxa, a trend also apparent in age- et al. 2001; Dalla Vecchia 2003, 2006) and can equivalent strata of the Chinle Group and German also be considered a Revueltian index taxon (but Keuper (Huber et al. 1993c; Lucas & Huber 2003). 470 S. G. LUCAS

The Cliftonian LVF is based on a low-diversity phytosaurs, aetosaurs, and rauisuchians provide a assemblage defined by the distribution of the proco- strong basis for assigning a Revueltian age to the lophonid Hypsognathus fenneri. This taxon is Lower Stubensandstein (Lucas & Hunt 1993a; common in the type area, from the middle (?Mettlars Hunt 1994; Lucas 1999). The younger, Middle and Member) to the upper (?Member TT) Passaic For- Upper Stubensandstein, produce a similar, but less mation of the northern Newark basin (e.g. Baird diverse assemblage, so I also assign them a Revuel- 1986). It is also known from the upper New Haven tian age. Whether or not the lowest occurrence of Arkose of the Hartford basin, central Connecticut, Mystriosuchus in the Middle Stubensandstein is of and the basal Blomidon Formation in the Fundy biochronologic significance is not clear. The assem- basin, Nova Scotia (Sues et al. 1997). The Fundy blages of the Upper Stubensandstein and Knollen- basin specimen of Hypsognathus was obtained mergel (Tro¨ssingen Formation) are almost entirely from pebble conglomerate at the base of the Blomi- dinosaurian – 95% or more of the fossils are of don Formation, which unconformably overlies the dinosaurs (Benton 1986, 1991). This contrasts Wolfville Formation. The only other vertebrates sharply with the Lower and Middle Stubensandstein that occur in the interval of Cliftonian age are inde- assemblages, in which dinosaurs are a much smaller terminate phytosaur remains (including the holotype percentage of the fossils collected. However, I of ‘Clepsysaurus pennsylvanicus’ Lea 1851) from regard this change to dinosaur domination as the Ukrainian Member of the Passaic Formation in largely a local facies/taphonomic effect, not a bio- the Newark basin, moderately diverse tetrapod foot- chronologically significant event (Hunt 1991). It print assemblages at many horizons in the Passaic seems likely, but not certain, that the Knollenmergel Formation (e.g. Szajna & Silvestri 1996; Lucas & assemblage is of Apachean age (see below). Sullivan 2006), and an indeterminate sphenodontid In the Lombardian Alps of northern Italy, after from the upper New Haven Arkose (Olsen 1980; the regional progradation of platform carbonates Sues & Baird 1993; Lucas & Huber 2003). during the early-middle Norian (Dolomia Princi- The Malmos Klint and overlying Ørsted Dal pale), extensional tectonism produced intraplatform Members of the Fleming Fjord Formation in depressions occupied by patch reefs, turbiditic eastern Greenland yield fossil tetrapods of Revuel- debris flows and lagoonal to freshwater facies tian age (Jenkins et al. 1994, 1997, 2008). The (Jadoul 1985; Jadoul et al. 1994). Tetrapods from Malmos Klint Member has produced fragmentary these intraplatform strata, the Zorzino Limestone at fossils of plagiosaurid amphibians, the amphibian the Cene and Endenna quarries in Lombardy, are Cyclotosaurus, possible phytosaur fragments and the the diapsids Endennasaurus and Vallesaurus, the prosauropod dinosaur Plateosaurus. The Ørsted Dal prolacertiform Longobardisaurus, the rhynchoce- Member assemblage is much more diverse: the phalian Diphydontosaurus, the drepanosaurids amphibians Gerrothorax and Cyclotosaurus, the Drepanosaurus and Megalancosaurus, the phyto- turtle cf. Proganochelys, unidentified sphenodon- saur Mystriosuchus, the aetosaur Aetosaurus, the tians, the aetosaurs Aetosaurus and Paratypothorax, pterosaurs Eudimorphodon and Peteinosaurus and the pterosaur Eudimorphodon, the prosauropod the placodont Psephoderma (e.g. Wild 1989; Pinna dinosaur ‘Plateosaurus’, a theropod dinosaur, ther- 1993; Renesto 2006). In Germany, Mystriosuchus opod dinosaur footprints (Grallator), and the is well known from the Middle Stubensandstein mammals Kuehneotherium, cf. ?Brachyzostrodon and Aetosaurus from the Lower-Middle Stubensand- and Haramiyavia. As Jenkins et al. (1994) argued, stein, so a Revueltian age of the Zorzino Limestone this assemblage shares many taxa with the German is certain. The Calcare di Zorzino also crops out Stubensandstein. More specifically, other than Pla- in Austria, where it yields specimens of Langobardi- teosaurus, most taxa from the Ørsted Dal Member saurus and the pterosaur Austriadactylus, a likely are known in the Lower Stubensandstein, to which synonym of Preondactylus (Dalla Vecchia 2009; I correlate the Greenland assemblage. S. Renesto, written commun. 2009). Also, in Austria, In Germany, the best known and most diverse unpublished specimens of Mystriosuchus are known Keuper tetrapod assemblage is that of the Lower fromTotesGebirge(possiblyDachstein)(S.Renesto, Stubensandstein (Lo¨wenstein Formation). This written commun. 2009). assemblage includes the amphibians Cyclotosaurus The other Italian Late Triassic tetrapod sites are and Gerrothorax, the earliest European turtles (Pro- in the Forni Dolomite (Dolomia di Forni) in the ganochelys and Proterochersis), Pseudopalatus- Veneto Prealps of northeastern Italy. They yield grade phytosaurs (Nicrosaurus), the aetosaurs the drepanosaurids Drepanosaurus and Megalanco- Aetosaurus and Paratypothorax, rauisuchians (Ter- saurus, and the pterosaurs Eudimorphodon and Pre- atosaurus), theropod dinosaurs, and the prosauro- ondactylus (Dalla Vecchia 1995) and a specimen of pod dinosaurs Sellosaurus and Thecodontosaurus Langobardisaurus under study by S. Renesto (e.g. Benton 1993; Hungerbu¨hler 1998; Lucas 1999; (written commun. 2009). The presence of Eudimor- Schoch & Werneburg 1999; Schoch 2007). The phodon supports a Revueltian age assignment. TRIASSIC TETRAPODS 471

Upper Triassic tetrapod assemblages from the biochronology in that the beginning of a LVF can Indian Subcontinent come from the Pranhita– be defined by a true species-level evolutionary Godavari Valley of south–central India. Several event, not the appearance of a genus-level taxon. summaries (Jain et al. 1964; Kutty 1969; Kutty & Aetosaurus is one of the most robust index fossils Roychowdhury 1970; Sengupta 1970; Jain & of the Triassic tetrapod timescale (Fig. 9). Lucas Roychowdhury 1987; Yadagiri & Rao 1987; Kutty et al. (1998b) presented a detailed taxonomic revi- et al. 1988; Kutty & Sengupta 1989; Bandyopadhyay sion based on study of all North American and Euro- & Roychowdhury 1996; Bandyopadhyay & Sen- pean specimens. Aetosaurus has a marine record in gupta 2006) have been published, but other than the middle Norian of northern Italy (Wild 1989), the lower Maleri assemblage (see above), relatively and all of its nonmarine records are Revueltian. Cri- few of the fossils have been adequately documented ticism of the use of Aetosaurus, typified by Rayfield in print, forcing me to rely largely on unsubstantiated et al. (2005, 2009), is based on the claim that genus-level identifications to evaluate the ages of the because Aetosaurus has been portrayed as the ple- tetrapod assemblages. A case in point is the Dhar- siomorphic sister taxon of other aetosaurs in cladis- maram Formation, which yields two stratigraphi- tic analyses (e.g. Heckert & Lucas 2000) it ‘must’ cally discrete vertebrate fossil assemblages (lower have a long ghost lineage that therefore renders it and upper). The stratigraphic range of the lower useless in biostratigraphy. I regard this as specious assemblage has not been published, and it includes cladotaxonomic reasoning (Lucas et al. 1999a, a phytosaur that Kutty & Sengupta (1989, table 2) 2007c, e). Thus, the position of a taxon on a clado- list as Nicrosaurus, aetosaurs, including a so-called gram has nothing to do with its biostratigraphic ‘Paratypothorax-like’ form, and prosauropod dino- utility unless all the assumptions of the cladogram – saurs. Based primarily on the supposed Nicrosaurus and the existence of a ghost lineage is nothing more record, I consider the lower assemblage of the Dhar- than an assumption – are brought into the biostrati- maram Formation a possible Revueltian correlative. graphic analysis. Indeed, an alternative interpret- ation of the Heckert & Lucas (2000) cladogram of Comments. Hunt & Lucas (1993c) suggested that, aetosaurs, one that views Aetosaurus as a highly perhaps along the lines of the Cliftonian- derived, dwarfed and simplified form, would Neshanician subdivision used in the Newark Super- produce a very different ‘ghost lineage’. group, the Revueltian merits subdivision, and Hunt Aetosaurus thus is a taxonomically stable and (1994, 2001) subdivided it into three sub-LVFs of robust Revueltian index fossil (e.g. Fraas 1877; regional utility. Two of these, the Barrancan (early Huene 1921; Walker 1961; Wild 1989; Parrish Revueltian) and Lucianoan (later Revueltian) are 1994; Heckert et al. 1996, 2007a; Heckert & Lucas readily correlated in the western USA using 1998; Small 1998; Lucas et al. 1998b, 1999a; various index fossils (e.g. Heckert & Lucas 2006). Heckert & Lucas 2000; Parker 2007). Some of the discussion of the Revueltian has Pseudopalatus-grade phytosaurs include Pseudopa- focused on whether or not it is readily distinguished latus, Nicrosaurus and Mystriosuchus, all taxa from the next younger Apachean LVF (Long & restricted to Revueltian time. Like the use of Murry 1995; Rayfield et al. 2005, 2009). These dis- Rutiodon-grade phytosaurs to identify the Adama- cussions are rooted in taxonomic arguments, as the nian, this is a convenient and concise way to refer type assemblages of the Revueltian and Apachean to a group of broadly contemporaneous phytosaur are stratigraphically superposed in east–central taxa whose stratigraphic ranges are well established, New Mexico, USA and thus are obviously time but whose genus- and species-level nomenclature successive. remain in flux (compare, e.g. the differing phytosaur Typothorax, Aetosaurus and Pseudopalatus- taxonomies of Ballew 1989; Hunt 1994; Long & grade phytosaurs were listed as Revueltian index Murry 1995; and Hungerbu¨hler 2002). fossils by Lucas (1998a). However, recognition of Heckert & Lucas (1997) suggested that Revuel- an older, Adamanian species of Typothorax, T. anti- tosaurus might serve as an index taxon of Revuel- quum, by Lucas et al. (2002b) has modified this; it is tian time. At that time Revueltosaurus, which was the species T. coccinarum that is a Revueltian index known solely from teeth, was considered to be an fossil, and this is part of what prompted Hunt et al. ornithischian dinosaur. Parker et al. (2005) docu- (2005) to redefine the beginning of the Revueltian as mented associated skulls and postcrania of Revuel- the FAD of T. coccinarum, a decision followed by tosaurus callenderi, demonstrating that that taxon Lucas et al. (2007e) and also used here. is actually a crurotarsan archosaur. However, they Typothorax coccinarum stands as a robust index noted that, following Hunt (1989), Padian (1990) fossil of the Revueltian across the Chinle Group. and others, the teeth are indeed diagnostic, and Indeed, its likely descent from T. antiquum as part the taxon is valid. Heckert & Lucas (2006) then of an anagenetic evolutionary lineage (Lucas et al. showed that in the Chinle Group Revuletosaurus is 2002b) is significant to the Triassic tetrapod restricted to strata of Revueltian age. 472 S. G. LUCAS

Newark Supergroup, eastern USA New Mexico- Italy sub- ammonoid Eastern STAGE Colorado Germany (Lombardian Biochron stage zones Hartford Newark Deep River Greenland Basin (CT) Basin (NJ) Basin (NC) USA Alps) Q Rock Point Zorzino Formation Limestone Himavatites columbianus Neshanic Member Aralalta middle

(holotype) Group Lithofacies Drepanites Aetosaurus arcuatus Perkasie Association rutherfordi Member III Middle (Alaunian)

LM Orsted/ Dal K Bull New Canyon Member Juvavites I Formation magnus Haven

Passaic Formation (part) Formation Passaic (part) Arkose Stubensandstein Graters Fleming Fjord Formation Member Steinmergel Keuper Dolomia NORIAN Principale Lithofacies Aetosaurus biochron Malayites ET Association Chinle Group (part) dawsoni II lower Early (Lacian)

Sitkinoceras Warford kerri Member Malmros Trujillo Klint Formation Member C

Aetosaurus occurrences

Fig. 9. Global correlation of Aetosaurus localities, which identify an Aetosaurus biochron of Revueltian age.

This demonstrates the irrelevance of the with the FAD of the phytosaur Redondasaurus. assumed position of a taxon in a phylogeny to bios- The end of Apachean time is the beginning of the tratigraphy. The changing phylogenetic position of Wassonian LVF, which is the FAD of the crocodylo- Revueltosaurus alters neither its biostratigraphic morph Protosuchus (Lucas & Huber 2003; Lucas & significance nor its biochronological utility. What Tanner 2007a, b). is biostratigraphically important about Revuelto- saurus is that it is distinctive (easily identified), Characteristic tetrapod assemblage. The character- relatively common and/or widespread, and known istic tetrapod assemblage of the Apachean LVF is from a restricted stratigraphic interval. Whether it from the Redonda Formation of the Chinle Group is an ornithischian (as previously supposed) or a in Guadalupe and Quay Counties, New Mexico, crurotarsan (the current phylogenetic hypothesis) USA. The following taxa are present: the amphibian is irrelevant to its biostratigraphic and biochronolo- Apachesaurus, a sphenodontid, a procolophonid, gical utility. the phytosaur Redondasaurus, the aetosaur Redon- dasuchus, the rauisuchian Redondavenator, the sphenosuchian Vancleavea, a rauisuchian, theropod Apachean dinosaurs and a ?cynodont (e.g. Hunt 1994; Hunt & Definition. The Apachean LVF is the time interval Lucas 1993b, 1997; Heckert et al. 2001; Hunt et al. between the FAD of the phytosaur Redondasaurus 2005; Spielmann et al. 2006a, b). and the FAD of the crocodylomorph Protosuchus Index fossils. The following tetrapod genera are (Fig. 1). Lucas & Hunt (1993a) introduced the restricted to Apachean time and are widespread term Apachean LVF to refer to the time equivalent and/or common enough to be useful as index to the vertebrate fossil assemblage of the Redonda fossils: the phytosaur Redondasaurus, the aetosaur Formation (Chinle Group) in east–central New Redondasuchus and the dinosaur Riojasaurus. Mexico, USA (Lucas et al. 1985; Hunt 1994; Hunt & Lucas 1997; Lucas 1998a; Lucas et al. 2001; Principal correlatives. Principal correlatives of the Spielmann et al. 2006a, b). Apachean time begins type Apachean assemblage are the Whitaker TRIASSIC TETRAPODS 473 quarry in the Rock Point Formation of the Chinle been considered Late Triassic. Lucas & Hancox Group at Ghost Ranch, New Mexico, the Cliftonian (2001) reviewed the age of this assemblage, which LVF assemblages (in part) of the Newark Super- is dominated by sauropodomorph dinosaurs, but group, the Knollenmergel (Tro¨ssingen Formation), also has rare amphibians (a large chigutisaurid), a time-equivalent upper Arnstadt Formation and the possible rauisuchian (Basutodon), the ornithischian ‘Rhaetian Bonebed’ of the Germanic Basin, the Col- dinosaur Eocursor, a traversodontid (Scaleno- oradan LVF of Argentina and the tetrapod assem- dontoides) and the characteristic Late Triassic foot- blage of the Lower Elliot Formation in South print ichnogenus Brachychirotherium (Kitching & Africa. Some of the fissure-fill assemblages in the Raath 1984; Lucas & Hancox 2001; Butler et al. uppermost Mercia Mudstone Group and/or lower- 2007). This is the ‘Euskelosaurus range zone’ of most Penarth Group of the United Kingdom Kitching & Raath (1984), the youngest Triassic tet- (Fraser 1994; Benton & Spencer 1995; Whiteside & rapod assemblage in the Karoo basin. Yates (2003) Marshall 2008) may be Apachean correlatives. re-evaluated the prosauropods of the Lower Elliott Some of the so-called Rhaetian vertebrate sites in Formation and concluded that most are indetermi- France, such as Saint-Nicolas-de-Port, may be Apa- nate sauropodomorphs or basal sauropods. He chean correlatives as well (Lucas & Huber 2003). noted similarities of indeterminate prosauropods At Ghost Ranch in New Mexico, the Whitaker from the Lower Elliott Formation to Riojasaurus quarry bone bed is dominated by skeletons of the from the Los Colorados Formation of Argentina, theropod dinosaur Coelophysis bauri (Colbert and similarities between the basal sauropod Anteto- 1989). Nevertheless, it also includes the sphenodont nitrus from South Africa and Lessemsaurus from Whitakersaurus, at least one drepanosaur, a rauisu- Argentina (Yates & Kitching 2003). These con- chian skeleton (cf. Postosuchus), the sphenosuchians clusions suggest a Lower Elliott–Los Colorados Hesperosuchus and Vancleavea, the chatterjeeid correlation, and thus a tentative Apachean age Shuvosaurus (¼ Effigia) and the phytosaur Redonda- assignment. saurus (e.g. Hunt & Lucas 1993b; Clark et al. 2000; In the United Kingdom, fissure fills such as Harris & Downs 2002; Hungerbu¨hler 2002; Hunt Durdham Down in Clifton yield fossils that include et al. 2002; Lucas et al. 2003; Rinehart et al. 2004; phytosaurs, aetosaurs, dinosauriforms and dinosaurs Nesbitt 2007; Lucas et al. 2005, 2007e; Heckert (e.g. Fraser 1994; Fraser et al. 2002; Galton 2005, et al. 2008; Renesto et al. 2009). 2007a, b; Whiteside & Marshall 2008). Unfortu- In Argentina, the Los Colorados Formation con- nately, other than a tentative record of Aetosaurus sists of siliciclastic red beds approximately 800 m based on a single osteoderm (Lucas et al. 1999b), thick. Near its base, a single tetrapod fossil – a dicy- the fissure fill tetrapods are mostly endemic taxa of nodont skull, the holotype of ‘Jachaleria’ colorata no biochronological significance or cosmopolitan Bonaparte 1970 – was collected. The remainder of taxa with long age ranges, such as the sphenodontian the tetrapod fossils from the Los Colorados Clevosaurus. Recently, Whiteside & Marshall Formation are from its middle and upper parts but (2008), based primarily on the palynoflora, assigned have not been stratigraphically organized. The the Tytherington fissure fill a Rhaetian age, and assemblage includes the turtle Palaeochersis, the extrapolated this age to the other fissures. If this ornithosuchid Riojasuchus, the aetosaur Neo- Rhaetian age is correct, then the fissure fill tetrapods aetosauroides, the rauisuchid Fasolasuchus, the are of Apachean age. However, as Lucas & Hunt crocodylomorphs Hemiprotosuchus and Pseudhe- (1994, p. 340) noted, ‘a single age should not necess- sperosuchus, the prosauropod dinosaurs Riojasaurus arily be assigned to the fossils from one fissure and Coloradisaurus, the theropod dinosaur Zupay- and .... individual fossils from the fissures may saurus and the tritheledontid cynodont Chaliminia range in age from middle Carnian to Sinemurian’. (e.g. Bonaparte 1970, 1971, 1978, 1980, 1997; Therefore, I continue to regard as problematic the Lucas & Hunt 1994; Rougier et al. 1995; Arcucci precise age of the Triassic tetrapod assemblages et al. 2004). The correlative Quebrada del Barro from the British fissure fills. and El Tranquilo formations also produce prosauro- pods (e.g. Riojasaurus,‘Mussaurus’) (Casamiquela Comments. The Apachean is the most difficult 1980; Bonaparte & Vince 1979; Bonaparte & Triassic LVF to correlate globally. This almost cer- Pumares 1995). The Los Colorados assemblage tainly reflects a provincialization of the global tetra- clearly is of Late Triassic age (Arcucci et al. 2004) pod fauna near the end of the Triassic. Some of the and must be post-Ischigualastian. However, its apparent endemism of Apachean land-vertebrate endemism makes it difficult to correlate precisely. assemblages may also be due to facies, sampling I tentatively consider it an Apachean correlative and taphonomic biases. Thus, rather than recognize based primarily on its abundant prosauropods. a global Apachean LVF, it may ultimately be The age of the tetrapod assemblage from the necessary to recognize two or more provincial Lower Elliott Formation in South Africa has long LVFs during this time interval. 474 S. G. LUCAS

There is no evidence that any part of the Apa- some basis for correlation of the LVFs to the standard chean is of Jurassic age. The FAD of the crocody- global chronostratigraphic scale (Fig. 10). Neverthe- lomorph Protosuchus, which defines the beginning less, reliable data for this correlation remain rela- of the next LVF, the Wassonian, appears to corre- tively sparse, so the correlation of the LVFs to the spond closely to the beginning of the Jurassic SGCS is still imprecise in many time intervals. (Lucas & Tanner 2007a, b). Thus, Protosuchus occurs in units assigned an Early Jurassic based on Lootsbergian diverse evidence: the McCoy Brook Formation (Newark Supergroup), the upper Stormberg Group The base of the Triassic (¼ Permo-Triassic bound- of South Africa and the upper part of the Dinosaur ary [PTB], ¼ base of Induan Stage) has been for- Canyon Member of the Moenave Formation in mally defined by the FAD of the conodont Utah-Arizona (Colbert & Mook 1951; Sues et al. Hindeodus parvus at a global stratotype section 1996; Lucas et al. 2005; Lucas & Tanner 2007a, and point (GSSP) located at Meishan in southern b). The Moenave record of Protosuchus is stratigra- China (Yin et al. 2001). This means it is possible phically superposed above Apachean body fossil to attempt to correlate a potential Triassic base in assemblages of the uppermost Chinle Group the nonmarine section to a fixed, agreed-upon (Lucas et al. 1997b, 2005; Lucas & Tanner 2007a, point in the marine timescale. b). Furthermore, it is correlative to the Lower Juras- It is important to ask how the Lootsbergian cor- sic conchostracan assemblages from the Whitmore relates to the marine PTB in order to establish the Point Member of the Moenave Formation (Lucas synchrony or diachroneity of marine and nonmarine & Tanner 2007a; Kozur & Weems 2010). Relatively events across the PTB. However, such correlation is recent recognition that Apachean-age strata extend not simple, because no sections are known where above the Chinle Group into part of the strata bearing terrestrial tetrapods can be directly Moenave–Wingate (lower Glen Canyon Group) correlated (say by interfingering lithostratigraphy) lithosome has been based, in part, on the occurrence to the marine record across the PTB. Thus, magne- of a Redondasaurus skull in the lower part of the tostratigraphy, isotope stratigraphy, conchostracan Wingate Sandstone in southeastern Utah (Lucas biostratigraphy and palynostratigraphy have been et al. 1997b; Lucas & Tanner 2007a, b). (Note used to correlate the nonmarine and marine that Spielmann et al. 2007a, fig. 8A–B illustrated records across the PTB. Lucas (2009) provides a a cast of this skull and mistakenly attributed it to detailed discussion of this correlation, which is the Revueltian Owl Rock Formation). briefly reviewed here. Lucas (1998a) listed three Apachean index There is a well documented negative d13C excur- fossils: the aetosaur Redondasuchus, the phytosaur sion at the PTB in marine sections that closely Redondasaurus and the dinosaur Riojasaurus. coincides with the major extinction that precedes Riojasaurus is known from Argentina and may be the formally-defined PTB (e.g. Payne et al. 2004; present in the Lower Elliott Formation in South Yin et al. 2005, 2007; Richoz 2006). Diverse ana- Africa. The Apachean is readily distinguished in lyses indicate that the marine PTB is within the North America by its primary index fossils, Redon- lower third of a long normal-polarity chron (e.g. dasaurus and Redondasuchus. However, some Ogg 2004; Steiner 2006; Hounslow & Muttoni workers (Long & Murry 1995; Martz 2002) have 2010). Palynostratigraphy has also been used by questioned the validity of Redondasaurus and some to correlate marine to nonmarine sections at Redondasuchus, proclaiming the former a synonym the PTB (e.g. Morante 1996; Looy et al. 1999, of Pseudopalatus and the latter a synonym of 2001; Twitchett et al. 2001; Collinson et al. 2006), Typothorax. Nevertheless, Heckert et al. (2001) particularly the fungal abundance spike documented and Spielmann et al. (2006a, b) reaffirmed the in marine and nonmarine sections that some have distinctiveness and validity of Redondasuchus and considered to correspond to the PTB marine mass Redondasaurus, and Hungerbu¨hler (2002) also extinction (e.g. Eshet et al. 1995; Visscher et al. recognized Redondasaurus as distinct from 1996; Steiner et al. 2003). Pseudopalatus. In the conchostracan biostratigraphy, which is well correlated with the marine scale, the PTB Correlation of the LVFs to the Marine coincides with the boundary between the Falsisca postera Zone and the Falsisca verchojanica Zone SGCS (Kozur 1998a, b, 1999; Bachmann & Kozur 2004; Introduction Kozur & Weems 2010). As in the marine section, this conchostracan zonal boundary lies in the lower Records of nonmarine Triassic tetrapods in marine third of a long normal magnetostratigaphic zone that strata (Lucas & Heckert 2000), palynostratigraphy, straddles the PTB, and it is characterized by a magnetostratigraphy and radioisotopic ages provide minimum in d13C in continental beds (Bachmann & TRIASSIC TETRAPODS 475

J Hettangian Wassonian 201 Rhaetian Apachean

210 Norian Revueltian Aetosaurus, Drepanosaurus, Edennasaurus, Eudimorphodon, Macrocnemus, Megalancosaurus, Mystriosuchus,Peteinosaurus, Preonodactylus, Sikkanisuchus

220 Adamanian Late

Carnian

230 Otischalkian Metoposaurus, Parasuchus TRIASSIC

Berdyankian cf. Cyclotosaurus, aff. Dinodontosaurus, Ladinian Mastodonsaurus 240

Eocyclotosaurus, Lotosaurus Perovkan Anisian Middle Aphaneramma, Lyrocephaliscus, Parotosuchus, Peltostega, Sasenisaurus, Tertrema 250 Olenekian Nonesian

Early Induan Luzocephalus, Stoschiosaurus, 252 Lootsbergian Permian Changxingian Platbergian Tupilakosaurus, Wetlugasaurus

Fig. 10. Marine records of Triassic nonmarine tetrapod correlated to the marine SGCS and the Triassic land–vertebrate faunachrons. See Lucas & Heckert (2000) for details. Restoration of Eocyclotosaurus by Matt Celeskey.

Kozur 2004; Korte & Kozur 2005b). This minimum At the Meishan section in southern China, a in d13C occurs in continental lakes without facies sharp drop in d13C values coincides with the changes, and the conchostracan boundary occurs maximum amount of marine extinction, and this in the Dalongkou section in northwestern China mass extinction and carbon isotope excursion are close to the HO of Dicynodon (Kozur 1998a, b; older than the PTB defined by the lowest occurrence Metcalfe et al. 2009; Kozur & Weems 2010). of the conodont Hindeodus parvus (Fig. 11). 476 S. G. LUCAS

Fig. 11. Magnetostratigraphic correlation of marine PTB section at Meishan, China (based on Yin et al. 2005) to PTB tetrapod extinction interval in Karoo basin of South Africa (based on Ward et al. 2005). The sections are correlated at the base of the normal polarity magnetozone that encompasses the PTB. However, they are not scaled to each other based on time intervals or section thickness, so the only certain point of correlation indicated is the base of the normal polarity magnetozone that encompasses the PTB. From Lucas (2009). Restoration of Lystrosaurus by Matt Celeskey.

However, in sections without a weathered boundary The PTB marine extinction took place during clay (e.g. Shahreza, Iran, and Gerenava´r, Bu¨kk Mts, a relatively long interval of normal magnetic Hungary) or without a boundary clay, the d13C polarity that straddles the PTB, well documented minimum lies around the FAD of H. parvus, at the in a variety of marine sections (Ogg 2004; Steiner PTB (Korte & Kozur 2005a). In the Karoo basin 2006) as well as in continental sections (Szurlies of South Africa, d13C isotope data through the 2004; Bachmann & Kozur 2004). In the Karoo PTB have been used to correlate to the marine basin there is an interval of normal polarity that d13C excursion (MacLeod et al. 2000; Ward et al. encompasses the highest occurrence of Dicynodon 2005). However, these isotope data do not convin- and is part of the stratigraphically thick (c.60m) cingly support the conclusion that the highest occur- interval of low d13C values (Schwindt et al. 2003; rence of Dicynodon in the Karoo basin is equivalent De Kock & Kirschvink 2004; Ward et al. 2005; to the PTB. Indeed, Tabor et al. (2007) recently pub- Steiner 2006) (Fig. 11). These magnetostratigraphic lished an analysis of d13C across the PTB in the data indicate that the lowest occurrence of Lystro- Karoo basin and argued that changes in that record saurus (in an interval of reversed polarity) is older are driven by local facies changes and are not a than the PTB (as already suggested by King & reflection of atmospheric carbon values. Therefore, Jenkins 1997; Kozur 1998a, b; and Botha & Smith the d13C record in the Karoo basin cannot be reliably 2007, among others), and that the highest occur- correlated to the d13C record in marine strata across rence of Dicynodon is closer to the PTB (Fig. 11). the PTB. However, at Dalongkou in northwestern I favour the magnetostratigraphic correlation of China the HO of Dicynodon is close to the PTB the Meishan and Karoo sections, and it is consistent defined and correlated to the marine scale by with all other available correlation data. The corre- conchostracans. Therefore, the minimum in d13C lation indicates that in the Karoo basin the base of in the Karoo basin may be a primary signal, and the Lootsbergian (LO of Lystrosaurus) pre-dates the HO of Dicynodon is close to the PTB. the main marine extinction event. The LO of TRIASSIC TETRAPODS 477

Lystrosaurus cannot be used to place the PTB in reported a generically-indeterminate trematosaurid nonmarine sections; the highest occurrence of jaw from the South African Lootsbergian strata Dicynodon is a much better proxy for the PTB. and claimed it extends Lootsbergian time up to the Thus, the earliest Lootsbergian is of latest late Olenekian, largely because of its resemblance Permian (Changshingian) age. Correlation of the to Olenekian Trematosaurus. An equally likely rest of the Lootsbergian to at least part of the possibility is that Damiani et al. (2000) simply marine Induan Stage is clear (Lucas 1998a; Lucas documented an Induan-age trematosaurid. Thus, et al. 2007e). However, whether the Lootsbergian the possibility exists that Lootsbergian time is as equates to part, all or more than Induan time is not young as early Olenekian, but no reliable data possible to determine with the available data. The are known to support a Lootsbergian–Olenekian Wordy Creek Formation in eastern Greenland has correlation. a record of Lootsbergian amphibians interbedded with marine late Griesbachian–early Dienerian Nonesian (middle Induan) age strata. Thus, the stratigraphi- cally lowest record of Luzocephalus here is in the Cross correlation of the Nonesian to at least part of Ophiceras commune ammonite zone, and the genus the Olenekian is clear because of the occurrence of extends up through the ‘Proptychites rosenkrantzi the Nonesian index temnospondyl Parotosuchus in Zone’. Most of the Wordy Creek Formation amphi- marine upper Olenekian (Spathian) strata in the bians come from the younger ‘Anodontophora Mangyshlak Peninsula of western Kazakstan. Thus, fassarensis beds’, which are the youngest Lower from Mangyshlak, Lozovsky & Shishkin (1974) Triassic strata in this section (Nielsen 1935; documented Parotosuchus sequester from marine Sa¨ve-So¨derbergh 1935). This indicates a range of upper Olenekian (Spathian) strata that yield Tiro- Luzocephalus from late Griesbachian through early lites and other ammonites. Parotosuchus is an Dienerian (middle Induan), but the other temnospon- index taxon of Nonesian time, and the Kazak dyl taxa are of middle or late Dienerian (late Induan) record thus provides a direct Nonesian-late Olene- age (Tru¨mpy 1961; Silberling & Tozer 1968; Tozer kian correlation. Furthermore, a Spathian conchos- 1994). Luzocephalus, Tupilakosaurus, and Wetlu- tracan fauna of the Germanic Basin in the gasaurus occur in the Vokhmian Horizon of the Hardegsen Formation (with Parotosuchus) is well Vetlugian Series of the Russian Urals. This fauna correlated with marine beds in Hungary (with includes Lystrosaurus, an index taxon of the Loots- Spathian ammonoids) and northern Siberia (Kozur bergian land–vertebrate faunachron, so the amphi- & Weems 2010). bian records from Greenland establish a middle– In the western USA, the Nonesian Torrey For- late Induan age for at least part of Lootsbergian time. mation of the Moenkopi Group overlies the early In northwestern Madagascar the upper part of Olenekian (Smithian) ammonite-bearing Sinbad the marine Andavakoera Formation (Dienerian) Formation, whereas the Nonesian Wupatki yields a diverse assemblage of temnospondyls: Member of the Moenkopi Formation is clearly ?Benthosuchus,?Wetlugasaurus, Mahavisaurus, younger than the late Olenekian (Spathian) Virgin Aphaneramma, Ifasaurus, Tertrema, Tertremoides, Limestone (e.g. McKee 1954; Blakey 1974; Trematosaurus, Wantzosaurus and Deltacephalus Morales 1987; Steiner et al. 1993; Lucas & Schoch (Swinton 1956; Lehman 1961, 1966, 1979). The 2002; Goodspeed & Lucas 2007; Lucas et al. Benthosuchus and Wetlugasaurus identifications 2007a). This suggests a Smithian–Spathian (Olene- are not reliable (Cosgriff 1984; Damiani 2001), kian) age for the Moenkopi Nonesian tetrapods, and though the amphibians from the Andavakoera supports a broad Nonesian–Olenekian correlation. Formation may be of Lootsbergian age. This may The Sticky Keep Formation in Svalbard yields indicate correlation of part of the Lootsbergian amphibians that co-occur here with early Olenekian and the Dienerian. (Smithian) ammonites (Buchanen et al. 1965; Tozer Shishkin (2000, p. 65) asserted that the Lootsber- 1967). The amphibians are: Sasenisaurus, Peltos- gian includes assemblages younger than Induan, but tega, Aphaneramma, Lyrocephaliscus, Teretrema no credible data support his claim. For example, he and Boreaosaurus (Wiman 1910, 1915; Nilsson stated (p. 65) that ‘the Hesshanggou assemblage of 1942, 1943; Cox & Smith 1973). These tremato- China [which Lucas 1998a assigned a Lootsbergian saurs are believed to have been euryhaline amphi- age] ...is actually latest Spathian or Spathian– bians that may have actually lived in marine Anisian in age’. However, there is no direct way environments. They also reflect a high diversity to correlate Hesshanggou Formation red beds in and abundance of trematosaurs characteristic of Shanxi (long correlated by Chinese workers to the the Nonesian. However, the Svalbard trematosaur ‘Procolophon zone’ of the Karoo: Cheng 1981) to taxa are mostly endemic and thus only provide the SGCS (Lucas 1993a, 1998a, 2001; Lucas et al. stage-of-evolution evidence for an Olenekian– 2007e). In another case, Damiani et al. (2000) Nonesian cross-correlation. 478 S. G. LUCAS

Perovkan from the Upper Muschelkalk through the Letten- keuper, strata of late Ladinian (Longobardian) age A fairly direct correlation can be made of some (Schoch 1999). Perovkan tetrapod assemblages to the SGCS The lower Ladinian Partnach Formation of (Lucas & Schoch 2002). Thus, strata of the Ro¨t For- western Austria yielded a temnospondyl jaw frag- mation (Upper Buntsandstein) in southwestern ment that Sander & Meyer (1991) identified as Germany–eastern France are lower Anisian mar- cf. Cyclotosaurus sp. However, this specimen ginal marine to interbedded nonmarine/marine could just as well belong to Mastodonsaurus facies of well-established age because of their (cf. Schoch 1999), so it is of limited biochrono- close physical relationship to the Lower Muschelk- logical significance. alk. Indeed, marine facies of the lower Ro¨t contain Stur (1873) reported ?Mastodonsaurus giganteus early Anisian conodonts, the early Anisian from the Lunz Sandstone in the Austrian Alps. This (Aegean) ammonoid Beneckeia tenuis and age- is an early Carnian (Julian) record, broadly correla- diagnostic holothurian sclerites (e.g. Kozur 1993), tive to the German Schilfsandstein. However, I and magnetostratigraphic correlation of the Ro¨t have examined the material Stur described, and it Formation to marine magnetostratigraphy indicates is not diagnostic of M. giganteus; it could just as an early Anisian age (Szurlies 2007; Hounslow et al. well belong to Cyclotosaurus. Therefore, this record 2008). Furthermore, conchostracans of the Ro¨t also is of limited biochronologic significance. correlate with Aegean and lower Bithynian marine The Brazilian and Argentinian Dinodontosaurus intercalations (Kozur & Weems 2010). The assemblages are unambiguously correlated to each common amphibian from the Ro¨t Formation, Eocy- other, and have generally been considered Ladinian clotosaurus, is an index taxon of the Perovkan found based on flimsy palynostratigraphic evidence (Lucas in both Europe and the western United States (e.g. & Harris 1996; Lucas 2002). Tetrapod evidence to Ortlam 1970; Morales 1987; Lucas & Schoch correlate the Dinodontosaurus assemblages to the 2002). The Ro¨t records of Eocyclotosaurus thus European Berdyankian is also not robust; it consists provide a Perovkan–early Anisian correlation. of fragmentary remains of Dinodontosaurus-grade The Gogolin Formation (lowermost Muschel- and Stahleckeria-grade dicynodonts from the kalk) in Polish Silesia yields fragmentary temnos- German Muschelkalk and Russian Bukobay For- pondyl and archosaur fossils that include the types mation, respectively, not on shared alpha taxa of Mentosaurus waltheri, Eurycervix posthumus, (Lucas & Wild 1995; Lucas 1998a; Lucas et al. and ‘Xestorrhytias perrini’, all of which are indeter- 2007b). At present, this South American–European minate mastodonsaurids, and the rauisuchian Zan- correlation remains weakly supported and merits clodon silesiacus Jaekel, based on a single tooth. further study. This is one area where magnetostrati- Ammonite biostratigraphy places the Gogolin For- graphy (in South America) will be of assistance. mation in the lower Anisian (e.g. Kaim & Niedz´- Thus, all available robust data for correlating wiedzki 1999). The tetrapod material, however, is the Berdyankian to the SGCS indicate that it is too fragmentary to be of much biochronological equivalent to the late Ladinian. This may indicate utility. Nevertheless, the available material closely that there is a global gap equivalent to the early resembles some of the tetrapods from the Upper Ladinian in the Triassic tetrapod record. Buntsandstein (Ro¨t Formation) of southwestern Germany–eastern France, and thus supports a Otischalkian Perovkan–early Anisian correlation. Magentostratigraphic correlation of the Perov- There are two records of Otischalkian tetrapod kan Otter Sandstone in Great Britain indicates it is index taxa in marine strata in Austria that support of late Anisian age (Hounslow & McIntosh 2003). an Otischalkian–Carnian correlation: Abdala et al. (2005) assigned the Perovkan Cynog- 1. Raibler Schichten, Austria: Koken (1913) nathus zone C in the Karoo basin to the late Anisian described Metoposaurus santaecrucis from a based largely on the palynological content of its conglomeratic sandstone in the upper part of probable correlatives, such as the Wianamatta the Raibler Schichten. This is an early Carnian Group in the Sydney basin of Australia. Thus, (Julian) record, and thus correlates part of the there is good evidence that the Perovkan is equival- Otischalkian (index taxon ¼ Metoposaurus) to the ent to most of Anisian time. early Carnian. 2. Opponitzer Schichten, Austria: Huene (1939) Berdyankian described a skull fragment of the phytosaur Parasu- chus (¼ Francosuchus) from the lower part of the The German section provides the best data for a Opponitzer Schichten (Kalk) near Lunz, Austria. Berdyankian–Ladinian correlation. Thus, the Ber- The occurrence is of late Carnian (Tuvalian) age dyankian taxon Mastodonsaurus giganteus ranges (Janoscheck & Matura 1980), but it cannot be tied TRIASSIC TETRAPODS 479 precisely to a particular ammonite zone (Hunt & 1993a). Correlations to the Chinle Group based on Lucas 1991). palynomorphs, conchostracans and tetrapods indi- In Germany, Otischalkian tetrapods from the cate that the Adamanian LVF is older than the Schilfsandstein are as old as early Carnian (late base of the Passaic Formation. Based on counting Julian). Palynostratigraphy assigns a late Carnian cycles in the Newark, the estimated age of the age to the lower Chinle Group, including the strata Passaic Formation base (and the base of the of Otischalkian age, and an early Carnian age to Norian) is about 217 Ma (Kent & Olsen 1999), but the oldest Sanfordian strata of the Newark Super- in this counting a complete Rhaetian was assumed. group (Litwin et al. 1991, 1993; Cornet 1993). However, according to Kozur & Weems (2005, Sequence stratigraphy of the Chinle Group advo- 2007), most of the Rhaetian is missing in the cated by Lucas (1991, 1993b), Lucas & Marzolf Passaic Formation, where only the uppermost pre- (1993) and Lucas & Huber (1994) assigns lower cession cycle of c. 20 000 years yielded uppermost Chinle Group strata to a single sequence, the Shinar- Rhaetian conchostracans, whereas below these ump–Blue Mesa sequence. This sequence can be beds late Norian conchostacans are present. correlated to a late Carnian marine sequence in Recent correlations of Newark magnetostratigra- Nevada (Lupe & Silberling 1985; Lucas & Huber phy, however, have been used to argue for a much 1994), and recent studies of detrital zircons are con- older Norian base in the Newark section (Muttoni sistent with these correlations (Dickinson & Gehrels et al. 2004), one that would be close to the base of 2008; Dickinson et al. 2009). Magnetostratigraphy the Lockatong Formation, with an estimated age correlates lower Chinle Group strata to the late of c. 228 Ma based on Newark cycle counting. Fur- Carnian portion of the Newark Supergroup magne- thermore, in an abstract, Irmis & Mundil (2008) tostratigraphy (Kent et al. 1995; Molina-Garza reported a 206Pb/238U age of 219.2 + 0.7 Ma for et al. 1996; Muttoni et al. 2004). Therefore, the an Adamanian horizon of the Chinle Group in Otischalkian clearly is Carnian, equivalent to the west–central New Mexico. On face value, the early Carnian and part of the late Carnian. Chinle date and the interpretation of Newark mag- netostratigraphy of Muttoni et al. (2004) indicate Adamanian that the Adamanian is Norian. Nevertheless, the correlations Muttoni et al. I have long considered the Adamanian to be of late (2004) propose between the Newark and the marine Carnian age based on palynostratigraphy, sequence Late Triassic magnetostratigraphy from Pizzo Mon- stratigraphy and magnetostratigraphy (see refer- dello are fraught with problems, mostly because the ences cited above under marine cross-correlation marine section contains far fewer magnetochrons of the Otischalkian). In West Texas, Otischalkian than does the presumed age-equivalent interval of and Adamanian tetrapod assemblages are stratigra- the Newark. Furthermore, the correlation has aban- phically superposed (Lucas 1993b; Lucas & doned the only well-documented biostratigraphic Anderson 1993a, b, 1994, 1995; Lucas et al. 1993, datum in the Newark that allows a correlation to 1994). Therefore, Adamanian time is younger than marine strata: the Carnian–Norian boundary at the the Otischalkian. Revueltian vertebrates are strati- approximate base of the Passaic Formation (see graphically above Adamanian vertebrates in above). Thus, the proposed Pizzo Mondello-Newark Arizona, New Mexico and Texas. Therefore, Ada- magnetostratigraphic correlation lacks an indepen- manian vertebrates are either the youngest Carnian dent biostratigraphic datum by which to correlate. vertebrates known or the oldest Norian vertebrates Furthermore, the Pizza Mondelo marine section is known (or both). thin (c. 430 m of limestone-dominated section rep- Kozur & Weems (2007, 2010) discussed at length resent late Carnian and much of Norian time) in com- the biostratigraphic evidence to support a late parison to the more than 4-km-thick Newark section. Carnian (Tuvalian) correlation of the Adamanian. Therefore, it is not surprising that the Pizzo Mon- This is the concordance of three biostratigraphies dello section yields a magnetostratigraphic record – palynostratigraphy, conchostracan biostratigra- that does not directly correspond, in both reversal phy and vertebrate biostratigraphy – that all indi- frequency and pattern, to the Newark section. I cate that the Adamanian is Tuvalian. Particularly thus believe there is real reason to question the significant is the record in the Newark Supergroup reliability of the magnetostratigraphic correlations of eastern North America, where, for decades, paly- advocated by Muttoni et al. (2004). nostratigraphy placed the Carnian–Norian bound- I maintain a late Carnian (Tuvalian) age for the ary at or just above the base of the Passaic Adamanian, choosing biostratigraphic data over Formation (at the Warford Member), a placement what I judge to be less reliable correlations based supported by conchostracan and tetrapod biostrati- on magnetostratigraphy. As for the date reported in graphy (and by megafossil plant biostratigraphy: an abstract by Irmis & Mundil (2008), without sup- Ash 1980, 1987) (see summary by Huber et al. porting data its reliability cannot be fully evaluated. 480 S. G. LUCAS

However, if it is a reliable age, it dates part of the late Muttoni et al. 2004; Hounslow & Muttoni 2010, Carnian to c. 219 Ma, which means the base of the this volume), also suggests the Apachean is latest Norian would be younger than 219 Ma, in agreement Triassic (‘Norian–Rhaetian’). with the 217 Ma age suggested by Kent & Olsen Earlier arguments that the Apachean is equival- (1999), and the Norian is not as long as concluded ent to the Rhaetian (Hunt 1993; Lucas 1993b, by Muttoni et al. (2004). 1998a) cannot be sustained in the light of new data. These arguments were largely based on a Revueltian stage-of-evolution assessment of the Apachean phytosaur Redondasaurus. This phytosaur is more Two Italian records are critical to correlation of the derived than the Knollenmergel (late Norian) phyto- Revueltian to part of the Norian: saurs of the German Keuper, so Redondasaurus was 1. Zorzino Limestone, Lombardian Alps, Italy: therefore assigned a Rhaetian age. However, the The Zorzino Limestone (Calcare de Zorzino) has Norian aetosaur Aetosaurus occurs in Rock Point been correlated to the mid-Norian (uppermost Alau- strata in Colorado (Small 1998) and Rock Point nian) Himavatites columbianus ammonite zone strata in New Mexico, and the Rock Point palyno- (Jadoul et al. 1994; Roghi et al. 1995). Nonmarine morphs suggest a Norian age (Litwin et al. 1991). tetrapods from this unit at the Ceˆne and Endenna Clearly, the Apachean is younger than the Revuel- quarries in Lombardy include the Revueltian index tian (early–middle Norian), so I regard it as late taxa Mystriosuchus, Aetosaurus and Eudimorpho- Norian to Rhaetian in age (Lucas et al. 2005, don (Wild 1989; Renesto 2006). 2007e; Lucas & Tanner 2007a, b). 2. Forni Dolomite, Veneto Prealps, Italy: the The stratigraphically highest Apachean assem- Forni Dolomite (Dolomia di Forni) in northeastern blage from the American Southwest is in the Dino- Italy is the same age as the Zorzino Limestone, mid- saur Canyon Member of the Moenave Formation Norian (Roghi et al. 1995). Its nonmarine tetrapods and laterally equivalent Wingate Sandstone (Lucas include the Revuletian index taxon Eudimorphodon et al. 2005, 2006; Lucas & Tanner 2007a, b). (Dalla Vecchia 1995). The Italian records thus There are several compelling reasons to assign a provide direct evidence that at least part of the Late Triassic age to this assemblage: (1) the Apa- Revueltian ¼ middle Norian (Alaunian). I consider chean phytosaur Redondasaurus is present, and the Revueltian to correlate approximately with the no phytosaur is known from Jurassic strata; (2) early-middle Norian, which is consistent with the the footprint ichnogenus Brachychirotherium is Italian data (Lucas 1997a). present and not known anywhere from Jurassic Palynostratigraphy, magnetostratigraphy and strata; (3) the lower Dinosaur Canyon Member is sequence stratigraphy suggest the characteristic laterally equivalent to strata of well established Revueltian tetrapod assemblage in the Chinle Late Triassic age (upper Rock Point Formation of Group of New Mexico, USA, is of Norian age the Chinle Group); (4) the Wingate Formation (Lucas 1997a, 1998a). Based on stratigraphic pos- basal contact is gradational with underlying Upper ition (Huber et al. 1993b; Lucas & Huber 2003), Triassic strata of the Rock Point Formation; and magnetostratigraphy (Witte et al. 1991; Kent et al. (5) magnetostratigraphy of the Dinosaur Canyon 1995; Muttoni et al. 2004), and palynomorphs interval is reasonably correlated to the magnetostra- (Cornet 1977), the Neshanician LVF in the Newark tigraphy of uppermost Triassic strata of the Newark Supergroup of eastern North America is of early to Supergroup in eastern North America (Molina- middle Norian or just of middle Norian age. Strati- Garza et al. 2003). graphic position (Huber et al. 1993b; Lucas & Although it is possible to assign the Dinosaur Huber 1993), magnetostratigraphy (Witte et al. Canyon assemblage to the Late Triassic, its precise 1991; Kent et al. 1995; Muttoni et al. 2004), and correlation to the marine timescale is uncertain. palynomorphs (Cornet 1977; Fowell & Olsen Probably it equates to part or all of Rhaetian time, 1993; Lucas & Tanner 2007b) indicate the Cliftonian simply because the Dinosaur Canyon interval is the LVF is of late Norian–Rhaetian age. Thus, a Norian youngest Triassic interval on the Colorado Plateau correlation of the Revueltian is certain, with well and is conformably overlain by strata that apparently supported correlation to the early and middle Norian. correlate to the earliest part of the Early Jurassic (Hettangian) (Lucas & Tanner 2007a, b; Kozur & Apachean Weems 2010). This supports a correlation of the Apachean with the late Norian–Rhaetian. Apachean time is post-Revueltian (c. mid-Norian) and pre-Jurassic. Magnetostratigraphy of the upper- Triassic Footprint Biostratigraphy most Chinle Group in the Four Corners and in eastern New Mexico (Reeve & Helsley 1972; Molina-Garza In this volume, Klein & Lucas (2010) present a et al. 1996, 2003), correlated to the Newark Super- Triassic footprint biostratigraphy and biochronol- group magnetostratigraphy (Kent et al. 1995; ogy that build on, revize and synthesize previous TRIASSIC TETRAPODS 481 efforts, including those of Haubold (1969, 1971, LVFs; (3) changes and additions to the stratigraphic 1984, 1986), Demathieu & Haubold (1972, 1974), ranges of some index taxa; and (4) perceived pro- Olsen (1980, 1983), Lockley & Hunt (1995), Hunt blems of correlation to the SGCS. & Lucas (2007a, b), Lucas (2003, 2007a) and Klein Taxonomic disagreements lie at the heart of & Haubold (2007). Triassic tetrapod footprints many arguments over biostratigraphy, but I have a Pangaea-wide distribution; they are known believe that the extensive taxonomies developed from North and South America, Greenland, Europe, for many of the Triassic index taxa, especially tem- North Africa, China, Australia, Antarctica and nospondyls, phytosaurs, aetosaurs, dicynodonts and South Africa. They often occur in nonmarine Triassic cynodonts, provide a sound basis for their use in strata that lack well-preserved body fossils, so their biostratigraphy. Shifting opinions about taxonomy biostratigraphic utility has been of some interest. of these tetrapods will remain, and that will In Triassic strata, several characteristic footprint always affect correlations based on tetrapod fossils. assemblages and ichnotaxa have restricted strati- Lucas et al. (2007e) resolved the problems of graphic ranges and thus represent distinct time inter- potential overlap or gaps around the Nonesian–Per- vals. Key Triassic footprint ichnotaxa are archosaur ovkan boundary by redefining the beginning of the tracks: Rotodactylus, the chirotherian ichnotaxa Perovkan to obviate such problems. Stratigraphic Protochirotherium, Synaptichnium, Isochirother- range extensions and changes are the regular out- ium, Chirotherium, Brachychirotherium and gralla- growth of collecting and careful biostratigraphic torids (theropod dinosaur tracks). Nevertheless, study in the field. They always force adjustments non-archosaur footprints are common, especially to any biochronological scheme rooted in sound the ichnotaxa Rhynchosauroides, Procolophonich- biostratigraphy. Problems with correlation of the nium, Capitosauroides and several dicynodont- Triassic LVFs to the SGCS persist largely because related or mammal-like forms that dominate some in much of the nonmarine Triassic section few footprint assemblages. reliable data are available for correlation to the From the temporal distribution pattern Klein & marine timescale. Lucas (2010) identified five distinct tetrapod- Clearly, we need a nonmarine Triassic tetrapod footprint-based biochrons: (1) dicynodont tracks biochronology with which to correctly sequence (Lootsbergian); (2) Protochirotherium (Synaptich- the history of tetrapod evolution on land. Advances nium): also includes Rhynchosauroides and Pro- in the scheme proposed in the 1990s have come colophonichnium (Nonesian); (3) Chirotherium from new fossil discoveries, more detailed biostrati- barthii, also includes C. sickleri, Isochirotherium, graphy and additional alpha taxonomic studies Synaptichnium, Rotodactylus, Rhynchosauroides, based on sound evolutionary taxonomic principles. Procolophonichnium, dicynodont tracks and As the work reviewed here demonstrates, the Capitosauroides (Nonesian–Perovkan); (4) Atrei- global Triassic timescale based on tetrapod biochro- pus–Grallator (‘Coelurosaurichnus’), which also nology remains a robust tool for both global and includes Synaptichnium, Isochirotherium, regional age-assignment and correlation. Sphingopus, Parachirotherium, Rhynchosauroides, Procolophonichnium (Perovkan–Berdyankian); The synthesis reported here is based on work funded by and (5) Brachychirotherium, which also inclu- the National Geographic Society, DAAD and the Janet des Atreipus–Grallator, Grallator, Eubrontes, Stearns Memorial Trust. I am grateful for the collaboration Apatopus, Rhynchosauroides, dicynodont tracks of A. Heckert, P. Huber, A. Hunt, J. Spielmann and L. Tanner on many problems of Triassic tetrapod taxonomy (Otischalkian–Apachean). and biostratigraphy. I thank A. Heckert, H. Kozur, J. Ogg, Tetrapod footprints are thus useful for Triassic S. Renesto, J. Spielmann and R. Weems for their careful biostratigraphy and biochronology, but, compared and helpful reviews of the manuscript. to the tetrapod body fossil record with eight bio- chrons, the five footprint-based biochrons provide less temporal resolution. 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