Institut Català de Paleontologia Miquel Crusafont

Transmarine distribution and gigantism in the Late Cretaceous North American crocodyliform Deinosuchus has been difficult to reconcile with consistently inferred phylogenetic relationships to alligatorids, an otherwise freshwater and smaller-bodied group. We present an expanded phylogeny with increased spatiotemporally coherence that reinterprets species of Deinosuchus as stem-group crocodylians together with further putative alligatoroids, Leidyosuchus canadensis and the European Diplocynodon spp. (closely related to North American Borealosuchus). The novel topology elucidates the evolution of osmoregulation in Crocodylia and its close relatives by inferring plesiomorphic saltwater tolerance for Deinosuchus and the crown-group as well as secondary loss already in stem-group alligatorids. Divergence of Alligatoroidea coincided with extreme mid-Cretaceous sea level highs and the distribution of Deinosuchus across the Western Interior Seaway can be best explained by marine dispersal. Phylogenetic body-length analysis using a head-width proxy reveals phyletic dwarfism early in alligatoroid evolution and a reasonable total length estimate for the most complete specimen of Deinosuchus riograndensis. We find that gigantism in crocodyliforms is correlated with high-productive extensive aquatic ecosystems in the present and the past.

Scandentians, commonly known as treeshrews (tree shrews), are the sister group of primates and colugos within the Euarchonta, a clade with an evolutionary history rooted in the Cretaceous (Roberts et al., 2011; Melin et al., 2016). Scandentians are, however, extremely rare in the fossil record (Ni and Qiu, 2012; Li and Ni, 2016) and, other than the Oligocene species Ptilocercus kylin Li and Ni, 2016, their earliest undoubted representatives are not older than 18 Ma (Li and Ni, 2016, and references therein).

Functional morphologists have long noted that skeletal adaptations in primate phalanges reflect locomotor behavior. While most studies have successfully used two‐dimensional measurements to quantify general features of phalanx shape, a whole‐bone three‐dimensional analysis may better capture more subtle aspects of phalanx morphology that have not been quantifiable but are functionally meaningful. Here, we compare linear measurement (LM) and weighted spherical harmonic/sliding semilandmark (SPHARM‐sliding) analyses of the manual third proximal phalanx (PP3) in extant hominoids (Homo, Pan, Gorilla, Pongo, Symphalangus, Hylobates; n = 292) and specimens attributed to Australopithecus afarensis (n = 2) and Homo neanderthalensis (n = 2). Morphological variation was summarized using principal component (PC) analysis. Differences between extant taxa were tested for using non‐parametric MANOVAs (LM) and Procrustes distance resampling (SPHARM‐sliding). Linear discriminant analyses (LDA) were performed using PC scores to assess whether the SPHARM‐sliding or LM analysis better predicts group memberships of extant and fossil specimens. In both analyses, PC1 separates taxa along a locomotor gradient, and all extant genera are significantly different from one another (p ≤ 0.01) aside from Pongo versus Symphalangus in the LM analysis (p = 0.053). Only the SPHARM‐sliding analysis found significant differences between taxa within each genus (p ≤ 0.04), and differences were even significant among Gorilla subspecies (p < 0.001). LDAs indicated that accuracy, separation effectiveness, and confidence were greater for the SPHARM‐sliding analysis in predicting group membership among extant specimens, as well as fossil memberships to an extant group. Overall, results demonstrate that whole‐bone, high‐density landmark analyses can highlight nuanced features of PP3 morphology and may serve better for making inferences about fossils.

Senèze has yielded two primate fossils, neither from the recent field work. A partial ulna can be identified as cf. Macaca sylvanus. A nearly complete female skull is the holotype of Dolichopithecus arvernensis Depéret, 1928, now included in Paradolichopithecus. Emended and differential diagnoses are provided for this genus and species for the first time, and the Senèze skull is described and briefly compared to those of other species of the genus. Dental and cranial measurements are provided, along with comparative data. The macaque and Paradolichopithecus are moderately to highly terrestrial, respectively, and they suggest a woodland to more open environment. Senèze may be the youngest known occurrence of Paradolichopithecus arvernensis. A comparison of previous analyses results in retention of a Paradolichopithecus-Macaca link and of Paradolichopithecus and Procynocephalus as distinct genera.

The Senèze paleontological site (Haute-Loire, France) is situated on the banks of a maar lake, in the Auvergne volcanic region. It was first reported by M. Boule in 1892, following the chance discovery of a proboscidean skeleton reported by local scholars. Pierre Philis, a local peasant, became interested in fossil mammals and collected them in the fields around the hamlet over more than 40 years, from the end of the nineteenth century until the start of World War II. He sold them to museum curators, especially to H. G. Stehlin (Naturhistorisches Musem Basel), C. Depéret (Faculté des Sciences de Lyon), C. Gaillard (Muséum d’Histoire naturelle de Lyon) and M. Boule (Muséum national d’Histoire naturelle, Paris). The resulting collections, including mounted skeletons exhibited in museums, made the site famous. New species (and genera) of ruminant artiodactyls, primate and carnivores were defined from Senèze, which became a biochronological reference site for the late Villafranchian. After the war, only a few short field campaigns (prospecting, survey, mapping and minimal excavation) were undertaken, and it became clear that renewed research was required to better understand this important site. The Franco-American fieldwork led by the authors began with a survey and request for excavation permits in 2000, followed by intensive mapping, prospection and excavation from 2001–2006. After a brief review of previous work at and about Senèze, each year of our research is summarized and illustrated. The succeeding chapters of this volume are also “previewed”.

Our fieldwork at Senèze from 2000–2006 was designed to place the fauna from this important site in a precise stratigraphic and geochronologic framework, as well as to seek new elements of known taxa and possibly additional taxa. The analyses reported in this volume led to the recognition of six new taxa: Dinofelis sp., Canis sp., cf. Hemitragus, two indeterminate bovids and an indeterminate hyaenid; Ovis claudiusguerini n. sp. was newly named for a specimen previously indicated as Ovis sp. Hystrix refossa and Hyaenidae indet. (both recovered in our excavations) are new taxa for the site, as is Bison (Eobison) sp., a specimen of which was collected by A. Consigny on the surface and presented to the team. Senèze is the type-locality for 13 mammalian taxa: three genera, seven species and three subspecies. Fifty-eight vertebrates have now been recorded for the site: 39 mammals (one lagomorph, four rodents, 12 carnivores, two primates, 16 cetartiodactyls, three perissodactyls and one proboscidean), 17 birds and two fishes; three additional rare equids, if accepted, would raise the number of vertebrates to 61 and of mammals to 42. The major mammalian and avian taxa in the assemblage are reviewed, especially in terms of their chronological implications. The faunal list is compared to those from other sites in the Massif Central and the nearby Saint-Vallier, resulting in Senèze being distinguished from all others as representing the MNQ 18 biochronological unit, for which it is considered to be a reference locality. The definition of that unit is discussed and clarified. Further comparison was made with selected MNQ 17–18 localities from Spain, Italy, the Netherlands and Germany. The Senèze fauna (other than Bison and perhaps some rare equids) is relatively homogeneous, with no evidence for a second assemblage as suggested by some previous authors. Its base age of ca. 2.20 Ma suggests this date for the beginning of MNQ 18, but the age of the end of this unit is still under discussion.

Three genera of Plio-Pleistocene monodactyl equines are recognized and distinguished by cranial proportions: Equus, Plesippus and Allohippus. The rich material of equid fossils from Senèze is not homogeneous. In addition to a few caballine teeth and limb bones (possibly cataloguing errors or intrusive specimens), there is evidence of at least two species. The bulk of the material may be referred to Allohippus senezensis senezensis, which by size and proportions is intermediate between A. senezensis guthi of La Puebla de Valverde (Spain) and A. senezensis mygdoniensis of Gerakarou (Greece). Its relatively short muzzle and deep, not very robust, metapodials are usually found in equids living in rather dry conditions. Two partial skeletons and a few other possibly associated hindlimb elements were recovered in the new excavations close to the bottom of the local sequence, thus ca 2.2 Ma. One upper cheek tooth and 16 limb bones belong to a very large Allohippus. There are moreover a few fossils larger than the average of A. stenonis vireti of Saint-Vallier (France) and a few others as small as ?Allohippus of Pyrgos (Greece). The affinities and ages of various Pliocene and Pleistocene equid species are discussed and illustrated.

The age of the Senèze mammalian fauna has been discussed since it was first reported in 1892. 40Ar/39Ar ages reported by Nomade et al. (2014) and recalculated here to agree with current standards placed the deposits between 2.20 and 2.07 Ma. Paleomagnetic data collected in 2001 and 2004 help to narrow the age range, especially of the levels yielding fossils in 2001–2006. In the western sector, fossils can be securely dated between 2.10 and 2.08 Ma, while in the southeastern sector, they are slightly older, between 2.20 and 2.18 Ma. Senèze is one of the few later Cenozoic European sites dated by both argon geochronometry and paleomagnetism, which makes these ages so precise. Experiments with ESR/U-series dating on teeth proved unsuccessful as a result of the early U-uptake and high natural dose rate in the sediments of Senèze.

The new fieldwork at Senèze ran from 2000 to 2006, with the goals of clarifying the age, stratigraphy and taphonomy of Senèze, as well as finding additional remains, especially of the less well-known taxa. Here we summarize the findings of each chapter and discuss their broader implications. Four geological chapters consider field methods, stratigraphy, volcanology and tephra mineralogy and dating. Following a chapter on palynology, two chapters discuss non-mammalian paleontology: ichthyology and ornithology. Eight chapters cover work on the fossil mammals. The chapter on biochronology places Senèze among other sites at the start of MNQ 18. Based on that work and the mammal chapters, it is possible to review the relative frequency of mammalian families in the total Senèze assemblage. Of some 2200 specimens, over half are cervids, with bovids, rhinocerotids and equids far behind. According to data from palynology and the habitat preferences of the more common mammals, the paleoenvironment around the Senèze maar would have included forest, woodland and grassland, perhaps in a warmer and moister climate than today. Taphonomic studies revealed that bones often rested a long time under water, lacked any indication of carnivore attack and often displayed pathologies in their joints. Combining these results and those from stratigraphy, it is proposed that most associated skeletons were preserved after large mammals fell into the lake and drowned without being disturbed.

Objectives: The functional interpretation of postcranial remains of Middle Miocene great apes from Europe (dryopithecines) suggests a combination of quadrupedalism and orthograde behaviors without modern analogs. We provide further insights based on an isolated dryopithecine talus (IPS85037) from the Middle Miocene (11.7Ma) Abocador de Can Mata locality ACM/C8-B* (Vallès-Penedès Basin, NE Iberian Peninsula), which represents the most complete one known to date. Material and Methods: We compare the specimen with an extant anthropoid sample (n=68) and the stem hominoid Ekembo heseloni (KMN RU 2036, ~18Ma, Kenya) using 3D geometric morphometrics. For the two fossil tali, we assess their phenetic affinities using a between-group principal components analysis (bgPCA), estimate body mass based on centroid size, and make locomotor inferences using a partial least-squares regression (PLSR) between talar shape and locomotor repertoire. Results: Its large inferred body mass (~38kg) and the possession of several modern hominoid-like features (albeit combined with more plesiomorphic traits) support the attribution of IPS85037 to a male dryopithecine. The bgPCA indicates that IPS85037 falls close to the extant hominoid variation and is less cercopithecoid-like than that of Ekembo, whose inferred locomotor repertoire is vastly dominated by quadrupedalism (81%). In contrast, the locomotor repertoire inferred from IPS85037 combines important quadrupedal (32%) and vertical climbing/clambering (50%) components with only moderate suspension (10%). Discussion: Our results align with previous inferences derived from other postcranial elements of Middle Miocene dryopithecines and, given their classification as crown hominoids, support the hypothesis that certain suspensory adaptations shared by extant hylobatids and hominids likely evolved independently.

Primates are often considered to have a poor sense of smell. While all studies identify small olfactory bulbs (OB; the region of the brain responsible for processing scent) among haplorhines, whether or not strepsirrhines also possess small OBs is less clear, as is the evolutionary backdrop from which these patterns emerged. Here, we examine the relative size of the olfactory bulbs in cranial endocasts of living and fossil primates and their kin (Euarchontoglires [Primates, Dermoptera, Scandentia, Rodentia, Lagomorpha]), testing previous hypotheses. Regression analyses of OB volume and mass relative to endocranial volume (ECV) and body mass (BM), and ANOVAS of residuals, were performed on a dataset of 181 extant and 41 extinct species. Analyses show clear differences in the relative size of the OBs, with haplorhines possessing distinctly smaller OBs relative to all other clades. Pairwise tests indicate haplorhine OBs are significantly smaller than those of all other clades, including strepsirrhines; when the haplorhines are removed from analyses, strepsirrhines are significantly smaller than all other clades. This suggests that a reduction in OB size occurred at the crown primate node, a pattern also seen in ancestral state reconstruction (ASR) analyses. The ASR analyses suggest multiple iterations of olfactory bulb size decrease occurred in Haplorhini, reflecting large amounts of parallelism. These results likely differ from previous studies due to the inclusion of additional fossils and more appropriate outgroups based on up‐to‐date phylogenetic hypotheses.

The collection of fossils supports a thriving market that is largely unknown to the academic community. The fossil market is characterised by fairs, online shops, physical stores and auction sales that attract hundreds of thousands of people each year. The sale of fossils to private individuals is often perceived as a threat by professional palaeontologists, who fear that valuable, maybe unique, specimens are being removed from scientific inquiry. In this paper we explore the global fossil market based on data shared online by international auction houses. We analysed the sales of fossils made by 115 auction houses over a period of 12 years (January 2010 to December 2022) whose auction results were still available online in 2022. Thirty parameters were collected for each auctioned item, including taxonomic identification, provenance, estimates and realised prices, etc. A total of 10.471 records were analysed. The database also includes high profile auction results, such as the Stan auction ( Tyrannosaurus rex sold by Christie's in 2020 for around 30 million US dollars). Even if many (54%) of the auctioned fossils come from Africa and Asia, only a small minority of the auction houses operates from these continents. These results are useful not only to learn more about the international fossil trade and to provide relevant information to auction houses, fossil preparators and sellers, but also to better assess the economic value of fossils from an insurance point of view, which is useful information for museum and university collections. They also provide a basis to address legal and policy issues of the private fossil trade.

Flying squirrels (Rodentia, Sciuridae, Sciurinae, Pteromyini) have a long and complex history in North America. First recorded during the Late Eocene, they vanished during the early Late Miocene (at about 9 Ma) only to re-appear in the Pliocene and Pleistocene. The first flying squirrels to be recorded after this Late Miocene gap are surprisingly attributed to the Eurasian genus of giant flying squirrel Miopetaurista. These are just two specimens from Florida that purportedly belong to Miopetaurista webbi, an endemic species. In this work we review these occurrences and further describe a new specimen from the Early Pliocene (latest Hemphillian or early Blancan) Gray Fossil Site in Tennessee, which may represent the oldest record of the genus in North America. We ascribe this new material to Miopetaurista webbi and find that this species is probably closely related to Miopetaurista thaleri, the only known Pliocene Eurasian species. The occurrence of Miopetaurista in eastern North America is puzzling, as it is distant from the known geographical range of the genus and of that of its sister taxon, the extant Petaurista. We hypothesize that Miopetaurista, which was linked to warm forested environments, dispersed into North America via the Bering Land Bridge during the warm phases of the Early Pliocene in the context of a major faunal dispersal event involving many other taxa. Later climatic cooling isolated these squirrels in warmer refuges, such as Florida, until they finally became extinct during the Early Pleistocene.

Revealing the evolutionary processes which resulted in the derived morphologies that characterize the Neanderthal clade has been an important task for paleoanthropologists. One critical method to quantify evolutionary changes in the morphology of hominin populations is through evaluating morphological phenotypic diversity (i.e., disparity) in phylogenetically informative bones as a close proxy to neutral evolutionary processes. The goal of this study is to quantify the degree of disparity in the Neanderthal clade. We hypothesize that a reduction in bony labyrinth disparity is indicative of the underlying genetic variation resulting from bottleneck events. We apply a deformation-based geometric morphometric approach to investigate semicircular canal and vestibule shape of a chronologically broad sample of individuals belonging to the Neanderthal lineage. Our results identify a significant reduction in disparity after the start of Marine Isotope Stage 5 supporting our hypothesis of a late bottleneck, possibly leading to the derived morphology of Late Pleistocene Neanderthals.

We describe a new species of Apus (Aves, Apodidae) from the Early Pliocene of Langebaanweg (Cape Province, South Africa). A single swift humerus was reported from Langebaanweg’s extraordinary rich fossil bird association, but it was never analyzed in detail. More recently, we located a second bone, a carpometacarpus, and completed the comparison of these two specimens with the corresponding bones of as many species of Apodidae as possible, leading to the description of the new species Apus boanoi sp. nov. The new species represents the largest species of the genus Apus known so far and the oldest from Africa; it represents an important calibration point for the split between Apus and Tachymarptis. The rarity of the new species in the very rich Langebaanweg fossil bird assemblage fits with the current paleoenvironmental reconstruction of a flat fluvial and coastal environment not suitable as breeding area for Apus swifts.

The analysis of incremental marks in the enamel, dentine and cementum of extant and extinct species provides important information about the rate and pattern of tooth growth, which permits inferences about key life history traits. Traditionally, such research has mainly focused on primates, while other mammalian groups have remained relatively unexplored. In some cases, this has led to the misidentification of incremental markings and the miscalculation of dental growth parameters in non‐primate taxa, which has highlighted the importance of obtaining more reliable comparative frameworks. Here, we partially fill this gap by providing a detailed analysis of the dental microstructure in the extant giraffe Giraffa camelopardalis . We specifically studied the histology of the different cusps (i.e. protoconid, metaconid, hypoconid, entoconid and hypoconulid) of two first lower molars and two third lower molars with different degree of wear to identify the different incremental markings and to calculate dental growth parameters such as daily secretion rate and enamel formation front angle for each cusp and tooth. Our results show that incremental markings in enamel were more apparent as compared to those in dentine and/or cementum and have permitted a deeper analysis of the former tissue. Enamel laminations, which had a daily periodicity, were the most common incremental lines in all teeth. Supradaily Retzius lines and subdaily cross‐striations and laminations were also recognised in dental enamel, revealing multiple secretory pulses of the ameloblasts in the giraffe. Generally, values of enamel growth parameters (i.e. daily secretion rate and enamel formation front angle) obtained for the first lower molar were comparable to those reported for closely related taxa, while those calculated for the third lower molar present a higher degree of variation that may be linked to differences in general somatic rates of growth. Nevertheless, enamel growth parameters were highly variable within each tooth, suggesting caution when making general (palaeo)biological inferences from dental histology. The giraffe dentine and cementum also register incremental lines. In the dentine, most of these features were classified as daily von Ebner's lines and their counting and measurement revealed values of secretion rates that agree with those previously reported in other artiodactyls. The age calculated from the incremental lines in the dental cementum matches that deduced from dental wear, suggesting that the counting of yearly lines in this tissue is a reliable tool to estimate individual age in giraffids. This study further suggests ways to refine future analyses of dentine and cementum and sets the stage for dental palaeohistology of extinct giraffids and closely related ungulates for which life history information is still unknown.

The small‐sized cervid Procervulus is considered as the most basal member of the Cervidae and one of the earliest ruminants bearing antler‐like appendages. The Iberian Miocene record of this stem‐cervid is extensively documented and largely overlaps with the Miocene Climatic Optimum (MCO), a transient period of global warming of particular interest when comparing present and near future conditions. Despite receiving a substantial amount of attention, histological studies on Procervulus are very scarce and only limited to postcranial remains of P rocervulus praelucidus from Germany (MN3). Here we focus for the first time on the dental histology of Procervulus ginsburgi from the Early Miocene Iberian site of Artesilla (MN4, 16.49 Ma), and examine its daily enamel secretion rate (DSR), enamel extension rate (EER) and crown formation time (CFT). Results reveal a brief CTF and high DSR and EER for P. ginsburgi and suggest a fast development at least early in its ontogeny. In addition, the pronounced growth rate of P. ginsburgi emerges as higher than that of the roe deer C. capreolus —documented as an r‐strategist and here examined as a possible extant analog. Overall, our findings point toward a fast life history strategy for P. ginsburgi , which unexpectedly contrasts with that of the 2 million‐year‐older P. praelucidus from Wintershof‐West, with a marked slower growth and maturation. When these results are analyzed together with other evidence, the somewhat drier and more open conditions of Artesilla as a result of the effects of the MCO seem to be the explanation for the different life history and ecology between these Procervulus species. More generally, this study illustrates that life histories within a single genus evolve in response not only to internal constraints but also to the environments, as predicted by the Life History Theory.

Ancient tooth enamel, and to some extent dentin and bone, contain characteristic peptides that persist for long periods of time. In particular, peptides from the enamel proteome (enamelome) have been used to reconstruct the phylogenetic relationships of fossil taxa. However, the enamelome is based on only about 10 genes, whose protein products undergo fragmentation in vivo and post mortem. This raises the question as to whether the enamelome alone provides enough information for reliable phylogenetic inference. We address these considerations on a selection of enamel-associated proteins that has been computationally predicted from genomic data from 232 primate species. We created multiple sequence alignments for each protein and estimated the evolutionary rate for each site. We examined which sites overlap with the parts of the protein sequences that are typically isolated from fossils. Based on this, we simulated ancient data with different degrees of sequence fragmentation, followed by phylogenetic analysis. We compared these trees to a reference species tree. Up to a degree of fragmentation that is similar to that of fossil samples from 1-2 million years ago, the phylogenetic placements of most nodes at family level are consistent with the reference species tree. We tested phylogenetic analysis on combinations of different enamel proteins and found that the composition of the proteome can influence deep splits in the phylogeny. With our methods, we provide guidance for researchers on how to evaluate the potential of paleoproteomics for phylogenetic studies before sampling valuable ancient specimens.

How populations adapt to their environment is a fundamental question in biology. Yet, we know surprisingly little about this process, especially for endangered species, such as nonhuman great apes. Chimpanzees, our closest living relatives, are particularly notable because they inhabit diverse habitats, from rainforest to woodland-savannah. Whether genetic adaptation facilitates such habitat diversity remains unknown, despite it having wide implications for evolutionary biology and conservation. By using newly sequenced exomes from 828 wild chimpanzees (388 postfiltering), we found evidence of fine-scale genetic adaptation to habitat, with signatures of positive selection in forest chimpanzees in the same genes underlying adaptation to malaria in humans. This work demonstrates the power of noninvasive samples to reveal genetic adaptations in endangered populations and highlights the importance of adaptive genetic diversity for chimpanzees.