Autopodial skeleton evolution in side-necked turtles (Pleurodira)

Carpal and tarsal anatomy was documented based on the observation of dry skeletons of adult specimens representing 25 species in 15 genera and on data taken from the literature. In addition, histological sections and cleared and double‐stained autopodia of recently hatched and juvenile specimens representing seven chelid and pelomedusoid species were studied. There is much more morphological diversity in the manus than in the pes. Variation in autopodial skeletons includes: the astragalus and calcaneum are either separated or fused; fusion of distal carpals 3–4−5 or just 4–5; number of centralia in the carpus; and presence/absence of a pisiform and of an accessory radial element. The widespread and probably basal phalangeal formula for Pleurodira is 2.3.3.3.3. Deviations are Pelomedusa subrufa, exhibiting a reduction to 2.2.2.2.2, Pelusios spp. with one phalanx less in digit I and for one species in digit V as well, and Acanthochelys pallidipectoris with an additional phalanx in the fourth finger. Six discrete characters itemizing some of the morphological variation observed were plotted on a composite pleurodire phylogeny, revealing not only homoplastic patterns but also the utility of some characters in supporting the monophyly of several clades. The pisiform is the last carpal element to ossify in Chelus fimbriatus. We hypothesize that the so‐called fifth hooked metatarsal represents the fusion of distal tarsal 5 with metatarsal V. The accessory radial element that was occasionally present in the turtles examined may represent an atavism of the otherwise lost radiale of turtles.     

The Late Permian herbivore Suminia and the early evolution of arboreality in terrestrial vertebrate ecosystems

Vertebrates have repeatedly filled and partitioned the terrestrial ecosystem, and have been able to occupy new, previously unexplored habitats throughout their history on land. The arboreal ecospace is particularly important in vertebrate evolution because it provides new food resources and protection from large ground-dwelling predators. We investigated the skeletal anatomy of the Late Permian (approx. 260 Ma) herbivorous synapsid Suminia getmanovi and performed a morphometric analysis of the phalangeal proportions of a great variety of extant and extinct terrestrial and arboreal tetrapods to discern locomotor function and habitat preference in fossil taxa, with special reference to Suminia. The postcranial anatomy of Suminia provides the earliest skeletal evidence for prehensile abilities and arboreality in vertebrates, as indicated by its elongate limbs, intrinsic phalangeal proportions, a divergent first digit and potentially prehensile tail. The morphometric analysis further suggests a differentiation between grasping and clinging morphotypes among arboreal vertebrates, the former displaying elongated proximal phalanges and the latter showing an elongation of the penultimate phalanges. The fossil assemblage that includes Suminia demonstrates that arboreality and resource partitioning occurred shortly after the initial establishment of the modern type of terrestrial vertebrate ecosystems, with a large number of primary consumers and few top predators.     

The anatomy of the basal ornithischian dinosaur Eocursor parvus from the lower Elliot Formation (Late Triassic) of South Africa

Ornithischia is a morphologically and taxonomically diverse clade of dinosaurs that originated during the Late Triassic and were the dominant large‐bodied herbivores in many Cretaceous ecosystems. The early evolution of ornithischian dinosaurs is poorly understood, as a result in part of a paucity of fossil specimens, particularly during the Triassic. The most complete Triassic ornithischian dinosaur yet discovered is Eocursor parvus from the lower Elliot Formation (Late Triassic: Norian–Rhaetian) of Free State, South Africa, represented by a partial skull and relatively complete postcranial skeleton. Here, the anatomy of Eocursor is described in detail for the first time, and detailed comparisons are provided to other basal ornithischian taxa. Eocursor is a small‐bodied taxon (approximately 1 m in length) that possesses a plesiomorphic dentition consisting of unworn leaf‐shaped crowns, a proportionally large manus with similarities to heterodontosaurids, a pelvis that contains an intriguing mix of plesiomorphic and derived character states, and elongate distal hindlimbs suggesting well‐developed cursorial ability. The ontogenetic status of the holotype material is uncertain. Eocursor may represent the sister taxon to Genasauria, the clade that includes most of ornithischian diversity, although this phylogenetic position is partially dependent upon the uncertain phylogenetic position of the enigmatic and controversial clade Heterodontosauridae. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 648–684.     

Pterosaur Stance and Gait and the Interpretation of Trackways

The tracks ascribed to pterosaurs from the Late Jurassic limestones at Crayssac, France, must be pterosaurian because the manus prints are so far outside those of the pes, the pes print is four times longer than wide, and the manus prints appear to preserve distinct traces of a posteromedially directed wing-finger. These tracks are different in important ways from previously described Pteraichnus trackways, which have been variably considered pterosaurian, crocodilian, or indeterminate. No Pteraichnus (sensu stricto: those not from Crayssac) tracks have diagnostic features of pterosaurs and in none can a complete phalangeal or digital formula be reconstructed; however, all published Pteraichnidae tracks fulfill the criteria of poor preservation, and some have some diagnostic features of crocodile tracks. Reconstructions of pterosaurs walking in pteraichnid tracks do not fit those tracks well, but crocodiles do. In contrast, the Crayssac tracks demonstrate the erect stance and parasagittal gait previously reconstructed for pterosaurs. They also demonstrate that the footfall pattern was not as in typical reptiles (LH-RF-RH-LF), but that the manus must have been raised before the next forward step of the ipselateral foot (LH-LF-RH-RF), suggesting that the quadrupedal pattern was secondary. The metatarsus in pterosaurs was set low at the beginning of a stride, as it is in crocodilians and basal dinosaurs. The diagnosis of the Ichnofamily Pteraichnidae comprises features of possible crocodilian trackmakers, but not of possible pterosaurian trackmakers. Trackways considered for attribution to pterosaurs should show (1) manus prints up to three interpedal widths from midline of body, and always lateral to pes prints, (2) pes prints four times longer than wide at the metatarso-phalangeal joint, and (3) penultimate phalanges longest among those of the pes.     

Pterosaur tracks and the terrestrial ability of pterosaurs

Unusual tracks of a quadrupedal animal with a three-digit (occasionally four-digit) manus print and four-digit pes print were first interpreted as those of pterosaurs in the 1950s. In the 1980s these tracks were reinterpreted as crocodilian, but new material shows that the original identification was correct. Two features: evidence for elongate penultimate phalanges in digits two to four of the pes, and manus trackways up to three times the width of pes trackways, can only be attributed to pterosaurs. Recent improvements in understanding of pterosaur anatomy and functional morphology explain remaining difficulties regarding the interpretation of ich-nites such as the orientation of the manus digits and the absence of some expected ichnological features. Pteraichnus and Pteraichnus-like tracks show that, when grounded, some, perhaps all, pterosaurs were plantigrade, quadrupedal, and had a semi-erect stance and gait. This is consistent with some functional interpretations of pterosaur anatomy and resolves a long-running debate regarding the terrestrial ability of this group.     

Anatomy and phylogenetic value of the mandibular and coronoid canals and their associated foramina in proboscideans (Mammalia)

Characters associated with the mandibular canal are differently distributed amongst proboscidean lineages and provide useful information on the systematics and relationships of proboscideans. The aim of this paper is to describe the pattern of the mandibular canal and its associated foramina in proboscideans in order to fully appreciate the extent of interspecific variation of these structures within the group and to discuss its systematic and phylogenetic value. Outgroup comparison indicates that the condition presented by the basal proboscidean Phosphatherium is morphotypic for proboscideans. Primitive proboscidean characters are: the low position of the mandibular foramen, and its crescent‐shaped outline, the occurrence of a coronoid foramen (canal), the occurrence of two lateral mental foramina, the posterior one at the level of (or slightly behind) the posterior margin of the symphysis, the anterior one in a more distal position, the absence of a medial mental foramen (MMF), the mandibular canal set just below the tooth row. The occurrence of a single lateral mental foramen may represent a shared derived character of Daouitherium, Numidotherium, and Barytherium. A unique derived feature of the Elephantinae mandible is the occurrence of a medial mental foramen on the medial side of the incisive part of the mandible. MMFs have never been observed in other proboscideans excluding elephantines. The very high frequency of MMFs observed in Mammuthus meridionalis–Mammuthus trogontherii–Mammuthus primigenius (>93 per cent of the studied specimens) could be considered a synapomorphy of this group. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 161 , 391–413.     

Postcranial Functional Morphology of <Emphasis Type="Italic">Hyracotherium</Emphasis> (Equidae,Perissodactyla) and Locomotion in the Earliest Horses

A nearly complete skeleton of Hyracotherium grangeri is described from the early Wasatchian (early Eocene) of the Clarks Fork Basin in northwestern Wyoming. This specimen includes a virtually complete, well-preserved pre-caudal vertebral column allowing the first thorough investigation of the locomotory abilities of these early Eocene horses. The posterior thorax (T) and lumbus (L) are divided into a facultatively dorsostable region (T15-L3) and an obligately dorsostable region (L3-L7). The facultatively dorsostable region is characterized by robust, cranially-oriented neural spines and well-developed attachment sites for epaxial musculature, whereas the obligately dorsostable region is characterized by embracing zygapophyses and less developed spines and attachment sites. Limb morphology exhibits a mixture of primitive characters and derived features associated with cursoriality. As described by previous authors, limb mobility is restricted at the elbow and ankle due to interlocking humero-radial and tibio-astragalar articulations, respectively. In contrast, the presence of a rounded femoral head and well-developed pelvic and femoral attachment sites for hindlimb adductor muscles suggests a highly mobile hip joint. Limited inversion of the manus and pes may have been possible via differential movement between the proximal and distal carpal elements and at the transverse tarsal joint, respectively. The obligately dorsostable region of the vertebral column likely resisted variably-oriented torque applied by hip musculature, thereby preventing dislocation of the pelvis and lumbus. The facultatively dorsostable region would have facilitated dynamic positioning of the center of mass during rapid acceleration. Brief comparisons with Orohippus and Mesohippus postcrania are used to discuss the selective pressures imposed by changes in body mass and habitat on the postcranial evolution of Paleogene horses.     

Evolutionary and developmental aspects of phalangeal formula variation in pig-nose and soft-shelled turtles (Carettochelyidae and Trionychidae)

In order to examine the evolution of the phalangeal formula in a diverse clade of turtles, including hyperphalangy as a rare condition in this group, we studied 210 specimens representing all extant genera of Trionychidae and their sister taxon, Carettochelyidae. Both groups consist of highly aquatic species with elongated autopods that are either paddle-like (Trionychidae) or transformed to flippers (Carettochelyidae). Phalangeal formulae were obtained mostly by radiographs of alcohol-preserved or dry specimens, as well as by direct counts from skeletons. All trionychids and Carettochelys are pentadactylous, but their phalangeal formulae differ. Carettochelys exhibits the turtle-plesiomorphic state (manus and pes: 2-3-3-3-3), with no variation in adults. Trionychids exhibit intraspecific variation, ranging from 2-3-3-3-2 to 2-3-3-6-5 for the manus, and from 2-3-3-3-2 to 2-3-3-5-3 for the pes. The extant Carettochelys as well as the Middle Eocene Allaeochelys crassesculpta are characterized by an elongation of phalanges, whereas trionychids consistently have shorter phalanges. All trionychid genera exhibit some degree of hyperphalangy in digits IV and V, in both the manus and pes. Phalanges of the clawed digits I–III are very robust compared to phalanges of the non-clawed digits IV and V. The latter contribute significantly to the enlargement of the paddle by their additional phalanges. We hypothesize that this phalangeal pattern is coupled with prolongation of growth processes in the non-clawed digits. The differences in autopod morphology between carettochelyids and trionychids reflect different locomotor patterns related to different natural histories (elongated flippers for high-speed escape in the mainly herbivorous Carettochelys; broad paddles for rapid turns during hunting in the mainly carnivorous trionychids). The autopod of Pelodiscus sinensis is proposed as an experimental model to examine the developmental basis of adult autopod variation.     

On Asian ornithopods (Dinosauria: Ornithischia). 4. Probactrosaurus Rozhdestvensky, 1966

The genus Probactrosaurus was first established for material discovered by a joint Russian/Chinese expedition to the Chinese autonomous region of Neimongol (Inner Mongolia). Fossils were collected at a site named Maorty (= Maortu). Material attributable to ornithopod dinosaurs was considered sufficiently distinct to permit the definition of two species of the new genus: Probactrosaurus gobiensis and P. alashanicus. The former species was based on a considerable quantity of skeletal material, much of which is still to be found in the collections of the Palaeontological Institute (PIN), Moscow. The latter was based on far less well‐preserved specimens, including a holotype (a posterior skull roof) that can no longer be found in the collections of the PIN and which, along with other materials, was reportedly returned to the Institute of Vertebrate Palaeontology and Palaeoanthropology, Beijing. Some remnants of the original material attributed to P. alashanicus have been located in the PIN collections. Both taxa, established by A. K. Rozhdestvensky, are re‐described using all of the available material collected during the early Sino‐Soviet expeditions. Probactrosaurus alashanicus is considered to be a junior subjective synonym of P. gobiensis. Further comparisons are made with the recently described species Probactrosaurus mazongshanensis Lu, 1997. The latter does not appear to be referable to the genus Probactrosaurus. Probactrosaurus is a gracile ornithopod (ranging between 4 and 6 m in length). The skull is unadorned by any form of cranial crest; however, the premaxillary beak is deflected ventrally and the dentition is similar to that seen in more derived hadrosaurid ornithopods. The postcranial skeleton is notable for its gracility, in particular the elongate forearm and manus, and the retention of a small, conical pollex spine. Systematic analysis suggests that P. gobiensis is a derived non‐hadrosaurid iguanodontian ornithopod and the basal sister‐taxon to the clade Hadrosauridae. The phylogeny of currently known iguanodontians is reviewed. © 2002 The Linnean Society of London, Zoological Journal of the Linnean Society, 2002, 136 , 113–144.     

At the foot of the shrew: manus morphology distinguishes closely‐related Cryptotis goodwini and Cryptotis griseoventris (Mammalia: Soricidae) in Central America

Small‐eared shrews (Mammalia, Soricidae) of the New World genus Cryptotis are distributed from eastern North America to the northern Andes of South America. One well‐defined clade in this genus is the Central American Cryptotis mexicana group, whose members are set off from other species in the genus by their variably broader fore feet and more elongate and broadened fore claws. Two species in the C. mexicana group, Cryptotis goodwini Jackson and Cryptotis griseoventris Jackson, inhabit highlands in Guatemala and southern Mexico and are presumed to be sister species whose primary distinguishing feature is the larger body size of C. goodwini. To better characterize these species and confirm the identification of recently‐collected specimens, we obtained digital X‐ray images of the manus from large series of dried skins of both species. Measurements of the metacarpals and phalanges successfully separated most specimens of C. goodwini and C. griseoventris. These measurements also show that the fore feet of C. griseoventris from Chiapas, Mexico, are morphologically distinct from those of members of the species inhabiting Guatemala. Univariate, bivariate, and multivariate analyses indicate that fore foot characters are more conservative within species of the C. mexicana group than are cranio‐mandibular characters. Patterns of evolution of fore foot characters that superficially appear to be linear gradations are actually more complex, illustrating individual evolutionary trajectories. No claim to original US government works. Journal compilation © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 118–134.     

Was the Oligo‐Miocene Australian metatherian Yalkaparidon a ‘mammalian woodpecker’?

One of the most striking examples of convergent evolution within mammals is the suite of anatomical specializations shared by the primate Daubentonia of Madagascar and the marsupial Dactylopsila of Australia and New Guinea. Having last shared a common ancestor over 125 million years ago, these two genera have independently evolved extremely similar adaptations for feeding on xylophagous (wood-boring) insect larvae. These include enlarged incisors to gouge holes in wood, cranial modifications to strengthen the skull against the stresses generated by wood gouging and elongate manual digits that are used as probes to extract the larvae. Elsewhere in the world, the same ecological niche is filled by birds (woodpeckers or morphologically convergent forms) that use their beaks for wood gouging. An extinct group of eutherian mammals, the apatemyids, exhibit very similar craniodental and postcranial adaptations to Daubentonia and Dactylopsila and presumably also occupied the woodpecker niche. A qualitative analysis of characters of the skull and dentition of the enigmatic Oligo-Miocene Australian metatherian Yalkaparidon – specifically its combination of very large, open-rooted incisors, zalambdodont molars and features to strengthen the skull against rostral bending – supports the hypothesis that it is probably a fourth 'mammalian woodpecker'. Discovery of the (as yet unknown) manus of Yalkaparidon will test this hypothesis by revealing whether any of its digits are elongate.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 1–17.     

A new specimen of Biseridens qilianicus indicates its phylogenetic position as the most basal anomodont

A new well-preserved basal therapsid skull from the Xidagou Formation, Middle Permian of China, is identified as Biseridens qilianicus. The following synapomorphies distinguish Biseridens as an anomodont and not an eotitanosuchian as previously described: short snout; dorsally elevated zygomatic arch and septomaxilla lacking elongated posterodorsal process between nasal and maxilla. The presence of a differentiated tooth row; denticles on vomer, palatine and pterygoid; contact between tabular and opisthotic; lateral process of transverse flange of pterygoid free of posterior ramus and absence of mandibular foramen exclude it from other anomodonts. Our cladistic analysis indicates Biseridens to be the most basal anomodont, highlights separate Laurasian and Gondwanan basal anomodont clades and suggests that dicynodonts had their origins in the Gondwanan clade. The co-occurrence of the most basal anomodont (Biseridens) together with the most basal therapsid (Raranimus), basal anteosaurid dinocephalians, bolosaurids and dissorophids suggests that the earliest therapsid faunas are from China.     

A Revised Systematic Ichnotaxonomy and Review of the Vertebrate Footprint Ichnofamily Chirotheriidae from the British Triassic

Since 1838, many discoveries of the tetrapod footprint taxon Chirotherium and closely similar forms have been reported from the British Middle Triassic. These have been assigned to dozens of ichnotaxa, so that the identification and interpretation of members of the ichnofamily Chirotheriidae have become confused.

Part 1 of this article deals with ichnotaxonomy. Revision of all the available material shows that there are at least three valid ichnospecies of Chirotherium, one of Synaptichnium, and one of Isochirotherium. The presence of a fourth related ichnogenus, Brachychirotherium, cannot yet be confirmed in Britain. Emended diagnoses are proposed for all four ichnogenera; they may be readily distinguished by the shape of the hand (manus) and foot (pes) impression and especially by the phalangeal counts. The natural cast of a probable tail impression, associated with Chirotheriidae footprints from the British Triassic is figured for the first time, as is a rare detailed impression of skin and the tarsal region of an Isochirotherium lomasi pes cast.

Part 2 provides a critical history of the earliest discoveries of each of these ichnogenera in the British Triassic. It is subdivided into divisions, depending on the quality of data and existence of specimens, the localities in Britain which have yielded these ichnogenera; a detailed history and discussion is provided of the earliest discoveries made at each locality. The stratigraphic distribution and paleological significance of the ichnogenera/species are assessed.     

Derived appendicular morphology in Early Cretaceous aquatic turtles evidenced by the track record

A well‐preserved isolated manus impression evidences the presence of derived aquatic adaptations in the forelimb morphology of Berriasian turtles. Size and the abundant co‐occurring turtle fauna indicate that the track was left by a large‐sized, bottom‐walking basal pancryptodiran. The footprint shows an interesting mosaic of primitive and derived features. The basic turtle pattern is reflected by a short and broad autopodium but thin phalanges, claw reduction and extensive webbing enclosing the digits and lateral and medial margins of the distal metapodium are clearly specialized features related to the aquatic environment. Some characters including the proportional elongation of the first digit, which reached as far distally as the other digits and was probably highly mobile at the metacarpal‐phalangeal joint, indicate an even higher degree of specialization. The specimen gives evidence on a hitherto unknown range of appendicular modification in Early Cretaceous turtles, and redundantly proves the aquatic habitat and locomotion of large‐sized turtles from the Berriasian of northwestern Germany.     

Postcranial morphology of the extinct caviine rodent Microcavia criolloensis (late Pleistocene,South America)

A comparative study of the appendicular skeletal morphology, with a particular emphasis on the autopodial elements (manus and pes), of the extinct caviine rodent Microcavia criolloensis (Late Pleistocene, Uruguay), together with that of living species of Microcavia and some allied caviines is performed. Burrow‐digging and above‐ground behaviour by M. criolloensis could have evolved in the Late Pleistocene, as with its relative M. australis in the Recent. This is suggested based on the morphology of preserved articulated skeletons along with fossil burrow‐like structures. The most remarkable features are: in its forelimb, where the humerus has a structure that would have allowed it to perform similar activities to M. australis, based on humeral width across the epicondiles relative to total humerus length index and a good resistance as indicated by high values relating the diameter of the diaphysis to its total length. Qualitative comparison shows that M. criolloensis had a stout, wide manus with relatively short digits including short, wide phalanges, despite its large size. In its hind limb there is a stout hind‐foot with relatively short and wide metatarsals and phalanges, as compared with those of the recent species, that could arguably be considered a useful tool for shovelling out displaced soil. The generalized morphology suggests above‐ground behaviour together with digging ability. The environmental adaptations of M. criolloensis are also briefly discussed, which seem to differ from those of its extant relatives. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154 , 795–806.     

The systematic importance of anatomical data in Gagea (Liliaceae) from the Flora Iranica area

Anatomical features of basal leaves, pedicels and tepals of 22 species of Gagea belonging to four sections (Platyspermum, Plecostigma, Gagea and Didymobulbos) are investigated. Anatomical characters are mapped onto a molecular phylogenetic tree and their evolution is evaluated. The systematic importance of the anatomical characters is discussed. Anatomical characters are more systematically useful in the Irano‐Turanian taxa than in Euro‐Siberian taxa. The presence of collenchyma and/or sclerenchyma and the pentagonal outline of the transverse section of the basal leaf is found in Irano‐Turanian taxa of section Platyspermum and is mostly absent in Euro‐Siberian taxa. A diagnostic key based on combined anatomical characters is provided. The level of variation in anatomical characters is greater in the basal leaf than the pedicel and lowest in the tepals. Convergent evolution in anatomical characters is associated with ecological shifts between sunny, open, dry habitats and closed, humid habitats. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164 , 155–177.     

The musculature of the papuan frog Phrynomantis stictogaster (Anura,Microhylidae)

The musculature of Phrynomantis stictogaster, a burrowing Papuan microhylid frog, of the subfamily Asterophryinae, is described and compared with accounts of other frogs. P. stictogaster exhibits unusual characters: dense musculature investing an entirely adherent tongue; exceptionally massive jaw musculature; and hitherto underscribed attachments of some muscles in the manus and pes. The presence of an accessory tendon to the M. glutaeus magnus and the pattern of distal thigh tendons confirm previous diagnosis of the Microhylidae, but the presence of an accessory head to M. adductor magnus is a condition previously not noted in the family. Features of the hyoid, pectoral, and thigh muscles resemble those of members of the subfamilies Dyscophinae, Microhylinae, and Spenophryninae.     

A new Hyperodapedon (Archosauromorpha,Rhynchosauria) from the Upper Triassic of India: implications for rhynchosaur phylogeny

A new species of the rhynchosaur genus Hyperodapedon, namely H. tikiensis, is described from well‐preserved skeletal elements that were collected from the Upper Triassic Tiki Formation of India. Hyperodapedon tikiensis is diagnosed on the basis of several cranial and postcranial features including longer than wide basipterygoid process, crest‐shaped maxillary cross section lateral to the main longitudinal groove, deeply excavated neural arches of mid‐dorsal vertebrae, long scapular blade, a pronounced deltopectoral crest, proximal humeral end much broader than distal end, iliac length greater than iliac height, equal pre‐ and postacetabular iliac lengths and circular femoral cross section. Two distinct morphotypes of the maxillary tooth plates can be discerned, which are attributed to ontogenetic variations. A maximum‐parsimony analysis was carried out to show that the order Rhynchosauria is characterized by nine cranial and one postcranial character states. The analysis reveals that Otischalkia elderae is invalid and the basal forms, Howesia and Mesosuchus, are closely related. The Mid‐Triassic genus Ammorhynchus is more derived and forms a sister group to the Late Triassic subfamily Hyperodapedontinae. Isalorhynchus and Teyumbaita are basal to the pandemic genus Hyperodapedon. Twenty‐four characters that are not homoplasious document major patterns of skeletal evolution in rhynchosaurs. From laterally oriented scapula and slender propodials, the postcranial skeleton evolved into a more robust form as is evident from nearly vertical scapula and increase in the robustness of the propodials. Shortening of the femur is noted in the derived Late Triassic forms as exemplified in Hyperodapedon gordoni, Hyperodapedon huxleyi and H. tikiensis.     

Probable crocodilian tracks and traces from the Morrison Formation (Upper Jurassic) of eastern Utah

A tetradactyl pes impression and tridactyl manus impression are described as the type specimen of Hatcherichnus sanjuanensis ichnogen. et ichnosp. nov., a probable large crocodilian ichnite from the Salt Wash Member of the Upper Jurassic Morrison Formation in eastern Utah. A similar pes track from the Morrison Formation at Garden Park, Colorado, may also belong to this ichnogenus. The type specimen from Utah consists of plaster replicas of natural casts of a left pes impression and a left manus impression. Associated with the type specimen were possible tail and body drag impressions. The tracks do not appear to be part of a walking trackway and may be swim tracks associated with an animal in shallow water. The tracks occur at a visible contact between slightly fining‐upward channel sandstone units.