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Goniopholis sp. (Fig. 18) DESCRIPTION In addition to fragmentary cranial remains and numerous isolated teeth, a nearly complete skull and mandibular elements (ANG1 8-5914, 5920, 5921 and 5925) of a single individual have been excavated and prepared in 2018 (Fig. 18). The skull is broken transversally in front of the orbits and the rostrum is split longitudinally along the nasals and premaxillae. The associated mandible preserves both dentaries. In comparison with other goniopholidids from Europe, this skull is notable for its exquisite preservation, being minimally compacted dorsoventrally. Based on recent works (Salisbury et al. 1999; Schwarz 2002; Andrade et al. 2011; Buscalioni et al. 2013; Puértolas-Pascual et al. 2015; Martin et al. 2016a), several diagnostic characters used in combination permit assignment of this specimen to Goniopholididae, including: 1) a mesorostrine condition with a rostrum to skull ratio c. 0.65; 2) constricted outline of the upper jaw at the level of the premaxillary-maxillary suture; 3) double-caniniform maxillary teeth 4+5; 4) confluent dentary alveoli 3+4; 5) orbits smaller than supratemporal fenestrae and slightly smaller than frontal width at interorbital level; 6) skull table and dorsal surface of quadratojugal and jugal densely covered by large circular cupules; 7) internal choanae anteriorly bound by palatines; 8) semi-arched and widely expanded premaxillae with fifth alveolus more laterally placed than the rest of the premaxillary tooth row; and 9) absence of contact between ectopterygoid and posterior margin of maxillary tooth row. The Angeac-Charente specimen (ANG18-5914, 5920, 5921 and 5925) also exhibits several characters that permit provisional assignment to the genus Goniopholis. Among these, antorbital and interorbital ornamentation is informative. The presence of periorbital crests on the prefrontal and lacrimal are present in Goniopholis (Andrade & Hornung 2011) but absent in Anteophthalmosuchus and Hulkepholis (Salisbury & Naish 2011; Buscalioni et al. 2013; Martin et al. 2016a). The interorbital ridge is restricted to the frontal in Hulkepholis (Salisbury & Naish 2011; Buscalioni et al. 2013). In the Angeac-Charente specimen, the presence of both antorbital and interorbital crests is also a character shared with Goniopholis. A short postorbital spine is present in the Angeac-Charente specimen, a condition similar to other specimens of the genus Goniopholis (Salisbury et al. 1999; Schwarz 2002; Andrade et al. 2011) but unlike Anteophthalmosuchus in which this spine is longer, almost contacting the anterior margin of the orbits (Salisbury & Naish 2011; Martin et al. 2016a). The Angeac-Charente specimen is also similar in its maxillary tooth count (20) to Goniopholis kiplingi (Andrade et al. 2011) and with Anteophthalmosuchus, which possesses 19 alveoli (Martin et al. 2016a). The maxillary alveolar count has not been established for Goniopholis simus (Salisbury et al. 1999). However, this count is less than that of the goniopholidid Hulkepholis willetti, in which at least 24 maxillary alveoli are present (Salisbury & Naish 2011). Obvious differences in rostrum proportions are observed with Vectisuchus and Hulkepholis, which possess long and narrow rostra (Buffetaut & Hutt 1980; Salisbury & Naish 2011; Buscalioni et al. 2013). As in Anteophthalmosuchus and Goniopholis, the palpebral in the specimen from Angeac-Charente is small and not large and triangular as in Hulkepholis (Salisbury & Naish 2011; Buscalioni et al. 2013; Martin et al. 2016a). Comparison with Nannosuchus will not be discussed here as this taxon has been viewed either as a juvenile of Goniopholis (Salisbury 2002) or as a valid taxon (Andrade et al. 2011). Therefore, considering the characters mentioned above, we tentatively refer the Angeac-Charente specimens to Goniopholis sp. A detailed description of the Angeac-Charente material will be provided elsewhere. In the meantime, a study of the numerous goniopholid specimens from Cherves-de-Cognac and assigned to Goniopholis (Raslan-Loubatié 2007) will provide a useful basis for discussing morphological variability within a goniopholidid population. Preliminary observations concerning the Angeac-Charente specimen do not reveal any obvious morphological differences between the goniopholidids at both localities but further study is required., Published as part of Allain, Ronan, Vullo, Romain, Rozada, Lee, Anquetin, Jérémy, Bourgeais, Renaud, Goedert, Jean, Lasseron, Maxime, Martin, Jeremy E., Pérez-García, Adán, Fabrègues, Claire Peyre De, Royo-Torres, Rafael, Augier, Dominique & Bailly, Gilles, 2022, Vertebrate paleobiodiversity of the Early Cretaceous (Berriasian) Angeac-Charente Lagerstätte (southwestern France): implications for continental faunal turnover at the J / K boundary, pp. 683-752 in Geodiversitas 44 (25) on pages 704-706, DOI: 10.5252/geodiversitas2022v44a25, http://zenodo.org/record/6902033, {"references":["SALISBURY S. W., WILLIS P. M. A., PEITZ S. & SANDER P. M. 1999. - The crocodilian Goniopholis simus from the lower Cretaceous of north-western Germany. Special Papers in Palaeontology 60: 121 - 148.","SCHWARZ D. 2002. - A New Species of Goniopholis from the Upper Jurassic of Portugal. Palaeontology 45: 185 - 208. https: // doi. org / 10.1111 / 1475 - 4983.00233","ANDRADE M. B. & HORNUNG J. J. 2011. - A new look into the periorbital morphology of Goniopholis (Mesoeucrocodylia: Neosuchia) and related forms. Journal of Vertebrate Paleontology 31 (2): 352 - 368. https: // doi. org / 10.1080 / 02724634.2011.550353","BUSCALIONI A. D., ALCALA L., ESPILEZ E. & MAMPEL L. 2013. - European Goniopholididae from the Early Albian Escucha Formation in Arino (Teruel, Aragon, Spain). Spanish Journal of Palaeontology 28: 103 - 122. https: // doi. org / 10.7203 / sjp. 28.1.17835","MARTIN J. E., DELFINO M. & SMITH T. 2016 a. - Osteology and affinities of Dollo's goniopholidid (Mesoeucrocodylia) from the Early Cretaceous of Bernissart, Belgium. Journal of Vertebrate Paleontology 36 (6): e 1222534. https: // doi. org / 10.1080 / 0272 4634.2016. 1222534","SALISBURY S. W. & NAISH D. 2011. - Crocodilians, in EnglishWealden Fossils. The Palaeontological Association, London: 305 - 369.","BUFFETAUT E. & HUTT S. 1980. - Vectisuchus leptognathus, n. g. n. sp., a slender-snouted goniopholid crocodilian from the Wealden of the Isle of Wight. Neues Jahrbuch fur Geologie und Palaontologie Monatshefte 7: 385 - 390. https: // doi. org / 10.1127 / njgpm / 1980 / 1980 / 385","SALISBURY S. W. 2002. - Crocodilians from the lower Cretaceous (Berriasian) Purbeck Limestone Group of Dorset, Southern England. Special Papers in Palaeontology 68: 121 - 144."]}

Although goniopholidids constitute the most abundant remains of the Lower Cretaceous record of the Iberian Peninsula crocodilians, few specimens have been found complete enough to provide precise taxonomic determinations. The palaeontological site of Mina Santa Maria in Arino (Teruel, Early Albian Escucha Formation) has yielded several neosuchian monotaxic concentrations. The skulls discovered at the Arino Mine show a character combination linking them to the latter European species. The Arino skulls share apomorphies with Anteophthalmosuchus hooleyi, Goniopholis willetti (e.g., loss of the transverse frontal crest and of the prefrontal-lachrymal crest, supratemporal fossa larger than the orbit and supratemporal fenestra subequal, among other derived characters). The new taxa, Hulkepholis plotos gen. n., sp. n. and Anteophthalmosuchus escuchae sp. n., are herein erected, and they constitute the most recent record of their clade in Europe. These sympatric species lived in a coastal marsh system with barrier islands and lagoon

The evolution of thermometabolism in pseudosuchians (Late Triassic to the present) remains a partly unsolved issue: extant taxa (crocodilians) are ectothermic, but the clade was inferred ancestrally endothermic. Here we inferred the thermometabolic regime of two neosuchian groups, Goniopholididae (Early Jurassic to Late Cretaceous) and Dyrosauridae (middle Cretaceous to late Eocene), close relatives of extant crocodilians, in order to elucidate the evolutionary pattern across Metasuchia (Early Jurassic to the present), a clade comprising Neosuchia (Early Jurassic to the present) and Notosuchia (Middle Jurassic until the late Miocene). We propose a new integrative approach combining geochemical analyses to infer body temperature from the stable oxygen isotope composition of tooth phosphate and paleohistology and phylogenetic comparative methods to infer resting metabolic rates and red blood cell dimensions. †Dyrosaurus and †Goniopholis share with extant crocodilians similar lifestyles, body forms, bone tissue organization, body temperatures, metabolic rates, and red blood cell dimensions. Consistently, we infer ectothermy for †Dyrosaurus and †Goniopholis with the parsimonious implication of neosuchians and metasuchians being primitively ectothermic.

BackgroundThe neosuchian crocodyliform genusHulkepholisconstitutes the longirostral lineage of the European Goniopholididae. It comprises two species ranging from the Valanginian of southern England to the lower Albian of the northern Teruel (Spain). A new species ofHulkepholisis described based on a partially complete skull from the lower Barremian Camarillas Formation. We investigate its phylogenetic position and the palatal patterns among members of Goniopholididae and the closely related Thalattosuchia and Tethysuchia.MethodsPhylogenetic relationships were investigated with two matrices using a previously published dataset as the basis: the first differed only by the addition of the new species, the second had newly discovered states for 11 characters, the new species plus several additional specimens ofHulkepholisandAnteophthalmosuchus. Both matrices were processed using TNT v. 1.1, in a heuristic analysis of maximum parsimony, with tree bisection and reconnection 1,000 random addition replicates and saving the 10 most parsimonious trees per replicate, and up to 10 suboptimal trees to calculate Bremer supports. The skull geometry of nine species from Thalattosuchia, Tethysuchia and Goniopholididae was explored to test shape variation between the rostral and postrostral modules, and to visualize the differences on the secondary palate. A set of 18 landmarks was used to delimit significant anatomical features, and the skulls were isotropically scaled using Adobe Illustrator, with the longest skull (Sarcosuchus imperator) as the baseline for comparison.ResultsThe European lineages of goniopholidids are two clades (Nannosuchus+Goniopholis) plus (Hulkepholis+Anteophthalmosuchus). The new species,Hulkepholis rorisp. nov, shares with the latter clade the following apormorphies: a long anterolateral postorbital process, postorbital process almost reaching the anterior jugal ramus, and basioccipital tubera with lateral edges turned posteriorly.Anteophthalmosuchuswas found to be monophyletic, andHulkepholisparaphyletic due to the poor preservation ofH. willetti.Hulkepholis roriis distinguished by having vascular fossae and a mid-protuberance on the ventral surface of the basioccipital, and wide internal fossae in the quadrate. Among Goniopholididae differences on the secondary palate are the presence of a palatal cleft, the narrowness of the secondary choana, and a wide foramen of the median pharyngeal tube.ConclusionsThe new species is the earliestHulkepholisfrom the Iberian Peninsula. New characters have been recognized in the organization of the palate and in the occipital region raising unexpected questions on the evolution of Goniopholididae. The set of palatal characters is discussed as part of a singular palatogenesis in Goniopholididae. The protruding occipital areas suggest that the longirostralHulkepholiswould have had an aquatic lifestyle with particular neck and skull movements.

Goniopholidids and pholidosaurids are common crocodilians in Late Jurassic and Early Cretaceous freshwater environments in many parts of the world, although evidence that goniopholidids ever reached Gondwanan regions needs to be substantiated. In the ‘mid’-Cretaceous, some pholidosaurids (Terminonaris, Oceanosuchus) became adapted to life in the sea, a domain that goniopholidids apparently never occupied. Whereas goniopholidids were mainly mesorostral forms, pholidosaurids were longirostral. There is currently no consensus about the phylogenetic relationships between goniopholidids and pholidosaurids. They have been considered as closely related on the basis of various shared characters (Buffetaut, 1982), but in some recent phylogenetic analyses (e.g., Jouve et al., 2006; Jouve, 2009; Pol et al., 2009) at least some pholidosaurids cluster with other longirostral forms such as dyrosaurids and thalattosuchians, which may suggest that the family Pholidosauridae is not monophyletic. Nevertheless, a number of characters uniting Pholidosauridae have recently been proposed (Hua et al., 2007), but were not tested in a phylogenetic framework. On the other hand, only one unambiguous character diagnoses the clade Goniopholididae in phylogenetic studies: the presence of a depression on the posterior end of the maxilla, described as the ‘depression maxillaire’ (maxillary depression) by Buffetaut (1982; see also Lauprasert et al., 2007). On this basis, Goniopholididae is accepted as a monophyletic entity comprising the generaCalsoyasuchus,Goniopholis,Eutretauranosuchus, Siamosuchus, and Sunosuchus (Tykoski et al., 2002; Lauprasert et al., 2007; Smith et al., 2010), thus agreeing with pre-cladistic views (Buffetaut, 1982). Functionally, the maxillary depression may have housed a sensory organ helping to localize and capture prey in the aquatic environment (Brandalise de Andrade, 2009). Here, we report for the first time the presence of a maxillary depression in Pholidosauridae. Its morphology and position is for the first time described in taxa where the maxillojugal area is preserved. Implications for a close relationship between Goniopholididae and Pholidosauridae are discussed on the basis of this previously overlooked character. Institutional Abbreviations—AMNH, American Museum of Natural History, New York, U.S.A.; BYU, Brigham Young University, Provo, Utah, U.S.A.; CHE, Universite Lyon 1, Lyon, France, specimen from Cherves de Cognac; CMNH, Cleveland Museum of Natural History, Cleveland, Ohio, U.S.A.; DORCM, Dorset County Museum, Dorchester, United Kingdom; MNHN, Museum National d’Histoire Naturelle, Paris, France;MNN, Musee National du Niger, Niamey, Republique du Niger; NHMUK, The Natural History Museum, London, United Kingdom.

A new species, Goniopholis kiplingi sp. nov., based on an exceptionally preserved skull from the Lower Cretaceous of England is described in detail. It shows great similarity with Goniopholis simus and Goniopholis baryglyphaeus, but can be distinguished by the presence of longer lachrymals, smooth (not edged) dorsal surface of the quadrate, and proportionally longer rostrum. A comprehensive phylogenetic analysis of Mesoeucrocodylia (104 taxa; 486 characters) focused on goniopholidids (14 species) places G. kiplingi as sister-group of G. simus, and as part of a monophyletic group also containing G. baryglyphaeus. The relationships of Nannosuchus gracilidens and three undescribed European taxa are explored, and preliminary analyses of Denazinosuchus kirtlandicus (Upper Cretaceous, USA) and ‘Goniopholis’ phuwiangensis (Lower Cretaceous, Thailand) are presented. The assignment of taxa to the genus Goniopholis is discussed. Goniopholis, in its traditional sense, is considered paraphyletic and a restricted updated definition is proposed, with comments on the evolution of other goniopholidids. Morphological characteristics of fragmentary material attributed to Goniopholis are not considered sufficient to secure their generic/specific assignment, and provide no support for the presence of Goniopholis in Gondwanan and/or Upper Cretaceous sedimentary units. Currently Goniopholis is restricted to the Upper Jurassic−Lower Cretaceous of Europe. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163, S66–S108.

The periorbital morphology of goniopholidids is discussed, exploring the diversity of patterns and the relevance of the data for phylogenetic studies. Revision of material is focused on Goniopholis spp. and aff. Goniopholis spp., from England, Germany, and Belgium, providing a comparative description of their interorbital morphology. Traditional interpretation of the interorbital elements in species of Goniopholis (G. simus, G. baryglyphaeus), where the frontal is interpreted as excluded from the orbit by a prefrontal-postorbital contact in the skull roof, is contested and clarified through the analysis of new specimens, including a morphometric analysis. In Goniopholis, failure to identify the palpebral and its subtle contact with the prefrontal has lead to misinterpretation of elements and structures near the orbit, and the differential preservation/loss of palpebrals explains variability of the orbit in shape and orientation. In all European goniophilidids the frontal reaches the primary orbit...

The Crocodyliformes are the most represented vertebrate clade in the Upper Cretaceous sequences of the Bauru Group, Paraná Basin. However, some of the species described have an uncertain taxonomic status and phylogenetic position. For instance, "Goniopholis" paulistanus has been assigned as a nomem dubium, due to its description being based on scarce material. The "G". paulistanus specimens (i.e. teeth and a left tibia) were discovered in two different localities in São Paulo state: Mirandópolis and Valparaíso municipalities; where the upper interval of the Adamantina Formation (Early Maastrichtian of Bauru Group) crops out. Revisiting these specimens, we observed multicrenulated teeth in middle dentary toot- row, a remarkable feature only shared with teleosaurids Machimosaurus hugii (Upper Jurassic of Laurasia) and M. rex (Lower Cretaceous of Tunisia). This apomorphy was also recognized in new material from the Alfredo Marcondes municipality (Presidente Prudente Formation), which are here also referred to "G". paulistanus. We recognized the teeth of "G." paulistanus as the lectotype, however the tibia cannot be assigned to a species as it was not collected in association with the teeth. We performed a phylogenetic analysis with a data matrix composed of 388 characters and 86 taxa, analyzed in TNT. The strict consensus tree recovered Neosuchia and Ziphosuchia (Notosuchia + Sebecia) within Mesoeucrocodylia. The species "G" paulistanus is valid, as a distinct and new genus within Sebecia, in a polytomy with Barreirosuchus, Pepesuchus, Itasuchus and Peirosaurus, forming the clade Itasuchidae. Stolokrosuchus is the sister taxon to Itasuchidae, the sister group of all other Sebecia (Peirosauridae (Mahajangasuchidae + Sebecidae and taxa affinis)). The clades Ziphosuchia, Sebecia and Itasuchidae are here redefined, and we find the last two clades to be more closely related to terrestrial notosuchids than to semiaquatic neosuchians.

The crocodyliform faunas of the lowermost Cretaceous Rabekke and Jydegard Formations on the Baltic island of Bornholm, Denmark, and the Annero Formation of Skane, southernmost Sweden, are represented by isolated teeth, osteoderms, and vertebrae. The rich Berriasian assemblage of the Rabekke Formation includes at least three distinctive taxa: Bernissartia sp., Theriosuchus sp., and Goniopholis sp., an association that is also known from several other contemporaneous European vertebrate localities. In contrast to this fauna, the Jydegard and Annero Formations have yielded only rare mesoeucrocodylian remains, which are assigned to Theriosuchus sp. and an undetermined mesoeucrocodylian taxon, possibly Pholidosaurus. Geographically, the Scandinavian localities represent the easternmost and northernmost distribution of typical continental Jurassic-Cretaceous crocodyliform communities in Europe. (C) 2009 Elsevier Ltd. All rights reserved. (Less)

International audience; New and nearly complete cranial remains of Pholidosaurus purbeckensis are described on the basis of specimens recovered from the Berriasian locality of Cherves-de-Cognac, France. Two skulls, associated mandibles and a set of dorsal osteoderms are available and allow a refinement of the anatomy of the genus, known otherwise from coeval deposits in Germany and England. Because of its longirostrine morphology, convergent with other crocodylomorph lineages, the phylogenetic relationships of Pholidosaurus are likely to be affected by Long Branch Attraction problems. Various tests of removing/excluding longirostrine lineages confirm that Dyrosauridae have a labile position and that their affinities with Pholidosauridae are weakly supported. Results from comparative anatomy and phylogenetic analyses recover Pholidosaurus as the basalmost member of Pholidosauridae, a group closely related to Goniopholididae. Pholidosaurus is recovered together with abundant remains of Goniopholis and with more limited remains of Theriosuchus. The paleoecology of Pholidosaurus and more generally, of Pholidosauridae, is discussed.

Two specimens originally referred by Louis Dollo to Goniopholis simus from the Lower Cretaceous of Bernissart, Belgium, are described. They consist of fully articulated skeletons, one missing the skull and mandible. Comparison of these specimens with recently revised specimens from the Wealden of England allows confirmation that the Belgian specimens are referable to the goniopholidid Anteophthalmosuchus hooleyi. The Belgian specimens are the most completely known representatives not only of this species but also of any Goniopholididae. Study of the postcranial skeletons from Bernissart reveals that the appendicular skeleton closely resembles that of derived neosuchians. The dorsal and ventral shields present a morphology directly comparable to other goniopholidids and pholidosaurids. Such observations stress the necessity to gather an osteological database of postcranial elements to test relationships of the various neosuchian lineages. Goniopholididae were relatively diverse during the Early Cretaceous of Europe, and depending on taxonomic opinion, three to five genera are recognized: Anteophthalmosuchus, Goniopholis, Hulkepholis, and possibly Vectisuchus and Nannosuchus.

A new taxon of a broad-snouted goniopholidid, Siamosuchus phuphokensis, gen. et sp. nov., was discovered in the Sao Khua Formation (ante-Aptian), northeastern Thailand. The specimen is described on the basis of an incomplete skeleton showing the premaxillae, maxillae, nasals, left lacrimal, frontal, left jugal, right postorbital, caudal part of the cranial table, most of the right part of the postcranial skeleton, and several heavily sculptured osteoderms. S. phuphokensis is distinguished from the other broad-snouted goniopholidids on the basis of the following characters: penetration of the nasals in the narial border; presence of a robust and high median ridge on the interorbital surface; lateral margin of the frontal forming the medial edge of the orbit; supraoccipital exhibiting a wide pentagonal shape. A phylogenetic analysis using 101 characters for 29 mesoeucrocodylian taxa and 3 outgroup taxa supports the monophyly of the family Goniopholididae. In addition, the Thai specimen forms a clade with the European Goniopholis. Based on our strict consensus tree, the generic name Goniopholis should be restricted to the European forms only.

Dyrosaurus phosphaticus (Thomas 1893) has been described several times, but the poor preservation and the incomplete preparation of the material left this species inadequately described. Further preparation of a specimen referred to this species now allows a complete description of the braincase and its various cranial nerves. Dyrosaurid taxonomy is mainly based on mandibular characters and the complete description of the skull of D. phosphaticus, and comparison with the other known species of the family enables enhancement of other taxonomic characters. An emended diagnosis is given for D. phosphaticus, the only known species of this genus. With this new description, new characteristics that enable comparisons with other crocodyliforms are available. The presence of a distinct anterolateral postorbital process is observed in dyrosaurs, Elosuchus, Terminonaris robusta, some Goniopholis, and, although smaller, in Sarcosuchus imperator. In the dyrosaurids D. phosphaticus and Rhabdognathus, the postorbital participates largely in the dorsal margin of the infratemporal fenestra, as in Elosuchus, S. imperator, and Thalattosuchia. The laterodorsal position of the lateral eustachian foramen in dyrosaurids is similar to those of Elosuchus and thalattosuchians. These characters are not shared by Stolokrosuchus lapparenti (included in the family Elosuchidae with Elosuchus), suggesting that without clear phylogenetic analysis, the creation of the Elosuchidae was premature. A preliminary phylogenetic analysis based on comparisons of D. phosphaticus with other dyrosaurids shows that Phosphatosaurus gavialoides and Sokotosuchus ianwilsoni are the most primitive dyrosaurids, and Rhabdognathus is more closely related to Hyposaurus than to D. phosphaticus.

The new goniopholidid crocodile Goniopholis baryglyphaeus sp. nov. from the Kimmeridgian of Guimarota/Leiria, Portugal is based on the oldest known relatively complete skeleton of Goniopholis from Europe and consists of a nearly complete skull together with a mandible and various postcranial remains. G. baryglyphaeus shows characters that hitherto have only been described for Goniopholis simus: a caudal expansion of the prefrontal that excludes the frontal from the medial margin of the orbit; and the loss of contact between the paroccipital process and the lateral surface of the squamosal and the quadrate, which exposes the cranioquadrate canal laterally. This new species can be distinguished from other European Goniopholis species by an articular facet placed ventral to the retroarticular process, and directed slightly medioventrally; a caudal margin of the mandible forming almost a right angle at its caudoventral corner; an almost square rostrolateral corner of the cranial table; a straight transverse suture between the parietal and the frontal with a median rostral process; a rostrally tapering wedge-like lacrimal, and a regular heavy skull-sculpturing. The new find of Goniopholis in Portugal shows that this genus was distributed in a larger area, and was present much earlier, than hitherto reported in south-western Europe.

The East African Rift separates the northwestern and southeastern Ethiopian high plateaus, which are capped by massive Cenozoic volcanics overlying thick deposits of marine and nonmarine Mesozoic sediments. During geological mapping projects of the 1920s-1930s, a few Mesozoic vertebrate fossils were found on the southeastern plateau. in contrast, beginning in 1976, and then from 1993 to the present, paleontological field work in the Abay (Blue Nile) River gorge along the eastern edge of the northwestern plateau resulted in the discovery of fossil chondrichthyans (Priohybodus, Hybodus, Rhinobatos), osteichthyans (Lepidotes, cf. Pycnodus), dipnoans (Asiatoceratodus), chelonians (Pelomedusidae, Plesiochelyidae, Pleurosternidae), crocodylians (Goniopholis), dinosaurs, (cf. Acrocanthosaurus Hypsilophodontidae), pollen and other microfossils documenting a coastal biota in part, if not entirely, of latest Jurassic (Tithonian) age. These fossils include new biogeographic records for Africa and document biostratigraphic range extensions. The Ethiopian Mesozoic fauna adds to the growing evidence of limited interchange of vertebrates between Africa and Western Europe during the transition from the Jurassic into the Cretaceous.

A new species of Goniopholis (Order Crocodylia , Suborder Mesosuchia , Family Goniopholididae ) is described as G. phuwiangensis on the basis of a dentary from the Sao Khua Formation at Phu Wiang, in north-eastern Thailand. It is closely related to forms from the Upper Jurassic of North America and Europe and the Lower Cretaceous of Europe, and suggests a Late Jurassic to Early Cretaceous age for the Sao Khua Formation. The occurrence of Goniopholis in Thailand is indicative of Laurasian faunal affinities.

An examination of the crocodilian skull exhibited last session by Mr. Willett, and of an undescribed skull in the British Museum, No. 41,098*, has brought to light a peculiarity in the posterior nares which, so far as I am aware, had been previously recognized only in the small crocodilian skull from Brook, Isle of Wight, in the Jermyn Street Museum, described by Prof. Huxley in a paper communicated to this Society on 28th April, 1875, and figured in the Quart. Journ. Geol. Soc. vol. xxxi. pl. xix. fig. 3. This, and the great probability of the correctness of Mr. Willett9s reference of his acquisition to Goniopholis crassidens , the skull of which was before quite unknown, have seemed to me to invest these skulls with sufficient interest to warrant me in offering a short account of them to the Fellows. I. Mr. Willett9s Crocodilian Skull (Goniopholis crassidens, Owen ?). (P1. XV. figs. 1 & 2.) Viewed from above, the outline of this skull has the form of a slender isosceles triangle, the base of which, taken between the extreme outer posterior angles of the quadrate bones, measures about 13 inches, and the nearly straight sides from the same points to the premaxillary suture 25 inches. The uniform convergence of the sides is broken by a slight inbend in front of the orbits, by a small outbend at the 6th, 7th, and 8th maxillary teeth, reckoned from before backwards, and by the abrupt expansion of the praemaxillae occasioning a deep lateraI notch

(1878). Monograph on the Fossil Reptilia of the Wealden and Purbeck Formations. Supplement No. VIII. Pages 1–15; Plates I—VI. Crocodilia (Goniopholis, Petrosuchus, and Suchosaurus). Issued in the Volume for the year 1878. Monographs of the Palaeontographical Society: Vol. 32, No. 149, pp. 1-15.

I. History of The Descovery of The Specimen. In the late autumn of 1904, at a place locally called ‘Tie Pits,’ near Atherfield Point (Isle of Wight), a large mass of the cliff, comprising many thousand tons of the Wealden Shales, subsided, pushing its foot across the beach, until below low-water line. As the sea washed away the base, the mass continued sinking, and fresh horizons were denuded. In 1905 a series of heavy ‘ground-seas’ cast up blocks of limestone and ironstone, containing crocodilebones, which were discovered on the sand, between high and low water marks. The skull came ashore in six pieces, and on as many different occasions. Scutes and fragments of bones were constantly picked up. One block found by a local fisherman was ibrwarded to the Sedgwick Museum, at Cambridge. After correspondence with Mr. Henry Keeping, the well-known Curator, I was put in communication with Prof. T. McK. Hughes, F.R.S., who, with great liberality and courtesy, wrote that in 9 the interest of scientific progress9 he thought that their block should be handed over to me, and this was accordingly done. The collection of the skull, etc. would have been impossible, but for the aid of Mr. Walter White, the coxswain of the Atherfield lifeboat, who obtained the separate portions by visiting the beach at every tide. The horizon whence the specimen was derived is 80 to 90 feet below the top of the Wealden Shales. II. History of The British GoniopholidAE. The genus.

The description of new material from the Lower Cretaceus of the Galve Syncline (province of Teruel, Spain) belonging to several outcrops, confirms the presence of the following Metamesosuchian crocodiles: Goniopholis, Thenosuchus and Bemissartia . These taxa had been identified based on isolated teeth (Estes & Sanchiz, 1982; Buscalioni & Sanz, 1984) and by means of a whole specimen Bemissartia (Buscalioni et al. , 1984). The postcraneal remains coming from the outcrop called Rocha-Pelejon (RPAH) (unit 3/4, Diaz et al. , 1984; Diaz & Yebenes, in this volume) have been identified as Mesosuchia indel. Proximal and distal epiphyseal rests of stylopods and zeugopods compose the studied material. Morphological comparations between Mesosuchia and Eusuchia are also discused. A few features are proposed to be shared among some Mesosuchia, such as Trematochampsidae and Goniopholididae, that are not present in the Eusuchia. These characters are: lesser inclination, in those Mesosuchia, of the proximal end, lesser development of the proximal femoral end respect to the shaft; the presence of an anterior prominence in the proximal outline of the tibia and the humeral head with a marked steps like outline. The described craneal rest from Galve's outcrops, Cabezo Sta. Barbara (CBP) (unit 5, Diaz et al. , 1984) and Rocha-Pelejon (RPAH), are identified as cf. Goniopbolis sp. and Goniopbolis cf crassidens , respectively. The present apomorphies which are exposed as the presence of the character (considering Tecodonts Proterosuchia+Erythrosuchia as the out-group) are interpreted from the material available: a) hypapophises of the first dorsal centra; b) ventral scutes; e) exoccipitals getting the occipital condyle; d) a dorso-ventral ridge in the basioccipital along the sagital plane; e) basisphenoids hidden in palatal view; f) maxillary wall depth low and g) finally, maxillary festooning. These apomorphic character-states are shared with some Mesosuchia: Goniopholididae, Bemissartidae and Trematochampsidae. A phenetic comparation between the studied material and the Lower Cretaceous British specimens, belonging to Goniopbolis , shows that Galve exemplars are closer to Goniopholidids. 64 isolated vertebral centra found in most oC Galve's outcrops (Pelejon, PH, unit 3/4, Diaz et al. , 1984; Colladico-Blanco, CBH, unit 4; Barranco Poyales-Cerrada Roya, PBCH, unit 511; Rocha-Pelejon, RPAH; Partida-Poyales, PYH, unit 5II; Cerrada Roya Mina, CR, unit 5I y Cabezo Sta. Barbara fit well with the identifications that had been proposed with isolated teeth. Three diferent types of centra are described: procoelous and semi-procoelous which have been atributed to the Metamesosuchia Theriosuchus or/and Bemissartia . Anphicoelous are separated into several morphotypes. None of the features proposed seems to be significative, and perphaps either the appreciated diversity or variability could be supported in ontogenetical and/or individual factors.