Despite the vast crocodyliform fossil record, little is known about the reproductive biology and ... more Despite the vast crocodyliform fossil record, little is known about the reproductive biology and nesting strategies of the extinct members of the group. Here we report a large accumulation of crocodilian fossil eggs from the type-locality of the baurusuchid Pissarrachampsa sera. Sedimentary facies and architectural elements of the site support a palaeoenvironmental model with a shallow lacustrine, playa-lake system interacting to ephemeral braided fluvial channels, with aeolian influence and development of sandy soils. The presence of pedogenic calcretes in the palaeosols indicates arid to semi-arid conditions. The crocodilian affinity of the eggs is supported by the thin eggshell which bears wedge-shaped shell units with tabular microstructures. Furthermore, taphonomic data support an autochthonous assemblage of eggs and skeletal remains, hinting at a monotypical stratigraphic horizon and suggesting P. sera as the egg-laying taxon. The repeated pattern of four (eventually five) eggs per clutch at the site demonstrates that P. sera laid fewer eggs compared to modern crocodilians, indicating that k-selected reproductive strategy pattern is pervasive in the fossil record of Notosuchia. In the crocodyliform phylogenetic framework, the k-strategy and the " egg clutch sizes " optimization of Notosuchia is opposite to the strategy with larger clutches consistently occurring in modern Crocodylia and Neosuchia, the sister clade to Notosuchia. Yet, the lack of data on more early-branching taxa renders unclear which pattern is plesiomorphic for Crocodyliformes as a whole.
The postcranial anatomy of Crocodyliformes has historically been neglected, as most descriptions ... more The postcranial anatomy of Crocodyliformes has historically been neglected, as most descriptions are based solely on skulls. Yet, the significance of the postcranium in crocodyliforms evolution is reflected in the great lifestyle diversity exhibited by the group, with members ranging from terrestrial animals to semi-aquatic and fully marine forms. Recently, studies have emphasized the importance of the postcranium. Following this trend, here we present a detailed description of the postcranial elements of Pissarrachampsa sera (Mesoeucrocodylia, Baurusuchidae), from the Adamantina Formation (Bauru Group, Late Cretaceous of Brazil). The preserved elements include dorsal vertebrae, partial forelimb, pelvic girdle, and hindlimbs. Comparisons with the postcranial anatomy of baurusuchids and other crocodyliforms, together with body-size and mass estimates, lead to a better understanding of the paleobiology of Pissarrachampsa sera, including its terrestrial lifestyle and its role as a top predator. Furthermore, the complete absence of osteoderms in P. sera, a condition previously known only in marine crocodyliforms, suggests osteoderms very likely played a minor role in locomotion of baurusuchids, unlike other groups of terrestrial crocodyliforms. Finally, a phylogenetic analysis including the newly recognized postcranial features was carried out, and exploratory analyses were performed to investigate the influence of both cranial and postcranial characters in the phylogeny of Crocodyliformes. Our results suggest that crocodyliform relationships are mainly determined by cranial characters. However, this seems to be a consequence of the great number of missing entries in the data set with only postcranial characters and not of the lack of potential (or synapomorphies) for this kind of data to reflect the evolutionary history of Crocodyliformes. How to cite this article Godoy et al. (2016), Postcranial anatomy of Pissarrachampsa sera (Crocodyliformes, Baurusuchidae) from the Late Cretaceous of Brazil: insights on lifestyle and phylogenetic significance. PeerJ 4:e2075
ABSTRACT This article may be used for research, teaching, and private study purposes. Any substan... more ABSTRACT This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae, and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand, or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material.
The rhynchosaur previously referred to as the ''Mariante Rhynchosaur'' is here formally described... more The rhynchosaur previously referred to as the ''Mariante Rhynchosaur'' is here formally described as a new genus and species based on two specimens: a complete skull (without the lower jaw) articulated with the three first cervical vertebrae and a set of right maxilla and dentary. Both specimens were collected at the same site in Rio Grande do Sul, Brazil, from deposits of the Santa Maria Formation considered of Ladinian (Middle Triassic) age. Diagnostic characters include the contact between pre-frontal and postfrontal, a pair of deep frontal grooves, and a very deep skull. A new phylogenetic analysis recovered the new taxon as a member of the Stenaulorhynchinae, a relatively diverse clade of Middle Triassic rhynchosaurs, with records in India, east Africa, and the Americas. Evidence suggests that the extinction of that clade took place in the context of a faunal turnover across the Ladinian-Carnian boundary, when it was replaced by the much more abundant Late Triassic hyperodapedontine rhynchosaurs.
The unique outer ear of crocodylians consists of a large meatal chamber (MC) concealed by a pair ... more The unique outer ear of crocodylians consists of a large meatal chamber (MC) concealed by a pair of muscular earlids that shape a large part of the animal's head. This chamber is limited medially by the enlarged tympanic membrane. Yet, the anatomy of this distinctive and complex region is underexplored and its evolutionary history untraced. The osteology and soft tissues of the MC in extant crocodylians was analysed to describe it and establish osteological correlates within this region. A broad survey of the osteological correlates was conducted in major clades of fossil crocodyliforms to estimate evolutionary trends of the MC. The reorganization of the MC at the origin of crocodyliforms includes characters also present in more basal taxa such as 'sphenosuchians' as well as unique traits of crocodyliforms. Three major patterns are recognized in the MC of basal mesoeucrocodylians. The distinct 'thalattosuchian pattern' indicates that extensive modifications occurred in this clade of aquatic fossil crocodyliforms, even when multiple alternative phylogenetic positions are taken into account. Some traits already established in putative closely related clades are absent or modified in this group. The 'basal notosuchian/sebecian pattern' is widespread among basal metasuchians, and establishes for the first time characters maintained later in neosuchians and extant forms. The 'advanced notosuchian pattern' includes modifications of the MC possibly related to a terrestrial lifestyle and potentially a structure analogous to the mammalian pinna. The main variation in the MC of neosuchians is associated with the homoplastic secondary opening of the cranioquadrate passage. The inferred phylogenetic trends in the crocodyliform MC suggest the great anatomical disparity in this region followed a complex evolutionary pattern, and tympanic hearing played an important role in the origin and diversification of Crocodyliformes.
Pan-Podocnemididae turtles are ubiquitous in Late Cretaceous rocks of the Bauru Group in southeas... more Pan-Podocnemididae turtles are ubiquitous in Late Cretaceous rocks of the Bauru Group in southeastern Brazil. This group of side-necked turtles is particularly abundant in a turtle-bearing site of the Presidente Prudente Formation known as Tartaruguito. Here, we describe the first turtle egg (LPRP-USP 0052) from the Tartaruguito site. LPRP-USP 0052 is nearly complete but misses a pole and measures 5,1 and 2,9–2,2 centimeters due to its flattened minor axis. The egg morphology and microstructure were analyzed by observations performed with CT, Optic Microscopy, Scanning Electronic Microscopy and Wave Dispersion Energy analyses. The eggshell ranges from 145 to160 micrometers thick. Considering the matching morphology of the new specimen and its provenance from the stratigraphic horizon that yielded only the podocnemidids Bauruemys and Roxochelys, it is most likely that LPRP-USP 0052 was produced by a podocnemidid turtle.
In contrast to the rich record of eggs from non-avian dinosaurs, complete eggs attributable to Me... more In contrast to the rich record of eggs from non-avian dinosaurs, complete eggs attributable to Mesozoic birds are relatively scarce. Nevertheless, several well-preserved specimens have been discovered over the last three decades revealing functional and phylogenetic characters that shed light on the breeding strategies of extinct birds. Here we report the first fossil avian egg from Brazil, which was discovered in Upper Cretaceous strata of São Paulo in the southeastern part of the country. The taxonomic identity and structural features of the biomineralized tissues were determined using a combination of Scanning Electron Microscopy, Wave Dispersion Energy analyses and Computed Tomography. These show that the 125.5-μm-thick shell of the 31.4 × 19.5mm egg incorporates three structural layers of similar thickness with both prismatic and aprismatic boundaries. Close similarity between the Brazilian bird egg and those of enantiornithines from the Upper Cretaceous Bajo de la Carpa Formation (RíoColorado Subgroup) of Argentina advocates affinity with basal Ornithothoraces. Furthermore, coherency of their depositional contexts might imply a compatible preference for breeding and nesting environments.
Despite the vast crocodyliform fossil record, little is known about the reproductive biology and ... more Despite the vast crocodyliform fossil record, little is known about the reproductive biology and nesting strategies of the extinct members of the group. Here we report a large accumulation of crocodilian fossil eggs from the type-locality of the baurusuchid Pissarrachampsa sera. Sedimentary facies and architectural elements of the site support a palaeoenvironmental model with a shallow lacustrine, playa-lake system interacting to ephemeral braided fluvial channels, with aeolian influence and development of sandy soils. The presence of pedogenic calcretes in the palaeosols indicates arid to semi-arid conditions. The crocodilian affinity of the eggs is supported by the thin eggshell which bears wedge-shaped shell units with tabular microstructures. Furthermore, taphonomic data support an autochthonous assemblage of eggs and skeletal remains, hinting at a monotypical stratigraphic horizon and suggesting P. sera as the egg-laying taxon. The repeated pattern of four (eventually five) eggs per clutch at the site demonstrates that P. sera laid fewer eggs compared to modern crocodilians, indicating that k-selected reproductive strategy pattern is pervasive in the fossil record of Notosuchia. In the crocodyliform phylogenetic framework, the k-strategy and the " egg clutch sizes " optimization of Notosuchia is opposite to the strategy with larger clutches consistently occurring in modern Crocodylia and Neosuchia, the sister clade to Notosuchia. Yet, the lack of data on more early-branching taxa renders unclear which pattern is plesiomorphic for Crocodyliformes as a whole.
The postcranial anatomy of Crocodyliformes has historically been neglected, as most descriptions ... more The postcranial anatomy of Crocodyliformes has historically been neglected, as most descriptions are based solely on skulls. Yet, the significance of the postcranium in crocodyliforms evolution is reflected in the great lifestyle diversity exhibited by the group, with members ranging from terrestrial animals to semi-aquatic and fully marine forms. Recently, studies have emphasized the importance of the postcranium. Following this trend, here we present a detailed description of the postcranial elements of Pissarrachampsa sera (Mesoeucrocodylia, Baurusuchidae), from the Adamantina Formation (Bauru Group, Late Cretaceous of Brazil). The preserved elements include dorsal vertebrae, partial forelimb, pelvic girdle, and hindlimbs. Comparisons with the postcranial anatomy of baurusuchids and other crocodyliforms, together with body-size and mass estimates, lead to a better understanding of the paleobiology of Pissarrachampsa sera, including its terrestrial lifestyle and its role as a top predator. Furthermore, the complete absence of osteoderms in P. sera, a condition previously known only in marine crocodyliforms, suggests osteoderms very likely played a minor role in locomotion of baurusuchids, unlike other groups of terrestrial crocodyliforms. Finally, a phylogenetic analysis including the newly recognized postcranial features was carried out, and exploratory analyses were performed to investigate the influence of both cranial and postcranial characters in the phylogeny of Crocodyliformes. Our results suggest that crocodyliform relationships are mainly determined by cranial characters. However, this seems to be a consequence of the great number of missing entries in the data set with only postcranial characters and not of the lack of potential (or synapomorphies) for this kind of data to reflect the evolutionary history of Crocodyliformes. How to cite this article Godoy et al. (2016), Postcranial anatomy of Pissarrachampsa sera (Crocodyliformes, Baurusuchidae) from the Late Cretaceous of Brazil: insights on lifestyle and phylogenetic significance. PeerJ 4:e2075
ABSTRACT This article may be used for research, teaching, and private study purposes. Any substan... more ABSTRACT This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae, and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand, or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material.
The rhynchosaur previously referred to as the ''Mariante Rhynchosaur'' is here formally described... more The rhynchosaur previously referred to as the ''Mariante Rhynchosaur'' is here formally described as a new genus and species based on two specimens: a complete skull (without the lower jaw) articulated with the three first cervical vertebrae and a set of right maxilla and dentary. Both specimens were collected at the same site in Rio Grande do Sul, Brazil, from deposits of the Santa Maria Formation considered of Ladinian (Middle Triassic) age. Diagnostic characters include the contact between pre-frontal and postfrontal, a pair of deep frontal grooves, and a very deep skull. A new phylogenetic analysis recovered the new taxon as a member of the Stenaulorhynchinae, a relatively diverse clade of Middle Triassic rhynchosaurs, with records in India, east Africa, and the Americas. Evidence suggests that the extinction of that clade took place in the context of a faunal turnover across the Ladinian-Carnian boundary, when it was replaced by the much more abundant Late Triassic hyperodapedontine rhynchosaurs.
The unique outer ear of crocodylians consists of a large meatal chamber (MC) concealed by a pair ... more The unique outer ear of crocodylians consists of a large meatal chamber (MC) concealed by a pair of muscular earlids that shape a large part of the animal's head. This chamber is limited medially by the enlarged tympanic membrane. Yet, the anatomy of this distinctive and complex region is underexplored and its evolutionary history untraced. The osteology and soft tissues of the MC in extant crocodylians was analysed to describe it and establish osteological correlates within this region. A broad survey of the osteological correlates was conducted in major clades of fossil crocodyliforms to estimate evolutionary trends of the MC. The reorganization of the MC at the origin of crocodyliforms includes characters also present in more basal taxa such as 'sphenosuchians' as well as unique traits of crocodyliforms. Three major patterns are recognized in the MC of basal mesoeucrocodylians. The distinct 'thalattosuchian pattern' indicates that extensive modifications occurred in this clade of aquatic fossil crocodyliforms, even when multiple alternative phylogenetic positions are taken into account. Some traits already established in putative closely related clades are absent or modified in this group. The 'basal notosuchian/sebecian pattern' is widespread among basal metasuchians, and establishes for the first time characters maintained later in neosuchians and extant forms. The 'advanced notosuchian pattern' includes modifications of the MC possibly related to a terrestrial lifestyle and potentially a structure analogous to the mammalian pinna. The main variation in the MC of neosuchians is associated with the homoplastic secondary opening of the cranioquadrate passage. The inferred phylogenetic trends in the crocodyliform MC suggest the great anatomical disparity in this region followed a complex evolutionary pattern, and tympanic hearing played an important role in the origin and diversification of Crocodyliformes.
Pan-Podocnemididae turtles are ubiquitous in Late Cretaceous rocks of the Bauru Group in southeas... more Pan-Podocnemididae turtles are ubiquitous in Late Cretaceous rocks of the Bauru Group in southeastern Brazil. This group of side-necked turtles is particularly abundant in a turtle-bearing site of the Presidente Prudente Formation known as Tartaruguito. Here, we describe the first turtle egg (LPRP-USP 0052) from the Tartaruguito site. LPRP-USP 0052 is nearly complete but misses a pole and measures 5,1 and 2,9–2,2 centimeters due to its flattened minor axis. The egg morphology and microstructure were analyzed by observations performed with CT, Optic Microscopy, Scanning Electronic Microscopy and Wave Dispersion Energy analyses. The eggshell ranges from 145 to160 micrometers thick. Considering the matching morphology of the new specimen and its provenance from the stratigraphic horizon that yielded only the podocnemidids Bauruemys and Roxochelys, it is most likely that LPRP-USP 0052 was produced by a podocnemidid turtle.
In contrast to the rich record of eggs from non-avian dinosaurs, complete eggs attributable to Me... more In contrast to the rich record of eggs from non-avian dinosaurs, complete eggs attributable to Mesozoic birds are relatively scarce. Nevertheless, several well-preserved specimens have been discovered over the last three decades revealing functional and phylogenetic characters that shed light on the breeding strategies of extinct birds. Here we report the first fossil avian egg from Brazil, which was discovered in Upper Cretaceous strata of São Paulo in the southeastern part of the country. The taxonomic identity and structural features of the biomineralized tissues were determined using a combination of Scanning Electron Microscopy, Wave Dispersion Energy analyses and Computed Tomography. These show that the 125.5-μm-thick shell of the 31.4 × 19.5mm egg incorporates three structural layers of similar thickness with both prismatic and aprismatic boundaries. Close similarity between the Brazilian bird egg and those of enantiornithines from the Upper Cretaceous Bajo de la Carpa Formation (RíoColorado Subgroup) of Argentina advocates affinity with basal Ornithothoraces. Furthermore, coherency of their depositional contexts might imply a compatible preference for breeding and nesting environments.
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