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A portion of a left humerus from the Upper Maastrichtian of Vratsa district (NW Bulgaria)
is shown to be from a non-avian theropod dinosaur: this is the first record of a
dinosaur from Bulgaria. We describe this bone, suggest that it most likely pertains to an
ornithomimosaur, and discuss the fossil record of other similar taxa of Late Cretaceous
age that have been reported from Europe. To investigate the taphonomy of this fossil,
rare earth element (REE) analysis is combined with strontium (Sr) isotope data to confirm
that this Bulgarian dinosaur bone was initially fossilized in a terrestrial environment,
then later re-worked into late Maastrichtian marine sediments.

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Despite more than a century of collecting, resulting in one of the best-studied vertebrate fossil records anywhere in the world, the Upper Jurassic Morrison Formation has produced surprisingly few examples of dinosaur eggs associated with embryonic remains. Even more puzzling, none of these seem to pertain to the theropod Allosaurus, one of the most common and best-understood dinosaur taxa in the formation. Here we
report on a dinosaur nest site that has produced both abundant prismatoolithid eggshell and embryonic (or perinatal) bones of Allosaurus from Fox Mesa, Wyoming. This represents the first such discovery for any theropod in the Jurassic of North America. The nest is heavily weathered but contains a few ellipsoid eggshell clusters that suggest an egg size of about 8 x 6.5 cm. Study of the eggshell morphology and microstructure confirms that a single egg type is present throughout, which is indistinguishable from Prismatoolithus coloradensis. All of the identifiable embryonic materials pertain to theropods, and two premaxillae specimens show the five alveoli diagnostic for Allosaurus among Morrison theropods. This confirms the theropod origin of Prismatoolithus eggs and implicates Allosaurus as the specific Morrison parent taxon. As a result, it is now possible to assign several previous discoveries of dinosaur eggs and potential nests to Allosaurus, including the isolated egg from the Cleveland-Lloyd Quarry. This discovery
also calls into question prior assignments of Prismatoolithus eggs to ornithopods, and suggests that more detailed study of such sites is warranted. Prismatoolithus eggshells are also associated with the Upper Jurassic theropod Lourinhanosaurus from Portugal, along with larger embryos that exhibit four premaxillary alveoli.

Despite more than a century of collecting, resulting in one of the best-studied vertebrate fossil records anywhere in the world, the Upper Jurassic Morrison Formation has produced surprisingly few examples of dinosaur eggs associated with embryonic remains. Even more puzzling, none of these seem to pertain to the theropod Allosaurus, one of the most common and best-understood dinosaur taxa in the formation. Here we report on a dinosaur nest site that has produced both abundant prismatoolithid eggshell and embryonic (or perinatal) bones of Allosaurus from Fox Mesa, Wyoming. This represents the first such discovery for any theropod in the Jurassic of North America. The nest is heavily weathered but contains a few ellipsoid eggshell clusters that suggest an egg size of about 8 x 6.5 cm. Study of the eggshell morphology and microstructure confirms that a single egg type is present throughout, which is indistinguishable from Prismatoolithus coloradensis. All of the identifiable embryonic materials pertain to theropods, and two premaxillae specimens show the five alveoli diagnostic for Allosaurus among Morrison theropods. This confirms the theropod origin of Prismatoolithus eggs and implicates Allosaurus as the specific Morrison parent taxon. As a result, it is now possible to assign several previous discoveries of dinosaur eggs and potential nests to Allosaurus, including the isolated egg from the Cleveland-Lloyd Quarry. This discovery also calls into question prior assignments of Prismatoolithus eggs to ornithopods, and suggests that more detailed study of such sites is warranted. Prismatoolithus eggshells are also associated with the Upper Jurassic theropod Lourinhanosaurus from Portugal, along with larger embryos that exhibit four premaxillary alveoli.

The non-avian saurischians that have associated eggshells and embryos are represented only by the sauropodomorph Massospondylus and Coelurosauria (derived theropods), thus missing the basal theropod representatives. We report a dinosaur clutch containing several crushed eggs and embryonic material ascribed to the megalosaurid theropod Torvosaurus. It represents the first associated eggshells and embryos of megalosauroids, thus filling an important phylogenetic gap between two distantly related groups of saurischians. These fossils represent the only unequivocal basal theropod embryos found to date. The assemblage was found in early Tithonian fluvial overbank deposits of the Lourinhã Formation in West Portugal. The morphological, microstructural and chemical characterization results of the eggshell fragments indicate very mild diagenesis. Furthermore, these fossils allow unambiguous association of basal theropod osteology with a specific and unique new eggshell morphology.

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Clavicles and interclavicles are plesiomorphically present in Reptilia. However, several groups show reduction or even loss of these elements. Crocodylimorpha, e.g., lost the clavicles, whereas dinosaurs are generally interpreted to only preserve the clavicles, the theropod furcula representing an unique case of fused clavicles. In sauropods, reports of clavicles are relatively frequent in non-titanosauriforms. These elements are elongated, curved, and rather stout bones with a spatulate and a bifurcate end. However, they were always found as single bones, and differ from the relatively short and unbifurcated clavicles found articulated with the scapulae of basal sauropodomorphs.
Elements from the Howe Quarry (Late Jurassic; Wyoming, USA) shed new light on these interpretations. Besides the elongated, curved bones (herein named morphotype A), also pairs of symmetric, L-shaped bones were recovered (morphotype B), associated with diplodocid dorsal and cervical vertebrae. Elements resembling morphotype B - articulated between the scapulae - have recently been reported from a diplodocid found near Tensleep, Wyoming. Taphonomic evidence, as well as the fact that they were preserved in symmetrical pairs, therefore implies that morphotype B represents the true sauropod clavicles.
Contrary to earlier reports, morphotype A elements from the Howe Quarry, as well as of previously reported specimens show a symmetry plane following the long axis of the elements. It is thus possible that the morphotype A elements were single bones from the body midline. The only such element present in the pectoral girdle of tetrapods are the interclavicle and the furcula. Comparison with crocodilian and lacertiform interclavicles indicates that the bifurcate end of the sauropod elements might represent the reduced transverse processes of the anterior end, and the spatulate end would have covered the coracoids or sternal plates ventrally.
The presence of both clavicles and interclavicles in the pectoral girdle stiffens the anterior trunk, and enhances considerably its stability. Such an enforcement might have been needed in diplodocids due to the strong lateral forces induced to the fore-limbs by the posteriorly placed center of mass (due to shorter fore- than hind-limbs), as well as lateral movements of the enormously elongated necks and tails. The absence of clavicles and interclavicles in titanosauriforms coincides with the development of wide-gauge locomotion style.
The presence of interclavicles in sauropods supports the recently proposed homology of the furcula with the interclavicle, instead of representing fused clavicles. Interclavicles were thus not lost, but may have remained cartilaginous or have yet to be found in basal dinosauriforms.

Clavicles and interclavicles are plesiomorphically present in Reptilia. However, several groups show reduction or even loss of these elements. Crocodylimorpha, e.g., lost the clavicles, whereas dinosaurs are generally interpreted to only preserve the clavicles, the theropod furcula representing an unique case of fused clavicles. In sauropods, reports of clavicles are relatively frequent in non-titanosauriforms. These elements are elongated, curved, and rather stout bones with a spatulate and a bifurcate end. However, they were always found as single bones, and differ from the relatively short and unbifurcated clavicles found articulated with the scapulae of basal sauropodomorphs. Elements from the Howe Quarry (Late Jurassic; Wyoming, USA) shed new light on these interpretations. Besides the elongated, curved bones (herein named morphotype A), also pairs of symmetric, L-shaped bones were recovered (morphotype B), associated with diplodocid dorsal and cervical vertebrae. Elements resembling morphotype B - articulated between the scapulae - have recently been reported from a diplodocid found near Tensleep, Wyoming. Taphonomic evidence, as well as the fact that they were preserved in symmetrical pairs, therefore implies that morphotype B represents the true sauropod clavicles. Contrary to earlier reports, morphotype A elements from the Howe Quarry, as well as of previously reported specimens show a symmetry plane following the long axis of the elements. It is thus possible that the morphotype A elements were single bones from the body midline. The only such element present in the pectoral girdle of tetrapods are the interclavicle and the furcula. Comparison with crocodilian and lacertiform interclavicles indicates that the bifurcate end of the sauropod elements might represent the reduced transverse processes of the anterior end, and the spatulate end would have covered the coracoids or sternal plates ventrally. The presence of both clavicles and interclavicles in the pectoral girdle stiffens the anterior trunk, and enhances considerably its stability. Such an enforcement might have been needed in diplodocids due to the strong lateral forces induced to the fore-limbs by the posteriorly placed center of mass (due to shorter fore- than hind-limbs), as well as lateral movements of the enormously elongated necks and tails. The absence of clavicles and interclavicles in titanosauriforms coincides with the development of wide-gauge locomotion style. The presence of interclavicles in sauropods supports the recently proposed homology of the furcula with the interclavicle, instead of representing fused clavicles. Interclavicles were thus not lost, but may have remained cartilaginous or have yet to be found in basal dinosauriforms.

The geochemical signatures of dinosaur eggshells represent well-established proxies in paleoenvironmental and paleobiological research. The variable sampling procedures reported in the literature, however, deserve attention. In order to evaluate the impact of different sampling methodologies on carbon and oxygen isotope and elemental concentrations, grinding was contrasted with drilling to extract powder samples from eggshell fragments collected at several locations. Eggshell data were further contrasted with surface materials, encasing matrix and compared with independent proxies using petrographic and elemental techniques. Iron and manganese elemental concentrations revealed an enrichment sequence depending on the sampling strategy for the same eggshell fragment. This pattern can be mistaken for a variable state of preservation. In contrast, carbon and oxygen isotope values exhibited only subtle differences and lacked clear trends. This suggests that isotope data are less susceptible to different methodological approaches. It is shown that drilling offers a wider range of possibilities compared to grinding (e.g., faster and less destructive). Additionally, drilled powder samples can confidently be used for elemental and isotope analysis, excluding contamination, thus providing a more accurate set of proxy data from eggshell archives.

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In this paper we present a method for estimating soil pCO2 in ancient environments using the measured carbon-isotope values of pedogenic carbonates and plant-derived organic matter. The validity of soil pCO2 estimates proves to be highly dependent on the organic δ13C values used in the calculations. Organic matter should be sourced from the same paleosol profiles as sampled carbonates to yield the most reliable estimates of soil pCO2. In order to demonstrate the potential use of soil pCO2 estimates in paleoecological and paleoenvironmental studies, we compare samples from three Upper Jurassic localities. Soil pCO2 estimates, interpreted as a qualitative indicator of primary paleoproductivity, are used to rank the Late Jurassic terrestrial environments represented by the Morrison Formation in western North America, the informally named Lourinhã formation in Western Europe, and the Stanleyville Group in Central Africa. Because modern terrestrial environments show a positive correlation between primary productivity and faunal richness, a similar relationship is expected in ancient ecosystems. When the relative paleoproductivity levels inferred for each study area are compared with estimates of dinosaur generic richness, a positive correlation emerges. Both the Morrison and Lourinhã formations have high inferred productivity levels and high estimated faunal richness. In contrast, the Stanleyville Group appears to have had low primary productivity and low faunal richness. Paleoclimatic data available for each study area indicate that both productivity and faunal richness are positively linked to water availability, as observed in modern terrestrial ecosystems.