Abstract Fossil vertebrates from the Cabao Formation discovered in the area of Nalut in northwest... more Abstract Fossil vertebrates from the Cabao Formation discovered in the area of Nalut in northwestern Libya include the hybodont shark Priohybodus, the crocodilian Sarcosuchus, an abelisaurid, a baryonichine spinosaurid and a large sauropod with spatulate teeth. The Cabao Formation may be Hauterivian to Barremian in age, although an earlier Berriasian to Valanginian age cannot be excluded. Its dinosaur assemblage is reminiscent of that of the El Rhaz and Tiouraren formations of Niger and strongly differs from both the Cenomanian ...
The important question of early angiosperm growth habit (i.e., trees, shrubs or herbs?) remains u... more The important question of early angiosperm growth habit (i.e., trees, shrubs or herbs?) remains unanswered. Various theories have been based on data from both living and fossil plants. The Early Cretaceous fossil wood record, however, was seldom used to investigate early angiosperm habit. We set up a database for the Early Cretaceous and Cenomanian of Europe, as this area has the most complete and stratigraphically well-constrained record. The database has 170 entries, based on a bibliographical survey and on the examination of more than 600 new fossil wood specimens from a wide range of palaeoenvironments. In our record the woody characteristic in angiosperms appeared during the Albian, whereas most of the angiosperm's early evolution took place earlier, during the earliest Cretaceous. From the European fossil wood record for the Early Cretaceous and Cenomanian, the global extension and dominance of angiosperms in the Cenomanian is concomitant with a sharp increase in heteroxylous wood diversity. It appears that small stature and weak wood limited the angiosperm ecological radiation for some time. (J.-P. Billon-Bruyat), [email protected] (M. Guiomar), [email protected] (J.-L. Latil), [email protected] (J. Le loeuff), [email protected] (D. Néraudeau), [email protected] (D. Olivero), [email protected] (J. Schlögl).
1 Seccion de Geologia, Facultad de Ciencias, Universidad del Pais Vasco (Euskal Herriko Unibertsi... more 1 Seccion de Geologia, Facultad de Ciencias, Universidad del Pais Vasco (Euskal Herriko Unibertsitatea), Apdo 644, 48080 Bilbao, Euskadi. 2 CNRS, Laboratoire de Paléontologie des Vertébrés, Université Paris VI, 4 place Jussieu, 75252 Paris Cedex 05, France. 3 Unidad ...
169 coprolites were found at Huai Nam Aun in Chaiyaphum province (Huai Hin Lat Formation, Upper
T... more 169 coprolites were found at Huai Nam Aun in Chaiyaphum province (Huai Hin Lat Formation, Upper Triassic) and are sub-divided into seven different morphotypes. Four groups of spiral coprolites are interpreted as being produced by fish-eating sharks and lungfish, whereas non-spiral coprolites containing numerous bone fragments and fish scales were probably produced by other carnivorous vertebrates. Liassocopros hawkinsi and Saurocopros bucklandi are recognized in this study for the first time in Southeast Asia; this discovery supports palynological studies suggesting a Carnian-Norian age for the Huai Hin Lat Formation.
The global Late Maastrichtian non-avian dinosaur apparent biodiversity is extensively surveyed fo... more The global Late Maastrichtian non-avian dinosaur apparent biodiversity is extensively surveyed for the first time. It amounts to 104 species (including unnamed forms) in 2010. The real biodiversity being obscured by taphonomical biases and the scarcity of the continental fossil record, a species-area relationship is used to estimate it. The results show that several hundreds (between 628 and 1078) non-avian dinosaur species were alive in the Late Maastrichtian, which is almost an order of magnitude above previous estimates. Because of the complex Late Cretaceous palaeobiogeography, discussions about dinosaur extinction should be based on this estimated real global biodiversity, not on the apparent biodiversity of a single area. Given the mean duration of dinosaur genera (7.7 Ma), the presence of so many dinosaur species in the Latest Cretaceous is consistent with the termination of most lineages at the
Cretaceous-Tertiary boundary (the Late Maastrichtian sub-stage is 2.8 m.y. long). The Late Maastrichtian dinosaurian biodiversity is therefore consistent with the sudden extinction of the group following the Chicxulub impact.
A sauropod from the Albian of Normandy (northwestern France) is described as a new genus and spec... more A sauropod from the Albian of Normandy (northwestern France) is described as a new genus and species of basal Titanosauria, Normanniasaurus genceyi. N. genceyi shares several primitive characters with slightly younger basal Titanosauria such as Epachthosaurus and Andesaurus. It can be distinguished from other Albian sauropods of Western Europe. The late Early Cretaceous radiation of basal Titanosauria in South America, Europe and Africa is discussed.
New Mexico Museum of Natural History and Science Bulletin, 2012
In the midst of the “coprolite mining rush” of the early part of the second half of the nineteent... more In the midst of the “coprolite mining rush” of the early part of the second half of the nineteenth century, the amateur geologist Ferdinand Panescorse prospected for phosphatic nodules in his native region of the Var in Provence (southeastern France). He published in 1872 a catalogue of his finds, that he interpreted as coprolites, coining 70 morphotypes distributed among 19 groups. It is shown that most if not all of his “coprolites” are nodules of sedimentary origin.
The main Early Cretaceous vertebrate localities of Africa are listed and tentatively correlated. ... more The main Early Cretaceous vertebrate localities of Africa are listed and tentatively correlated. A Late Jurassic (Kimmeridgian−Tithonian: Tendaguru, Tanzania) to earliest Cretaceous (Valanginian: Algoa Basin, South Africa) assemblage with stegosaurids, brachiosaurids and diplodocids is replaced by a pre-Aptian (Hauterivian−Barremian?) assemblage with baryonychine spinosaurids, the pholidosaurid crocodile Sarcosuchus and large iguanodontids recorded from Niger (El Rhaz Formation), Cameroon (Koum Formation), and partially from Libya (Cabao Formation) and Tunisia (Douiret Formation) with the shark Priohybodus arambourgi. Aptian? to Early Albian assemblages still include iguanodontids but spinosaurine spinosaurids replace baryonychines. Early Cenomanian assemblages (Bahariya, Kem Kem) are characterized by the association of dinosaurs (Spinosaurus, Carcharodontosaurus), sharks (Onchopristis numidus) and crocodiles (libycosuchids, stomatosuchids). It is suggested that some supposedly Early Cretaceous formations from central Africa (Galula Formation of Tanzania, dinosaur beds of Malawi) are Late Cretaceous in age. Fossil vertebrates seem to be essential to correlate those continental formations.
Gargantuavis philoinos was described as a giant terrestrial bird on the basis of various postcran... more Gargantuavis philoinos was described as a giant terrestrial bird on the basis of various postcranial elements (synsacrum and pelvis, femur) from Late Cretaceous (Campanian-Maastrichtian) localities in Southern France. It has recently been suggested that these remains in fact belong to giant pterosaurs. A detailed comparison between bones referred to Gargantuavis and the corresponding skeletal elements of pterosaurs reveals considerable differences and confirms the avian nature of Gargantuavis. The broad pelvis of Gargantuavis is similar to that of various extinct graviportal terrestrial birds.
"Geologist Yves Milon, the Dean of the Faculty of Sciences of Rennes, hired the painter Mathurin ... more "Geologist Yves Milon, the Dean of the Faculty of Sciences of Rennes, hired the painter Mathurin Me´heut in 1941 to produce a large mural decoration for the new Geological Institute. This resulted in a little known 130 m2 artwork that includes a Mesozoic triptych, the genesis of which is described here. The work was executed during the World War II, when Milon’s illegal activities in several English intelligence services-led Resistance movements possibly prevented him from supervising the artist’s work and which led to some anatomical inaccuracies. This
decoration has survived several threats and constitutes a unique example of a large decorative palaeontological artwork in France. It has a special place in the history of dinosaur reconstructions as the choice of a decorative painting style is far from the usual forms of natural history illustration."
Fossils of vertebrates have been found in great abundance in the continental and marine early Lat... more Fossils of vertebrates have been found in great abundance in the continental and marine early Late Cretaceous sediments of Southeastern Morocco for more than 50 years. About 80 vertebrate taxa have so far been recorded from this region, many of which were recognised and diagnosed for the first time based on specimens recovered from these sediments. In this paper, we use published data together with new field data to present an updated overview of Moroccan early Late Cretaceous vertebrate assemblages. The Cretaceous series we have studied encompasses three Formations, the Ifezouane and Aoufous Formations, which are continental and deltaic in origin and are often grouped under the name ‘‘Kem Kem beds”, and the Akrabou Formation which is marine in origin. New field observations allow us to place four recognised vertebrate clusters, corresponding to one compound assemblage and three assemblages, within a general temporal framework. In particular, two ammonite bioevents characterise the lower part of the Upper Cenomanian (Calycoceras guerangeri Zone) at the base of the Akrabou Formation and the upper part of the Lower Turonian (Mammites nodosoides Zone), that may extend into the Middle Turonian within the Akrabou Formation, and allow for more accurate dating of the marine sequence in the study area. We are not yet able to distinguish a specific assemblage that characterises the Ifezouane Formation when compared to the similar Aoufous Formation, and as a result we regard the oldest of the four vertebrate ‘‘assemblages” in this region to be the compound assemblage of the ‘‘Kem Kem beds”. This well-known vertebrate assemblage comprises a mixture of terrestrial (and aerial), freshwater and brackish vertebrates. The archosaur component of this fauna appears to show an intriguingly high proportion of large-bodied carnivorous taxa, which may indicate a peculiar trophic chain, although collecting biases alter this palaeontological signal. A small and restricted assemblage, the OT1 assemblage, possibly corresponds to a specific, localised ecosystem within the Kem Kem beds compound assemblage. Microfossils and facies from the Aoufous Formation, corresponding to the top of the compound assemblage, provide evidence of extremely abiotic conditions (hypersalinity), and thus of great environmental instability. At the base of the Akrabou Formation the first ammonite bioevent, Neolobites, corresponds to the onset of the marine transgression in the early Late Cenomanian while the Agoult assemblage (Late Cenomanian?) contains a variety of small fish species that have Central Tethyan affinities. Finally, the youngest Mammites bioevent in the late Early Turonian corresponds to a deepening of the marine environment: this sequence is isochronous with the Goulmima assemblage, a diverse collection of fish and other marine taxa, and shows affinities with taxa from the South Atlantic, the Central Tethys and the Western Interior seaway of North America, and further highlights the biogeographical importance of these North African Late Cretaceous assemblages.
The holotype of Phuwiangosaurus sirindhornae MARTIN, BUFFETAUT and SUTEETHORN 1994, a 10 % compl... more The holotype of Phuwiangosaurus sirindhornae MARTIN, BUFFETAUT and SUTEETHORN 1994, a 10 % complete sauropod skeleton, was excavated in the early 1980s by a Thai-French expedition. In 1993, the site was re-opened and yielded two additional vertebrae: one posterior dorsal and one proximal caudal vertebra. On the basis of the size and shape of the vertebrae and the absence of identical bones from the type locality, we conclude that these vertebrae belong to the same individual as the holotype of P. sirindhornae and consider them as topotypes. A phylogenetic analysis, which for the first time includes recently described cranial and postcranial material, indicates that Phuwiangosaurus is a basal member of Titanosauriformes.
The Aptian-Albian locality of Huai Dan Chum in northeastern Thailand has yielded several trackway... more The Aptian-Albian locality of Huai Dan Chum in northeastern Thailand has yielded several trackways of crocodylians, together with ornithopod and theropod trackways. The crocodyliform trackways are described and some are referred to the ichnofamily Batrachopodidae.
Fossil vertebrates from the Cabao Formation discovered in the area of Nalut in northwestern Libya... more Fossil vertebrates from the Cabao Formation discovered in the area of Nalut in northwestern Libya include the hybodont shark Priohybodus, the crocodilian Sarcosuchus, an abelisaurid, a baryonichine spinosaurid and a large sauropod with spatulate teeth. The Cabao Formation may be Hauterivian to Barremian in age, although an earlier Berriasian to Valanginian age cannot be excluded. Its dinosaur assemblage is reminiscent of that of the El Rhaz and Tiouraren formations of Niger and strongly differs from both the Cenomanian assemblages of Morocco and Egypt and the Late Aptian to Albian fauna of Tunisia. Fossil vertebrates may be an important tool to establish the stratigraphical framework of the poorly dated Early Cretaceous continental deposits of Africa.
Vertebrate footprints have been discovered in recent years from seven Mesozoic formations of Thai... more Vertebrate footprints have been discovered in recent years from seven Mesozoic formations of Thailand and Laos dating from the Late Triassic (Kuchinari Group) and the Early Cretaceous (Khorat Group). The sites are reviewed here in chronological order. The ichnological record reflects fairly well the broad picture of the evolution of continental vertebrates in Asia known from the skeletal record. Norian basal archosaurs are replaced by Rhaetian dinosaurs although both footprint morphotypes look different from the contemporaneous European and North American forms. Two successive ornithopod radiations can be observed in the Early Cretaceous, with primitive small tetradactyl Hypsilophodon-like dinosaurs in the Earliest Cretaceous followed by advanced iguanodontoids with tridactyl fleshy footprints in the Aptian. Late Early Cretaceous dinosaur footprints from NE Thailand, however, do not validate previous hypotheses on the geographical distribution of Cretaceous ornithopod tracks in Asia. The ichnological record also reveals a hitherto unsuspected high diversity of theropods in the early Cretaceous with many different morphotypes.
A new skeleton of a sauropod dinosaur has been discovered in the Early Cretaceous Sao Khua Format... more A new skeleton of a sauropod dinosaur has been discovered in the Early Cretaceous Sao Khua Formation at Ban Na Khrai in Changwat Kalasin (NE Thailand). All sauropod bones from Ban Na Khrai share all their characteristics with the type specimen of Phuwiangosaurus sirindhornae Martin, Buffetaut & Suteethorn 1994. The 60% complete skeleton is very well preserved and includes cranial elements (a tooth, a frontal, a postorbital, a squamosal, both quadrates, and the braincase), whereas the type specimen is only 10% complete and consists of postcranial bones only. The material from Ban Na Khrai belongs to a single subadult individual of Phuwiangosaurus, as attested by the unfused neurocentral sutures of the vertebrae, which are firmly fused and larger in size in the holotypic specimen.
Abstract Fossil vertebrates from the Cabao Formation discovered in the area of Nalut in northwest... more Abstract Fossil vertebrates from the Cabao Formation discovered in the area of Nalut in northwestern Libya include the hybodont shark Priohybodus, the crocodilian Sarcosuchus, an abelisaurid, a baryonichine spinosaurid and a large sauropod with spatulate teeth. The Cabao Formation may be Hauterivian to Barremian in age, although an earlier Berriasian to Valanginian age cannot be excluded. Its dinosaur assemblage is reminiscent of that of the El Rhaz and Tiouraren formations of Niger and strongly differs from both the Cenomanian ...
The important question of early angiosperm growth habit (i.e., trees, shrubs or herbs?) remains u... more The important question of early angiosperm growth habit (i.e., trees, shrubs or herbs?) remains unanswered. Various theories have been based on data from both living and fossil plants. The Early Cretaceous fossil wood record, however, was seldom used to investigate early angiosperm habit. We set up a database for the Early Cretaceous and Cenomanian of Europe, as this area has the most complete and stratigraphically well-constrained record. The database has 170 entries, based on a bibliographical survey and on the examination of more than 600 new fossil wood specimens from a wide range of palaeoenvironments. In our record the woody characteristic in angiosperms appeared during the Albian, whereas most of the angiosperm's early evolution took place earlier, during the earliest Cretaceous. From the European fossil wood record for the Early Cretaceous and Cenomanian, the global extension and dominance of angiosperms in the Cenomanian is concomitant with a sharp increase in heteroxylous wood diversity. It appears that small stature and weak wood limited the angiosperm ecological radiation for some time. (J.-P. Billon-Bruyat), [email protected] (M. Guiomar), [email protected] (J.-L. Latil), [email protected] (J. Le loeuff), [email protected] (D. Néraudeau), [email protected] (D. Olivero), [email protected] (J. Schlögl).
1 Seccion de Geologia, Facultad de Ciencias, Universidad del Pais Vasco (Euskal Herriko Unibertsi... more 1 Seccion de Geologia, Facultad de Ciencias, Universidad del Pais Vasco (Euskal Herriko Unibertsitatea), Apdo 644, 48080 Bilbao, Euskadi. 2 CNRS, Laboratoire de Paléontologie des Vertébrés, Université Paris VI, 4 place Jussieu, 75252 Paris Cedex 05, France. 3 Unidad ...
169 coprolites were found at Huai Nam Aun in Chaiyaphum province (Huai Hin Lat Formation, Upper
T... more 169 coprolites were found at Huai Nam Aun in Chaiyaphum province (Huai Hin Lat Formation, Upper Triassic) and are sub-divided into seven different morphotypes. Four groups of spiral coprolites are interpreted as being produced by fish-eating sharks and lungfish, whereas non-spiral coprolites containing numerous bone fragments and fish scales were probably produced by other carnivorous vertebrates. Liassocopros hawkinsi and Saurocopros bucklandi are recognized in this study for the first time in Southeast Asia; this discovery supports palynological studies suggesting a Carnian-Norian age for the Huai Hin Lat Formation.
The global Late Maastrichtian non-avian dinosaur apparent biodiversity is extensively surveyed fo... more The global Late Maastrichtian non-avian dinosaur apparent biodiversity is extensively surveyed for the first time. It amounts to 104 species (including unnamed forms) in 2010. The real biodiversity being obscured by taphonomical biases and the scarcity of the continental fossil record, a species-area relationship is used to estimate it. The results show that several hundreds (between 628 and 1078) non-avian dinosaur species were alive in the Late Maastrichtian, which is almost an order of magnitude above previous estimates. Because of the complex Late Cretaceous palaeobiogeography, discussions about dinosaur extinction should be based on this estimated real global biodiversity, not on the apparent biodiversity of a single area. Given the mean duration of dinosaur genera (7.7 Ma), the presence of so many dinosaur species in the Latest Cretaceous is consistent with the termination of most lineages at the
Cretaceous-Tertiary boundary (the Late Maastrichtian sub-stage is 2.8 m.y. long). The Late Maastrichtian dinosaurian biodiversity is therefore consistent with the sudden extinction of the group following the Chicxulub impact.
A sauropod from the Albian of Normandy (northwestern France) is described as a new genus and spec... more A sauropod from the Albian of Normandy (northwestern France) is described as a new genus and species of basal Titanosauria, Normanniasaurus genceyi. N. genceyi shares several primitive characters with slightly younger basal Titanosauria such as Epachthosaurus and Andesaurus. It can be distinguished from other Albian sauropods of Western Europe. The late Early Cretaceous radiation of basal Titanosauria in South America, Europe and Africa is discussed.
New Mexico Museum of Natural History and Science Bulletin, 2012
In the midst of the “coprolite mining rush” of the early part of the second half of the nineteent... more In the midst of the “coprolite mining rush” of the early part of the second half of the nineteenth century, the amateur geologist Ferdinand Panescorse prospected for phosphatic nodules in his native region of the Var in Provence (southeastern France). He published in 1872 a catalogue of his finds, that he interpreted as coprolites, coining 70 morphotypes distributed among 19 groups. It is shown that most if not all of his “coprolites” are nodules of sedimentary origin.
The main Early Cretaceous vertebrate localities of Africa are listed and tentatively correlated. ... more The main Early Cretaceous vertebrate localities of Africa are listed and tentatively correlated. A Late Jurassic (Kimmeridgian−Tithonian: Tendaguru, Tanzania) to earliest Cretaceous (Valanginian: Algoa Basin, South Africa) assemblage with stegosaurids, brachiosaurids and diplodocids is replaced by a pre-Aptian (Hauterivian−Barremian?) assemblage with baryonychine spinosaurids, the pholidosaurid crocodile Sarcosuchus and large iguanodontids recorded from Niger (El Rhaz Formation), Cameroon (Koum Formation), and partially from Libya (Cabao Formation) and Tunisia (Douiret Formation) with the shark Priohybodus arambourgi. Aptian? to Early Albian assemblages still include iguanodontids but spinosaurine spinosaurids replace baryonychines. Early Cenomanian assemblages (Bahariya, Kem Kem) are characterized by the association of dinosaurs (Spinosaurus, Carcharodontosaurus), sharks (Onchopristis numidus) and crocodiles (libycosuchids, stomatosuchids). It is suggested that some supposedly Early Cretaceous formations from central Africa (Galula Formation of Tanzania, dinosaur beds of Malawi) are Late Cretaceous in age. Fossil vertebrates seem to be essential to correlate those continental formations.
Gargantuavis philoinos was described as a giant terrestrial bird on the basis of various postcran... more Gargantuavis philoinos was described as a giant terrestrial bird on the basis of various postcranial elements (synsacrum and pelvis, femur) from Late Cretaceous (Campanian-Maastrichtian) localities in Southern France. It has recently been suggested that these remains in fact belong to giant pterosaurs. A detailed comparison between bones referred to Gargantuavis and the corresponding skeletal elements of pterosaurs reveals considerable differences and confirms the avian nature of Gargantuavis. The broad pelvis of Gargantuavis is similar to that of various extinct graviportal terrestrial birds.
"Geologist Yves Milon, the Dean of the Faculty of Sciences of Rennes, hired the painter Mathurin ... more "Geologist Yves Milon, the Dean of the Faculty of Sciences of Rennes, hired the painter Mathurin Me´heut in 1941 to produce a large mural decoration for the new Geological Institute. This resulted in a little known 130 m2 artwork that includes a Mesozoic triptych, the genesis of which is described here. The work was executed during the World War II, when Milon’s illegal activities in several English intelligence services-led Resistance movements possibly prevented him from supervising the artist’s work and which led to some anatomical inaccuracies. This
decoration has survived several threats and constitutes a unique example of a large decorative palaeontological artwork in France. It has a special place in the history of dinosaur reconstructions as the choice of a decorative painting style is far from the usual forms of natural history illustration."
Fossils of vertebrates have been found in great abundance in the continental and marine early Lat... more Fossils of vertebrates have been found in great abundance in the continental and marine early Late Cretaceous sediments of Southeastern Morocco for more than 50 years. About 80 vertebrate taxa have so far been recorded from this region, many of which were recognised and diagnosed for the first time based on specimens recovered from these sediments. In this paper, we use published data together with new field data to present an updated overview of Moroccan early Late Cretaceous vertebrate assemblages. The Cretaceous series we have studied encompasses three Formations, the Ifezouane and Aoufous Formations, which are continental and deltaic in origin and are often grouped under the name ‘‘Kem Kem beds”, and the Akrabou Formation which is marine in origin. New field observations allow us to place four recognised vertebrate clusters, corresponding to one compound assemblage and three assemblages, within a general temporal framework. In particular, two ammonite bioevents characterise the lower part of the Upper Cenomanian (Calycoceras guerangeri Zone) at the base of the Akrabou Formation and the upper part of the Lower Turonian (Mammites nodosoides Zone), that may extend into the Middle Turonian within the Akrabou Formation, and allow for more accurate dating of the marine sequence in the study area. We are not yet able to distinguish a specific assemblage that characterises the Ifezouane Formation when compared to the similar Aoufous Formation, and as a result we regard the oldest of the four vertebrate ‘‘assemblages” in this region to be the compound assemblage of the ‘‘Kem Kem beds”. This well-known vertebrate assemblage comprises a mixture of terrestrial (and aerial), freshwater and brackish vertebrates. The archosaur component of this fauna appears to show an intriguingly high proportion of large-bodied carnivorous taxa, which may indicate a peculiar trophic chain, although collecting biases alter this palaeontological signal. A small and restricted assemblage, the OT1 assemblage, possibly corresponds to a specific, localised ecosystem within the Kem Kem beds compound assemblage. Microfossils and facies from the Aoufous Formation, corresponding to the top of the compound assemblage, provide evidence of extremely abiotic conditions (hypersalinity), and thus of great environmental instability. At the base of the Akrabou Formation the first ammonite bioevent, Neolobites, corresponds to the onset of the marine transgression in the early Late Cenomanian while the Agoult assemblage (Late Cenomanian?) contains a variety of small fish species that have Central Tethyan affinities. Finally, the youngest Mammites bioevent in the late Early Turonian corresponds to a deepening of the marine environment: this sequence is isochronous with the Goulmima assemblage, a diverse collection of fish and other marine taxa, and shows affinities with taxa from the South Atlantic, the Central Tethys and the Western Interior seaway of North America, and further highlights the biogeographical importance of these North African Late Cretaceous assemblages.
The holotype of Phuwiangosaurus sirindhornae MARTIN, BUFFETAUT and SUTEETHORN 1994, a 10 % compl... more The holotype of Phuwiangosaurus sirindhornae MARTIN, BUFFETAUT and SUTEETHORN 1994, a 10 % complete sauropod skeleton, was excavated in the early 1980s by a Thai-French expedition. In 1993, the site was re-opened and yielded two additional vertebrae: one posterior dorsal and one proximal caudal vertebra. On the basis of the size and shape of the vertebrae and the absence of identical bones from the type locality, we conclude that these vertebrae belong to the same individual as the holotype of P. sirindhornae and consider them as topotypes. A phylogenetic analysis, which for the first time includes recently described cranial and postcranial material, indicates that Phuwiangosaurus is a basal member of Titanosauriformes.
The Aptian-Albian locality of Huai Dan Chum in northeastern Thailand has yielded several trackway... more The Aptian-Albian locality of Huai Dan Chum in northeastern Thailand has yielded several trackways of crocodylians, together with ornithopod and theropod trackways. The crocodyliform trackways are described and some are referred to the ichnofamily Batrachopodidae.
Fossil vertebrates from the Cabao Formation discovered in the area of Nalut in northwestern Libya... more Fossil vertebrates from the Cabao Formation discovered in the area of Nalut in northwestern Libya include the hybodont shark Priohybodus, the crocodilian Sarcosuchus, an abelisaurid, a baryonichine spinosaurid and a large sauropod with spatulate teeth. The Cabao Formation may be Hauterivian to Barremian in age, although an earlier Berriasian to Valanginian age cannot be excluded. Its dinosaur assemblage is reminiscent of that of the El Rhaz and Tiouraren formations of Niger and strongly differs from both the Cenomanian assemblages of Morocco and Egypt and the Late Aptian to Albian fauna of Tunisia. Fossil vertebrates may be an important tool to establish the stratigraphical framework of the poorly dated Early Cretaceous continental deposits of Africa.
Vertebrate footprints have been discovered in recent years from seven Mesozoic formations of Thai... more Vertebrate footprints have been discovered in recent years from seven Mesozoic formations of Thailand and Laos dating from the Late Triassic (Kuchinari Group) and the Early Cretaceous (Khorat Group). The sites are reviewed here in chronological order. The ichnological record reflects fairly well the broad picture of the evolution of continental vertebrates in Asia known from the skeletal record. Norian basal archosaurs are replaced by Rhaetian dinosaurs although both footprint morphotypes look different from the contemporaneous European and North American forms. Two successive ornithopod radiations can be observed in the Early Cretaceous, with primitive small tetradactyl Hypsilophodon-like dinosaurs in the Earliest Cretaceous followed by advanced iguanodontoids with tridactyl fleshy footprints in the Aptian. Late Early Cretaceous dinosaur footprints from NE Thailand, however, do not validate previous hypotheses on the geographical distribution of Cretaceous ornithopod tracks in Asia. The ichnological record also reveals a hitherto unsuspected high diversity of theropods in the early Cretaceous with many different morphotypes.
A new skeleton of a sauropod dinosaur has been discovered in the Early Cretaceous Sao Khua Format... more A new skeleton of a sauropod dinosaur has been discovered in the Early Cretaceous Sao Khua Formation at Ban Na Khrai in Changwat Kalasin (NE Thailand). All sauropod bones from Ban Na Khrai share all their characteristics with the type specimen of Phuwiangosaurus sirindhornae Martin, Buffetaut & Suteethorn 1994. The 60% complete skeleton is very well preserved and includes cranial elements (a tooth, a frontal, a postorbital, a squamosal, both quadrates, and the braincase), whereas the type specimen is only 10% complete and consists of postcranial bones only. The material from Ban Na Khrai belongs to a single subadult individual of Phuwiangosaurus, as attested by the unfused neurocentral sutures of the vertebrae, which are firmly fused and larger in size in the holotypic specimen.
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Papers by Jean Le Loeuff
Triassic) and are sub-divided into seven different morphotypes. Four groups of spiral coprolites are interpreted as
being produced by fish-eating sharks and lungfish, whereas non-spiral coprolites containing numerous bone
fragments and fish scales were probably produced by other carnivorous vertebrates. Liassocopros hawkinsi and
Saurocopros bucklandi are recognized in this study for the first time in Southeast Asia; this discovery supports
palynological studies suggesting a Carnian-Norian age for the Huai Hin Lat Formation.
Cretaceous-Tertiary boundary (the Late Maastrichtian sub-stage is 2.8 m.y. long). The Late Maastrichtian dinosaurian biodiversity is therefore consistent with the sudden extinction of the group following the Chicxulub impact.
assemblage with stegosaurids, brachiosaurids and diplodocids is replaced by a pre-Aptian (Hauterivian−Barremian?) assemblage with baryonychine spinosaurids, the pholidosaurid crocodile Sarcosuchus and large iguanodontids recorded from Niger (El Rhaz Formation), Cameroon (Koum Formation), and partially from Libya (Cabao Formation) and Tunisia
(Douiret Formation) with the shark Priohybodus arambourgi. Aptian? to Early Albian assemblages still include iguanodontids but spinosaurine spinosaurids replace baryonychines. Early Cenomanian assemblages (Bahariya, Kem Kem)
are characterized by the association of dinosaurs (Spinosaurus, Carcharodontosaurus), sharks (Onchopristis numidus) and crocodiles (libycosuchids, stomatosuchids). It is suggested that some supposedly Early Cretaceous formations from central Africa (Galula Formation of Tanzania, dinosaur beds of Malawi) are Late Cretaceous in age. Fossil vertebrates seem to be essential to correlate those continental formations.
considerable differences and confirms the avian nature of Gargantuavis. The broad pelvis of Gargantuavis is similar to that of various extinct graviportal terrestrial birds.
decoration has survived several threats and constitutes a unique example of a large decorative palaeontological artwork in France. It has a special place in the history of dinosaur reconstructions as the choice of a decorative painting style is far from the usual forms of natural history illustration."
data to present an updated overview of Moroccan early Late Cretaceous vertebrate assemblages. The Cretaceous series we have studied encompasses three Formations, the Ifezouane and Aoufous Formations, which are continental and deltaic in origin and are often grouped under the name ‘‘Kem Kem beds”, and the Akrabou Formation which is marine in origin. New field observations allow us to place four recognised vertebrate clusters, corresponding to one compound assemblage and three assemblages, within a general temporal framework. In particular, two ammonite bioevents characterise the lower part of the Upper Cenomanian (Calycoceras guerangeri Zone) at the base of the Akrabou Formation and the upper part of the Lower Turonian (Mammites nodosoides Zone), that may extend into the Middle Turonian within the Akrabou Formation, and allow for more accurate dating of the marine sequence in the study area. We are not yet able to distinguish a specific assemblage that characterises the Ifezouane Formation when compared to the similar Aoufous Formation, and as a result we regard the oldest of the four vertebrate ‘‘assemblages” in this region to be the compound assemblage of the ‘‘Kem Kem beds”. This
well-known vertebrate assemblage comprises a mixture of terrestrial (and aerial), freshwater and brackish vertebrates. The archosaur component of this fauna appears to show an intriguingly high proportion of large-bodied carnivorous taxa, which may indicate a peculiar trophic chain, although collecting biases alter this palaeontological signal. A small and restricted assemblage, the OT1 assemblage, possibly corresponds to a specific, localised ecosystem within the Kem Kem beds compound assemblage. Microfossils and facies from the Aoufous Formation, corresponding to the top of the compound assemblage, provide evidence of extremely abiotic conditions (hypersalinity), and thus of great environmental instability. At the base of the Akrabou Formation the first ammonite bioevent, Neolobites, corresponds to the onset of
the marine transgression in the early Late Cenomanian while the Agoult assemblage (Late Cenomanian?) contains a variety of small fish species that have Central Tethyan affinities. Finally, the youngest Mammites bioevent in the late Early Turonian corresponds to a deepening of the marine environment: this sequence is isochronous with the Goulmima assemblage, a diverse collection of fish and other marine
taxa, and shows affinities with taxa from the South Atlantic, the Central Tethys and the Western Interior seaway of North America, and further highlights the biogeographical importance of these North African Late Cretaceous assemblages.
expedition. In 1993, the site was re-opened and yielded two additional vertebrae: one posterior dorsal and one proximal caudal vertebra. On the basis of the size and shape of the vertebrae and the absence of identical bones from the type locality, we conclude that these vertebrae belong to the same individual as the holotype of P. sirindhornae and consider them as topotypes. A phylogenetic analysis, which for the first time includes recently described cranial and postcranial material, indicates that Phuwiangosaurus is a basal member of Titanosauriformes.
baryonichine spinosaurid and a large sauropod with spatulate teeth. The Cabao Formation may be Hauterivian to Barremian in age, although an earlier Berriasian to Valanginian age cannot be excluded. Its dinosaur assemblage is reminiscent of that of the El Rhaz and Tiouraren formations of Niger and strongly differs from both the Cenomanian assemblages of Morocco and Egypt and the Late Aptian to Albian fauna of Tunisia. Fossil vertebrates may be an important tool to establish the stratigraphical framework of the poorly dated Early Cretaceous continental deposits of Africa.
Ban Na Khrai share all their characteristics with the type specimen of Phuwiangosaurus sirindhornae Martin, Buffetaut & Suteethorn 1994. The 60% complete skeleton is very well preserved and includes cranial elements (a tooth, a frontal, a postorbital, a squamosal, both quadrates, and the braincase), whereas the type specimen is only 10% complete and consists of postcranial bones only. The material from Ban Na Khrai belongs to a single subadult individual of Phuwiangosaurus,
as attested by the unfused neurocentral sutures of the vertebrae, which are firmly fused and larger in size in the holotypic specimen.
Triassic) and are sub-divided into seven different morphotypes. Four groups of spiral coprolites are interpreted as
being produced by fish-eating sharks and lungfish, whereas non-spiral coprolites containing numerous bone
fragments and fish scales were probably produced by other carnivorous vertebrates. Liassocopros hawkinsi and
Saurocopros bucklandi are recognized in this study for the first time in Southeast Asia; this discovery supports
palynological studies suggesting a Carnian-Norian age for the Huai Hin Lat Formation.
Cretaceous-Tertiary boundary (the Late Maastrichtian sub-stage is 2.8 m.y. long). The Late Maastrichtian dinosaurian biodiversity is therefore consistent with the sudden extinction of the group following the Chicxulub impact.
assemblage with stegosaurids, brachiosaurids and diplodocids is replaced by a pre-Aptian (Hauterivian−Barremian?) assemblage with baryonychine spinosaurids, the pholidosaurid crocodile Sarcosuchus and large iguanodontids recorded from Niger (El Rhaz Formation), Cameroon (Koum Formation), and partially from Libya (Cabao Formation) and Tunisia
(Douiret Formation) with the shark Priohybodus arambourgi. Aptian? to Early Albian assemblages still include iguanodontids but spinosaurine spinosaurids replace baryonychines. Early Cenomanian assemblages (Bahariya, Kem Kem)
are characterized by the association of dinosaurs (Spinosaurus, Carcharodontosaurus), sharks (Onchopristis numidus) and crocodiles (libycosuchids, stomatosuchids). It is suggested that some supposedly Early Cretaceous formations from central Africa (Galula Formation of Tanzania, dinosaur beds of Malawi) are Late Cretaceous in age. Fossil vertebrates seem to be essential to correlate those continental formations.
considerable differences and confirms the avian nature of Gargantuavis. The broad pelvis of Gargantuavis is similar to that of various extinct graviportal terrestrial birds.
decoration has survived several threats and constitutes a unique example of a large decorative palaeontological artwork in France. It has a special place in the history of dinosaur reconstructions as the choice of a decorative painting style is far from the usual forms of natural history illustration."
data to present an updated overview of Moroccan early Late Cretaceous vertebrate assemblages. The Cretaceous series we have studied encompasses three Formations, the Ifezouane and Aoufous Formations, which are continental and deltaic in origin and are often grouped under the name ‘‘Kem Kem beds”, and the Akrabou Formation which is marine in origin. New field observations allow us to place four recognised vertebrate clusters, corresponding to one compound assemblage and three assemblages, within a general temporal framework. In particular, two ammonite bioevents characterise the lower part of the Upper Cenomanian (Calycoceras guerangeri Zone) at the base of the Akrabou Formation and the upper part of the Lower Turonian (Mammites nodosoides Zone), that may extend into the Middle Turonian within the Akrabou Formation, and allow for more accurate dating of the marine sequence in the study area. We are not yet able to distinguish a specific assemblage that characterises the Ifezouane Formation when compared to the similar Aoufous Formation, and as a result we regard the oldest of the four vertebrate ‘‘assemblages” in this region to be the compound assemblage of the ‘‘Kem Kem beds”. This
well-known vertebrate assemblage comprises a mixture of terrestrial (and aerial), freshwater and brackish vertebrates. The archosaur component of this fauna appears to show an intriguingly high proportion of large-bodied carnivorous taxa, which may indicate a peculiar trophic chain, although collecting biases alter this palaeontological signal. A small and restricted assemblage, the OT1 assemblage, possibly corresponds to a specific, localised ecosystem within the Kem Kem beds compound assemblage. Microfossils and facies from the Aoufous Formation, corresponding to the top of the compound assemblage, provide evidence of extremely abiotic conditions (hypersalinity), and thus of great environmental instability. At the base of the Akrabou Formation the first ammonite bioevent, Neolobites, corresponds to the onset of
the marine transgression in the early Late Cenomanian while the Agoult assemblage (Late Cenomanian?) contains a variety of small fish species that have Central Tethyan affinities. Finally, the youngest Mammites bioevent in the late Early Turonian corresponds to a deepening of the marine environment: this sequence is isochronous with the Goulmima assemblage, a diverse collection of fish and other marine
taxa, and shows affinities with taxa from the South Atlantic, the Central Tethys and the Western Interior seaway of North America, and further highlights the biogeographical importance of these North African Late Cretaceous assemblages.
expedition. In 1993, the site was re-opened and yielded two additional vertebrae: one posterior dorsal and one proximal caudal vertebra. On the basis of the size and shape of the vertebrae and the absence of identical bones from the type locality, we conclude that these vertebrae belong to the same individual as the holotype of P. sirindhornae and consider them as topotypes. A phylogenetic analysis, which for the first time includes recently described cranial and postcranial material, indicates that Phuwiangosaurus is a basal member of Titanosauriformes.
baryonichine spinosaurid and a large sauropod with spatulate teeth. The Cabao Formation may be Hauterivian to Barremian in age, although an earlier Berriasian to Valanginian age cannot be excluded. Its dinosaur assemblage is reminiscent of that of the El Rhaz and Tiouraren formations of Niger and strongly differs from both the Cenomanian assemblages of Morocco and Egypt and the Late Aptian to Albian fauna of Tunisia. Fossil vertebrates may be an important tool to establish the stratigraphical framework of the poorly dated Early Cretaceous continental deposits of Africa.
Ban Na Khrai share all their characteristics with the type specimen of Phuwiangosaurus sirindhornae Martin, Buffetaut & Suteethorn 1994. The 60% complete skeleton is very well preserved and includes cranial elements (a tooth, a frontal, a postorbital, a squamosal, both quadrates, and the braincase), whereas the type specimen is only 10% complete and consists of postcranial bones only. The material from Ban Na Khrai belongs to a single subadult individual of Phuwiangosaurus,
as attested by the unfused neurocentral sutures of the vertebrae, which are firmly fused and larger in size in the holotypic specimen.