GEOMETRIC MORPHOMETRICS OF THE SKULL ROOF OF STEREOSPONDYLS (AMPHIBIA: TEMNOSPONDYLI)

Abstract:  Geometric morphometric analysis using relative warps is applied to the skull roof of 62 species of stereospondyls and their closest outgroups (i.e. basal archegosauriforms) from among temnospondyl amphibians. Twenty-one landmarks and five taxonomic groups are used for comparisons. Their skull evolution is quantified in a morphospace defined by two relative warps axes. The majority of groups show poor concordance between morphological and phylogenetic distances. The only exception is represented by Yates and Warren's study of stereospondyl relationships, in which concordance is high. Only basal archegosauriforms and rhinesuchids show significant overlap in morphospace, although this might be due to low sample sizes. Regression of estimated mean disparity against taxon sample size shows that species within both the trematosauroid and the rhytidostean groups are more widely dispersed in morphospace than species belonging to any of the remaining stereospondyl groups. Stereospondyl skull evolution was characterized by divergence between major clades and convergence within those clades. Changes in patterns of morphospace occupation through time agree with the hypothesis of an 'explosive' radiation in the early Early Triassic, after the extinction of basal archegosauriforms at the end of the Permian.     

A new stereospondyl from the German Middle Triassic, and the origin of the Metoposauridae

Recent finds of well-preserved temnospondyl skeletons from the Lower Keuper (Ladinian, Middle Triassic) in southern Germany are assigned to a new genus and species, Callistomordax kugleri . This taxon is characterized by the following autapomorphies: (1) wide unpaired frontal; (2) vomerine fangs greatly enlarged to occupy entire width of element; (3) intercentra elongated and massive, anterior face being convex; (4) humerus semilunar with enlarged deltopectoral crest; (5) cleithrum strongly curved and bow-shaped; (6) trunk extremely elongated to reach three times the length of the skull. Callistomordax shares with the Metoposauridae the pattern of dermal ornamentation, the proportion of both posterior skull table and snout, the position of the lacrimal, the morphology of the basicranial region, and the structure of the clavicle and interclavicle. Phylogenetic analysis suggests Callistomordax to be the sister taxon of the Metoposauridae, nested within a grade formed by various trematosaurian taxa. In this assemblage, Lyrocephaliscus and a clade formed by Almasaurus , Rileymillerus , Callistomordax , and the Metoposauridae are sister taxa. In all variants of the cladistic analysis, Callistomordax and the Metoposauridae form immediate sister groups. According to the present findings, neither plagiosaurids nor brachyopoids and rhytidosteids are closely related to this 'trematosaurian' monophylum, although these taxa share a range of homoplasies.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 152 , 79–113.     

The skull and the palaeoecological significance of Labidosaurus hamatus, a captorhinid reptile from the Lower Permian of Texas

The cranial skeleton of the large captorhinid reptile Labidosaurus hamatus , known only from the Lower Permian of Texas, is described on the basis of new, undescribed specimens. Labidosaurus is distinguished from other captorhinids by the more extreme sloping of the ventral (alveolar) margin of the premaxilla, a low dorsum sellae of the parabasisphenoid, a reduced prootic, a narrow stapes, and a relatively small foramen intermandibularis medius. Despite the presence of a single row of teeth in each jaw, the skull of Labidosaurus resembles most closely those of moradisaurines, the large multiple-tooth-rowed captorhinids of the latest Early and Middle Permian. A phylogenetic analysis confirms that the single-tooth-rowed L. hamatus is related most closely to moradisaurines within Captorhinidae, a relationship that supports the hypothesis of a diphyletic origin for multiple rows of marginal teeth in captorhinids (in the genus Captorhinus and in the clade Moradisaurinae). In view of the close relationship between L. hamatus and moradisaurines, which are regarded to have been herbivorous, L. hamatus is a critical taxon for studies of the evolution of herbivory in early tetrapods. L. hamatus shares several trademark features of herbivorous adaptation with moradisaurines, which suggest that this captorhinid species was omnivorous. As such, it represents a transitional taxon between faunivorous basal reptiles and the herbivorous moradisaurines.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 149 , 237–262.     

A new gomphodont cynodont (Traversodontidae) from the Middle–Late Triassic Dinodontosaurus Assemblage Zone of the Santa Maria Supersequence,Brazil

The tetrapod faunas from the terrestrial Middle–Late Triassic basins of Africa and South America are among the richest in the world, especially in non‐mammalian cynodonts. Despite the great abundance of cynodont specimens found in these basins, there are few known taxa that exhibit interbasinal distributions. Here we describe a new species of traversodontid cynodont of the genus Scalenodon from the Triassic Dinodontosaurus Assemblage Zone of the Santa Maria Supersequence, from the state of Rio Grande do Sul, Brazil. Scalenodon ribeiroae sp. nov. is based on a partial skull that possesses a combination of features not observed in any other South American traversodontid: ellipsoid upper postcanines with the transverse crest formed by three cusps, lacking a mesiobuccal accessory cusp, and with lingual cusp projected lingually creating a concave lingual surface on the upper postcanines; the paracanine fossa is positioned medially to the upper canine, and jugal lacks a suborbital process. A phylogenetic analysis places the new taxon in a basal position within the Family Traversodontidae, with the African Scalenodon angustifrons as sister‐taxon. The new specimen of Scalenodon represents the first record of this genus outside of the Manda Beds of Tanzania, and reinforces the biostratigraphical and biogeographical connection between Gondwanan Middle–Late Triassic tetrapod faunas. Although recent advances have been made, our current knowledge of these faunas is limited by the lack of absolute dates for most units and by uncertainties in the taxonomy and stratigraphical provenance of key fossils.     

A primitive ornithischian dinosaur from the Late Triassic of South Africa, and the early evolution and diversification of Ornithischia

Although the group played an important role in the evolution of Late Mesozoic terrestrial ecosystems, the early evolutionary history of the ornithischian dinosaurs remains poorly understood. Here, we report on a new primitive ornithischian, Eocursor parvus gen. et sp. nov., from the Late Triassic (?Norian) Lower Elliot Formation of South Africa. Eocursor is known from a single specimen comprising substantial cranial and postcranial material and represents the most complete Triassic member of Ornithischia, providing the earliest evidence for the acquisition of many key ornithischian postcranial characters, including an opisthopubic pelvis. A new phylogenetic analysis positions this taxon near the base of Ornithischia, as the sister taxon to the important and diverse clade Genasauria. The problematic clade Heterodontosauridae is also positioned basal to Genasauria, suggesting that an enlarged grasping manus may represent a plesiomorphic ornithischian condition. This analysis provides additional phylogenetic support for limited ornithischian diversity during the Late Triassic, and suggests that several major ornithischian clades may have originated later than generally believed. There are few morphological differences between Late Triassic and Early Jurassic ornithischians, supporting previous suggestions that the Early Jurassic ornithischian radiation may simply represent the filling of vacant ecological space following Late Triassic terrestrial extinctions.     

A NEW PARASEMIONOTID-LIKE FISH FROM THE LOWER TRIASSIC OF JURONG, JIANGSU PROVINCE, SOUTH CHINA

Abstract:  Based on a well-preserved specimen from the Early Triassic Lower Qinglong Formation exposed at Qingshan Quarry, Jurong, Jiangsu Province of China, the new taxon Peia jurongensis gen. et sp. nov. is named and described with anatomical details of its dermal skull, neurocranium and postcranial skeleton. The new taxon is characterized by the following set of characters: shallow V-shaped, tuber-like rostral; anterior part of frontal widened to roughly same width as its posterior part; roughly squared parietal; supraorbital absent; wide and slightly posteriorly inclined preopercle; five small ossifications present in dorsal half of preopercle; preopercular canal shifting anteriorly in ventral half of the preopercle, and reaching the anteroventral corner of this bone; dermosphenotic joining the skull roof and with a descending lamina; numerous branchiostegal rays. Comparisons are made between the new taxon and several parasemionotids from the same locality; a discussion is carried out on the distribution and evolution of several characters in halecomorphs and other closely related neopterygians.     

A new species of basal rhynchosaur (Diapsida: Archosauromorpha) from the early Middle Triassic of South Africa,and the early evolution of Rhynchosauria

Rhynchosauria was an important clade of herbivorous archosauromorph reptiles during the Triassic, with a worldwide distribution. We describe a new genus and species of early rhynchosaur, E ohyosaurus wolvaardti gen. et sp. nov. , from the early Middle Triassic (early Anisian) Cynognathus Assemblage Zone (Subzone B) of the Karoo Supergroup, South Africa. Eohyosaurus wolvaardti is known from a single skull, and is recovered as the sister taxon of Rhynchosauridae in a new phylogenetic analysis. Cynognathus Subzone B has previously yielded the stratigraphically oldest well‐understood rhynchosaur species, Mesosuchus browni and Howesia browni. Eohyosaurus wolvaardti increases the rhynchosaur diversity within this stratigraphical horizon to three species. Intriguingly, all currently confirmed rhynchosaur occurrences from the Early Triassic to earliest Middle Triassic are from South Africa. This may suggest a relatively restricted palaeogeographical distribution for early rhynchosaurs, followed by a global dispersal of rhynchosaurids during the Middle Triassic. © 2015 The Linnean Society of London     

A NEW SPECIES OF CLEVOSAURUS (LEPIDOSAURIA: RHYNCHOCEPHALIA) FROM THE UPPER TRIASSIC OF RIO GRANDE DO SUL, BRAZIL

Abstract:  Well-preserved cranial remains of a small sphenodontian lepidosaur from the Upper Triassic Caturrita Formation of Rio Grande do Sul, Brazil, are the first record of the genus Clevosaurus Swinton, 1939 from South America. They represent a new species, Clevosaurus brasiliensis , which is distinguished by a very short antorbital region of the skull (corresponding to about 20 per cent of skull length) and the presence of teeth in addition to two longitudinal rows on the pterygoid. C. brasiliensis most closely resembles C. bairdi from the Lower Jurassic of Nova Scotia (Canada) and C. mcgilli from the Lower Jurassic of Yunnan (China). The discovery of Clevosaurus in the Upper Triassic of southern Brazil provides a significant range extension of this widely distributed sphenodontian genus. Along with other recent finds, it also suggests that there may have been less biotic provincialism among terrestrial vertebrates during the Late Triassic than has previously been assumed.     

FIRST RECORD OF A HETERODONT BIVALVE (MOLLUSCA) FROM THE EARLY TRIASSIC: PALAEOECOLOGICAL SIGNIFICANCE AND IMPLICATIONS FOR THE 'LAZARUS PROBLEM'

Abstract:  The heterodont bivalve Sinbadiella pygmaea gen. et sp. nov. is described from the Sinbad Limestone Member (Olenekian, Smithian) of the Lower Triassic Moenkopi Formation of Utah (USA). The new taxon is tentatively assigned to the Lucinidae on the grounds of its external morphology, larval shell morphology and hinge characters. It is the only definite representative of the Superorder Heterodonta in the Olenekian, removing this major clade of bivalves from the long list of Early Triassic Lazarus taxa. Although at least locally very abundant, the new taxon has been overlooked by earlier workers, probably due to its minute size. Dwarfism is a widespread phenomenon in Early Triassic faunas, which has been attributed to ongoing reduction of primary production. If S .  pygmaea was a lucinid, a chemosymbiotic feeding mode can be inferred, which might explain why it was able to thrive in conditions that suppressed filter feeding heterodonts.     

New information about <Emphasis Type="Italic">Irajatherium hernandezi</Emphasis> Martinelli,Bonaparte, Schultz &; Rubert 2005 (Eucynodontia,Tritheledontidae) from the upper triassic (Caturrita Formation,Paraná Basin) of Brazil

In this contribution, new specimens of the tritheledontid eucynodont Irajatherium hernandezi, from the Late Triassic (Caturrita Formation) of southern Brazil, are analyzed. The new material provides significant information about incisor count, canine size and shape, basicranial morphology, and other previously unknown aspects of skull and dentition. A cladistic analysis with inclusion of the new data supports the assignment of Irajatherium to Tritheledontidae, basal to Chalimininae and Pachygenelinae. Previously unknown characters of Irajatherium revealed by the new material include: the presence of three lower incisors; the first lower incisor is enlarged; the presence of large upper and lower canines with deep paracanine fossa on the maxilla; almost complete upper and lower postcanine tooth row with a pattern similar to that of other tritheledontids (e.g. Pachygenelus and Chaliminia); there is a conspicuous crest on the inner surface of the maxilla for the attachment of the inferred maxillary turbinates; partially confluent jugular foramen and fenestra rotunda within the jugular fossa, separated by a finger-like projection of the posterolateral wall of the opisthotic; and hypoglossal foramina located outside the jugular fossa. Irajatherium is a key taxon for understanding the early evolution of ictidosaurs, a group of cynodonts closely related to mammaliaforms, during the cynodont–mammal transition from the Late Triassic to Early Jurassic.     

The early Triassic rhynchosaur Mesosuchus browni and the interrelationships of basal archosauromorph reptiles

Restudy of the unique diapsid reptile Mesosuchus browni Watson, from the Cynognathus Assemblage Zone (late Early Triassic to early Middle Triassic) of the Burgersdorp Formation (Tarkastad Subgroup; Beaufort Group) of South Africa, confirms that it is the most plesiomorphic known member of the Rhynchosauria. A new phylogenetic analysis of basal taxa of Archosauromorpha indicates that Choristodera falls outside of the Sauria, Prolacertiformes is a paraphyletic taxon with Prolacerta sharing a more recent common ancestor with Archosauriformes than with any other clade, Megalancosaurus and Drepanosaurus are sister taxa in the clade Drepanosauridae within Archosauromorpha, and are the sister group to the clade Tanystropheidae composed of Tanystropheus, Macrocnemus, and Langobardisaurus. Combination of the phylogenetic relationships of basal archosauromorphs and their known stratigraphic ranges reveals significant gaps in the fossil records of Late Permian and Triassic diapsids. Extensions of the temporal ranges of several lineages of diapsids into the Late Permian suggests that more groups of terrestrial reptiles survived the end-Permian mass extinction than thought previously.     

A New Genus of Rhynchonellid Brachiopod from the Lower Triassic of South China and Implications for Timing the Recovery of Brachiopoda After the End-Permian Mass Extinction

A new genus, Meishanorhynchia , is proposed based on new material from the Lower Triassic of the Meishan section, South China. It is of a late Griesbachian age based on both associated biozones (ammonoids and bivalves) and radiometric dates of the intercalated volcanic ash beds. Comparison with both Palaeozoic and Mesozoic–Cenozoic-related genera suggests that it may represent the first radiation of progenitor brachiopods in the aftermath of the end-Permian extinction. The lowest brachiopod horizon that contains the genus is estimated to be about 250.1 ± 0.3 Ma. This implies that the initial stage of recovery of Brachiopoda in the Early Triassic was probably about 1.3 ± 0.3 myr after the major pulse of the end-Permian mass extinction (dated as 251.4 ± 0.3 Ma). This is in agreement with Hallam's expectancy that biotic recovery typically begins within one million years or so of major mass extinctions, in contrast to current views on the end-Permian extinction event which propose that the recovery of most if not all biotic groups in the Early Triassic was severely delayed and only began about five million years after the end-Permian extinction.     

A new brittle star (Ophiuroidea: Ophiodermatina) from the Early Triassic Paris Biota (Bear Lake County,Idaho, USA)

Knowledge on the morphology of Early Triassic ophiuroids is very limited, in spite of the relatively high number of described species. This hampers attempts to use morphology-based phylogenetic analyses in order to explore the early diversification of the ophiuroid modern clades between the latest Paleozoic and the early Mesozoic. Here, a new ophiuroid from the Early Triassic of southern Idaho (USA) is described and assigned to the new taxon Shoshonura brayardi Thuy nov. gen., nov. sp. It is part of an exceptional Early Triassic faunal assemblage recently described as the Paris Biota. On account of the arm morphology and the shape of the arm spines and arm spine articulations, Shoshonura brayardi nov. gen., nov. sp. is considered a basal member of the suborder Ophiodermatina.     

A Basal Lithostrotian Titanosaur (Dinosauria: Sauropoda) with a Complete Skull: Implications for the Evolution and Paleobiology of Titanosauria

We describe Sarmientosaurus musacchioi gen. et sp. nov., a titanosaurian sauropod dinosaur from the Upper Cretaceous (Cenomanian—Turonian) Lower Member of the Bajo Barreal Formation of southern Chubut Province in central Patagonia, Argentina. The holotypic and only known specimen consists of an articulated, virtually complete skull and part of the cranial and middle cervical series. Sarmientosaurus exhibits the following distinctive features that we interpret as autapomorphies: (1) maximum diameter of orbit nearly 40% rostrocaudal length of cranium; (2) complex maxilla—lacrimal articulation, in which the lacrimal clasps the ascending ramus of the maxilla; (3) medial edge of caudal sector of maxillary ascending ramus bordering bony nasal aperture with low but distinct ridge; (4) ‘tongue-like’ ventral process of quadratojugal that overlaps quadrate caudally; (5) separate foramina for all three branches of the trigeminal nerve; (6) absence of median venous canal connecting infundibular region to ventral part of brainstem; (7) subvertical premaxillary, procumbent maxillary, and recumbent dentary teeth; (8) cervical vertebrae with ‘strut-like’ centroprezygapophyseal laminae; (9) extremely elongate and slender ossified tendon positioned ventrolateral to cervical vertebrae and ribs. The cranial endocast of Sarmientosaurus preserves some of the most complete information obtained to date regarding the brain and sensory systems of sauropods. Phylogenetic analysis recovers the new taxon as a basal member of Lithostrotia, as the most plesiomorphic titanosaurian to be preserved with a complete skull. Sarmientosaurus provides a wealth of new cranial evidence that reaffirms the close relationship of titanosaurs to Brachiosauridae. Moreover, the presence of the relatively derived lithostrotian Tapuiasaurus in Aptian deposits indicates that the new Patagonian genus represents a ‘ghost lineage’ with a comparatively plesiomorphic craniodental form, the evolutionary history of which is missing for at least 13 million years of the Cretaceous. The skull anatomy of Sarmientosaurus suggests that multiple titanosaurian species with dissimilar cranial structures coexisted in the early Late Cretaceous of southern South America. Furthermore, the new taxon possesses a number of distinctive morphologies—such as the ossified cervical tendon, extremely pneumatized cervical vertebrae, and a habitually downward-facing snout—that have rarely, if ever, been documented in other titanosaurs, thus broadening our understanding of the anatomical diversity of this remarkable sauropod clade. The latter two features were convergently acquired by at least one penecontemporaneous diplodocoid, and may represent mutual specializations for consuming low-growing vegetation.     

A NEW TEMNOSPONDYL FROM THE PERMO-TRIASSIC BUENA VISTA FORMATION OF URUGUAY

Abstract:  A partial skull recovered from conglomerates of the Permo-Triassic Buena Vista Formation in Uruguay belongs to a new species, Uruyiella liminea gen. et sp. nov. This species is characterized by a broadly triangular skull with laterally projecting posterior corners, rhytidosteid-like dermal sculpturing, and orbits positioned close to the skull margin. Uruyiella liminea is distinguished from other temnospondyls by a combination of primitive and derived character states, such as the anterior extent of the palatine ramus of the pterygoid, which excludes the ectopterygoid and most of the palatine from the lateral border of the interpterygoid vacuity, and the absence of both tabular horns and otic notches. A phylogenetic analysis places Uruyiella and the enigmatic Early Triassic Laidleria in a clade to which we attach the family name Laidleriidae. The Plagiosauridae and the Laidleriidae form a clade at the base of Dvinosauria, which is the sister group of a clade that includes Stereospondyli and Archegosauroidea. This result is unexpected because Laidleria and Plagiosauridae are nested deeply within Stereospondyli in most phylogenies. The sister-group relationship of Uruyiella and Laidleria suggests that a ghost lineage for the latter genus extends down into the earliest Triassic and perhaps even into the Late Permian, which in turn would suggest survivorship of the Laidleriidae through the Permo-Triassic extinction event.     

Arizonasaurus and its implications for archosaur divergence

The Moenkopi Formation has yielded partial and isolated remains of important archosaurs including rauisuchian skull fragments and isolated poposaur centra and pelvic girdle elements. A recently discovered skeleton referable to Arizonasaurus babbitti shows that most of these archosaurian remains belong to one taxon. Characteristics of the skeleton of Arizonasaurus show that it belongs to a poorly known group of Middle Triassic (240-230 Myr ago) archosaurs called the ctenosauriscids, and that ctenosauriscids are or are closely related to poposaurs. Furthermore, many characteristics of Arizonasaurus provide evidence that poposaurids and ctenosauriscids are derived rauisuchians. The presence of a poposaurid in the early Middle Triassic suggests that the divergence of birds and crocodiles occurred earlier than previously thought. Middle Triassic ctenosauriscids also allow the correlation of Triassic faunas in Europe, Asia, North America and Africa. The Moenkopi Formation fauna represents a transitional fauna between Early and Late Triassic faunas.     

THE TYPE SERIES OF 'SINEMYS'WUERHOENSIS, A PROBLEMATIC TURTLE FROM THE LOWER CRETACEOUS OF CHINA, INCLUDES AT LEAST THREE TAXA

Abstract:  We re-examine the type series of ' Sinemys ' wuerhoensis Yeh (at least 20 specimens, including several shells and skulls on three slabs of matrix and one isolated skull) from the Early Cretaceous Tugulu Group of China. Our study shows that the type series of ' S. ' wuerhoensis is actually a chimera made up of at least three distinct taxa. The holotype of this taxon should be assigned to the basal eucryptodire genus Xinjiangchelys Yeh. As there are no characters that distinguish ' S. ' wuerhoensis from Xinjiangchelys species, we consider it to be a nomen dubium . This new assignment of ' S. ' wuerhoensis expands the temporal range of Xinjiangchelys from the Late Jurassic into the Early Cretaceous in Asia. The majority of the paratypes of ' S. ' wuerhoensis (several shells in dorsal and ventral aspect and skulls) are referred to the basal eucryptodire genus Ordosemys Brinkman and Peng. We establish a new name for these specimens, Ordosemys brinkmania sp. nov. One additional specimen in the type series of ' S. ' wuerhoensis , a skull, is referred to cf. Pantrionychia Joyce, Parham and Gauthier indet.