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Notobatrachus degiustoi. A, anterior portion of skull in dorsal aspect (cast of CTES-Pz 5739B); B, anterior portion of skull in ventral aspect (cast of CTES-Pz 5739A); C, dorsal aspect of juvenile (cast of MLP 55-VI-1-4) / A, parte anterior del cráneo en vista dorsal (molde de CTES-Pz 5739B); B, parte anterior del cráneo en vista ventral (molde de CTES-Pz 5739A); C, vista dorsal de juvenil (molde de MLP 55- VI-1-4).  

Notobatrachus degiustoi. A, anterior portion of skull in dorsal aspect (cast of CTES-Pz 5739B); B, anterior portion of skull in ventral aspect (cast of CTES-Pz 5739A); C, dorsal aspect of juvenile (cast of MLP 55-VI-1-4) / A, parte anterior del cráneo en vista dorsal (molde de CTES-Pz 5739B); B, parte anterior del cráneo en vista ventral (molde de CTES-Pz 5739A); C, vista dorsal de juvenil (molde de MLP 55- VI-1-4).  

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Notobatrachus degiustoi is the most completely known Jurassic frog and has been recorded in many outcrops of the La Matilde Formation of the Deseado Massif area in southern Patagonia. Herein, we erect a new species of the genus based on partially articulated remains collected from the Callovian Las Chacritas Member of the Canadon Asfalto Formation,...

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... Tympanic hearing is known to be a primitive condition in anurans. Complete middle ears have been found in multiple fossil taxa, including the early Triassic stem anuran Triadobatrachus massinoti (Rage & Roček, 1989), the early Jurassic anuran Prosalirus bitis (Shubin & Jenkins, 1995), the middle Jurassic anurans Notobatrachus degiustoi and N. reigi (Báez & Basso, 1996;Báez & Nicoli, 2008), and the Jurassic-Cretaceous anurans Callobatrachus sanyanensis and Mesophryne beipiaoensis (Gao & Wang, 2001). Throughout the evolutionary history of anurans, the TME has been lost partially or completely at least 38 times, and at least 10 of those losses occurred in the family Bufonidae (Pereyra et al., 2016). ...
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The tympanic middle ear (TME) is a complex morphological adaptation that facilitates auditory reception in multiple clades of tetrapods, including anurans. Despite its utility, it has been partially or completely lost at least 38 times in anurans, and there is no clear explanation for how or why this loss occurred. We used micro-CT scans and 3D segmentation to catalog presence/absence and shape of the columella throughout the Central American toad genus Incilius. We found that the TME has been lost multiple times in Incilius and our outgroup Rhinella, with at least one regaining event. Absence of the columella and the different shapes of the columella both have high phylogenetic signal. Incilius species have a different distribution of columella shapes than many other genera of anurans that have been surveyed, including more blade-shaped columellae than found in other genera and one species with vestigial remnants of a columella. The columella shape most common in Incilius, a blade, may be derived from the rod shape also found in many other bufonids; the adaptive significance of this trait remains unknown. This study succeeds in providing a fine-scale review of earlessness and columellar morphology in Incilius, but the processes underlying the patterns we observe remain unclear. The tympanic middle ear (TME) plays an important role in sound detection and is thus important in location of prey, detection of predators, and social communication among tetrapods. The TME is a morphological complex formed by four main elements: the tympanic membrane, which captures sound pressure in the air and is the most lateral (or superficial) element of the middle ear; the tympanic an-nulus, a cartilaginous ring that attaches to the tympanic membrane and provides support for it; the middle ear cavity ; and the columella, a structure of cartilage and bone that crosses the middle ear cavity and connects the tym-panic membrane to the inner ear, where sound waves produce fluid motion that is transduced by hair cells into electrical signals (Pereyra et al.
... In the Puesto Almada Member spinicaudatans (Gallego et al., 2010;Monferran et al., 2013), bivalves (Martínez et al., 2007), gastropods , ostracods (Musacchio, 1995;Ballent and Díaz, 2011), and Trichoptera (Caddisflies) were identified. The vertebrate content includes tetrapods, amphibians, turtles, dinosaurs, and mammals (Bonaparte, 1979a, b;Rich et al., 1999;Sterli, 2008;Rauhut et al., 2002Rauhut et al., , 2005Martin and Rauhut, 2005;Rougier et al., 2007;Ba ez and Nicoli, 2008;Escapa et al., 2008;Pol and Rauhut, 2012), and fish remains (Bochino, 1967;Bordas, 1943;L opez Arbarello et al., 2013). The member also includes megaplants (equisetaleans, ferns, and conifers -Frenguelli, 1949;Cort es and Baldoni, 1984;Escapa et al., 2008) and gymnosperm and cheirolepidiacean pollen grains . ...
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The Cañadón Asfalto basin, central Chubut, Argentina, comprises a volcano-sedimentary sequence related to the opening of the Atlantic Ocean during Mesozoic times. The Lonco Trapial, Cañadón Asfalto and Cañadón Calcáreo formations are the main units related to the evolution of this basin. The Las Chacritas and Puesto Almada members are distinguished in the Cañadón Asfalto Formation. LA-HR-ICP-MS U-Pb and Lu-Hf data on zircon were obtained on these units. The Lonco Trapial Formation gave a weighted average age of 172.3 ± 1.8 Ma. A pyroclastic level from the Las Chacritas Member gave a weighted average age of 168.2 ± 2.2 Ma. Two U-Pb concordant ages of 160.3 ± 1.7 Ma on a laminated tuffite and 158.3 ± 1.3 Ma on a pyroclastic level were obtained for the Puesto Almada Member. Two maximum depositional ages constrain the sedimentary provenance areas for the basin: 1) A sample from the Sierra de la Manea range, where a controversial unit related either to the Cañadón Asfalto or to the Cañadón Calcáreo formation occurs, gave an age of 176.6 ± 1.0 Ma. Two younger zircon crystals indicate that this unit may be related to the Cañadón Calcáreo Formation. 2) A sandstone with cross-stratification from the Puesto Almada Member gave a maximum depositional age of 173.6 ± 6.4 Ma. In terms of U-Pb and Lu-Hf isotopes, two magmatic events are identified in central Patagonia: the Mamil Choique magmatic event characterized by negative εHf values around −5.0 and representing recycling during Permian times of Mesoproterozoic crust (TDM of ∼1.5 Ga), and the Cañadón Asfalto magmatic event with negative (−8.2) to positive (+4) εHf values and Meso- to Neoproterozoic TDM between 1.5 and 0.8 Ga. The younger event is characterized by three main cycles: C1 related to the Lonco Trapial magmatism, C2 to the Las Chacritas volcanism, and C3 to the Puesto Almada volcanism. These cycles are related with Marifil, Chon Aike and El Quemado formations volcanics events of Patagonia and the Neuquén Basin during the Mesozoic.
... The dorsal tubercle (tuber superior) is present as a small process rather than a distinct protuberance anterodorsal to the supraacetabulum crest (PKUP V0402-V0404, and V0407), for the attachment of m. gluteus magnus (Duellman and Trueb, 1986). In living frogs, the dorsal tubercle is either absent or low, but the lack of a distinct dorsal tubercle is a plesiomorphic condition for Anura, as evidenced by the condition in the Jurassic stem anurans Prosalirus and Notobatrachus (Shubin and Jenkins, 1995;Báez and Nicoli, 2008). The supraacetabulum crest is present, but not noticeably high. ...
... The Triassic proanuran Triadobatrachus has as many as 14 presacral vertebrae (Rage and Roček, 1989), whereas the number of vertebrae is unknown for the closely related Czatkobatrachus from the Triassic of Poland (Evans and Borsuk-Bialynicka, 1998). One Jurassic stem anuran, Vieraella, has 10 presacrals, and Notobatrachus has nine (Báez and Basso, 1996;Báez and Nicoli, 2008). The Early Jurassic Prosalirus from North America is based on a composite of isolated elements from different individuals; thus, the actual number of presacral vertebrae cannot be determined (Shubin and Jenkins, 1995). ...
... The rigid ilio-sacral articulation of Triadobatrachus suggests the incapacity of jumping locomotion (Shubin and Jenkins, 1995). The modern frog pelvis appeared first in the Jurassic Prosalirus and Notobatrachus (Shubin and Jenkins, 1995;Báez and Nicoli, 2008), characterized by a nonexpanded sacral diapophysis directed posterolaterally, smooth dorsal surface of ilium and urostyle without a ridge, and a bicondylar sacral-urostyle articulation. This set of characters is also seen in the most primitive living frogs Ascaphus + Leiopelma, all Early Cretaceous fossil discoglossids and pipoids from Spain (Báez and Sanchiz, 2007;Báez, 2013), and the Early Cretaceous fossil neobatrachians from Brazil (Báez et al., 2009). ...
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Based on 12 well-preserved skeletons of postmetamorphic individuals, a new crown-group frog taxon is named and described from the Lower Cretaceous Guanghua (upper part of Longjiang) Formation (stratigraphic equivalent of the world-famed Yixian Formation) exposed in Dayangshu Basin, Hulunbuir, in the far northeast of Inner Mongolia, China. The new taxon, Genibatrachus baoshanensis, documents another Early Cretaceous anuran having reduction of the presacral vertebrae to eight in number, similar to several frog taxa of roughly the same age from Spain and Brazil. The new frog also displays several features that are ontogenetically and phylogenetically informative, including ontogenetic fusion of the palatine to the sphenethmoid, and ontogenetic fusion of ribs to the diapophyses of the posterior trunk vertebrae. In addition, the new discovery extends the geographic range of Early Cretaceous frogs of the Jehol Biota northward to near the 50th parallel north in East Asia.
... The dorsal tubercle (tuber superior) is present as a small process rather than a distinct protuberance anterodorsal to the supraacetabulum crest (PKUP V0402-V0404, and V0407), for the attachment of m. gluteus magnus (Duellman and Trueb, 1986). In living frogs, the dorsal tubercle is either absent or low, but the lack of a distinct dorsal tubercle is a plesiomorphic condition for Anura, as evidenced by the condition in the Jurassic stem anurans Prosalirus and Notobatrachus (Shubin and Jenkins, 1995;Baez and Nicoli, 2008). The supra¬ acetabulum crest is present, but not noticeably high. ...
... The Triassic proanuran Triadobatrachus has as many as 14 presacral vertebrae (Rage and Rocek, 1989), whereas the number of vertebrae is unknown for the closely related Czatkobatrachus from the Triassic of Poland (Evans and Borsuk-Bialynicka, 1998). One Jurassic stem anuran, Vieraella, has 10 presacrals, and Notobatrachus has nine (Baez and Basso, 1996;Baez and Nicoli, 2008). The Early Jurassic Prosalirus from North America is based on a composite of isolated elements from different individuals; thus, the actual number of presacral vertebrae cannot be determined (Shubin and Jenkins, 1995). ...
... The modern frog pelvis appeared first in the Jurassic Prosalirus and Notobatrachus (Shubin and Jenkins, 1995;Baez and Nicoli, 2008), characterized by a nonexpanded sacral diapophysis directed posterolaterally, smooth dorsal surface of ilium and urostyle without a ridge, and a bicondylar sacral-urostyle articulation. This set of characters is also seen in the most primitive living frogs Ascaphus + Leiopelma, all Early Cretaceous fossil discoglossids and pipoids from Spain (Baez and Sanchiz, 2007;Baez, 2013), and the Early Cretaceous fossil neobatrachians from Brazil (Baez et al., 2009). ...
... The slightly later stem anuran Czatkobatrachus was smaller (about 5 cm) but no hind limb elements are so far known (Evans andBorsuk-Bialynicka 1998, Roček andRage 2000). However, the exquisitely preserved Jurassic Notobatrachus degiustoi REIG, 1956(Baez and Nicoli 2004, 2008 was much larger than Prosalirus and Czatkobatrachus suggesting that the basal-most anurans may have rapidly evolved larger body sizes. Overall, the available fossil data suggest a relative decrease in body size but increase in relative limb length from Triadobatrachus to more crown-ward stem-anurans like Prosalirus (see Evans and Borsuk-Bialynicka 1998) followed by a subsequent evolution of taxa with larger body size, such as Notobatrachus REIG, 1956. ...
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Frogs are the most species-rich and ecologically diverse group of amphibians and are characterized by a unique body plan including long legs, elongated ilia, and fused caudal vertebrae. Stem anurans such as Triadobatrachus or Czatkobatrachus have been suggested to have used jumping or hopping as part of their locomotor repertoire based on their anatomy. The earliest known true frog, Prosalirus bitis was suggested to have been a proficient jumper. However, data on jumping performance in frogs have never been used to attempt reconstruction of ancestral features at the base of the radiation. Here we provide data on jumping performance (forces and acceleration) in 20 species of extant frogs including representatives of most of the early radiating clades. We next use ancestral character value inferences to assess ancestral features. Our analyses suggest that frog ancestors were of small to medium size, had relatively short limbs, produced rather low jump forces, yet were capable of relatively high acceleration. Given the short limbs and low forces, the unique frog bauplan with a reduced vertebral column and a mobile ilio-sacral joint may not have been an adaptation for powerful jumping.
... The slightly later stem anuran Czatkobatrachus was smaller (about 5 cm) but no hind limb elements are so far known (Evans andBorsuk-Bialynicka 1998, Roček andRage 2000). However, the exquisitely preserved Jurassic Notobatrachus degiustoi REIG, 1956(Baez and Nicoli 2004, 2008 was much larger than Prosalirus and Czatkobatrachus suggesting that the basal-most anurans may have rapidly evolved larger body sizes. Overall, the available fossil data suggest a relative decrease in body size but increase in relative limb length from Triadobatrachus to more crown-ward stem-anurans like Prosalirus (see Evans and Borsuk-Bialynicka 1998) followed by a subsequent evolution of taxa with larger body size, such as Notobatrachus REIG, 1956. ...
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Full-text available
Frogs are the most species rich and ecologically diverse group of amphibians and are characterized by a unique body plan including long legs, elongated ilia, and fused caudal vertebrae. Stem anurans such as Triadobatrachus or Czatkobatrachus have been suggested to use jumping or hopping as part of their locomotor repertoire based on their anatomy. The earliest known true frog, Prosalirus bitis has been suggested to be a proficient jumper. Yet, data on actual jumping ability in frogs have never been used to attempt reconstructions of ancestral features at the base of the radiation. Here we provide data on jumping performance (forces and accelerations) in 20 species of extant frogs including representatives of most basal clades. We next use ancestral character state reconstructions to infer ancestral features. Our analyses suggest that frog ancestors were of small to medium size, had relatively short limbs, produced low jump forces, yet were capable of relatively high accelerations. This suggests that the unique frog bauplan with a reduced vertebral column and a mobile ilio-sacral joint may also have evolved in the context of other locomotor behaviors rather than being an adaptation for long distance jumping.
... Eodiscoglossus oxoniensis from the Middle Jurassic (Bathonian) of England (Evans et al., 1990) is the oldest crowngroup frog in the fossil record. Early Jurassic and other Middle Jurassic frogs (known from the Americas) are considered to be outside the crown-group clade (B aez and Nicoli, 2008;Dong et al., 2013). If our assignment of the atlantal centrum from Berezovsk Quarry to Eodiscoglossus is correct, then this find extends the known geographical range of the oldest crown frogs in the Middle Jurassic and demonstrates their wide Laurasian distribution at this time. ...
Article
During the last 15 years our knowledge of Jurassic vertebrates from Siberia has greatly increased through the discovery by S.A. Krasnolutskii in 2000 of the Berezovsk Quarry vertebrate locality in the Middle Jurassic (Bathonian) Itat Formation, in the southern part of Krasnoyarsk Territory, western Siberia (Alifanov et al., 2001). The vertebrate assemblage of Berezovsk Quarry is one of the most taxonomically diverse Jurassic vertebrate assemblages in Asia and includes hybodont sharks, palaeonisciform and amiiform actinopterygians, dipnoans, stem- and crown-group salamanders, xinjiangchelid turtles, basal lepidosauromorphs, scincomorph lizards, choristoderes, goniopholidid crocodyliforms, ornithischian and saurischian dinosaurs, pterosaurs, tritylodontids, and diverse mammaliaforms (e.g., Alifanov et al., 2001; Averianov et al., 2005, 2010, 2014; Kuzmin et al., 2013; Skutschas, 2013; Martin et al., 2014). The composition of this vertebrate assemblage is generally similar to that of the contemporary Forest Marble Formation (Great Britain), especially in the mammalian and salamander components (Evans and Milner, 1994; Kielan-Jaworowska et al., 2004; Benton et al., 2005; Averianov et al., 2010; Skutschas, 2013). Here we describe an atlantal centrum of a frog (Anura) from Berezovsk Quarry of the Bathonian Itat Formation of Western Siberia. This find represents a new faunal component from that locality and is the first unambiguous record for Anura in the Jurassic of Asia.
... Most of the Jurassic eusauropod remains discovered in Argentina come from the Cañadón Asfalto Formation [27], which crops out in north-central Chubut Province, Argentinean Patagonia, and has recently been dated as ranging from the latest Early to the early Middle Jurassic (Toarcian to Aalenian-Bajocian; [28]). Vertebrate fossils are very abundant in this unit and include anurans [29], lepidosaurs [30], turtles [31,32], mammals [33][34][35][36][37], pterosaurs [38,39], and ornithischian [40][41][42], theropod (e.g. [19,20,43,44]), and eusauropod dinosaurs [19,21,25], contributing to a relatively complete Middle Jurassic ecosystem [45]. ...
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The early Middle Jurassic is regarded as the period when sauropods diversified and became major components of the terrestrial ecosystems. Not many sites yield sauropod material of this time; however, both cranial and postcranial material of eusauropods have been found in the Cañadón Asfalto Formation (latest Early Jurassic–early Middle Jurassic) in Central Patagonia (Argentina), which may help to shed light on the early evolution of eusauropods. These eusauropod remains include teeth associated with cranial and mandibular material as well as isolated teeth found at different localities. In this study, an assemblage of sauropod teeth from the Cañadón Asfalto Formation found in four different localities in the area of Cerro Condor (Chubut, Argentina) is used as a mean of assessing sauropod species diversity at these sites. By using dental enamel wrinkling, primarily based on the shape and orientation of grooves and crests of this wrinkling, we define and describe three different morphotypes. With the exception of one taxon, for which no cranial material is currently known, these morphotypes match the local eusauropod diversity as assessed based on postcranial material. Morphotype I is tentatively assigned to Patagosaurus, whereas morphotypes II and III correspond to new taxa, which are also distinguished by associated postcranial material. This study thus shows that enamel wrinkling can be used as a tool in assessing sauropod diversity.
... The Cañadón Asfalto Formation of Chubut Province, Argentina, has yielded the most diverse and important Middle Jurassic terrestrial biota of Gondwana (Escapa et al. 2008). Vertebrate groups reported so far include basal members of every major lineage of terrestrial vertebrates to be expected in the Jurassic (Escapa et al. 2008), including a wealth of dinosaur remains (Bonaparte 1979Bonaparte , 1986 Rauhut 2003a Rauhut , 2005 Escapa et al. 2008), pterosaurs (Rauhut et al. 2001; Cordoniú et al. 2010), turtles (Sterli 2008), mammals (Martin and Rauhut, 2005; Rauhut et al. 2002; Rougier et al. 2007a, b) and anurans (Báez and Nicoli 2008). Until recently, the dinosaur fauna was exclusively composed of sauropod and theropod saurischians, with only a single reported non-diagnostic specimen of an ornithischian (Rauhut and Lopez-Arbarello 2008). ...
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Heterodontosauridae is a morphologically divergent group of dinosaurs that has recently been interpreted as one of the most basal clades of Ornithischia. Heterodontosaurid remains were previously known from the Early Jurassic of southern Africa, but recent discoveries and studies have significantly increased the geographical and temporal range for this clade. Here, we report a new ornithischian dinosaur from the Middle Jurassic Cañadón Asfalto Formation in central Patagonia, Argentina. This new taxon, Manidens condorensis gen. et sp. nov., includes well-preserved craniomandibular and postcranial remains and represents the only diagnostic ornithischian specimen yet discovered in the Jurassic of South America so far. Derived features of its anatomy indicate that Manidens belongs to Heterodontosauridae, as the sister taxon of Heterodontosaurus and other South African heterodontosaurids. The presence of posterior dentary teeth with high crowns but lacking extensive wear facets in Manidens suggests that this form represents an intermediate stage in the development of the remarkable adaptations to herbivory described for Heterodontosaurus. The dentition of Manidens condorensis also has autapomorphies, such as asymmetrically arranged denticles in posterior teeth and a mesially projected denticle in the posteriormost teeth. At an estimated total length of 60-75 cm, Manidens furthermore confirms the small size of basal heterodontosaurids.
... The results of these projects have provided not only new dinosaur taxa but also new taxa from other taxonomic groups, including amphibians, mammals, turtles, and pterosaurs. Amphibians: Frog remains recently collected from the Zitarroza site in the Cañadón Bagual locality have been interpreted as a new species of Notobatrachus, N. reigi Báez and Nicoli (2008). The holotype is represented by a single specimen preserved as partially articulated cranial (mandible, suspensorium and maxillary arch remains) and postcranial impressions. ...
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The plant and tetrapod fossil record of the Cañadón Asfalto Formation (Middle to Late Jurassic) found in Cerro Cóndor area (Chubut Province) is summarized here. The flora is dominated by conifers (Araucariaceae, Cupressaceae sensu lato) but also includes ferns and equisetaleans. The tetrapod fauna is composed of dinosaur taxa described in the 70's as well as other remains recently described and other vertebrate groups such as amphibians, turtles, and mammals. The amphibian remains have been interpreted as representatives of a new species of Notobatrachus, considered one of the most basal members of the anuran lineage. Similarly, turtle remains have been recently recognized as a new species of basal turtle, bringing valuable information about the early evolution of this group. The dinosaur remains are largely dominated by saurischian taxa, represented by basal forms of Eusauropoda and Tetanurae. In addition, three different mammalian species have been identified and considered as early representatives of an endemic Gondwanan mammalian fauna. The fossil record of this formation represents the most completely known biota from the continental Middle to Late Jurassic of the Southern Hemisphere and one of the most complete of the entire world.