Adult dinosaurs preserved attending their nests in brooding positions are among the rarest verteb... more Adult dinosaurs preserved attending their nests in brooding positions are among the rarest vertebrate fossils. By far the most common occurrences are members of the dinosaur group Oviraptorosauria. The first finds of these were specimens recovered from the Djadokhta Formation at the Mongolian locality of Ukhaa Tolgod and the Chinese locality of Bayan Mandahu. Since the initial discovery of these specimens, a few more occurrences of nesting oviraptors have been found at other Asian localities. Here we report on a second nesting oviraptorid specimen (IGM 100/1004) sitting in a brooding position atop a nest of eggs from Ukhaa Tolgod, Omnogov, Mongolia. This is a large specimen of the ubiquitous Ukhaa Tolgod taxon Citipati osmolskae. It is approximately 11% larger based on humeral length than the original Ukhaa Tolgod nesting Citipati osmolskae specimen (IGM 100/979), yet eggshell structure and egg arrangement are identical. No evidence for colonial breeding of these animals has been recovered. Reexamination of another " nesting " oviraptorosaur, the holotype of Oviraptor philoceratops (AMNH FARB 6517) indicates that in addition to the numerous partial eggs associated with the original skeleton that originally led to its referral as a protoceratopsian predator, there are the remains of a tiny theropod. This hind limb can be provisionally assigned to Oviraptoridae. It is thus at least possible that some of the eggs associated with the holotype had hatched and the perinates had not left the nest.
A new troodontid dinosaur, Almas ukhaa, from the Late Cretaceous deposits of the Djadokhta Format... more A new troodontid dinosaur, Almas ukhaa, from the Late Cretaceous deposits of the Djadokhta Formation at Ukhaa Tolgod, Mongolia, is described here. The holotype specimen (IGM 100/1323) comprises an almost complete and articulated cranium and partial articulated postcranial skeleton. This specimen has a small body size and a short snout as in basal paravians, but it exhibits a number of derived troodontid features that differentiate Almas ukhaa from the Early Cretaceous troodontids reported from China and unite this new taxon with other Late Cretaceous troodontids. Relative to other troodontids, Almas ukhaa is autapomorphic in the presence of a posteriorly curved pterygoid flange, absence of a lateral groove on the anterior part of the dentary, presence of a distinct spikelike process on the ischium, and elongate chevrons. The eggshell associated with IGM 100/1323 can be assigned to Prismatoolithidae indet. based on the smooth surface, eggshell thickness, and micro-structural characteristics, and also preserves attributes similar to Protoceratopsidovum minimum. A unique relationship between Byronosaurus and the perinate troodontids IGM 100/972 and IGM 100/974 is no longer supported based on the new observations of Almas ukhaa and Gobivenator.
The origin of the avian hand, with its reduced and fused carpals and digits, from the five-finger... more The origin of the avian hand, with its reduced and fused carpals and digits, from the five-fingered hands and complex wrists of early dinosaurs represents one of the major transformations of manus morphology among tetrapods. Much attention has been directed to the later part of this transition, from four-to three-fingered taxa. However, earlier anatomical changes may have influenced these later modifications, possibly paving the way for a later frameshift in digit identities. We investigate the five-to four-fingered transition among early dinosaurs, along with changes in carpus morphology. New three-dimensional reconstructions from computed tomography data of the manus of the Triassic and Early Jurassic theropod dinosaurs Coelophysis bauri and Megapnosaurus rhodesiensis are described and compared intra-and interspecifically. Several novel findings emerge from these reconstructions and comparisons, including the first evidence of an ossified centrale and a free intermedium in some C. bauri specimens, as well as confirmation of the presence of a vestigial fifth metacarpal in this taxon. Additionally, a specimen of C. bauri and an unnamed coelophysoid from the Upper Triassic Hayden Quarry, New Mexico, are to our knowledge the only theropods (other than alvarezsaurs and birds) in which all of the distal carpals are completely fused together into a single unit. Several differences between the manus of C. bauri and M. rhodesiensis are also identified. We review the evolution of the archosauromorph manus more broadly in light of these new data, and caution against incorporating carpal characters in phylogenetic analyses of fine-scale relationships of Archosauromorpha, in light of the high degree of observed polymorphism in taxa for which large sample sizes are available, such as the theropod Coelophysis and the sauropodomorph Plateosaurus. We also find that the reduction of the carpus and ultimate loss of the fourth and fifth digits among early dinosaurs did not proceed in a neat, stepwise fashion, but was characterized by multiple losses and possible gains of carpals, metacarpals and phalanges. Taken together, the high degree of intra-and interspecific variability in the number and identities of carpals, and the state of reduction of the fourth and fifth digits suggest the presence of a 'zone of developmental variability' in early dinosaur manus evolution, from which novel avian-like morphologies eventually emerged and became channelized among later theropod clades.
Although Mesozoic fossils are quite common in the Gobi Desert of Central Asia, it is often diffic... more Although Mesozoic fossils are quite common in the Gobi Desert of Central Asia, it is often difficult to correlate among different localities because of a dearth of rocks amenable to absolute dating. Specifically, correlating between the eastern Gobi Desert and more western localities has been challenging. Here we give a Santonian-Campanian age for the enigmatic Zos Canyon beds in the Nemegt basin. This is based on the occurrence of the primitive ornithopod dinosaur Haya griva at both eastern Gobi exposures of the Javkhlant Formation and the Zos Canyon locality.
In 1991, Sabath described “larger avian eggs” from the Upper Cretaceous Barun Goyot and Djadokhta... more In 1991, Sabath described “larger avian eggs” from the Upper Cretaceous Barun Goyot and Djadokhta Formations of Mongolia. These were later included in the ootaxon Gobioolithus major. Here we recognize the larger avian eggs of Sa- bath as a distinct ootaxon, Styloolithus sabathi, oogen. et oosp. nov. These eggs differ from those of Gobioolithus in being larger (70 by 32 mm) and more elongate. Microscopically, the shell bears a third layer (possible external zone) thicker than the mammillary layer and nearly as thick as the second layer (possible squamatic zone); the continuous layer (in- cluding layers two and three) to mammillary layer thickness ratio is 3.1:1. Within the clutch, the tightly spaced eggs stand with their long axes steeply inclined. Adult remains are associated with two clutches, suggesting an incubation mode similar to that of troodontid maniraptorans, where adults sat atop largely buried eggs. S. sabathi provides evidence that relative egg size in Mesozoic non-ornithuromorph birds had increased markedly from the non-avian theropod condition in oviraptorids and troodontids, but had not yet reached the modern egg-adult proportions of Neornithes. Sediment-bound upright eggs appear common to Enantiornithes and more basal avians, suggesting that like non-avian theropods, these birds lacked chalazae, the chords of albumen allowing egg rotation in modern birds. Absence of this simple structure may have restricted these basal birds to ground nesting in areas with appropriate substrates and not permitted the type of nesting diversity found in Neornithes. Neornithes are the only Mesozoic clade of Dinosauria to nest completely free of sediment; this may have played a crucial role in their surviving the K-Pg mass extinction event.
Numerous discoveries in the Tiantai basin of Zhejiang Province, China, enrich our understanding o... more Numerous discoveries in the Tiantai basin of Zhejiang Province, China, enrich our understanding of the parataxonomy, paleobiology and taphonomic histories of fossil eggs from a diverse array of Cretaceous oofamilies. We describe the most abundant of these egg types catalogued in the Zhejiang Museum of Natural History, Spheroolithus cf. zhangtoucaoensis (oofamily Spheroolithidae). Scanning electron microscopy, here utilised for the first time on Spheroolithus eggs from Tiantai, and petrographic microscopy reveal 0.81–1.37-mm thick eggshell composed of a single structural layer of calcite with slightly flaring shell units, irregular pores, horizontal accretion lines and a sweeping extinction pattern under crossed polars. This contradicts previous reports of the presence of two structural layers in Tiantai Spheroolithus. Clutches consist of 2–13 eggs arranged in an irregular, single-layered pattern. As these eggs are among the oldest Asian examples of Spheroolithus currently known, they may shed light on the early evolution of this oogenus. This study establishes the definitive presence of the Spheroolithidae in the Tiantai basin, contra recent reports, and provides the framework for ongoing examination of egg diversity and taphonomy in the Tiantai basin.
Adult dinosaurs preserved attending their nests in brooding positions are among the rarest verteb... more Adult dinosaurs preserved attending their nests in brooding positions are among the rarest vertebrate fossils. By far the most common occurrences are members of the dinosaur group Oviraptorosauria. The first finds of these were specimens recovered from the Djadokhta Formation at the Mongolian locality of Ukhaa Tolgod and the Chinese locality of Bayan Mandahu. Since the initial discovery of these specimens, a few more occurrences of nesting oviraptors have been found at other Asian localities. Here we report on a second nesting oviraptorid specimen (IGM 100/1004) sitting in a brooding position atop a nest of eggs from Ukhaa Tolgod, Omnogov, Mongolia. This is a large specimen of the ubiquitous Ukhaa Tolgod taxon Citipati osmolskae. It is approximately 11% larger based on humeral length than the original Ukhaa Tolgod nesting Citipati osmolskae specimen (IGM 100/979), yet eggshell structure and egg arrangement are identical. No evidence for colonial breeding of these animals has been recovered. Reexamination of another " nesting " oviraptorosaur, the holotype of Oviraptor philoceratops (AMNH FARB 6517) indicates that in addition to the numerous partial eggs associated with the original skeleton that originally led to its referral as a protoceratopsian predator, there are the remains of a tiny theropod. This hind limb can be provisionally assigned to Oviraptoridae. It is thus at least possible that some of the eggs associated with the holotype had hatched and the perinates had not left the nest.
A new troodontid dinosaur, Almas ukhaa, from the Late Cretaceous deposits of the Djadokhta Format... more A new troodontid dinosaur, Almas ukhaa, from the Late Cretaceous deposits of the Djadokhta Formation at Ukhaa Tolgod, Mongolia, is described here. The holotype specimen (IGM 100/1323) comprises an almost complete and articulated cranium and partial articulated postcranial skeleton. This specimen has a small body size and a short snout as in basal paravians, but it exhibits a number of derived troodontid features that differentiate Almas ukhaa from the Early Cretaceous troodontids reported from China and unite this new taxon with other Late Cretaceous troodontids. Relative to other troodontids, Almas ukhaa is autapomorphic in the presence of a posteriorly curved pterygoid flange, absence of a lateral groove on the anterior part of the dentary, presence of a distinct spikelike process on the ischium, and elongate chevrons. The eggshell associated with IGM 100/1323 can be assigned to Prismatoolithidae indet. based on the smooth surface, eggshell thickness, and micro-structural characteristics, and also preserves attributes similar to Protoceratopsidovum minimum. A unique relationship between Byronosaurus and the perinate troodontids IGM 100/972 and IGM 100/974 is no longer supported based on the new observations of Almas ukhaa and Gobivenator.
The origin of the avian hand, with its reduced and fused carpals and digits, from the five-finger... more The origin of the avian hand, with its reduced and fused carpals and digits, from the five-fingered hands and complex wrists of early dinosaurs represents one of the major transformations of manus morphology among tetrapods. Much attention has been directed to the later part of this transition, from four-to three-fingered taxa. However, earlier anatomical changes may have influenced these later modifications, possibly paving the way for a later frameshift in digit identities. We investigate the five-to four-fingered transition among early dinosaurs, along with changes in carpus morphology. New three-dimensional reconstructions from computed tomography data of the manus of the Triassic and Early Jurassic theropod dinosaurs Coelophysis bauri and Megapnosaurus rhodesiensis are described and compared intra-and interspecifically. Several novel findings emerge from these reconstructions and comparisons, including the first evidence of an ossified centrale and a free intermedium in some C. bauri specimens, as well as confirmation of the presence of a vestigial fifth metacarpal in this taxon. Additionally, a specimen of C. bauri and an unnamed coelophysoid from the Upper Triassic Hayden Quarry, New Mexico, are to our knowledge the only theropods (other than alvarezsaurs and birds) in which all of the distal carpals are completely fused together into a single unit. Several differences between the manus of C. bauri and M. rhodesiensis are also identified. We review the evolution of the archosauromorph manus more broadly in light of these new data, and caution against incorporating carpal characters in phylogenetic analyses of fine-scale relationships of Archosauromorpha, in light of the high degree of observed polymorphism in taxa for which large sample sizes are available, such as the theropod Coelophysis and the sauropodomorph Plateosaurus. We also find that the reduction of the carpus and ultimate loss of the fourth and fifth digits among early dinosaurs did not proceed in a neat, stepwise fashion, but was characterized by multiple losses and possible gains of carpals, metacarpals and phalanges. Taken together, the high degree of intra-and interspecific variability in the number and identities of carpals, and the state of reduction of the fourth and fifth digits suggest the presence of a 'zone of developmental variability' in early dinosaur manus evolution, from which novel avian-like morphologies eventually emerged and became channelized among later theropod clades.
Although Mesozoic fossils are quite common in the Gobi Desert of Central Asia, it is often diffic... more Although Mesozoic fossils are quite common in the Gobi Desert of Central Asia, it is often difficult to correlate among different localities because of a dearth of rocks amenable to absolute dating. Specifically, correlating between the eastern Gobi Desert and more western localities has been challenging. Here we give a Santonian-Campanian age for the enigmatic Zos Canyon beds in the Nemegt basin. This is based on the occurrence of the primitive ornithopod dinosaur Haya griva at both eastern Gobi exposures of the Javkhlant Formation and the Zos Canyon locality.
In 1991, Sabath described “larger avian eggs” from the Upper Cretaceous Barun Goyot and Djadokhta... more In 1991, Sabath described “larger avian eggs” from the Upper Cretaceous Barun Goyot and Djadokhta Formations of Mongolia. These were later included in the ootaxon Gobioolithus major. Here we recognize the larger avian eggs of Sa- bath as a distinct ootaxon, Styloolithus sabathi, oogen. et oosp. nov. These eggs differ from those of Gobioolithus in being larger (70 by 32 mm) and more elongate. Microscopically, the shell bears a third layer (possible external zone) thicker than the mammillary layer and nearly as thick as the second layer (possible squamatic zone); the continuous layer (in- cluding layers two and three) to mammillary layer thickness ratio is 3.1:1. Within the clutch, the tightly spaced eggs stand with their long axes steeply inclined. Adult remains are associated with two clutches, suggesting an incubation mode similar to that of troodontid maniraptorans, where adults sat atop largely buried eggs. S. sabathi provides evidence that relative egg size in Mesozoic non-ornithuromorph birds had increased markedly from the non-avian theropod condition in oviraptorids and troodontids, but had not yet reached the modern egg-adult proportions of Neornithes. Sediment-bound upright eggs appear common to Enantiornithes and more basal avians, suggesting that like non-avian theropods, these birds lacked chalazae, the chords of albumen allowing egg rotation in modern birds. Absence of this simple structure may have restricted these basal birds to ground nesting in areas with appropriate substrates and not permitted the type of nesting diversity found in Neornithes. Neornithes are the only Mesozoic clade of Dinosauria to nest completely free of sediment; this may have played a crucial role in their surviving the K-Pg mass extinction event.
Numerous discoveries in the Tiantai basin of Zhejiang Province, China, enrich our understanding o... more Numerous discoveries in the Tiantai basin of Zhejiang Province, China, enrich our understanding of the parataxonomy, paleobiology and taphonomic histories of fossil eggs from a diverse array of Cretaceous oofamilies. We describe the most abundant of these egg types catalogued in the Zhejiang Museum of Natural History, Spheroolithus cf. zhangtoucaoensis (oofamily Spheroolithidae). Scanning electron microscopy, here utilised for the first time on Spheroolithus eggs from Tiantai, and petrographic microscopy reveal 0.81–1.37-mm thick eggshell composed of a single structural layer of calcite with slightly flaring shell units, irregular pores, horizontal accretion lines and a sweeping extinction pattern under crossed polars. This contradicts previous reports of the presence of two structural layers in Tiantai Spheroolithus. Clutches consist of 2–13 eggs arranged in an irregular, single-layered pattern. As these eggs are among the oldest Asian examples of Spheroolithus currently known, they may shed light on the early evolution of this oogenus. This study establishes the definitive presence of the Spheroolithidae in the Tiantai basin, contra recent reports, and provides the framework for ongoing examination of egg diversity and taphonomy in the Tiantai basin.
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