Hussam Zaher

A Cretaceous terrestrial snake with robust hindlimbs and a sacrum.

It has commonly been thought that snakes underwent progressive loss of their limbs by gradual diminution of their use. However, recent developmental and palaeontological discoveries suggest a more complex scenario of limb reduction, still poorly documented in the fossil record. Here we report a fossil snake with a sacrum supporting a pelvic girdle and robust, functional legs outside the ribcage. The new fossil, from the Upper Cretaceous period of Patagonia, fills an important gap in the evolutionary progression towards limblessness because other known fossil snakes with developed hindlimbs, the marine Haasiophis, Pachyrhachis and Eupodophis, lack a sacral region. Phylogenetic analysis shows that the new fossil is the most primitive (basal) snake known and that all other limbed fossil snakes are closer to the more advanced macrostomatan snakes, a group including boas, pythons and colubroids. The new fossil retains several features associated with a subterranean or surface dwelling life that are also present in primitive extant snake lineages, supporting the hypothesis of a terrestrial rather than marine origin of snakes.

Molecular phylogeny of the New World Dipsadidae (Serpentes: Colubroidea): a reappraisal

We present a phylogenetic analysis of the New World dipsadids based on an expanded data matrix that includes 246 terminal taxa including 196 dipsadids. The species are sampled for eight genes (12S, 16S, cytb, nd2, nd4, bdnf, c‐mos, rag2). The data are explored using two distinct optimality procedures—maximum parsimony and maximum likelihood—and two alignment strategies—dynamic homology and static homology. Two previously unsampled dipsadid genera, Sordellina and Rhachidelus, are now included in the analysis. The definitions of the genera, Erythrolamprus, Clelia, Hypsirhynchus, Philodryas and Phimophis, and the tribes Alsophiini, Echinantherini and Conophiini, are revised. In order to maintain monophyly, the genus Umbrivaga is synonymized with Erythrolamprus, and two new genera are erected to accommodate Phimophis iglesiasi and Clelia rustica, as well as their closely related species. The West Indian genera Schwartzophis, Darlingtonia, Antillophis and Ocyophis are resurrected.

A Complete Skull of an Early Cretaceous Sauropod and the Evolution of Advanced Titanosaurians

Advanced titanosaurian sauropods, such as nemegtosaurids and saltasaurids, were diverse and one of the most important groups of herbivores in the terrestrial biotas of the Late Cretaceous. However, little is known about their rise and diversification prior to the Late Cretaceous. Furthermore, the evolution of their highly-modified skull anatomy has been largely hindered by the scarcity of well-preserved cranial remains. A new sauropod dinosaur from the Early Cretaceous of Brazil represents the earliest advanced titanosaurian known to date, demonstrating that the initial diversification of advanced titanosaurians was well under way at least 30 million years before their known radiation in the latest Cretaceous. The new taxon also preserves the most complete skull among titanosaurians, further revealing that their low and elongated diplodocid-like skull morphology appeared much earlier than previously thought.

Redescription of the Cranial Morphology of Mariliasuchus Amarali, and Its Phylogenetic Affinities (crocodyliformes, Notosuchia)

Abstract The cranial morphology of Mariliasuchus amarali, a poorly known notosuchian from the Late Cretaceous of Southeastern Brazil, is redescribed based on new material. Its phylogenetic affinities within Crocodylomorpha are evaluated through a parsimony analysis involving 46 taxa and 198 characters. Mariliasuchus is nested well inside the clade Notosuchia, as the sister group of Comahuesuchus, a derived notosuchian from the Late Cretaceous of Argentina. Both taxa share the following unambiguous synapomorphies: ventral half of the lacrimal tapering posteroventrally, not contacting or only slightly contacting the jugal; presence of a large foramen on the lateral surface of the anterior part of the jugal; presence of procumbent premaxillary and anterior dentary alveoli; and ectopterygoids that do not participate of the palatine bar. The presence of procumbent premaxillary teeth, specialized tooth crown morphology, and fore–aft jaw movements suggests that this group presented complex jaw movements related to specialized feeding habits.

THE ANATOMY AND RELATIONSHIPS OF HAASIOPHIS TERRASANCTUS, A FOSSIL SNAKE WITH WELL-DEVELOPED HIND LIMBS FROM THE MID-CRETACEOUS OF THE MIDDLE EAST

Abstract The fossil snake species Haasiophis terrasanctus Tchernov, Rieppel, Zaher, Polcyn, and Jacobs, 2000, from the early Upper Cretaceous of the Middle East, is described and illustrated, following a review of the current debate on snake relationships and origins. The description and discussion presented here adds important detail to the knowledge of this taxon and its phylogenetic significance beyond the limited account presented in the original description of Haasiophis. The species is remarkable in that it shows the skull of a relatively advanced (i.e., macrostomatan) snake, yet preserves well-developed hind limbs. The hind limb includes a femur, tibia, fibula, astragalus, calcaneum, distal tarsal four, and remains of four metatarsals and two phalanges. Haasiophis cannot be considered a juvenile specimen of Pachyrhachis. The implications of the presence of well-developed hind limbs in Haasiophis, Pachyrhachis, and Podophis for the cladistic analysis of the phylogenetic interrelationships of these fossil snakes is discussed. The presence of well-developed hind limbs in Pachyrhachis and Haasiophis also creates methodological problems for the cladistic analysis of the phylogenetic relationships of these fossil snakes. Scenarios of snake origins are reviewed and found to be deficient in the absence of a well-corroborated hypothesis of snake relationships within Squamata.

A New Notosuchian from the Late Cretaceous of Brazil and the Phylogeny of Advanced Notosuchians

A new notosuchian crocodyliform from the Late Cretaceous Bauru Group found in the southeastern State of São Paulo (Brazil) is described here. The new taxon, Caipirasuchus stenognathus, is referred as a new species of the recently erected genus Caipirasuchus within the clade Sphagesauridae based on a phylogenetic analysis of basal mesoeucrocodylians. Caipirasuchus stenognathus is represented by an almost complete skull and lower jaw that has autapomorphic characters that distinguish it from other species of Sphagesauridae. These autapomorphies include: maxilla forming part of the orbital margin (absence of lacrimal-jugal contact), nasal with smooth depressions on the posterior region close to the contact with the maxilla and lacrimal, postorbital with posterior palpebral facet that extends posteriorly underneath the ear-flap groove, and a distinct anterior process of the medial flange of the retroarticular process. Additionally, the new taxon lacks autapomorphic features described in other sphagesaurids. The phylogenetic analysis results in a monophyletic genus Caipirasuchus, that is the sister group of a clade fomed by Sphagesaurus huenei, Caryonosuchus pricei, and Armadillosuchus arrudai. Sphagesaurids also include a basal clade formed by Adamantinasuchus navae and Yacarerani boliviensis. Other notosuchian taxa, such as Mariliasuchus amarali, Labidiosuchus amicum, Notosuchus terrestris, and Morrinhosuchus luziae are successive sister taxa of Sphagesauridae, forming a clade of advanced notosuchians that are restricted to the Late Cretaceous of South America. These results contrast with most previous phylogenetic hypotheses of the group that depicted some members of Sphagesauridae as more closely related to baurusuchids, or found Asian (e.g., Chimaerasuchus) or African (Malawisuchus, Pakasuchus) forms nested within advanced notosuchians that are, according to our analysis, endemic of the Late Cretaceous of South America.

Testing the phylogenetic relationships of the Pleistocene snake Wonambi naracoortensis Smith

Abstract The skeletal material of Wonambi naracoortensis has been re-examined, and used to test its phylogenetic relationships. The skeletal description, and the resulting data matrix, concentrates on characters that can be assessed by observation (on the associated material of Wonambi kept at the South Australian Museum). Macrostomatan affinities of Wonambi prove to be well supported. The taxon nests within Macrostomata, above Xenopeltis and Loxocemus.

A new species of Baurusuchus (Crocodyliformes, Mesoeucrocodylia) from the Upper Cretaceous of Brazil, with the first complete postcranial skeleton described for the family Baurusuchidae

The present work describes a new species of Baurusuchidae from Upper Cretaceous sediments of the Bauru Basin, and provides the first complete postcranial description for the family. Many postcranial features observed in the new species are also present in other notosuchian taxa, and are thus considered plesiomorphic for the genus. These are: long cervical neural spines; robust deltopectoral crest of the humerus; large proximal portion in the radiale that contacts the ulna; ulnare anterior distal projection; supra-acetabular crest well developed laterally; post-acetabular process posterodorsally deflected; presence of an anteromedial crest in the femur; fourth trocanter of femur posteriorly positioned; tibia with a laterally curved shaft; calcaneum tuber posteroventrally oriented; osteoderms ornamented with grooves and imbricated in the tail. On the other hand, we found the following sacral and carpal features to be unique among all mesoeucrocodylians analyzed: transverse processes of sacral vertebrae dorsolaterally deflected; presence of a longitudinal crest in the lateral surface of sacral vertebrae; pisiform carpal with a condyle-like surface. The majority of these cited features corroborates a cursorial locomotion for the new species described in the present study, suggesting that members of the family Baurusuchidae were also cursorial species.

The phylogenetic relationships of Pachyrhachis problematicus, and the evolution of limblessness in snakes (Lepidosauria, Squamata)

We have revisited the type material of Pachyrhachis problematicus, a fossil snake from the Cenomanian (Cretaceous) of the Middle East, and found a number of characters in need of revision with respect to its recent re-description. A new interpretation is given for several cranial characters. There is no identifiable regionalization of the presacral-vertebral column beyond that found in booid snakes. The identification of a free sacral rib in Pachyrhachis is more parsimoniously interpreted as the first lymphapophysis. It is questionable that the pelvic rudiment would have been suspended from the axial skeleton. We have also extensively reviewed the evidence that is believed to link Pachyrhachis, and with it all other snakes, to a mosasauroid ancestor. We were not able to corroborate this hypothesis, which involves characters related to the dentition, braincase morphology, and the intramandibular joint. Instead, re-analysis of the phylogenetic relationships of Pachyrhachis corroborates the hypothesis that it represents the sister-group of macrostomatan snakes, i.e., advanced snakes. Given that Pachyrhachis is nested within the hierarchy of snakes, it is entirely possible that this snake re-developed almost complete hind limbs. A less parsimonious hypothesis would be to consider the hind limbs to have been lost independently in the major basal clades of extant snakes (Scolecophidia, Anilioidea, and Macrostomata). The latter hypothesis is considered a serious possibility given the recently described findings on the mechanisms of limb development in Python, which could account for the highly variable pattern of hindlimb reduction found among basal snakes.

A new species of Mesoclemmys, from the open formations of northeastern Brazil (Chelonii, Chelidae)

A new species of South American side-necked turtle is described from the state of Piaui, Brazil. It is related to the Amazonian species of the genus Mesoclemmys s.l. (new acceptation) and is apparently a biogeographically relictual population of a species formerly living under a more humid climate in northeastern Brazil.