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Revista de la Asociación Geológica Argentina 63 (4): 557 - 585 (2008) 557

ARCHOSAUR EVOLUTION DURING THE : A SOUTHERN PERSPECTIVE

Oliver W. M. RAUHUT and Adriana LOPEZ-ARBARELLO

Bayerische Staatssammlung für Paläontologie und Geologie, München, Germany. E-mails: [email protected], [email protected]

ABSTRACT The record of - crocodylomorphs, and - from the Jurassic of the Southern Hemisphere is critically reviewed, and its evolutionary implications are evaluated. Although several important faunas and also isolated finds are known from , the record in total is still very patchy, and any evolutionary scenario based on this record should be seen as tentative. Compared to the Northern Hemisphere, southern archosaurs are much more poorly known, which is especially true for terrestrial crocodiles and pterosaurs. Marine crocodiles are rather well represented in south-western , whereas the report of terrestrial archosaurs is currently best for . However, in South America, important and especially promising faunas are known from the Cañadón Asfalto and the (?) Cañadón Calcáreo formations of Chubut province, Argentina. Early and Gondwanan archosaurs demonstrate that the faunas of that period still had a generally Pangean distribution, whereas first indications of differential archosaur evolution in the Northern and Southern Hemispheres are evident in Gondwanan faunas. Keywords: Crocodylomorphs, Pterosaurs, Dinosaurs, Gondwana, Fossil record

RESUMEN: La evolución de los arcosaurios durante el Jurásico: una perspectiva desde el Sur. El presente trabajo incluye una revisión crí- tica del registro fósil de los arcosaurios - cocodrilos, pterosaurios y dinosaurios - del Jurásico del Hemisferio Sur. Se evalúan además, sus implicancias evolutivas. Aunque se conocen varias faunas importantes y hallazgos aislados, el registro, en su tota- lidad, es todavía incompleto y cualquier escenario evolutivo basado en este registro debería verse como tentativo. En compa- ración con la situación en el Hemisferio Norte, es mucho menos lo que sabemos sobre los arcosaurios del Hemisferio Sur, y esto es especialmente cierto para los cocodrilos terrestres y pterosaurios. Los cocodrilos marinos están bastante bien repre- sentados en el sudoeste de Sudamérica, en tanto que el registro de arcosaruios terrestres es actualmente mejor en África. No obstante, en Sudamérica, se conocen faunas importantes, y especialmente prometedoras, del Caloviano de la Formación Cañadón Asfalto y del Titoniano (?) de la Formación Cañadón Calcáreo, en la provincia de Chubut, Argentina. Las faunas de arcosaurios del Jurásico Temprano y Medio de Gondwana muestran una distribución pangeica, en tanto que los primeros indi- cios de diferenciación en la evolución de los arcosaurios de los Hemisferios Norte y Sur se evidencian en las faunas gondwá- nicas del Jurásico Tardío.

Palabras clave: Cocodrilos, Pterosaurios, Dinosaurios, Gondwana, Registro fósil

INTRODUCTION crocodylomorphs, the Jurassic saw the almost none of which survived this pe- origin and diversification of the Mesoeu- riod. However, by the end of this period, Archosaurs - crocodylomorphs, ptero- crocodylia, the largest , which also basically all important lineages were esta- saurs, dinosaurs, and several basal, mainly includes the modern forms (Fig. 1, e.g. blished (Fig. 3, Sereno 1999a, Weisham- groups - are certainly one of the Benton and Clark 1988, Tykoski et al. pel et al. 2004). The Jurassic thus witnes- most prolific of Mesozoic terres- 2002, Pol and Gasparini 2007). In ptero- sed important faunal turnovers in all trial , and their living repre- saurs, the early "rhamphorhynchoid" fau- major clades of archosaurs and is there- sentatives, modern crocodiles and , nas reached their peak diversity during fore of greatest importance for our un- are important components of recent ver- this period, and the pterydactyloids, to derstanding of the evolutionary history tebrate faunas. All three derived groups which almost all pterosaurs of these groups. (crocodylomorphs, , dinosaurs) belong, originated and first diversified in It is still rather unclear what caused these originated in the later stages of the Trias- the Upper Jurassic (Fig. 2, Kellner 2003, important changes in Jurassic archosaur sic, but experienced their first and most Unwin 2003, 2006). Dinosaurs started faunas. In the Jurassic the initial break-up important radiations in the Jurassic. In with only a few clades into the Jurassic, of the supercontinent of Pangea took 558 O. W. M. RAUHUT AND A. LOPEZ-ARBARELLO

Jurassic archosaur evolution. In the follow- ing, the colloquial term "crocodile" is used to refer to crocodylomorphs in general. True members of the Crocodylia are so far unknown from the Jurassic and only appear in the Cretaceous. Institutional abbreviations: MACN: Museo Argentino de Ciencias Naturales "Bernardino Rivadavia", Buenos Aires, MB: Museum für Naturkunde, , MPEF: Museo Paleontológico Egidio Feruglio, Trelew, Argentina.

JURASSIC ARCHOSAURS FROM THE SOUTHERN HEMISPHERE

Jurassic terrestrial archosaurs from the Southern Hemisphere are known only from a handful of localities (Fig. 4), and most finds are isolated and/or fragmen- tary. In the following, the Gondwanan archosaur fossil record will be briefly outlined and compared with that from the Northern Hemisphere. Only the bo- dy fossil record will be dealt with in detail here. Although tracks can give important insights into faunal compositions and other aspects of archosaur biology, they are usually less diagnostic than body fos- Figure 1: Phylogeny of crocodiles showing the origin and first diversification of the Mesoeucroco- sils on a lower taxonomic level. Thus, dylia during the Jurassic. Based on Pol and Gasparini (2007). Dotted lines indicate uncertain limits the, for some stages, more numerous of the stratigraphic range of the bearing sediments of a certain taxon. track record is only taken into considera- place, during which the western Tethian Scotese et al. 1999). Archosaur radiations tion in cases where it inidcates the pre- and northern Atlantic oceans opened and faunal turnovers certainly have to be sence of otherwise not represented and finally separated Gondwana from seen in the context of these environmen- groups. the northern continents in the Late Ju- tal changes (see Forster 1999), but even rassic (Scotese 1991, Smith et al. 1994, the timing and the modes of these evolu- Scotese et al. 1999, Ford and Golonka tionary events are still largely unknown, 2003). Vicariance as a consequence of owing to the rather poor Jurassic fossil The Early Jurassic Gondwanan archo- this separation has been proposed as an record in many groups. Furthermore, our saur record is strongly biased towards important aspect of dinosaurian radia- current ideas about Jurassic faunal chan- Africa, with one important fauna known tions (Russell 1993, Sereno 1999a, b), ges are almost entirely based on the fos- from , and isolated occurrences and evidence for continent-wide vica- sil record of the Northern Hemisphere, from South America and (Fig. riance was found in empirical analyses of making it basically impossible to check 4a). dinosaurian biogeography (Upchurch et possible effects of the separation of The most important Lower Jurassic al. 2002). Furthermore, these changes in Gondwana on the evolutionary history Gondwanan faunas are cer- geography brought about other modifi- of the different groups. tainly those of the Upper Karoo Super- cations, such as the end of the mega- In the current paper, we critically evalu- group, namely the upper Stormberg monsoonal climate that had dominated ate the Jurassic archosaur fossil record of Group, comprising the Upper Elliot and large parts of Pangea in the Triassic and the Southern Hemisphere and its impor- Clarens formations [Fig. 4a (1), see Knoll early stages of the Jurassic (Parrish 1993, tance for our current understanding of 2005], and contemporaneous units in Archosaur evolution during the Jurassic: a southern perspective 559

sauropodomorph in the upper Stormberg, at least one other "prosauro- pod" and two basal sauropods are also present in this unit (Barrett 2004, Yates et al. 2004, 2007), and the basal sauropod has been described from stra- ta of Early Jurassic age in (Raath 1972, Cooper 1984). Theropod dinosaurs from the Early Jurassic of sou- thern Africa include the small coelophy- soid Coelo-physis rhodesiensis, which repre- sents one of the best-known Early Jurassic theropods (Raath 1969, 1977, 1985, 1990, Galton 1971, Chinsamy 1990, Bristowe and Ra-ath 2004). Most material of this theropod comes from the Forest of Zimbabwe (Raath 1969, 1977), but the taxon also seems to be represented in the Upper (Raath 1980). Recently, Yates (2005) described a new taxon of theropod from the Upper Elliot Formation, . Recent field- work indicates, however, that this speci- men might be derived from the basal part of the overlying (OR, pers. obs. 2008). The Upper Elliot Formation has yielded Figure 2: Phylogeny of pterosaurs showing the origin and first diversification of the two main linea- ges, Ramphorhynchidae (node 1) and the (node 2) during the Jurassic (based on the most diverse Early Jurassic ornithis- Unwin 2006). chian fauna known. Taxa represented in- clude the heterodontosaurids Heterodonto- Zimbabwe, such as the . (, and Zimbabwe). saurus, and (Haug- Several taxa of crocodylomorphs have Since the first description of the "pro- hton 1924, Crompton and Charig 1962, been described especially from the Up- sauropod" from the Upper Hopson 1975, see also Knoll 2005), the per Elliot and Clarens formations, but Elliot Formation by Owen (1854), a ple- probably basal thyreophoran Lesothosau- many of them seem to be junior subjec- thora of dinosaurs, mainly basal sauro- rus (Thulborn 1970, 1972, Galton 1978, tive synonyms of other taxa (see review podomorphs, have been described from Sereno 1991, Butler et al. 2008), and the in Knoll 2005). Taxa currently regarded these units. However, most of these taxa basal cerapodan Stormbergia (Knoll 2002a, as valid (Knoll 2005) include the 'spheno- are considered to be junior synonyms of b, Butler 2005). suchians' (Haughton 1915, Massospondylus (Cooper 1981), which Apart from the Karoo basin, Early Ju- Walker 1990) and Litargosuchus (Clark and remains the best known Early Jurassic rassic archosaurs from Africa have only Sues 2002), and the protosuchids Proto- "prosauropod" from the Southern He- been described from Morocco [Fig. 4a suchus (Gow 2000) and Orthosuchus (Nash misphere, and is well-documented from (2)]. Material from the Toundoute conti- 1968, Pol et al. 2004). Pterosaur remains the upper Stormberg (e.g. Gow 1990, nental series of the include are currently unknown from the Elliot Gow et al. 1990, Sues et al. 2004, Yates the basal sauropod (Allain and Clarens Formations, though this and Vasconcelos 2005, Barrett and Yates et al. 2004) and the probable abelisauroid group might be represented by tracks 2006), including remains of eggs and em- theropod Berberosaurus (Allain et al. 2007). (Knoll 2005). bryos (Reisz et al. 2005), but also known Furthermore, Taquet (1985) reported Dinosaurs are certainly the best-repre- from other Early Jurassic units in sou- and figured a partial theropod hindlimb sented and best-known archosaur group thern Africa, mainly Zimbabwe (Cooper from an unnamed Early Jurassic unit in from the upper and 1981). However, although Massospondylus Morocco. equivalent strata in certainly represents the most common Outside Africa, the most important Early 560 O. W. M. RAUHUT AND A. LOPEZ-ARBARELLO

Figure 3: Phylogenetic relationships and stratigraphic distribution of the main clades, showing the diversification of almost all main dinosaur lineages during the Jurassic. Minor clades or single taxa have been considered if of stratigraphic importance. Composite cladogram, (based on Rauhut 2003), on Wilson (2002) and Smith and Pol (2007), and on Weishampel, Dodson and Osmólska (2004) and Butler et al. (2008). 1) , 2) , 3) , 4) , 5) Spinosauroidea, 6) , 7) , 8) , 9) Oviraptorosauria, 10) Therizinosauroidea, 11) , 12) Dromaeosauridae, 13) Troodontidae, 14) , 15) , 16) + , 17) (sensu Upchurch et al. 2004), 18) Omeisauridae, 19) , 20) , 21) , 22) spp., 23) , 24) , 25) , 26) Emausaurus, 27) , 28) Ankylosauridae, 29) Nodosauridae, 30) , 31) Yinlong, 32) Psittacosauridae, 33) Centrosaurinae, 34) Chasmosaurinae, 35) "Hypsilophodontidae" (non-monophyletic group of basal ornithopods, the interrelationships of which are still unclear), 36) Dryosauridae, 37) Camptosaurus spp., 38) Iguanodontidae, 39) Hadrosaurinae, 40) Lambeosaurinae.

Jurassic Gondwanan archosaur localities ly Jurassic in age (Bandyopadhyay et al. and Manhas 2007). are found in the Prahita-Godavari valley 2002, Weishampel et al. 2004, Bandyo- Again, it was Owen (1854) who first re- of eastern India, mainly from the Kota padhyay and Sengupta 2006), there is ported archosaur remains from this area, Formation [Fig. 4a(3)]. However, it might some uncertainty about the dating of this from levels now assigned to the Kota be noted that, although the Kota For- unit, and it might be as young as Early Formation. The remains described by mation is generally considered to be Ear- Cretaceous (Prasad et al. 2004, Prasad Owen (1854) represent a marine, thalat- Archosaur evolution during the Jurassic: a southern perspective 561

Kota Formation of India, referred to an ankylosaurian dinosaur by Nath et al. (2002: pl. 1a-e), most probably represents a crocodilian, as indicated by the pitted bone surface and the general shape of the fragments. Judged by the general morphology, this fragment most proba- bly represents a terrestrial crocodile, but further interpretation has to await a de- tailed reanalysis of this specimen. Jain (1974) described the campylogna- thoid pterosaur Campylognathoides indicus from the upper levels of the Kota For- mation, which still represents the oldest pterosaur known from the Southern Hemisphere. Dinosaurs from the were described by Jain et al. (1975, 1979) and Yadagiri (1988, 2001), who named and described the sauropods and , respectively. The material referred to the latter taxon includes at least one further taxon of sauropod (OR, pers. obs. 2005). Nath et al. (2002) described fragmentary dino- saur remains from the Kota Formation and referred them to an ankylosaurian thyreophoran, which would represent the oldest record of this group. However, this identification is questionable, since more than one taxon is represented in the material (see comments on a proba- ble crocodylomorph included in this ma- terial above), but some of the scutes des- cribed and figured by Nath (2002: pl. 1f- j) might indeed represent a thyreopho- ran. Apart from these remains, thero- Figure 4: Palaeogeographic maps (modified from Mollewide Plate Tectonic Maps available at pods are represented in the Kota Forma- http://jan.ucc.nau.edu/~rcb7/globaltext2.html) indicating the main Jurassic archosaur localities in tion by undiagnostic material (Bandyo- and Gondwana. a) Early Jurassic: Elliot and Clarens formations, upper Stormberg Group, Upper , South Africa (1), Toundoute continental series, High Atlas, Morocco (2), padhyay et al. 2002, Prasad and Manhas Kota Formation, Prahita-Godavari valley, India (3), , , 2007). Antarctica (4), and and equivalent beds, USA (5), Newark Just recently, Kutty et al. (2007) described Supergroup, (6), in Germany and comparable strata in England and France (7), , (8). b) Middle Jurassic: Durham Downs, Queensland, two taxa of sauropodomorphs, the "pro- (1), Tilougguit Formation and equivalent beds of Morocco (2), Isalo III Formation, Madagascar (3), sauropod" Pradhania and the probable Cañadón Asfalto Formation, Chubut, Argentina (4), Lautaro Formation, Chile (5), various marine basal sauropod Lamplughasaura, and men- localities in England and France (6), Balabansai Formation, Kirghizia (7), Lower Shaximiao Formation, Sichuan, China (8). C. Late Jurassic: Formation and equivalent beds, tioned the presence of a third sauropo- Neuquén Basin, Argentina (1), Cañadón Calcáreo Formation, Chubut, Argentina (2): Tendaguru domorph taxon in the Upper Dharmaran Beds, Tanzania (3), Kadsi Formation, Zimbabwe (4), , USA (5), various locali- Formation (), which underlies ties in Portugal (6), and equivalent beds in Germany (7), Shishougou Formation, , China (8), Upper Shaximiao Formation of China (9). the Kota Formation. Bradmeister et al. (2006) described a ver- tosuchian crocodile (Kutty et al. 1987, from the Gondwana sequence of this tebrate fauna, including probable thero- Bandyopadhyay et al. 2002), and are still basin. However, it might be noted that a pod and sauropod teeth, from the Isalo the only crocodile remains described partial from the Lower Jurassic II Formation of Madagascar. Although 562 O. W. M. RAUHUT AND A. LOPEZ-ARBARELLO

they stated in the abstract that the verte- brate fauna indicates an Early Jurassic age for this fauna, the faunal association, including probable colobodontid and aetosaurian and phytosaurian archo- saurs, is more indicative of a Late Trias- sic age. Only fragmentary archosaur material has been described from the Early Jurassic of South America so far. Huene (1927) des- cribed Lower Jurassic crocodile remains from Mendoza province in Argentina, South America. The material consists of two dorsal vertebrae, which he made the type of a new of the thalattosu- chian , Steneosaurus gerthi, but this species is currently considered to be a nomen dubium (Pol and Gasparini 2007). An articulated partial metatarsus from the Sinemurian of Alto de Varas in the Cordillera de Domeyko in Chile was des- cribed by Chong-Diaz and Gasparini (1972) and also referred to a thalattosu- chian (see also Gasparini 1985, Gasparini et al. 2000). Early Jurassic dinosaur Figure 5: Schematic remains have been described from the stratigraphic column probably Lower Jurassic La Quinta For- of the archosaur- bearing units of the mation of Venezuela. Russell et al. (1992) Middle Jurassic and Sánchez-Villagra and Clark (1994) Cañadón Asfalto reported ornithischian material from this Formation and the formation, which they referred to the Late Jurassic Cañadón Calcáreo southern African , and Formation of Moody (1997) described some unidenti- Chubut province, fiable theropod teeth. However, a recent Argentina. revision of this material shows that the ornithischian material is not referable to fauna, including dinosaurs, from Hanson mains of Middle Jurassic age have been Lesothosaurus, and that a further taxon of Formation in the Transantarctic Moun- reported so far, and were named Ozraptor probable basal saurischian was present tains of Antarctica [Fig. 4a(4)] was made and , respectively (Longman (Barrett et al. 2008). Martínez (1999) refer- known by Hammer and Hickerson 1926, 1927, Long & Molnar 1998). Oz- red some material from the probably (1994). This southernmost archosaur raptor comes from the shallow marine Early Jurassic Cañón de Colorado For- fauna discovered so far includes a hume- Colalura Sandstone () of west- mation of the province of San Juan, Ar- rus of a pterosaur, the theropod Cryolo- ern Australia, whereas Rhoetosaurus was gentina, to Massospondylus, though reex- phosaurus, the basal sauropodomorph found in terrestrial deposits at Durham amination of this material indicates that , and an undescribed sauro- Downs in Queensland [Fig. 4b(1)]. it represents a new genus (Pol, pers. com. pod (Hammer and Hickerson 1994, Middle Jurassic strata in Africa have yield- 2008). Another basal sauropodomorph 1999, Smith and Pol 2007, Smith et al. ed a few dinosaur remains from the Ba- has recently been reported from the Las 2007a, b). thonian-Callovian Tilougguit Formation Leoneras Formation of the province of and equivalent beds of Morocco [Fig. 4b Chubut, Argentina (Pol and Garrido Middle Jurassic (2)], including the sauropods "Cetiosau- 2007), but these remains have not been rus" mogrebiensis and imelakei described in detail yet. From Australia, only a distal theropod (Lapparent 1955, Monbaron et al. 1999), A noteworthy Early Jurassic vertebrate tibia and some fragmentary sauropod re- as well as the sauropod Chebsaurus algerien- Archosaur evolution during the Jurassic: a southern perspective 563

sis from indeterminate Middle Jurassic rocks of Algeria (Mahammed et al. 2005). Isolated vertebrae from the of Morocco that were originally described as a theropod, mersensis, by Lapparent (1955), probably represent a thalattosuchian crocodile (Chabli 1985). However, theropods are represented in the Bathonian of Morocco by isolated and unidentified teeth (Monbaron et al. 1999). Remains of the thalattosuchian Steneosaurus are found in the Bathonian of Tunisia (Mueller-Töwe 2006). Figure 6: Pterosaur material from the Middle Jurassic Cañadón Asfalto Formation, Chubut provin- The Bathonian Isalo III Formation of ce, Argentina. a) Left in lateral view. b) Left mandible in medial view. c) Right humerus in Madagascar [Fig. 4b(3)] has yielded a anterior view. d) Proximal end of wing phalanx 1. Scale bars: 1 cm. rich, though mainly fragmentary archo- saur fauna. Taxa reported include a pos- Chubut, Argentina [Figs. 4b(4), 5]. This are marine crocodiles from western sible atoposaurid crocodilian, a "rham- Callovian unit has yielded so far uniden- Argentina and Chile, which include the phorhynchoid" pterosaur, indeterminate tified and unstudied crocodile remains, as oldest record of the genus Metriorhynchus ornithischians, theropods, all represented well as a diverse, but also still unstudied from the Bajocian Lautaro Formation of by teeth only (Flynn et al. 2006), and the pterosaur fauna, comprising at least three Chile [Fig. 4b(5), Gasparini et al. 2000]. sauropods madagascarien- taxa, possibly including the oldest ptero- Two species of Metriorhynchus were des- sis and Archaeodontosaurus descouensi (Bo- dactyloid known from Gondwana (Rau- cribed from the Callovian of Chile, Me- naparte 1986a, Buffetaut 2005, Maga- hut et al. 2001, Unwin et al. 2004, Rauhut, triorhynchus casamiquelai from an unnamed nuco et al. 2005). Flynn et al. (2006) also unpublished data). Most of the remains unit in the Quebrada Sajasa in northern listed the sauropod species Bothriospon- come from a single pterosaur bonebed, Chile (Gasparini and Chong Díaz 1977, dylus madagascariensis in the Isalo III For- which has yielded disarticulated, but very Gasparini 1985), and Metriorhynchus wester- mation, but the material of this taxon well-preserved and three-dimensional manni from the Mina Chica For-mation was referred to the new genus Lapparen- material (Fig. 6), which is very rare in Ju- of northern Chile (Gasparini 1980, 1985) tosaurus by Bonaparte (1986a, see also rassic pterosaurs, potentially making this [Fig. 4b(5)]. The latter was regarded as a Buffetaut 2005), and furthermore refer- the most important Middle Jurassic pte- synonym of the former by Gasparini et red several teeth to an unspecified neo- rosaur locality globally. Most noteworthy, al. (2000), but new material shows that it sauropod, though they noted that the however, is the dinosaur fauna, which is is a valid taxon (Gasparini et al. in press). apparent differences to the teeth of Ar- the second most diverse Middle Jurassic Further thalattosuchian oc-currences in chaeodontosaurus might be due to preserva- dinosaur fauna known, following the Ba- Chile were discussed by Gasparini et al. tion, so this assignment is questionable. thonian Laurasian fauna from the Lower (2000). Recently, Gaspa-rini et al. (2005) Thus, only two sauropod taxa are recog- Shaximiao Formation of China (Peng et reported a fragmentary skull of nized from this formation, Lapparen- al. 2005). Taxa described include the sau- Metriorhynchus from the Batho-nian of the tosaurus and Archaeodontosaurus. Maganuco ropod and , and of Neuquén pro- et al. (2005) referred theropod teeth to the theropods and Condor- vince, Argentina. several different groups of theropods, in- raptor (Bonaparte 1979, 1986b, Rauhut Apart from the fauna of the Cañadón cluding advanced coelurosaurs, but Flynn 2005a), and at least one further taxon of Asfalto Formation, Middle Jurassic dino- et al. (2006) considered all this material as sauropod is present (Rauhut 2003a). Or- saurs from South America are only Theropoda indet. In addition to the nithischian dinosaurs have not been des- known from two localities in Argentina. fauna from the Isalo III Formation, ma- cribed from this formation so far, but are Cabrera (1947) described fragmentary terial of the thalattosuchian Steneosaurus represented by an isolated ungual from sauropod remains from Pampa de Agnia, has been reported from the Callovian of the locality of Cerro Cóndor (Fig. 7) and Chubut, as patagonicus. The Madagascar (Mueller-Töwe 2006). some teeth from the microvertebrate lo- material comes from the Cerro Carne- Middle Jurassic archosaurs from South cality Queso Rallado. rero Formation, which is most probably America are mainly known as isolated Apart from the fauna of the Cañadón of - age, and thus re- finds, with the exception of the fauna Asfalto Formation, the most important presents the oldest Middle Jurassic ar- from the Cañadón Asfalto Formation of archosaur remains from South America chosaur-bearing unit in South America. 564 O. W. M. RAUHUT AND A. LOPEZ-ARBARELLO

Figure 7: : Isolated pedal pha- lanx of a probably basal cerapo- dan ornithischian from the Middle Jurassic Cañadón Asfalto Formation of Chubut, Argentina, MACN CH 223. a) Dorsal view. b) Lateral view. c) Ventral view. Scale bar: 1 cm.

Additional material from the same loca- Late Jurassic ments of Chile (Martill et al. 2000), al- lity was referred to this taxon and even though the latter might be Early Cre- thought to represent the same individual Although the Late Jurassic is often seen taceous in age. as the type by Casamiquela (1963), but as a well-known stage in the history of Dinosaurs were almost unknown from Rauhut (2003b) showed that more than terrestrial vertebrates, mainly due to nu- the Late Jurassic of South America until one individual is represented by these merous localities in the Northern Hemi- recently, with the exception of a single remains. Nevertheless, Amygdalodon still sphere, such as the famous Morrison sauropod from unspecified Late represents the oldest sauropod record Formation of the western USA, very lit- Jurassic sediments of northern Colombia from South America. The recent find of tle is still known about Late Jurassic te- (Langston and Durham 1955) and undes- a partial sauropod from the Up- rrestrial vertebrates from Gondwana, cribed dinosaur remains from the Late per Aalenian Remoredo Formation of and only few localities yielding archosaur Jurassic of Chile, reported by Chong Dí- Neuquén, Argentina (Salgado and Gas- material are known. az and Gasparini (1976). More recently, parini 2004), demonstrates that these ani- Late Jurassic archosaurs from South an isolated sauropod knee joint was re- mals were already widely distributed in America were mainly known from mari- ported from the Tordillo South America at the beginning of the ne crocodiles until recently. With the ex- Formation of Neuquén, Argentina (Gar- Middle Jurassic. ception of one vertebra from Chile, re- cía et al. 2003), and theropod teeth have Only isolated finds of fragmentary dino- ported by Gasparini (1985), all marine been found in the Tacurembó Formation saur material have been reported from crocodile remains from the Late Jurassic in Uruguay (Perea et al. 2003), which is the Middle Jurassic of India so far (see of South America come from the Neu- probably also Late Jurassic in age (Soto, overview in Moser et al. 2006). Most of quén Basin in Argentina, from the Ti- pers. comm. 2007). Salgado et al. (2008) the material reported, which comes from thonian Vaca Muerta Formation and described some theropod remains from different units in north-western India equivalent beds [Fig. 4C(1), Gasparini the Tithonian Toqui Formation of south- and ranges in age from the Aalenian-Ba- 1985, 1992, 1996, Gasparini and Fernan- ern Chile. This material, which includes jocian to the Callovian, seems to repre- dez 1997, Gasparini et al. 2006]. Taxa at least one taxon of tetanurans, thus sent sauropod dinosaurs, but only a few reported from these units include the represents the southernmost Late Juras- fragmentary remains from Khadir Island metriorhynchids Geosaurus araucanensis, sic dinosaur body known. have been described in detail so far (Mo- Metriorhynchus potens, and Dakosaurus The most important, but still rather ser et al. 2006). Mainly on the basis of the andiniensis. poorly studied Late Jurassic dinosaur fau- morphology of a proximal fibula, Moser Pterosaurs are still very poorly known na reported from South America so far is et al. (2006) concluded that some of this from the Late Jurassic of South America. that of the Cañadón Calcáreo Formation material represents a camarasauromorph Casamiquela (1975) reported a partial of Chubut province, Argentina [Figs. 4c sauropod, which would represent the skeleton of a supposed theropod dino- (2), 5]. The age of this unit is still some- oldest record of a neosauropod from the saur, Herbstosaurus pigmaeus, found in an what uncertain: whereas a radiometric Southern Hemisphere. However, better isolated boulder, which was later reinter- date puts its basal part in the Tithonian material is needed to substantiate this preted as a pterosaur, demonstrably from (see Rauhut 2006a), new palynological claim, especially since the shape of the Upper Jurassic (Tithonian) strata of the evidence seems to indicate a lowermost proximal articular surface of the element Vaca Muerta Formation in Neuquén pro- Cretaceous age for at least parts of the in question is rather reminiscent of sau- vince, Argentina (Bonaparte 1981, Co- formation (Silva Nieto et al. 2007). More ropod metacarpals II-IV (see e.g. Ostrom dorniú and Gasparini 2007). Further, geological and stratigraphic work is need- and McIntosh 1966: pl. 56-58, 60-61), as mainly fragmentary fossils have been ed to tie down the stratigraphic position is its size in comparison to other meta- described since, including Tithonian pte- of the dinosaurs from this formation. carpal fragments found at the same loca- rosaur remains from the same beds in The first dinosaur described from the lity (Moser et al. 2006: figs. 4.1, 4.2). Argentina (Gasparini et al. 1987, Codor- Cañadón Calcáreo Formation was Tehuel- niú et al. 2006) and from coastal sedi- chesaurus, though originally thought to be Archosaur evolution during the Jurassic: a southern perspective 565

from the Middle Jurassic Cañadón As- Middle Saurian Bed, to P. brancai and africana, and Australodocus bohetii (Aberhan falto Formation (Rich et al. 1999). This transferred this species to the Early Crea- et al. 2002, Remes 2006, 2007), the thero- taxon was first considered to be closely tceous Asian genus . Inte- pods bambergi and at least related to the Middle Jurassic Chinese restingly, Unwin and Heinrich (1999) four more taxa, including two tetanurans sauropod (Rich et al. 1999, described a pterosaurian jaw fragment and two ceratosaurs (Janensch 1925, Upchurch et al. 2004), but new data indi- from the Middle Saurian Beds as a new Rauhut, unpublished data). Both orni- cates that it is a member of the Macro- species, Tendaguripterus recki, which they thischians known from Tendaguru, Ken- naria and close to or within the Tita- also referred to the Dsungaripteroidea. trosaurus aethiopicus and Dysalotosaurus let- nosauriformes (Rauhut 2002, Rauhut et However, since the type and referred towvorbecki, are also present in the Middle al. 2005, Carballido et al. 2007). More specimens of "Dsungaripterus" brancai are Saurian Beds (Aberhan et al. 2002). The recently, the dicraeosaurid Brachytrache- tibiotarsi, more material is necessary to Upper Saurian Beds have yielded the sau- lopan mesai (Fig. 8) and an unidentified establish whether only one, or two taxa ropods brancai, brachiosaurid were reported from the of dsungaripteroids were present in the robusta, Tendaguria tanzaniensis, Dicraeosau- same formation (Rauhut et al. 2005, Rau- Tendaguru fauna. rus sattleri, and africana, at least hut 2006a). The dinosaur fauna from Tendaguru is two ceratosaurian and two tetanuran the- The most important Late Jurassic terres- dominated by sauropods, which are re- ropods (Rauhut 2005c, unpublished da- trial vertebrate fauna from Gondwana presented by at least seven species, the ta), and the stegosaur aethiopi- known so far comes undoubtedly from brachiosaurid Brachiosaurus brancai, the cus (Aberhan et al. 2002). the famous Tendaguru Beds of Tanzania probably basal titanosaurs Janenschia ro- Another dinosaur fauna reported from [Fig. 4C(3), Maier 2003]. Archosaurs busta and Tendaguria tanzaniensis (which the Upper Jurassic of Africa comes from from three different levels within the might be conspecific), the dicraeosaurids the Kadsi Formation of Zimbabwe [Fig. Tendaguru Beds include crocodile teeth, hansemanni and D. sattleri, 4c(4), Raath and McIntosh 1987], but the isolated pterosaur remains, and nume- and the diplodocids Tornieria africana and material collected so far is fragmentary rous taxa of dinosaurs (Janensch 1914, Australodocus bohetii (Janensch 1914, 1929, and identifications of taxa have to be tre- Russell et al. 1980, Aberhan et al. 2002, Bonaparte et al. 2000, Upchurch et al. ated with caution. Raath and McIntosh Maier 2003). The ages of the three diffe- 2004, Remes 2006, 2007). Theropod di- (1987) identified at least four, possibly rent beds are probably for the nosaurs are represented by the basal neo- five sauropod taxa from these beds, and Lower Saurian Bed, Kimmeridgian-Early ceratosaur Elaphrosaurus bambergi (Fig. 9) referred them to the genera Brachiosaurus, Tithonian for the Middle Saurian Bed, and at least six more taxa (Janensch 1920, Janenschia (=Tornieria in the cited work, and Late Tithonian to earliest Cretaceous 1925, Rauhut 2005b, c, 2006b). The the- for the convoluted taxonomic history of for the Upper Saurian Beds (Aberhan et ropod fauna seems to be dominated by this genus see Wild 1991), Dicraeosaurus, al. 2002). basal tetanurans and neoceratosaurs, , and, possibly, Camarasaurus. Crocodiles are so far only represented by whereas coelurosaur remains could not However, only the remains referred to isolated teeth from the Upper Saurian be identified so far (Rauhut 2005b, c, Brachiosaurus might be diagnostic on ge- Beds, which seem to be similar to those 2006b). Ornithischians are much less neric level, the rest of the material would of the neosuchian Ber- diverse and are represented only by the better be understood as basal titanosaur nissartia (Maier 2003: 315). Pterosaurs are stegosaur Kentrosaurus aethiopicus and the indet., two diplodocoids indet. (possibly represented by numerous, but mainly iso- small ornithopod Dysalotosaurus lettowvor- a dicraeosaurid and a diplodocid), and an lated and often fragmentary elements. becki (Hennig 1925, Janensch 1955). Di- unidentified sauropod. Two theropod fe- Most pterosaur remains come from the nosaur remains are not evenly distributed mora, tentatively identified as ?allosaurid Upper Saurian Beds, though they are also in all levels at Tendaguru. The Lower by Raath and McIntosh (1987: 109, 113 present in the Middle Saurian Beds (Reck Saurian Beds have only yielded a few the- fig. 4) might rather represent a large neo- 1931). Reck (1931) described four species ropod teeth attributed to "Megalosaurus" ceratosaur, as indicated by the low, ali- on the basis of this material, though ingens (Janensch 1925) and fragmentary form lesser trochanter with a well-deve- mainly based on uncomparable remains: sauropod remains, possibly representing loped trochanteric shelf. tendagurensis, Pterodactylus Brachiosaurus (Aberhan et al. 2002). The A further Late Jurassic unit from Africa maximus, Pterodactylus brancai, and Pterodac- Middle and Upper Saurian Beds are has yielded archosaur remains, the Mu- tylus arningi, all from the Upper Saurian much richer in vertebrate remains, but gher Formation of Ethiopia, Beds. However, the validity of all of the- differ slightly in their dinosaur fauna. which is of probably Tithonian age se species is highly questionable (Unwin The Middle Saurian Beds contain re- (Goodwin et al. 1999). Archosaur fossils and Heinrich 1999). Galton (1980a) refer- mains of the sauropods Brachiosaurus from this formation include a possibly red another specimen, probably from the brancai, Dicraeosaurus hansemanni, Tornieria goniopholid crocodile and theropod, 566 O. W. M. RAUHUT AND A. LOPEZ-ARBARELLO

sauropod, and probable ornithischian Middle Jurassic South American sauro- 2003) and protosuchids and the oldest teeth (Werner 1995, Goodwin et al. pod Tehuelchesaurus as evidence for a Mid- known goniopholid, , were 1999). One theropod fragment was dle Jurassic age, but, as noted above, Te- reported (Colbert and Mook 1951, identified as cf. by huelchesaurus neither comes from Middle Crompton and Smith 1980, Sues et al. Goodwin et al. (1999: 735-736), but the Jurassic sediments, nor is it closely rela- 1994, Clark and Sues 2002, Tykoski et al. only character given in support of this ted to Omeisaurus. However, Averianov et 2002). A similar faunal composition, with assignment (chisel-shaped denticles) ac- al. (2007) further mentioned protosuchids and "sphenosuchians" is tually represents the plesiomorphic con- evidence for a Middle Jurassic age the also found in the McCoy Brook Forma- dition in theropods, so this specimen Upper Shaximiao Fm. On the other tion of the Newark Supergroup of east- should be regarded as Theropoda indet. hand, the fauna of the Upper Shixamiao ern North America (Shubin et al. 1994, From slightly older levels close to the city Formation shares the dinosaur genera Sues et al. 1996) and the Lufeng Forma- of Harrar in Ethiopia (Oxfordian-Kim- and with the tion of China (Simmons 1965, Luo and meridgian), Huene (1938) described a Shishougou Formation of Xinjiang, Chi- Wu 1994, Wu and Sues 1996, Harris et al. partial mandibular symphysis as a new na, which is considered to be Oxfordian 2000). pliosaur, Simolestes nowackianus, but this in age (Xu et al. 2006a, Xu, pers. comm. Pterosaurs are known from a number of specimen was later assigned to the thalat- to OR 2006), so the age of this unit is Early Jurassic Laurasian localities, though tosuchian crocodyliform Machimosaurus best considered to be uncertain. most remains are fragmentary, such as by Bardet and Hua (1996), thus represen- The most important Late Jurassic faunas the type of Rhamphinion from the Ka- ting the oldest record of this group in are certainly those of the Morrison For- yenta Formation of the western USA Africa. mation of the western USA [Fig. 4c(5)], (Padian 1984, Sues et al. 1994). However, various Kimmeridgian units in Portugal such specimens might provide important JURASSIC ARCHOSAURS [Fig. 4c(6)], the Solnhofen Limestone in anatomical information as well (e.g. Clark FROM THE NORTHERN southern Germany [Fig. 4c(7)], and va- et al. 1998). Well-preserved pterosaur re- HEMISPHERE rious units in China, including the mains have been found in the Posidonia Shishougou Formation of Xinjiang [Fig. Shale of Germany (see Padian and Wild Jurassic archosaurs from the Northern 4c(8)]. 1992) and the Lias of Lyme Regis in Hemisphere are known from a number England (see Padian 1983). These re- of localities. Early Jurassic mains represent the families Dimorpho- In the Early Jurassic, the probably most dontidae and Campylognathidae. important Laurasian localities are the Ka- In the Northern Hemisphere, crocodiles Dinosaurs are also known from a variety yenta Formation and Navajo Sandstone are well known from marine forms, as in of Early Jurassic localities in the Nor- and equivalent beds of the western USA the Southern Hemisphere. The oldest thern Hemisphere. Taxa recorded inclu- [Fig. 4A(5)], the Newark Supergroup of northern records of thalattosuchians are de basal thyreophoran ornithischians eastern North America [Fig. 4a(6)], the contemporaneous to those in the Sou- from western North America (Colbert Posidonia Shale in Germany and compa- thern Hemisphere and were found in the 1981, Padian 1989), (Owen 1863, rable strata in England and France [Fig. Sinemurian of France (see Gasparini et Haubold 1990) and (Simmons 1965, 4a(7)], and the Lufeng Formation of al. 2000, Mueller-Töwe 2006). Thalat- Dong 2001, Norman et al. 2007). The- south-eastern China [Fig. 4a(8)]. tosuchians are especially well-represent- ropods are represented by coelophysoids In the Middle Jurassic, important faunas ed in the Toarcian of the Posidonia Sha- in North and Central America (Tykoski are known especially from various, main- le and contemporaneous sediments in and Rowe 2004, Munter and Clark 2005, ly marine units in England and France Europe, from where the genera Pelago- Tykoski 2005), Europe (Carrano and [Fig. 4b(6)] and the Lower Shaximiao saurus, Platysuchus and Steneosaurus are Sampson 2004) and Asia (Irmis 2004) Formation of Sichuan, China [Fig. 4b known, the latter with several species and "dilophosaurs" in North America (8)]. Furthermore, the Upper Shaximiao (Mueller-Töwe 2006). In this stage, frag- (Welles 1954, 1984) and Asia (Hu 1993, Formation of China [Fig. 4c(9)], often mentary thalattosuchian remains are also see also Smith et al. 2007). A therizino- considered to be Late Jurassic in age (e.g. known from North America and China sauroid theropod reported from the Weishampel et al. 2004, Peng et al. 2005), (Mueller-Töwe 2006). Lufeng Formation (Xu et al. 2001) is of might also be of late Middle Jurassic (Ca- Terrestrial Early Jurassic crocodiles are doubtful affinities, and might represent a llovian) age (Averianov et al. 2005, 2007). known from the western US, from where prosauropod (Rauhut 2003c). Basal sau- Averianov et al. (2005) cited the supposed several taxa of "sphenosuchians" (the ropodomorphs ("prosauropods") are close relationships of the Chinese sauro- monophyly of which is still controversial, well-documented from the Lufeng For- pod Omeisaurus with the supposedly see Clark and Sues 2002 and Sues et al. mation of China (Young 1941, 1951, Archosaur evolution during the Jurassic: a southern perspective 567

Zhang and Yang 1995), but they are also al. 2005), which includes stegosaurian Morrison Formation of western North present in both western and eastern (Maidment and Wei 2006) and basal cera- America (see Foster 2003). Crocodiles North America (Attridge et al. 1985, Gal- podan ornithischians (Barrett et al. 2005), from this unit include some of the last ton 1976, Yates 2004). Furthermore, Ear- theropods, and sauropods. Theropods "sphenosuchians" known (Walker 1970, ly Jurassic sauropods are known from include basal tetanurans, such as Xuan- Göhlich et al. 2005), a cursorial basal China (e.g. He et al. 1998, Barrett 1999, hanosaurus and "Szechuanosaurus" zigongen- mesoeucrocoylian and other cursorial Upchurch et al. 2007) and Europe (Wild sis (Dong 1984, Gao 1993), and the pos- crocodylomorphs of unspecified syste- 1978). sible basal coelurosaur Gasosaurus (Dong matic position, the probable atoposaurid and Tang 1985, Holtz et al. 2004). The Hoplosuchus, several species of , Middle Jurassic sauropod fauna is dominated by ma- and the goniopholid menchisaurids (Peng et al. 2005), but also (Tykoski et al. 2002, Foster 2003, Pol and As for the Southern Hemisphere, the includes the basal sauropod Gasparini 2007: fig. 5.5). Pterosaurs are Middle Jurassic archosaur record from and the possibly oldest known neosauro- poorly represented in the terrestrial the Northern Hemisphere is rather poor, pods (Upchurch et al. 2004). Another clo- deposits of the Morrison Formation, but and, most notably, only one skeletal re- se relative of neosauropods or a basal include at least one "rhamphorhynchoid" main of an archosaur from that period, member of this clade is also known from and several taxa of pterodactyloids (Fos- the poorly preserved type specimen of the Callovian Balabansai Formation of ter 2003). The dinosaur fauna of the the crocodyliform Entradasuchus, has Kirghizia [Fig. 4b(7), Alifanov and Ave- Morrison Formation is the most diverse been described from North America rianov 2003]. Ornithischians are repre- fauna known from the Jurassic, and one (Hunt and Lockley 1995), though dino- sented in the same formation by an inde- of the most diverse Mesozoic dinosaur saurs are known from track evidence (e.g. terminate stegosaur (Averianov et al. faunas known in general (see Weisham- Kvale et al. 2001). Middle Jurassic croco- 2007) and isolated teeth that were descri- pel et al. 2004). The dinosaur fauna inclu- diles from Europe are represented by bed as the oldest known pachycephalo- des several species of diplodocid sauro- isolated teeth attributed to atoposaurids saur (Averianov et al. 2005). However, the pods, the basal macronarian Camarasaurus and goniopholids in England (Evans and teeth are rather poorly preserved and the as the most abundant taxon among the Milner 1994) and France (Kriwet et al. identification is questionable. sauropods, the basal titanosauriform 1997) and marine thalattosuchians from Although some of the first dinosaurs to Brachiosaurus altithorax, , various countries (Mueller-Töwe 2006). be described were Middle Jurassic taxa which either represents a basal diplodo- The record of these marine crocodyli- from Europe (e.g. Buckland 1824), Mid- coid, or a basal macronarian, and the forms is poor in the Aalenian and Ba- dle Jurassic dinosaurs from this continent probably basal diplodocoid jocian, although the first metriorhynchids are still poorly understood, mainly due to (Wilson 2002, Foster 2003, Upchurch et are known from the latter stage (Mueller- the fragmentary nature of most remains. al. 2004). The theropod fauna includes Töwe 2006). In the Bathonian and However, the fauna includes non-neo- rare ceratosaurs (Marsh 1884, Galton Callovian, both teleosaurid and me- sauropodan sauropods (Upchurch and 1982, Madsen and Welles 2000, Chure triorhynchid thallatosuchians are well- Martin 2002, 2003), the basal spinosau- 2001) and basal spinosauroids (Britt represented especially in England and roids Eustroptospondylus (Walker 1964, 1991, Holtz et al. 2004), abundant allo- France, although some remains are also Holtz et al. 2004), (Allain saurids (Madsen 1976, Foster 2003), and known from central Asia (Mueller-Töwe 2002, 2005) and (Allain and a variety of coelurosaurs (Foster 2003, 2006). Middle Jurassic terrestrial crocodi- Chure 2002), and one of the oldest coe- Holtz et al. 2004, Carpenter et al. 2005). les from Asia include a goniopholid and lurosaurs, (Woodward Ornithischians are represented by rare the enigmatic Hsisosuchus from the Lower 1910, Holtz 2000, Rauhut 2003c). Or- heterodontosaurids (Galton 2005), seve- Shixaniao Formation (Peng et al. 2005), nithischian dinosaurs are represented by ral stegosaurian and ankylosaurian thyre- as well as a "sphenosuchian" from the stegosaurs (Galton and Powell 1983, ophorans (Kirkland and Carpenter 1994, Wuceiwan Formation (Clark et al. 2004) Galton 1990), ankylosaurs (Galton 1983) Carpenter et al. 1998, 2001, Galton and and a goniopholid from the Totunhe and basal ornithopods (Galton 1980b, Upchurch 2004) and diverse ornithopods Formation in China (Maisch et al. 2003), Evans and Milner 1994, Kriwet et al. (Foster 2003, Norman et al. 2004, Nor- and a report of a goniopholid from Kir- 1997, Ruiz-Omeñaca et al. 2005). man 2004, Galton 2005). ghizia (Averianov 2000). Another northern fauna of particular in- The most important Middle Jurassic di- Late Jurassic terest in biogeographic terms is that nosaur fauna from the Northern Hemis- from central America and the Caribbean phere is that of the (?) Bathonian Lower One of the most important Late Jurassic (see e.g. Gasparini and Iturralde-Vinent Shaximiao Formation of China (Peng et archosaur faunas is certainly that of the 2006). Taxa reported from that area in- 568 O. W. M. RAUHUT AND A. LOPEZ-ARBARELLO

clude metriorhynchid crocodiles (Gas- Mayr et al. 2005, Göhlich and Chiappe Beds in Africa, Morrison Formation in parini and Iturralde-Vinent 2001, Buchy 2006), and some other finds of similar North America, various units in Portugal et al. 2006, 2007) and rhamphorhynchid taxa from various localities (see Wei- and Solnhofen in Europe, pterosaurs (Colbert 1969, Gasparini et al. shampel et al. 2004). Shishougou Formation in Asia), Middle 2004). In China, Late Jurassic archosaurs are Jurassic archosaurs are almost entirely In Europe, several units of Kimmeridg- mainly known from the Upper Shaxi- known from isolated finds, often also ian age in Portugal have yielded a fauna miao Formation (though see comments fragmentary. This is especially true for that is closely comparable to that of the on the age of this formation above) and the early Middle Jurassic (Aalenian-Bajo- Morrison Formation. Furthermore, the the Shishougou Formation. Crocodiles cian), whereas at least one important Kimmeridgian-Tithonian lithographic reported include basal crocodyliforms fauna is known from the Bathonian-Ca- limestones of southern Germany have (Peng et al. 2005) and goniopholids (see llovian, that of the Lower Shaximiao yielded a rich fauna of mainly non-dino- Wu et al. 1996), as well as another species Formation of China (Peng et al. 2005). saurian archosaurs, marine crocodiles are of Hsisosuchus (Peng et al. 2005). Late This underscores the potential import- known from a number of Late Jurassic Jurassic pterosaurs from Asia are rare, ance of the Late Middle Jurassic Ca- formations, and several isolated archo- but include at least the ctenochasmatid ñadón Asfalto Formation of Chubut, Ar- saur remains have been reported from (Dong 1982). gentina. Although mainly dinosaurs and various localities. Dinosaurs from the Late Jurassic of Asia have been described from this European Late Jurassic crocodiles inclu- include sinraptorid allosauroids, mamen- formation so far (Bonaparte 1979, 1986, de goniopholids (e.g. Schwarz 2002), ato- chisaurid sauropods, and stegosaurian Rauhut et al. 2002, Rauhut 2003a, 2005a, posaurids (e.g. Wellnhofer 1971), teleo- and ornithopodan ornithischians (Wei- Rougier et al. 2007a, b), abundant re- saurid and metriorhynchid thalattosu- shampel et al. 2004, Peng et al. 2005, mains of other vertebrates have also chians (see Mueller-Töwe 2006), and Maidment and Wei 2006). Furthermore, been found, but not described in detail some mesoeucrocodylian taxa of uncer- the oldest ceratopsian ornithischians are yet, including well-preserved pterosaurs tain affinities, such as the enigmatic known from the Tithonian Tuchengzi (Unwin et al. 2004) and remains of cro- Lisboasaurus and Lusitanosuchus (Busca- and the Oxfordian Shishougou forma- codiles (Rauhut et al. 2001), potentially lioni et al. 1996, Krebs and Schwarz 2000, tions of China (Zhao et al. 1999, Xu et al. making this unit a southern counterpart Schwarz and Fechner 2004). Pterosaurs 2006b), and the latter unit has also yiel- of the Lower Shaximiao Formation. are especially represented by the rich fau- ded the basal tyrannosauroid However, on stage level, biases in strati- nas of the Solnhofen limestone (see (Xu et al. 2006a) and the first ceratosaurs graphic distribution are also found in especially Wellnhofer 1970, 1975), but from Asia (Xu and Clark 2006). other epochs of the Jurassic. Thus, Ox- are also known from fragmentary mate- fordian archosaurs are still very poorly rial from other localities (e.g. Wiechmann JURASSIC ARCHOSAUR known, with the Shishougou Formation and Gloy 2000, Fastnacht 2005). Taxa EVOLUTION: A SOUTHERN of China basically representing the only reported in-clude the enigmatic "frog- PERSPECTIVE productive formation of that age. Like- mouthed" anuroganthids (Wellnhofer wise, several stages of the Early Jurassic 1975, Bennett 2007), scaphognathine and Quality of the Jurassic archosaur fos- might be underrepresented, though in- rhamphor-hynchine rhamphorhynchids sil record terpretation is hampered here by the un- (Wellnhofer 1975), ctenochasmatoids certain dating of almost all known terres- (Wellnhofer 1970, Unwin 2003, Jouve From the above, it is clear that the Juras- trial vertebrate localities of this epoch. 2004), and dsungariptoroids (Wellnhofer sic archosaur fossil record is far from Concerning the geographic distribution 1970, Un-win 2003, Fastnacht 2005). complete, both stratigraphically and geo- of Jurassic archosaurs, their fossil record Late Jurassic dinosaurs from Europe in- graphically (Figs. 4, 10), and our under- is strongly biased towards faunas from clude ceratosaurian, basal spinosauroid, standing of Jurassic archosaur evolution the Northern Hemisphere, which be- allosauroid, and various coelurosaurian is actually based on surprisingly few loca- comes evident when looking at the dino- theropods, brachiosaurid and diplodocid lities (Fig. 4). Whereas several important saur record from the Jurassic (Fig. 11). sauropods, and stegosaurian and orni- faunas are known in the Early Jurassic For the entire Jurassic, 121 valid species thopodan ornithischians from Portugal (Upper Elliot and Clarens Formations in of dinosaurs are recognized from Lau- (Antunes and Mateus 2003, Mateus 2006, Africa, Navajo Sandstone, Kayenta For- rasian continents, whereas only 27 have Mateus et al. 2006), basal coelurosaurs mation and equivalent beds in North been described from Gondwana (as of and the basal from the America, Posidonia Shale and equivalent October 2007, only formally named taxa lithographic limestones of southern Ger- beds in Europe, Lufeng Formation in have been considered, OR & ALA, many (Ostrom 1978, Wellnhofer 1992, Asia) and the Late Jurassic (Tendaguru unpublished data). For the different Archosaur evolution during the Jurassic: a southern perspective 569

epochs of the Jurassic, the numbers are more even for the Early Jurassic (19 spe- cies in Laurasia, 10 species in Gondwa- na), but very strongly biased towards northern faunas in the Middle (34 to 8) and especially the Late Jurassic (68 to 9). Thus, most scenarios of Jurassic archo- saur evolution are largely or even entirely based on the fossil record of the Nor- thern Hemisphere. This makes interpre- tation of evolutionary events often very difficult, and especially the hypothesis of increased endemism in terrestrial archo- saurs in response to the initial breakup of Pangea is hard to test in the absence of suitable localities in the Southern Hemi- sphere. For example, until recently it was difficult to evaluate the observed diffe- Figure 8: mesai from the (?)Tithonian Cañadón Calcáreo Formation, Chubut provin- rences between the Laurasian Late Ju- ce, Argentina. a) Vertebral column as preserved (MPEF PV 1716). b) Outline reconstruction show- ing preserved elements. rassic dinosaur fauna and that from the Tendaguru Beds in Tanzania, since, in Cañadón Calcáreo Formation of Argen- archosaur faunas, Africa has the most the absence of other Late Jurassic dino- tina (Rich et al. 1999, Rauhut et al. 2005, complete record, with remains known saur faunas from Gondwana, it was im- Rauhut 2006a, c) might help to put the from almost the entire Early Jurassic, the possible to decide whether these diffe- Tendaguru fauna in a Gondwanan con- latter stages of the Middle Jurassic, and rences reflect differential evolution of text, once more material is excavated and the upper stages of the Late Jurassic, the faunas, or maybe just different ecolo- studied (see also below). though a lot of especially the Middle Ju- gical settings of the localities. Indeed, Taking both the stratigraphic and geo- rassic finds are isolated occurrences. In given this paucity in comparative faunas, graphic bias in Jurassic archosaur distri- South America, only the later Middle Ju- more emphasis has been laid on the sup- bution into consideration, especially the rassic is well represented by the fauna posed similarities of the Tendaguru fau- Gondwanan archosaur record of this ti- from the Cañadón Asfalto Formation, na and those of the Morrison Forma- me can only be called patchy (Figs. 4, 10). though with great potential for new dis- tion and Late Jurassic Portuguese locali- In terms of marine crocodiles, South coveries in both the Early and the Late ties, and a variety of biogeographical hy- America certainly has the most complete Jurassic in Argentina. All other Gondwa- potheses have been proposed to explain record of all Gondwanan continents, nan continents have only isolated occur- these similarities (e.g. Lull 1915, Galton with thalattosuchians known from most rences, in the Early Jurassic in India and 1977, 1982, Mateus 2006). Fortunately, stages from the Sinemurian to the Ti- Antarctica, and in the Middle Jurassic in the newly recognized fauna from the thonian. Concerning terrestrial Jurassic Australia. However, it must be noted that

Figure 9: Outline reconstruction showing pre- served elements of the ceratosaurian theropod Elaphrosaurus from the Middle Saurian Beds (Kimmeridgian) of Tendaguru, Tanzanía, based on the MB dd unnumbered. Scale bar: 50 cm. 570 O. W. M. RAUHUT AND A. LOPEZ-ARBARELLO

even for the Jurassic stages with faunas known in the Gondwanan continents, these are usually single localities or for- mations, and comparable material from other parts of the continent is usually absent.

Faunal composition, evolutionary considerations and biogeography

a) Marine crocodiles: The evolutionary and biogeographic significance of the thalat- tosuchian crocodiles, especially of South America, has been discussed at length by Zulma Gasparini and co-workers (e.g. Gasparini 1985, 1992, 1996, Gasparini and Fernandez 1997, Gasparini et al. 2000, 2005, 2006), so only the most im- portant aspects will be outlined here. The occurrence of thalattosuchian cro- codiles in the Sinemurian of Chile (Chong Diaz and Gasparini 1972) and western Argentina (Huene 1927), and, probably, the Early Jurassic of India (Bandyopadhyay et al. 2002) demonstra- tes that this clade did not only have a cir- cumtethyan distribution in the Early Ju- rassic, but also already entered the Pa- cific region. This dispersal most probably happened through the Hispanic corridor (Gasparini et al. 2000), which was spora- dically open since at least the Pliens- Figure 10: bachian (Aberhan 2001), though the con- Stratigraphic com- temporaneous occurrence of the first pleteness of the archosaur fossil thalattosuchians in the Sinemurian of record of Gondwana. Europe and Chile might indicate that a) Crocodiles. b) marine faunal exchanges may have been Pterosaurs. c) Dinosaurs. Dotted possible even before. lines indicate uncer- With the repeated opening and closing of tain limits of the the Hispanic corridor during the Middle stratigraphic range of a certain geological Jurassic (Westermann 1993, Moyne et al. unit. Some minor 2004), there might have been an exchan- localities with frag- ge of marine crocodiles between the Pa- mentary and uninfor- cific and the Tethys every time the corri- mative material have been omitted, in the dor was in effect. This exchange was pro- chart for dinosaurs, bably not unidirectional; whereas the only localities that oldest representative of Metriorhynchus is have yielded at least two identifiable taxa found in Chile (Gasparini et al. 2000), and are shown. India the genus only later appears in Europe includes Madagascar, (Mueller-Töwe 2006), Geosaurus is first since these two areas were geographically known in Europe, and only subsequently united in the Jurassic. in South America (Pol and Gasparini Archosaur evolution during the Jurassic: a southern perspective 571

2007). The evolution of thalattosuchians in the Middle Jurassic might thus be cha- racterized by periods of dispersal separa- ted by periods of cladogeneses in separa- tion in the Tethys and the Pacific. Thus, the Hispanic corridor might have been a "motor" of thalattosuchian evolution in the Middle Jurassic, though this hypothe- sis needs further testing by new finds and rigorous cladistic and biogeographic ana- lyses of all species of Middle and Late Jurassic thalattosuchians. The exchange between the Tethyan re- gion and the Pacific continued during the Figure 11: Valid dinosaur spe- Late Jurassic, when the Hispanic corridor cies from the Jurassic of the was permanently open (Ford and Go- Northern and Southern lonka 2003), as evidenced by shared ge- Hemisphere (as of October 2007, OR & ALA unpublished nera of metriorhynchids, such as Geosau- data). rus and Dakosaurus. It is interesting, though, that, whereas all three genera of dylian lineages and Go- Jurassic. First, both groups might have metriorhynchids (Metriorhynchus, Geosau- niopholidae in the Early Jurassic, indicate been genuinely rare during the Jurassic rus and Dakosaurus) are present in both that the major radiation of mesoeucro- and were maybe restricted to a small re- areas, no teleosaurids have been reported codylians already happened during this gion, and only experienced their great from the Late Jurassic of western South period (Pol and Gasparini 2007), but radiation in the Cretaceous. Alternatively, America so far, although they are present most lineages are poorly represented glo- both groups might have been largely res- both in the northern and southern part bally before the Late Jurassic. The pre- tricted to the Southern Hemisphere al- of the Tethys during this period (e.g. sence of possible atoposaurids in the ready in the Jurassic, but not necessarily Krebs 1967, Bardet and Hua 1996). This Middle Jurassic in Madagascar and a pos- been rare, and have not been reported so might be due to the more fully marine sible goniopholid in the Late Jurassic of far due to the extremely poor record of environments represented by the thalat- Tanzania, if the identity of these frag- terrestrial vertebrates in the Jurassic of tosuchian-bearing sediments in Chile and mentary remains can be confirmed, Gondwana. Which of these scenarios, Argentina (Gasparini et al. 1999), and fur- might show that these more advanced which might not be completely exclusive, ther finds of marine from nears- mesoeucrocodylians were also globally is correct can only be demonstrated by hore environments will have to show distributed before the final separation of more material of terrestrial crocodiles whether this group was truly absent from the continents of the Northern and from the Southern Hemisphere. the Pacific coast of South America. Southern Hemisphere. Furthermore, the c) Pterosaurs: As in the case of the terres- b) Terrestrial crocodiles: Interpretation of atoposaurid record from Madagascar trial crocodiles, the extremely poor Ju- the evolution of terrestrial crocodiles in (Flynn et al. 2006) is roughly contempo- rassic Gondwanan record of pterosaurs Gondwana during the Jurassic is hampe- raneous with the oldest records of this severely restricts any interpretation of red by the abysmal fossil record of these group in the Northern Hemisphere (e.g. their evolutionary history in the Sou- in the Middle and Late Jurassic Kriwet et al. 1997), indicating that derived thern Hemisphere. From the scarce re- of the Southern Hemisphere. In the Ear- members of this lineage might originated mains, it is evident that pterosaurs were ly Jurassic, the first successful radiations already considerably earlier. already present in Gondwana in the Early of the crocodylomorph lineage, the However, the lack of two important me- Jurassic. However, pterosaur remains are "sphenosuchians" and basal crocodyli- soeucrocodylian lineages, which are espe- unknown from the best known Early Ju- forms ("protosuchians"), had a Pangean cially widely distributed in the Southern rassic vertebrate fauna from the South- distribution, as evidenced by the occu- Hemisphere during the Cretaceous, the ern Hemisphere, that of the Upper rrence of these groups in southern notosuchians and the peirosaurids (Fig. Stormberg Group of South Africa, des- Africa, North America and China during 1, see Pol and Gasparini 2007), during pite the fact that the Upper Elliot For- this time. Phylogenetic relationships the Jurassic is striking. Two possible sce- mation frequently preserves small verte- among crocodylomorphs, especially the narios might explain the lack of these brate remains in often exquisite detail. presence of the advanced mesoeucroco- clades in the fossil record during the This might indicate that these animals 572 O. W. M. RAUHUT AND A. LOPEZ-ARBARELLO

might have been very rare in southern tainly have the best Jurassic fossil record Marsicano 2001), and southern Africa Gondwana at that time. That pterosaurs in the Southern Hemisphere, though (Butler et al. 2007), indicates that the ori- had colonized even southern Gondwana their record is certainly far from comple- gin and early diversification of ornithis- by the Early Jurassic is shown by the iso- te, and even very poor if compared with chians is probably found in southern lated humerus found in the Hanson For- that of the Northern Hemisphere (see Gondwana. The phylogenetic position of mation of Antarctica (Smith and Pol above). Thus, all interpretations of Juras- Lesothosaurus as the basalmost thyreopho- 2007). sic dinosaur evolution in Gondwana ran known and of Stormbergia as the If confirmed, the occurrence of a ptero- should be regarded as tentative, and basalmost cerapodan (Butler et al. 2008) dactyloid in the Callovian Cañadón As- might easily change in the light of new is in general accordance with this inter- falto Formation of Argentina would be discoveries. pretation (Fig. 12). the earliest record of this clade globally, Early Jurassic dinosaurs faunas, especially Thus, the poor Gondwanan fossil record which, in the Northern Hemisphere ap- that of the Elliot and Clarens formations in the Middle Jurassic might account for pears abruptly in the fossil record with and equivalent beds in Zimbabwe in sou- our still rather poor understanding of the many different clades in the Kimmeridg- thern Africa highlight the close faunal re- early Mesozoic radiation of ornithis- ian-Tithonian (Unwin 2006). Once pro- lationships in different parts of Pangea at chians, since the major radiation of this perly studied, the pterosaur fauna from that time. Basal thyreophoran ornithis- group, in which basically all important the Cañadón Asfalto Formation might chians are present in basically all impor- lineages were established, took place dur- prove to be of great importance for our tant Early Jurassic dinosaur localities, in ing this time (Sereno 1999a, Butler et al. understanding of pterosaur evolution, southern Africa (Butler et al. 2008), the 2008). It is interesting, however, to note since it represents one of the very few Kota Formation of India (Nath et al. that ornithischians seem to have been strictly terrestrial faunas that have yielded 2002, see above), the Kayenta Formation rather rare in the Middle Jurassic of abundant and well-preserved pterosaur of western North America (Colbert Gondwana, where they have only been material (see Buffetaut 1995). 1981, Padian 1989), the Liassic of Eu- reported in the best known fauna, that of From the Late Jurassic, only pterodacty- rope (Owen 1863, Haubold 1990), and the Cañadón Asfalto Formation (see loid pterosaurs have been reported from the Lufeng Formation of China (Nor- above), and here they seem to be South America so far (Martill et al. 2000, man et al. 2007). This global distribution amongst the rarest vertebrate groups Codorniú and Gasparini 2007), but both gives testimony to the rapid evolution of represented in general (OR, pers. obs.). "rhamphorhynchoids" and pterodacty- thy-reophorans, which seem to represent Nevertheless, the wide distribution and loids are present in eastern Africa (Reck one of the first successful radiations of diversity of ornithischians on higher 1931, Galton 1980, Unwin and Heinrich ornithischians, soon after the origin of taxonomic level in the Late Jurassic indi- 1999). Again, the wide distribution of this group in the . cates that much of this radiation took pterydactyloids indicates that their evolu- Another group of ornithischians from place during Pangean times in the Early tion and radiation was well under way by southern Africa is of special interest as or Middle Jurassic, and so it should not the time the first members of this clade well, the heterodontosaurids. Heterodon- be too surprising to find similar groups appear in the fossil record. Furthermore, tosaurids represent the most diverse and of ornithischians in the faunas of the the occurrence of dsungaripteroids in abundant group of ornithischians in the Northern and Southern Hemisphere Europe (see Unwin 2003, Fastnacht Upper Elliot and Clarens Formations even after the final separation of the 2005), Africa (Unwin and Heinrich 1999) (Butler 2005, Knoll 2005), but they seem continents of Laurasia and Gondwana at and, possibly, South America (Martill et to be rare in the Northern Hemisphere, the Middle-Late Jurassic boundary. al. 2000) indicates a still global distribu- from where only one undescribed hete- Early Jurassic saurischians from Gond- tion of these animals in the Late Jurassic, rodontosaurid from the Kayenta Forma- wana are also indicative of a global Pan- as might be expected from clade of active- tion has been reported (Sereno pers. gean dinosaur fauna at that time (see also ly flying animals, and is also still found in com. in Butler 2005). The fact that hete- Smith et al. 2007b). In theropods, coe- the Cretaceous for several pterosaur cla- rodontosaurids probably represent the lophysoids are found on all major land- des, despite the separation of continental most basal clade and thus the earliest masses from which theropods have been masses (Unwin 2006). radiation of ornithischian dinosaurs (But- reported (Raath 1969, 1977, Rowe 1989, Again, more fossil finds from Gondwa- ler et al. 2008), together with the occur- Carrano and Sampson 2004, Irmis 2004, nan localities are needed to better un- rence of the basalmost ornithischians Tykoski 2005), with the exception of An- derstand pterosaur evolution during the known in the Late Triassic of South tarctica. Furthermore, the recently re- Jurassic. America (Casamiquela 1967, Ferigolo cognized dilophosaur clade (Smith et al. d) Dinosaurs: Compared to the other and Langer 2007), including the oldest 2007a, b) also shows a global distribu- groups discussed above, dinosaurs cer- known heterodontosaurid (Báez and tion, with representatives known from Archosaur evolution during the Jurassic: a southern perspective 573

Figure 12: Phylogeny of early ornitischians (based on Butler et al. 2007, 2008), showing the geographic distribution of early ornitischians. North America, China, South Africa, and Southern Hemisphere in the Late Juras- , the following discussion will Antarctica. Likewise, close relatives of sic and Cretaceous (see Tykoski and Ro- mainly consider . New dis- the southern African sauropodomorph we 2004, Rauhut 2005b), so its occurren- coveries and paleogeographic considera- Massospondylus are known from the Early ce already in the Early Jurassic would tions contradict the idea of restricted Jurassic of North (Attridge et al. 1985) represent a surprising case of endemism Early Jurassic faunas and an only Middle and South America (Martínez 1999), and of this group, especially considering the to Late Jurassic global distribution of Smith and Pol (2007) found close rela- northern Gondwanan (and thus more or sauropods. First, the discovery of proba- tionships of this genus with the Chinese less central Pangean) position of this ble sauropod remains in Antarctica and the Antarctic Glaciali- find and the global distribution of most (Smith and Pol 2007) might show that saurus, so that the Massospondylidae other groups. sauropods had reached low southern lati- (sensu Smith and Pol 2007) represent ano- Of special interest are furthermore the tudes by that time, although a detailed ther successful Pangean radiation of sauropods, which originated in the Trias- study of this material is needed to esta- dinosaurs in the early Mesozoic. Thus, sic (see Yates and Kitching 2003), but blish its sauropod affinities and answer especially if the radiation of dinosaurs seem to have experienced their first im- the question whether this material repre- really followed a Late Triassic portant radiation across the Triassic-Ju- sents a basal taxon, or a eusauropod. Fur- event, as argued by Olsen et al. (2002), rassic boundary. Gillette (2003: 686) sug- thermore, the occurrence of Amygdalodon this Pangean distribution of many groups gested that Early Jurassic sauropods had at around the Early-Middle Jurassic already in the Early Jurassic indicates a distribution in eastern Gondwana and boundary in Argentina (Rauhut 2003b) high evolutionary dynamics and great southern and eastern Laurasia, and were and other fragmentary eusauropod re- dispersal potential of these animals. It restricted to middle latitudes, and that mains from the Aalenian of this country might be noted, however, that high dino- sauropods reached North America and (Salgado and Gasparini 2004) indicate saur diversity already in the Triassic in western Gondwana only in the Middle or that eusauropods were also present in other parts of the world, including South even Late Jurassic. Most of this argu- western Gondwana early in the Jurassic. America, and inferred ghost lineages ment was based on negative evidence, i.e. Considering palaeogeographic recons- argue against such a sudden radiation the lack of sauropod fossils from these tructions of the Early Jurassic (Scotese after a major extinction event at the T-J regions during the Early Jurassic. Al- 1991, Smith et al. 1994), it seems further- boundary. though some non-eusauropodan taxa more unlikely that a dispersal of eusauro- Interesting is the report of a probable were included in Gillette's analysis (e.g. pods from eastern Gondwana to eastern abelisauroid from the Early Jurassic of Vulcanodon), most of this scenario in- Laurasia (or vice versa) could have happen- Gondwana (Allain et al. 2007). This line- voves eusauropods, and, given the cur- ed withouth these animals entering age is almost entirely restricted to the rent confusion about the definition of North America as well. Indeed, the pre- 574 O. W. M. RAUHUT AND A. LOPEZ-ARBARELLO

sence of , which, according to tive character of the vertebrae might be and the same is true for those forms still Yates (2004), represents an early, non-eu- due to the juvenile state of the type ma- kept in the same genera as comparable sauropodan sauropod (see also Fedak terial. However, the latter notion seems taxa from the Morrison Formation. In and Galton 2007), in the Early Jurassic of to be unlikely, since similar sized, juve- the case of Brachiosaurus brancai, for North America indicates that immediate nile specimens referred to Patagosaurus example, a rigorous phylogenetic analysis eusauropodan outgroups already had a (MACN Ch 932, 934), for example, al- of the taxa referred to the family Bra- global distribution during this time. This ready show more complicated vertebral chiosauridae would be necessary to test is also in agreement with the find of the structures (OR pers. obs.). Thus, the phy- whether this species really is more closely probable primitve eusauropod Ohmdeno- logenetic position of this interesting related to Brachiosaurus altithorax from the saurus in central Europe (Wild 1978), a taxon has to remain unresolved, pending Morrison Formation, or whether the lat- region that was geographically close to a revision of the known, or the discovery ter might rather group with other Lau- the eastern coast of North America. The of new material. Being intercalated rasian brachiosaurids, such as Sauroposei- absence of eusauropods from western within Laurasian taxa, or having close re- don. The same is true for Dysalotosaurus let- North America might thus have pelaeoe- latives in Laurasia, these Middle Jurassic towvorbecki (currently kept in the North cological reasons, or might simply be due sauropods furthermore indicate a gene- American genus Dryosaurus, see Galton to incomplete sampling of these faunas. rally Pangean sauropod fauna in the late 1977, Norman 2004). In this case, the Thus, it seems most probable that eusau- Middle Jurassic. problem is even more acute, since Dysa- ropods also reached a Pangean distribu- Of special interest are, of course, the di- lotosaurus and Dryosaurus represent the tion already in the Early Jurassic. nosaur faunas of the Late Jurassic of the only well-known dryosaurids, wheras all In the Middle Jurassic, the change in the- Southern Hemisphere, since it is in these other taxa within the Dryosauridae are ropod faunas from coelophysoid/basal faunas that we would expect to find first based on isolated elements or very frag- theropod dominated Early Jurassic fau- consequences of the separation of mentary remains. Thus, no analysis of nas to basal tetanuran faunas seems to Gondwana from the continents of the the interrelationships of dryosaurids has have been paralleled in the Southern Northern Hemisphere. However, analy- ever been attempted. In the case of Hemisphere (Rauhut 2005a), though the sis of these effects has been hampered Elaphrosaurus, the North American speci- timing and mode of this change remains by the fact that, until very recently, only mens referred to this taxon (Galton uncertain due to the very poor Middle one dinosaur fauna had been reported 1982, Chure 2001) might represent cera- Jurassic theropod record of Gondwana, from this period in Gondwana, that of tosaurs, but cannot be referred to this ge- which, in terms of identifiable taxa, is the Tendaguru Beds in Tanzania, so that nus (Rauhut and Carrano in prep). These restricted to the Cañadón Asfalto Forma- material for comparisons was missing examples again highlight the need for tion. Basal, non-neosauropodan eusauro- (see above). This had another effect in rigorous phylogenetic analyses as the pods were widely distributed in Gond- the description of the fauna from Ten- basis for biogeographic interpretations, wana during the Middle Jurassic, and daguru: due to the lack of comparable as already emphasized by Forster (1999). seem to represent several lineages of an material from the Southern Hemisphere, In the absence of such phylogenetic ana- early radiation of eusauropods, some of the taxa from this locality have generally lyses for the taxa concerned, a faunistic which might go back to the early Jurassic. been compared and often even been approach might shed some light on the This notion is supported by the associa- referred to taxa known from the North- biogeographic affinities of the Tendagu- tion of archaic forms in the later Middle ern Hemisphere, which led to the ru fauna. Although similar in terms of Jurassic, such as Archaeodontosaurus (Buf- emphasis on faunal similarities between taxonomic composition at higher taxo- fetaut 2005) or Volkheimeria (Bonaparte the fauna of Tendaguru and especially nomic levels, the relative abundance of 1986b, 1999), with much more progressi- that of the Morrison Formation. How- taxa and overall composition of the fau- ve forms of sauropods, some of them ever, although this similarity certainly na from this locality is strikingly different already close to or within , exists on higher taxonomic level, this from that of the Morrison Formation. In such as Atlasaurus (Monbaron et al. 1999, might not be too significant, since most the latter, the most common sauropod is Upchurch et al. 2004). Lapparentosaurus is lineages represented probably reach back Camarasaurus, with its several species, and a taxon of problematic position, since it to the Middle Jurassic and thus to the diplodocids, whereas titanosauriforms shows a combination of plesiomorphic time before the separation of Gondwana (only known from brachiosaurids in this and apomorphic characters (Upchurch et (see Remes and Rauhut 2005, Remes formation) are very rare (Dodson et al. al. 2004). Upchurch (1998) found this 2006, 2007). Recent work has further- 1980, Foster 2003). In Tendaguru, in taxon to represent a brachiosaurid, which more shown that taxonomic assignment contrast, brachiosaurids and other basal would be the oldest record of a derived to Laurasian forms on lower taxonomic titanosauriforms are among the most neosauropod, and argued that the primi- level is often questionable (Remes 2006), common sauropods (Russell et al. 1980), Archosaur evolution during the Jurassic: a southern perspective 575

which would be in good accordance with the titanosauriform dominated Creta- ceous faunas in Gondwana. Although this difference might be assigned to pa- laeoecological reasons, the same seems to be true for the Kadsi Formation of Zim- babwe (Raath and McIntosh 1987) and, maybe, the Cañadón Calcáreo Formation of Argentina (with Tehuelchesaurus repre- senting a basal titanosauriform or an im- mediate titanosauriform outgroup, Rau- hut et al. 2005, Carballido et al. 2007), though more sampling is needed to con- firm this. In the latter case, the paleoeco- logical setting seems to be different from that of Tendaguru, and this locality was placed considerably further south than the latter in Late Jurassic times. In addition to this relative abundance of specimens, differences in the faunal com- position are also reflected in the taxono- Figure 12: Taxonomic composition of Late Jurassic sauropod faunas of the Northern and Southern Hemisphere. a) Taxa represented in the Morrison Formation, USA and various localities in Portugal. mic composition of the sauropod faunas b) Taxa represented in the Tendaguru Beds, Tanzania, and other Gondwanan localities. Basal titano- (Fig. 13). For this comparison, only the sauriforms in b) include Tehuelchesaurus, which might be an immediate titanosauriform outgroup. sauropod fauna of the Morrison For- mation (Foster 2003, Upchurch et al. sauriforms (or immediate titanosauri- of both camarasaurids and dicraeosau- 2004) and the Late Jurassic of Portugal form outgroups) account for almost half rids indicate that these groups must have (Antunes and Mateus 2003) have been of the species in the Southern Hemi- originated in Pangean times as well, their considered for the Northern Hemi- sphere (4 of 9 species), followed by di- absence in the Late Jurassic in one of the sphere, since eastern Asia was separated craeosaurids (3 of 9 species), whereas Hemispheres must either be due to the from western Laurasia and Gondwana by only two species of diplodocids have be- failure to disperse to the other Hemi- epicontinental seas from at least the Ba- en described from Gondwana. The most sphere in Middle Jurassic times, or to dif- thonian (Smith et al. 1994), and, thus, the remarkable aspect of this faunal compa- ferential extinction in Laurasia and Chinese Late Jurassic sauropod fauna rison, however, is the lack of one of the Gondwana (see also Harris and Dodson shows a rather high endemism and can most common groups of sauropods 2004). Although this question can only thus not be regarded as a "typical Lau- from the Morrison Formation, the cama- be answered by new discoveries, the lack rasian" fauna (Upchurch et al. 2002). For rasaurids, in Gondwana, and, in turn, the of these important faunal components in the Southern Hemisphere, the Tenda- lack of an important Gondwanan line- the respective other Hemisphere might guru fauna and the two named taxa from age, the dicraeosaurids, in Laurasia. Al- thus represent a first sign of differential the Cañadón Calcáreo Formation have though the former might still be accoun- evolution of the dinosaur faunas of the been considered together as representa- ted for by the poor Gondwanan fossil two Hemispheres due to limitations in tives of the Late Jurassic Gondwanan record and the very different sample dispersal potential, following the separa- fauna. Taxa have been counted on spe- sizes, the latter is especially noteworthy, tion of Gondwana. cies level. since dicraeosaurids have been reported Looking thus at sauropod faunal compo- from all three Gondwanan localities that CONCLUSIONS sition, diplodocids constitute more than have yielded identifiable Late Jurassic half of the known sauropod taxa from sauropods (Raath and McIntosh 1987, Jurassic archosaurs from the Southern He- the Northern Hemisphere (11 of 21 spe- Aberhan et al. 2002, Rauhut et al. 2005), misphere are still rather poorly known. cies), whereas basal titanosauriforms (2 even though they are geographically, lati- Although this is especially true for terres- species) account for only ten percent of tudinally (and thus probably climatically) trial crocodiles and pterosaurs, the dino- the species. Camarasurids are the second and, as far as can be said at the current saur record from Gondwana is also still most important faunal element, with state of knowledge, ecologically dispara- far from complete. The most important 14% (3 species). In contrast, basal titano- te. Since the phylogenetic relationships archosaur faunas from the Jurassic of the 576 O. W. M. RAUHUT AND A. LOPEZ-ARBARELLO

Southern Hemisphere are that of the invitation to participate in the Jurassic tions for the phylogeny of basal . -Sinemurian Elliot and Sine- Symposium in Mendoza and to contribu- Journal of Vertebrate 22(3): murian- Clarens forma- te to this special volume. Too many col- 548-563. tions of southern Africa, the (?) Hettan- leagues to mention everybody have con- Allain, R. 2005. The postcranial anatomy of the gian-Toarcian Kota Formation of India, tributed to this work, with access to spe- megalosaur Dubreuillosaurus valesdunensis (Di- the Early Jurassic Hanson Formation of cimens, discussions, and critical com- nosauria, Theropoda) from the Middle Juras- Antarctica, the Callovian Cañadón As- ments on ideas expressed in meetings or sic of Normandy, France. Journal of Verte- falto Formation of Argentina, and the personal discussions over the , but brate Paleontology 25(4): 850-858. Kimmeridgian-Tithonian Saurian Beds we are thankful to each and every one of Allain, R. and Chure, D.J. 2002. Poekilopleuron buc- of Tendaguru, Tanzania. Marine crocodi- them for this input. Special thanks are klandii, the theropod dinosaur from the Mid- les are especially well-represented in the due to the Museo Paleontológico Egidio dle Jurassic (Bathonian) of Normandy. Pa- western part of southern South America, Feru-glio, Trelew, for the support during laeontology 45(6): 1107-1121. from where different taxa are known fieldwork in the Middle and Late Jurassic Allain, R., Aquesbi, N., Dejax, J., Meyer, C., from units ranging from the Sinemurian of Chubut Province, Argentina, and to Monbaron, M., Montenat, C., Richir, P., to the Tithonian. Multiple events of fau- Wolf Volkheimer for many insights into Rochdy, M., Russell, D. and Taquet, P. 2004. A nal exchange through the sporadically the Jurassic history of South America. basal sauropod dinosaur from the Early Ju- open Hispanic corridor during the Early The manuscript greatly benefited from rassic of Morocco. Comptes Rendus Palevol and Middle Jurassic might have been an critical comments by Diego Pol and Max 3: 199-208. important factor in thalattosuchian evo- Lan-ger. Allain, R., Tykoski, R.S., Aquesbi, N., Jalil, N.-E., lution. Parts of the research that led to this Monbaron, M., Russell, D.A. and Taquet, P. Terrestrial archosaur faunas show a Pan- review paper were financially supported 2007. An abelisauroid (Dinosauria: Thero- gean distribution in the Early Jurassic, by a DAAD fellowship, grants from Fun- poda) from the Early Jurassic of the High and, most probably in the Middle Jurassic dación Antorchas, the Jurassic Foun- , Morocco, and the radiation (for which almost no terrestrial crocodi- dation, BBC Horizon, and German of ceratosaurs. Journal of Vertebrate Paleon- les have been reported from the Sou- Research Foundation (DFG) grants RA tology 27(3): 610-624. thern Hemisphere). The rather high di- 1012/1-1, 2-1, 8-1, and 9-1 to OR, and Antunes, M.T. and Mateus, O. 2003. Dinosaurs of versity of basal ornithischian dinosaurs DFG grant LO 1401/1-2 to ALA. Portugal. Comptes Rendus Palevol 2: 77-95. in the Early Jurassic of southern Africa Attridge, J., Crompton, A.W. and Farish A. Jen- might indicate that the early radiation of WORKS CITED IN THE TEXT kins, J. 1985. The southern African Liassic this clade took place in southern Gond- prosauropod Massospondylus discovered in wana. In the Late Jurassic, especially sau- Aberhan, M. 2001. Bivalve palaeobiogeograohy North America. Journal of Vertebrate Pa- ropod faunas might already show the and the Hispanic Corridor: time of opening leontology 5(2): 128-132. first indications of differential faunal and effectiveness of a proto-Atlantic seaway. Averianov, A.O. 2000. 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