Late Cretaceous ammonoids show that drivers of diversification are regionally heterogeneous.

Palaeontologists have long sought to explain the diversification of individual clades to whole biotas at global scales. Advances in our understanding of the spatial distribution of the fossil record through geological time, however, has demonstrated that global trends in biodiversity were a mosaic of regionally heterogeneous diversification processes. Drivers of diversification must presumably have also displayed regional variation to produce the spatial disparities observed in past taxonomic richness. Here, we analyse the fossil record of ammonoids, pelagic shelled cephalopods, through the Late Cretaceous, characterised by some palaeontologists as an interval of biotic decline prior to their total extinction at the Cretaceous-Paleogene boundary. We regionally subdivide this record to eliminate the impacts of spatial sampling biases and infer regional origination and extinction rates corrected for temporal sampling biases using Bayesian methods. We then model these rates using biotic and abiotic drivers commonly inferred to influence diversification. Ammonoid diversification dynamics and responses to this common set of diversity drivers were regionally heterogeneous, do not support ecological decline, and demonstrate that their global diversification signal is influenced by spatial disparities in sampling effort. These results call into question the feasibility of seeking drivers of diversity at global scales in the fossil record.

Chemical and palaeoentomological evidence of a relationship between early Eocene Belgian and Oise (France) ambers.

Of the early Eocene amber deposits known across the world, Belgian amber has been mostly absent from the relevant literature. We reinvestigated amber held in the palaeobotanical collection of the Royal Belgian Institute of Natural Sciences, Brussels, which derived from three localities in Belgium that originated from two geographical areas (Leval-Trahegnies and Orp-le-Grand). Using Fourier transform infrared (FTIR) spectroscopy we show the close chemical relationship of Belgian amber to the early Eocene Oise amber from the Paris Basin, and highlight the potential effect of weathering on the amber chemistry. The amber derives from a very similar botanical source as the Oise amber (Combretaceae or Leguminosae-Caesalpinioideae), but from different coeval basins. The two Leval-Trahegnies localities provided amber that exhibit different stages of weathering (heavily fissured and crazed, darkened) and lacking any inclusions. The Orp-le-Grand locality provided the least weathered amber, with one amber piece containing two inclusions: a mite and a new genus and species of hemipteran (Cativolcus uebruum gen. et sp. nov.), and a second one that preserved the impression of insect wings pressed into the surface.

Taxonomic review, palaeoecological, and palaeobiogeographical significances of Campanian Tethyan oysters from the North Eastern Desert, Egypt.

The Late Cretaceous was a time of high eustatic sea level that enabled extensive epicontinental seaways and carbonate platforms across the Tethyan Realm, providing favorable habitats for oyster communities to flourish. This study focuses on the Campanian Tethyan oysters from the North Eastern Desert of Egypt regarding taxonomy, palaeoecology, and palaeobiogeography. Three oyster species, Nicaisolopha nicaisei (Coquand, 1862), Pycnodonte (Phygraea) vesicularis (Lamarck, 1806), and Ambigostrea bretoni (Thomas and Peron, 1891), were identified from the Campanian succession in two studied sections. The sampled specimens of the genus Nicaisolopha have undergone a systematic palaeontological revision. As a result, N. tissoti (Thomas and Peron, 1891) is considered herein a junior synonym of N. nicaisei (Coquand, 1862). Palaeobiogeographically, the likely primary migration pattern of the studied oysters suggests an east-west trend along the Southern Tethys margin. All identified oysters in this study exhibit a Tethyan affinity and are primarily abundant in two main provinces: the Southern Tethys and the Western Tethys. The macrofaunal contents are categorized into two fossil associations: the Nicaisolopha nicaisei association of the middle-late Campanian age and the Pycnodonte vesicularis association of the late Campanian age. These macrofaunal associations indicate a deepening trend during the middle-late Campanian age, suggesting a transition from shallow inner neritic to middle neritic environments. Additionally, it is observed that Pycnodonteinae tend to grow larger under eutrophic conditions, low-energy environments, and nutrient-rich waters with high carbonate contents.

52,000 years of woolly rhinoceros population dynamics reveal extinction mechanisms.

The extinction of the woolly rhinoceros (Coelodonta antiquitatis) at the onset of the Holocene remains an enigma, with conflicting evidence regarding its cause and spatiotemporal dynamics. This partly reflects challenges in determining demographic responses of late Quaternary megafauna to climatic and anthropogenic causal drivers with available genetic and paleontological techniques. Here, we show that elucidating mechanisms of ancient extinctions can benefit from a detailed understanding of fine-scale metapopulation dynamics, operating over many millennia. Using an abundant fossil record, ancient DNA, and high-resolution simulation models, we untangle the ecological mechanisms and causal drivers that are likely to have been integral in the decline and later extinction of the woolly rhinoceros. Our 52,000-y reconstruction of distribution-wide metapopulation dynamics supports a pathway to extinction that began long before the Holocene, when the combination of cooling temperatures and low but sustained hunting by humans trapped woolly rhinoceroses in suboptimal habitats along the southern edge of their range. Modeling indicates that this ecological trap intensified after the end of the last ice age, preventing colonization of newly formed suitable habitats, weakening stabilizing metapopulation processes, triggering the extinction of the woolly rhinoceros in the early Holocene. Our findings suggest that fragmentation and resultant metapopulation dynamics should be explicitly considered in explanations of late Quaternary megafauna extinctions, sending a clarion call to the fragility of the remaining large-bodied grazers restricted to disjunct fragments of poor-quality habitat due to anthropogenic environmental change.

First evidence of marine turtle gastroliths in a fossil specimen: Paleobiological implications in comparison to modern analogues.

Semi-articulated remains of a large chelonioid turtle from the Turonian strata (Upper Cretaceous; ca. 93.9-89.8 Myr) near Sant'Anna d'Alfaedo (Verona province, northeastern Italy) are described for the first time. Together with the skeletal elements, the specimen also preserves pebbles inside the thoracic area which are lithologically distinct from the surrounding matrix. These allochthonous clasts are here interpreted as geo-gastroliths, in-life ingested stones that resided in the digestive tract of the animal. This interpretation marks the first reported evidence of geophagy in a fossil marine turtle. SEM-EDS analysis, together with macroscopic petrological characterization, confirm the presence of both siliceous and carbonatic pebbles. These putative geo-gastroliths have morphometries and size ranges more similar to those of gastroliths in different taxa (fossils and extant) than allochthonous "dropstone" clasts from the same deposit that were carried by floating vegetation A dense pitted pattern of superficial erosion is microscopically recognizable on the carbonatic gastroliths, consistent with surface etching due to gastric acids. The occurrence of a similar pattern was demonstrated by the experimental etching of carbonatic pebbles with synthetic gastric juice. Gut contents of modern green sea turtles (Chelonia mydas) were surveyed for substrate ingestion, providing direct evidence of geophagic behavior in extant chelonioids. Comparison with modern turtle dietary habits may suggests that the pebbles were ingested as a way to supplement calcium after or in preparation for egg deposition, implying that the studied specimen was possibly a gravid female.

Deep-water first occurrences of Ediacara biota prior to the Shuram carbon isotope excursion in the Wernecke Mountains, Yukon, Canada.

Ediacara-type macrofossils appear as early as ~575 Ma in deep-water facies of the Drook Formation of the Avalon Peninsula, Newfoundland, and the Nadaleen Formation of Yukon and Northwest Territories, Canada. Our ability to assess whether a deep-water origination of the Ediacara biota is a genuine reflection of evolutionary succession, an artifact of an incomplete stratigraphic record, or a bathymetrically controlled biotope is limited by a lack of geochronological constraints and detailed shelf-to-slope transects of Ediacaran continental margins. The Ediacaran Rackla Group of the Wernecke Mountains, NW Canada, represents an ideal shelf-to-slope depositional system to understand the spatiotemporal and environmental context of Ediacara-type organisms' stratigraphic occurrence. New sedimentological and paleontological data presented herein from the Wernecke Mountains establish a stratigraphic framework relating shelfal strata in the Goz/Corn Creek area to lower slope deposits in the Nadaleen River area. We report new discoveries of numerous Aspidella hold-fast discs, indicative of frondose Ediacara organisms, from deep-water slope deposits of the Nadaleen Formation stratigraphically below the Shuram carbon isotope excursion (CIE) in the Nadaleen River area. Such fossils are notably absent in coeval shallow-water strata in the Goz/Corn Creek region despite appropriate facies for potential preservation. The presence of pre-Shuram CIE Ediacara-type fossils occurring only in deep-water facies within a basin that has equivalent well-preserved shallow-water facies provides the first stratigraphic paleobiological support for a deep-water origination of the Ediacara biota. In contrast, new occurrences of Ediacara-type fossils (including juvenile fronds, Beltanelliformis, Aspidella, annulated tubes, and multiple ichnotaxa) are found above the Shuram CIE in both deep- and shallow-water deposits of the Blueflower Formation. Given existing age constraints on the Shuram CIE, it appears that Ediacaran organisms may have originated in the deeper ocean and lived there for up to ~15 million years before migrating into shelfal environments in the terminal Ediacaran. This indicates unique ecophysiological constraints likely shaped the initial habitat preference and later environmental expansion of the Ediacara biota.

Ultrastructure of the Jurassic serpulid tubes-phylogenetic and paleoecological implications.

The ultrastructural diversity of the Middle and Late Jurassic serpulid tubes from the Polish Basin has been investigated. The inspection of 12 taxa representing the two major serpulid clades allowed for the identification of three ultrastructure types-irregularly oriented prismatic structure (IOP), spherulitic prismatic structure (SPHP), and simple prismatic structure (SP). Six of the studied species are single-layered and six species possess two distinct layers. Ultrastructural diversity corresponds to certain serpulid clades. The members of Filograninae have single-layered tube walls composed of possibly plesiomorphic, irregularly oriented prismatic structure (IOP). Two-layered tubes occur solely within the clade Serpulinae, where the external, denser layer is built of either the ordered spherulitic (SPHP) or simple prismatic microstructure (SP), and the internal layer is composed of irregularly oriented prismatic structure (IOP). Apart from phylogenetic signals provided by the tube ultrastructure, it can be used in analyzing paleoecological aspects of tube-dwelling polychaetes. Compared to the more primitive, irregularly oriented microstructures of Filograninae, the regularly oriented microstructures of Serpulinae need a higher level of biological control over biomineralization. The advent of the dense outer protective layer (DOL) in serpulids, as well as the general increase in ultrastructure diversity, was likely a result of the evolutionary importance of the tubes for serpulids. Such diversity of the tube ultrastructural fabrics allowed for maximizing functionality by utilizing a variety of morphogenetic programs. The biomineralization system of serpulids remains more complex compared to other tube-dwelling polychaetes. Physiologically more expensive tube formation allows for mechanical strengthening of the tube by building robust, strongly ornamented tubes and firm attachment to the substrate. Contrary to sabellids, which perform a fugitive strategy, an increased tube durability allows serpulids a competitive advantage over other encrusters.

Laetoli, Tanzania: Extant terrestrial mollusc faunas shed new light on climate and palaeoecology at a Pliocene hominin site.

Laetoli, Tanzania is one of the most important palaeontological and palaeoanthropological localities in Africa. We report on a survey of the extant terrestrial gastropod faunas of the Laetoli-Endulen area, examine their ecological associations and re-examine the utility of Pliocene fossil molluscs in palaeoenvironmental reconstruction. Standardised collecting at 15 sites yielded 7302 individuals representing 58 mollusc species. Significant dissimilarities were found among the faunas of three broad habitat types: forest, woodland/bushland and open (grassland and scattered, xeric shrubland). Overall, more species were recorded in the woodland/bushland sites than in the forest sites. Open sites were less diverse. Environmental factors contributing most strongly to the separation of habitat types were aridity index and elevation. The results are supplemented with new mollusc data from the Mbulu Plateau south of Lake Eyasi, and compared to the list of species cumulatively recorded from the Ngorongoro area. Some regional variation is apparent and historical factors may explain the absence of some fossil taxa from Laetoli today. Differences in seasonality separated upland forest sites on the Mbulu plateau from those at Lemagurut at Laetoli. Indicator species were identified for each habitat. These included several large-bodied species analogous to the Laetoli Pliocene fossil species that were then used for palaeoenvironmental reconstruction. Based on the estimated aridity index, and adopting the widely used United Nations Environment Programme (UNEP) global climate classification, the four stratigraphic subunits of the Upper Laetolil Beds (3.6-3.85 Ma) would be placed in either the UNEP's Dry Sub-humid or Semi-arid climate classes, whereas the Upper Ndolanya Beds (2.66 Ma) and Lower Laetolil Beds (3.85-<4.36 Ma) would be assigned to the Humid and Semi-arid climate classes respectively. Pliocene precipitation at Laetoli is estimated as 847-965 mm per year, refining previous estimates. This is close or slightly higher than the present mean annual precipitation, and is likely to have corresponded to a mosaic of forest, woodland and bushland within a grassland matrix consistent with other reconstructions.

Paleoneurology of stem palaeognaths clarifies the plesiomorphic condition of the crown bird central nervous system.

Lithornithidae, an assemblage of volant Palaeogene fossil birds, provide our clearest insights into the early evolutionary history of Palaeognathae, the clade that today includes the flightless ratites and volant tinamous. The neotype specimen of Lithornis vulturinus, from the early Eocene (approximately 53 million years ago) of Europe, includes a partial neurocranium that has never been thoroughly investigated. Here, we describe these cranial remains including the nearly complete digital endocasts of the brain and bony labyrinth. The telencephalon of Lithornis is expanded and its optic lobes are ventrally shifted, as is typical for crown birds. The foramen magnum is positioned caudally, rather than flexed ventrally as in some crown birds, with the optic lobes, cerebellum, and foramen magnum shifted further ventrally. The overall brain shape is similar to that of tinamous, the only extant clade of flying palaeognaths, suggesting that several aspects of tinamou neuroanatomy may have been evolutionarily conserved since at least the early Cenozoic. The estimated ratio of the optic lobe's surface area relative to the total brain suggests a diurnal ecology. Lithornis may provide the clearest insights to date into the neuroanatomy of the ancestral crown bird, combining an ancestrally unflexed brain with a caudally oriented connection with the spinal cord, a moderately enlarged telencephalon, and ventrally shifted, enlarged optic lobes.

Exploring the effects of weighting against homoplasy in genealogies of palaeontological phylogenetic matrices.

Although simulations have shown that implied weighting (IW) outperforms equal weighting (EW) in phylogenetic parsimony analyses, weighting against homoplasy lacks extensive usage in palaeontology. Iterative modifications of several phylogenetic matrices in the last decades resulted in extensive genealogies of datasets that allow the evaluation of differences in the stability of results for alternative character weighting methods directly on empirical data. Each generation was compared against the most recent generation in each genealogy because it is assumed that it is the most comprehensive (higher sampling), revised (fewer misscorings) and complete (lower amount of missing data) matrix of the genealogy. The analyses were conducted on six different genealogies under EW and IW and extended implied weighting (EIW) with a range of concavity constant values (k) between 3 and 30. Pairwise comparisons between trees were conducted using Robinson-Foulds distances normalized by the total number of groups, distortion coefficient, subtree pruning and regrafting moves, and the proportional sum of group dissimilarities. The results consistently show that IW and EIW produce results more similar to those of the last dataset than EW in the vast majority of genealogies and for all comparative measures. This is significant because almost all of these matrices were originally analysed only under EW. Implied weighting and EIW do not outperform each other unambiguously. Euclidean distances based on a principal components analysis of the comparative measures show that different ranges of k-values retrieve the most similar results to the last generation in different genealogies. There is a significant positive linear correlation between the optimal k-values and the number of terminals of the last generations. This could be employed to inform about the range of k-values to be used in phylogenetic analyses based on matrix size but with the caveat that this emergent relationship still relies on a low sample size of genealogies.

Fate and preservation of the late pleistocene cave bears from Niedzwiedzia Cave in Poland, through taphonomy, pathology, and geochemistry.

This comprehensive study examines fossil remains from Niedzwiedzia Cave in the Eastern Sudetes, offering detailed insights into the palaeobiology and adversities encountered by the Pleistocene cave bear Ursus spelaeus ingressus. Emphasising habitual cave use for hibernation and a primarily herbivorous diet, the findings attribute mortality to resource scarcity during hibernation and habitat fragmentation amid climate shifts. Taphonomic analysis indicates that the cave was extensively used by successive generations of bears, virtually unexposed to the impact of predators. The study also reveals that alkaline conditions developed in the cave during the post-depositional taphonomic processes. Mortality patterns, notably among juveniles, imply dwindling resources, indicative of environmental instability. Skeletal examination reveals a high incidence of forelimb fractures, indicating risks during activities like digging or confrontations. Palaeopathological evidence unveils vulnerabilities to tuberculosis, abscesses, rickets, and injuries, elucidating mobility challenges. The cave's silts exhibit a high zinc concentration, potentially derived from successive bear generations consuming zinc-rich plants. This study illuminates the lives of late cave bears, elucidating unique environmental hurdles faced near their species' end.

Exploring the preservation of a parasitic trace in decapod crustaceans using finite elements analysis.

The fossil record of parasitism is poorly understood, due largely to the scarcity of strong fossil evidence of parasites. Understanding the preservation potential for fossil parasitic evidence is critical to contextualizing the fossil record of parasitism. Here, we present the first use of X-ray computed tomography (CT) scanning and finite elements analysis (FEA) to analyze the impact of a parasite-induced fossil trace on host preservation. Four fossil and three modern decapod crustacean specimens with branchial swellings attributed to an epicaridean isopod parasite were CT scanned and examined with FEA to assess differences in the magnitude and distribution of stress between normal and swollen branchial chambers. The results of the FEA show highly localized stress peaks in reaction to point forces, with higher peak stress on the swollen branchial chamber for nearly all specimens and different forces applied, suggesting a possible shape-related decrease in the preservation potential of these parasitic swellings. Broader application of these methods as well as advances in the application of 3D data analysis in paleontology are critical to understanding the fossil record of parasitism and other poorly represented fossil groups.

'Snakes and ladders' in paleoanthropology: From cognitive surprise to skillfulness a million years ago.

A paradigmatic account may suffice to explain behavioral evolution in early Homo. We propose a parsimonious account that (1) could explain a particular, frequently-encountered, archeological outcome of behavior in early Homo - namely, the fashioning of a Paleolithic stone 'handaxe' - from a biological theoretic perspective informed by the free energy principle (FEP); and that (2) regards instances of the outcome as postdictive or retrodictive, circumstantial corroboration. Our proposal considers humankind evolving as a self-organizing biological ecosystem at a geological time-scale. We offer a narrative treatment of this self-organization in terms of the FEP. Specifically, we indicate how 'cognitive surprises' could underwrite an evolving propensity in early Homo to express sporadic unorthodox or anomalous behavior. This co-evolutionary propensity has left us a legacy of Paleolithic artifacts that is reminiscent of a 'snakes and ladders' board game of appearances, disappearances, and reappearances of particular archeological traces of Paleolithic behavior. When detected in the Early and Middle Pleistocene record, anthropologists and archeologists often imagine evidence of unusual or novel behavior in terms of early humankind ascending the rungs of a figurative phylogenetic 'ladder' - as if these corresponded to progressive evolution of cognitive abilities that enabled incremental achievements of increasingly innovative technical prowess, culminating in the cognitive ascendancy of Homo sapiens. The conjecture overlooks a plausible likelihood that behavior by an individual who was atypical among her conspecifics could have been disregarded in a community of Hominina (for definition see Appendix 1) that failed to recognize, imagine, or articulate potential advantages of adopting hitherto unorthodox behavior. Such failure, as well as diverse fortuitous demographic accidents, would cause exceptional personal behavior to be ignored and hence unremembered. It could disappear by a pitfall, down a 'snake', as it were, in the figurative evolutionary board game; thereby causing a discontinuity in the evolution of human behavior that presents like an evolutionary puzzle. The puzzle discomforts some paleoanthropologists trained in the natural and life sciences. They often dismiss it, explaining it away with such self-justifying conjectures as that, maybe, separate paleospecies of Homo differentially possessed different cognitive abilities, which, supposedly, could account for the presence or absence in the Pleistocene archeological record of traces of this or that behavioral outcome or skill. We argue that an alternative perspective - that inherits from the FEP and an individual's 'active inference' about its surroundings and of its own responses - affords a prosaic, deflationary, and parsimonious way to account for appearances, disappearances, and reappearances of particular behavioral outcomes and skills of early humankind.

Preservation of proteins in the geosphere.

Deep-time protein preservation has attracted increasing interest and rapid research activity within the palaeobiological community in recent years, but there are several different viewpoints without a cohesive framework for the interpretation of these proteins. Therefore, despite this activity, crucial gaps exist in the understanding of how proteins are preserved in the geological record and we believe it is vital to arrive at a synthesis of the various taphonomic pathways in order to proceed forward with their elucidation. Here we take a critical look at the state of knowledge regarding deep-time protein preservation and argue for the necessity of a more nuanced approach to understanding the molecular taphonomy of proteins through the lens of diagenetic pathways. We also propound an initial framework with which to comprehend the chemical changes undergone by proteins via the concept of 'proteagen'.

A remarkable assemblage of petroglyphs and dinosaur footprints in Northeast Brazil.

The Serrote do Letreiro Site, found on the northwest periphery of the Sousa Basin, Brazil, presents a remarkable convergence of paleontological and archaeological elements. It is constituted of sub-horizontal "lajeiros", or rock outcrops, intermingled with endemic Caatinga vegetation. The three prominent outcrops feature fossilized footprints of theropod, sauropod, and iguanodontian dinosaurs from the Early Cretaceous Period. Adjacent to these dinosaur tracks, indigenous petroglyphs adorn the surface. The petroglyphs, mainly characterized by circular motifs, maintain a striking resemblance to other petroglyphs found in the states of Paraíba and Rio Grande do Norte. This study primarily endeavors to delineate the site's major characteristics while concentrating on the relationship between the dinosaur footprints and the petroglyphs. It concurrently assesses the preservation status of this invaluable record, shedding light on its implications for the realms of paleontology, archaeology, and cultural heritage studies.

Sex estimation of the adult Neandertal Regourdou 1 (Montignac, France): Implications for sexing human fossil remains.

Sex is a biological trait fundamental to the study of hominin fossils. Among the many questions that can be addressed are those related to taxonomy, biological variability, sexual dimorphism, paleoobstetrics, funerary selection, and paleodemography. While new methodologies such as paleogenomics or paleoproteomics can be used to determine sex, they have not been systematically applied to Pleistocene human remains due to their destructive nature. Therefore, we estimated sex from the coxal bone of the newly discovered pelvic remains of the Regourdou 1 Neandertal (Southwest France, MIS 5) based on morphological and metric data employing two methods that have been recently revised and shown to be reliable in multiple studies. Both methods calculate posterior probabilities of the estimate. The right coxal bone of Regourdou 1 was partially reconstructed providing additional traits for sex estimation. These methods were cross validated on 14 sufficiently preserved coxal bones of specimens from the Neandertal lineage. Our results show that the Regourdou 1 individual, whose postcranial skeleton is not robust, is a male, and that previous sex attributions of comparative Neandertal specimens are largely in agreement with those obtained here. Our results encourage additional morphological research of fossil hominins in order to develop a set of methods that are applicable, reliable, and reproducible.