Thousands of coprolites have been collected from the Upper Miocene (Tortonian/Pannonian) sands of the Pécs-Danitzpuszta sand pit, one of the most important mixed Neogene vertebrate localities in Hungary. Although, the locality has been known for centuries, the coprolites have not been investigated in detail. We describe these fossils and explore their palaeoecological significance. Coprolites were examined with thin sections, X-ray powder diffraction and computed tomography. Their surface is mostly smooth, desiccation cracks were not observed. All coprolites mainly consist of apatite, indicating most probably a carnivorous diet. No evidence of herbivory has been found so far. The coprolites contain remains belonging to several groups: ostracod carapaces, bivalve shells, vertebrae from bony fishes, fish scales, as well as teeth. The CT scans show inhomogeneities in the matrix; however, in most cases they do not reliably indicate inclusions. The taphonomical features suggest rapid burial and most likely an aquatic origin. The spiral morphotype could be attributed to fishes with spiral intestinal valves. Teeth of reef-associated fish taxa indicate Badenian age for some specimens, though the coprolite assemblage might include older and younger specimens as well.

Thousands of coprolites have been collected from the Upper Miocene (Tortonian/Pannonian) sands of the Pécs-Danitzpuszta sand pit, one of the most important mixed Neogene vertebrate localities in Hungary. Although, the locality has been known for centuries, the coprolites have not been investigated in detail. We describe these fossils and explore their palaeoecological significance. Coprolites were examined with thin sections, X-ray powder diffraction and computed tomography. Their surface is mostly smooth, desiccation cracks were not observed. All coprolites mainly consist of apatite, indicating most probably a carnivorous diet. No evidence of herbivory has been found so far. The coprolites contain remains belonging to several groups: ostracod carapaces, bivalve shells, vertebrae from bony fishes, fish scales, as well as teeth. The CT scans show inhomogeneities in the matrix; however, in most cases they do not reliably indicate inclusions. The taphonomical features suggest rapid burial and most likely an aquatic origin. The spiral morphotype could be attributed to fishes with spiral intestinal valves. Teeth of reef-associated fish taxa indicate Badenian age for some specimens, though the coprolite assemblage might include older and younger specimens as well.

Dortokidae is an endemic family of stem-pleurodiran turtles, known exclusively from the Cretaceous and early Paleogene of Europe. Here we describe a new dortokid taxon from the Upper Cretaceous (Maastrichtian) Sînpetru Formation of the southern Haţeg Basin, Romania. The type material of Dortoka vremiri sp. nov. comprises a well-preserved carapace and a nearly complete plastron as well as the in situ right scapula and right pubis. Phylogenetic analyses performed to assess the position and interrelationships of Dortoka vremiri sp. nov. within Dortokidae as well as within the wider Pleurodira recovered the new taxon firmly nested within Dortokidae, and together with other dortokids, placed along the stem lineage of pleurodires. Our analysis provides evidence for allopatric speciation in Dortoka through the presence of two distinct lineages – an eastern and a western European lineage, respectively. A similar east/west disjunct distribution pattern has also been established previously for several vertebrate groups such as other turtles, dinosaurs and mammals, probably resulting from isolation due to the particular palaeogeographical setting of the Late Cretaceous European Archipelago. The phylogeny demonstrates local survival of Dortoka across the K-Pg boundary as the sister-taxon of D. vremiri is D. botanica from the uppermost Paleocene deposits of Romania and points to geographical selectivity, as the western lineage of Dortoka went extinct before the Paleogene. Osteology and novel taphonomical data imply a semi-terrestrial lifestyle for Kallokibotion bajazidi, a turtle occurring in coeval faunas with the aquatic and smaller-sized D. vremiri and most likely going extinct at the K-Pg boundary. Altogether, this pattern is consistent with selective extinction of terrestrial taxa previously observed for other continental vertebrate assemblages across the end-Cretaceous mass extinction with only two other examples from turtles. http://zoobank.org/urn:lsid:zoobank.org:pub:871AD436-9448-40C6-85A4-0213CBC04A29

The sand pit of Pécs-Danitzpuszta is one of the most important mixed Neogene vertebrate localities in Hungary, with older Miocene (Langhian – Serravallian) terrestrial and marine animal remains re-deposited into the upper Miocene (Tortonian) Lake Pannon sediments. Due to the mixed status of the Pécs-Danitzpuszta vertebrate assemblages, limited information is available about the former depositional environment and age of these fossils. However, the rare earth element (REE) composition of the fossil apatite can help to determine the stratigraphic origin and the early depositional environments of unprovenanced vertebrate remains. In this study, the REE composition of more than 60 vertebrate fossils, among them reworked and in situ remains from the Mecsek Mountain, were analysed. The results allowed separation between the Badenian, Sarmatian, and Pannonian aged fossils. The REE content and their distribution also helped us characterise the ancient depositional conditions of the investigated reworked fossils. Furthermore, several different types of vertebrate fossils were analysed, resulting in a large database that helped clarify the likely timings and mechanisms of REE uptake at this locality. Apparent variation in REE chemistry within bones (cortical vs cancellous) and teeth (enamel vs dentine) are best linked to fractionation along the REE series during their incorporation.

The sand pit of Pécs-Danitzpuszta is one of the most important mixed Neogene vertebrate localities in Hungary, with older Miocene (Langhian – Serravallian) terrestrial and marine animal remains re-deposited into the upper Miocene (Tortonian) Lake Pannon sediments. Due to the mixed status of the Pécs-Danitzpuszta vertebrate assemblages, limited information is available about the former depositional environment and age of these fossils. However, the rare earth element (REE) composition of the fossil apatite can help to determine the stratigraphic origin and the early depositional environments of unprovenanced vertebrate remains. In this study, the REE composition of more than 60 vertebrate fossils, among them reworked and in situ remains from the Mecsek Mountain, were analysed. The results allowed separation between the Badenian, Sarmatian, and Pannonian aged fossils. The REE content and their distribution also helped us characterise the ancient depositional conditions of the investigated reworked fossils. Furthermore, several different types of vertebrate fossils were analysed, resulting in a large database that helped clarify the likely timings and mechanisms of REE uptake at this locality. Apparent variation in REE chemistry within bones (cortical vs cancellous) and teeth (enamel vs dentine) are best linked to fractionation along the REE series during their incorporation.

Dortokidae is an endemic family of stem-pleurodiran turtles, known exclusively from the Cretaceous and early Paleogene of Europe. Here we describe a new dortokid taxon from the Upper Cretaceous (Maastrichtian) Sînpetru Formation of the southern Haţeg Basin, Romania. The type material of Dortoka vremiri sp. nov. comprises a well-preserved carapace and a nearly complete plastron as well as the in situ right scapula and right pubis. Phylogenetic analyses performed to assess the position and interrelationships of Dortoka vremiri sp. nov. within Dortokidae as well as within the wider Pleurodira recovered the new taxon firmly nested within Dortokidae, and together with other dortokids, placed along the stem lineage of pleurodires. Our analysis provides evidence for allopatric speciation in Dortoka through the presence of two distinct lineages – an eastern and a western European lineage, respectively. A similar east/west disjunct distribution pattern has also been established previously for several vertebrate groups such as other turtles, dinosaurs and mammals, probably resulting from isolation due to the particular palaeogeographical setting of the Late Cretaceous European Archipelago. The phylogeny demonstrates local survival of Dortoka across the K-Pg boundary as the sister-taxon of D. vremiri is D. botanica from the uppermost Paleocene deposits of Romania and points to geographical selectivity, as the western lineage of Dortoka went extinct before the Paleogene. Osteology and novel taphonomical data imply a semi-terrestrial lifestyle for Kallokibotion bajazidi, a turtle occurring in coeval faunas with the aquatic and smaller-sized D. vremiri and most likely going extinct at the K-Pg boundary. Altogether, this pattern is consistent with selective extinction of terrestrial taxa previously observed for other continental vertebrate assemblages across the end-Cretaceous mass extinction with only two other examples from turtles. http://zoobank.org/urn:lsid:zoobank.org:pub:871AD436-9448-40C6-85A4-0213CBC04A29

We report on sporadic fossil vertebrates from Brunei Darussalam (Borneo). Most of these isolated remains are reworked and derive from Penanjong Beach known for former coastal cliffs used to be rich in marine molluscs. Previously, the only vertebrate remains reported were shark teeth. With new material, the fish fauna is now represented by six shark and a single ray taxa, while remains of three turtle families were discovered (Trionychidae, Cheloniidae, Geoemydidae). This fauna is compared to nearby upper Miocene fossiliferous beds from where remains of cheloniid and trionychid turtles were unearthed. To assess the origin of the reworked remains, rare earth element (REE) concentrations of the fossil bioapatite were compared between the reworked and the geographically closest in-situ Ambug Hill fauna. The obtained trend in REE variations are identical, revealing similar REE uptake mechanism and early diagenetic conditions. With the additional fact that the succession outcropping at Ambug Hill cuts the coastline imply that the majority of the reworked fossils derive from the local Miocene beds. This is supported by Sr-isotope ages from reworked calcite bivalve shells ranging from 9.74 to 6.62 Ma. These turtle remains thus represent the hitherto known first Neogene fossil tetrapods from Borneo.

We report on sporadic fossil vertebrates from Brunei Darussalam (Borneo). Most of these isolated remains are reworked and derive from Penanjong Beach known for former coastal cliffs used to be rich in marine molluscs. Previously, the only vertebrate remains reported were shark teeth. With new material, the fish fauna is now represented by six shark and a single ray taxa, while remains of three turtle families were discovered (Trionychidae, Cheloniidae, Geoemydidae). This fauna is compared to nearby upper Miocene fossiliferous beds from where remains of cheloniid and trionychid turtles were unearthed. To assess the origin of the reworked remains, rare earth element (REE) concentrations of the fossil bioapatite were compared between the reworked and the geographically closest in-situ Ambug Hill fauna. The obtained trend in REE variations are identical, revealing similar REE uptake mechanism and early diagenetic conditions. With the additional fact that the succession outcropping at Ambug Hill cuts the coastline imply that the majority of the reworked fossils derive from the local Miocene beds. This is supported by Sr-isotope ages from reworked calcite bivalve shells ranging from 9.74 to 6.62 Ma. These turtle remains thus represent the hitherto known first Neogene fossil tetrapods from Borneo.