Increase on environmental seasonality through the European Early Pleistocene inferred from dental enamel hypoplasia.

An in-depth study of the Early Pleistocene European remains of Hippopotamus has allowed the first detailed description of the incidence and types of dental alterations related to palaeopathologies and potentially linked to climatic and environmental factors. The results of a long-term qualitative and quantitative assessment highlight the importance of nutrient deficiencies on the development of dental enamel hypoplasia in Hippopotamus. Glacial cyclicity and the resulting changes in humidity and plant community structure conditioned the local environments critical for the survival of this taxon. Two main intervals of putative constrained nutritionally restrictions were detected at ca. 1.8 Ma and ca. 0.86 Ma (i.e., MIS63 and MIS21, respectively). Statistical comparisons show an increase in the frequency of dental hypoplasia between these two chronological periods, thus reinforcing the idea of increased seasonality in the circum-Mediterranean environments during the Early Pleistocene.

The earliest Ethiopian wolf: implications for the species evolution and its future survival.

In 2017, a hemimandible (MW5-B208), corresponding to the Ethiopian wolf (Canis simensis), was found in a stratigraphically-controlled and radio-isotopically-dated sequence of the Melka Wakena paleoanthropological site-complex, on the Southeastern Ethiopian Highlands, ~ 2300 m above sea level. The specimen is the first and unique Pleistocene fossil of this species. Our data provide an unambiguous minimum age of 1.6-1.4 Ma for the species' presence in Africa and constitutes the first empirical evidence that supports molecular interpretations. Currently, C. simensis is one of the most endangered carnivore species of Africa. Bioclimate niche modeling applied to the time frame indicated by the fossil suggests that the lineage of the Ethiopian wolf faced severe survival challenges in the past, with consecutive drastic geographic range contractions during warmer periods. These models help to describe future scenarios for the survival of the species. Projections ranging from most pessimistic to most optimistic future climatic scenarios indicate significant reduction of the already-deteriorating territories suitable for the Ethiopian Wolf, increasing the threat to the specie's future survival. Additionally, the recovery of the Melka Wakena fossil underscores the importance of work outside the East African Rift System in research of early human origins and associated biodiversity on the African continent.

The brain of the North American cheetah-like cat Miracinonyx trumani.

The cheetah Acinonyx jubatus, the fastest living land mammal, is an atypical member of the family Felidae. The extinct feline Miracinonyx trumani, known as the North American cheetah, is thought to have convergently evolved with Acinonyx to pursue fast and open-country prey across prairies and steppe environments of the North American Pleistocene. The brain of Acinonyx is unique among the living felids, but it is unknown whether the brain of the extinct M. trumani is convergent to that of Acinonyx. Here, we investigate the brain of M. trumani from a cranium endocast, using a comparative sample of other big cats. We demonstrate that the brain of M. trumani was different from that of the living A. jubatus. Indeed, its brain shows a unique combination of traits among living cats. This suggests that the case of extreme convergence between Miracinonyx and its living Old World vicar should be reconsidered.

Heterochronies and allometries in the evolution of the hominid cranium: a morphometric approach using classical anthropometric variables.

This article studies the evolutionary change of allometries in the relative size of the two main cranial modules (neurocranium and splanchnocranium) in the five living hominid species and a diverse sample of extinct hominins. We use six standard craniometric variables as proxies for the length, width and height of each cranial module. Factor analysis and two-block partial least squares (2B-PLS) show that the great apes and modern humans share a pervasive negative ontogenetic allometry in the neurocranium and a positive one in the splanchnocranium. This developmental constraint makes it possible to interpret the cranial heterochronies in terms of ontogenetic scaling processes (i.e., extensions or truncations of the ancestral ontogenetic trajectory) and lateral transpositions (i.e., parallel translations of the entire trajectory starting from a different shape for a given cranial size). We hypothesize that ontogenetic scaling is the main evolutionary modality in the australopithecines while in the species of Homo it is also necessary to apply transpositions. Both types of processes are coordinated in Homo, which result in an evolutionary trend toward an increase in brain size and in the degree of paedomorphosis from the earliest habilines.

Déjà vu: a reappraisal of the taphonomy of quarry VM4 of the Early Pleistocene site of Venta Micena (Baza Basin, SE Spain).

Venta Micena, an Early Pleistocene site of the Baza Basin (SE Spain), preserves a rich and diverse assemblage of large mammals. VM3, the main excavation quarry of the site, has been interpreted as a den of the giant hyaena Pachycrocuta brevirostris in the plain that surrounded the Baza palaeolake. Taphonomic analysis of VM3 has shown that the hyaenas scavenged the prey previously hunted by the hypercarnivores, transported their remains to the communal den, and consumed the skeletal parts according to their marrow contents and mineral density. In a recent paper (Luzón et al. in Sci Rep 11:13977, https://doi.org/10.1038/s41598-021-93261-1 , 2021), a small sample of remains unearthed from VM4, an excavation quarry ~ 350 m distant from VM3, is analysed. The authors indicate several differences in the taphonomic features of this assemblage with VM3, and even suggest that a different carnivore could have been the agent involved in the bone accumulation process. Here, we make a comparative analysis of both quarries and analyse more skeletal remains from VM4. Our results indicate that the assemblages are broadly similar in composition, except for slight differences in the frequency of megaherbivores, carnivores and equids according to NISP values (but not to MNI counts), the degree of bone weathering, and the intensity of bone processing by the hyaenas. Given that VM4 and VM3 were not coeval denning areas of P. brevirostris, these differences suggest that during the years when the skeletal remains were accumulated by the hyaenas at VM3, the rise of the water table of the Baza palaeolake that capped with limestone the bones was delayed compared to VM4, which resulted in their more in-depth consumption by the hyaenas.

The early hunting dog from Dmanisi with comments on the social behaviour in Canidae and hominins.

The renowned site of Dmanisi in Georgia, southern Caucasus (ca. 1.8 Ma) yielded the earliest direct evidence of hominin presence out of Africa. In this paper, we report on the first record of a large-sized canid from this site, namely dentognathic remains, referable to a young adult individual that displays hypercarnivorous features (e.g., the reduction of the m1 metaconid and entoconid) that allow us to include these specimens in the hypodigm of the late Early Pleistocene species Canis (Xenocyon) lycaonoides. Much fossil evidence suggests that this species was a cooperative pack-hunter that, unlike other large-sized canids, was capable of social care toward kin and non-kin members of its group. This rather derived hypercarnivorous canid, which has an East Asian origin, shows one of its earliest records at Dmanisi in the Caucasus, at the gates of Europe. Interestingly, its dispersal from Asia to Europe and Africa followed a parallel route to that of hominins, but in the opposite direction. Hominins and hunting dogs, both recorded in Dmanisi at the beginning of their dispersal across the Old World, are the only two Early Pleistocene mammal species with proved altruistic behaviour towards their group members, an issue discussed over more than one century in evolutionary biology.

The bears from Dmanisi and the first dispersal of early Homo out of Africa.

We report on the taxonomy and paleodiet of the bear population that inhabited the emblematic palaeoanthropological Early Pleistocene (1.8 Ma) site of Dmanisi (Georgia), based on a dual approach combining morphometrics and microwear of upper and lower teeth. Given that the teeth of Ursus etruscus Cuvier, 1823 from Dmanisi show considerable size variability, their systematic position has been debated. However, a comparative study of the coefficients of variation for tooth size measurements in several modern bear species shows that the variability in tooth size of the ursid population from Dmanisi could result from sexual dimorphism. The analysis of tooth microwear indicates that these bears inhabited a mixed environment of open plain with forest patches, where they had a browsing diet with a substantial contribution of meat and/or fish. Comparative tooth morphometric analyses of modern ursids and fossil U. etruscus indicate that this extinct species had an omnivorous behavior similar to that of extant brown bears. The ecological interactions of the Dmanisi bears with other members of the large mammals community, including the first hominins that dispersed out of Africa, are discussed in the light of this new evidence.

The earliest cut marks of Europe: a discussion on hominin subsistence patterns in the Orce sites (Baza basin, SE Spain).

Ancient evidence of human presence in Europe is recorded in several Early Pleistocene archaeopalaeontological sites from Spain, France and Italy. This is the case of Barranco León (BL) and Fuente Nueva-3 (FN-3), two localities placed near the town of Orce (depression of Baza and Guadix, SE Spain) and dated to ~1.4 Ma. At these sites, huge assemblages of Oldowan tools and evidence of defleshing, butchering and marrow processing of large mammal bones have been recovered together with a deciduous tooth of Homo sp. in the case of level BL-D. In this study, we: (i) describe in detail the anthropic marks found in the bone assemblages from these sites; (ii) analyse patterns of defleshment, butchery and marrow processing, based on the modifications identified in the cortical surface of the fossils; and (iii) discuss on the subsistence strategies of the first hominins that inhabited the European subcontinent during Early Pleistocene times.

Sixty-six million years along the road of mammalian ecomorphological specialization.

The fossil record of the large terrestrial mammals of the North American Cenozoic has previously been quantitatively summarized in six sequential episodes of faunal associations-"evolutionary faunas"-that correspond well with previously proposed qualitative "Chronofaunas." Here, we investigate the ecological spectrum of these faunas by classifying their major taxonomic components into discrete ecomorphological categories of diet, locomotion, and body size. To specifically address the potential influence of long-term climatic shifts on the ecomorphological composition of these faunas, we analyze via contingency tables and detrended correspondence analyses the frequency distribution of ecomorph types within faunas. We show that each evolutionary fauna has a unique, nonrandom association of ecomorphs, and we identify a long-term trend toward greater ecomorphological specialization over successive faunas during the past 66 My. Major vegetation shifts induced by climatic changes appear to underlie the ecomorphological dynamics of these six temporal associations that summarize Cenozoic North American mammalian evolutionary history.

Non-decoupled morphological evolution of the fore- and hindlimb of sabretooth predators.

Specialized organisms are useful for exploring the combined effects of selection of functional traits and developmental constraints on patterns of phenotypic integration. Sabretooth predators are one of the most interesting examples of specialization among mammals. Their hypertrophied, sabre-shaped upper canines and their powerfully built forelimbs have been interpreted as adaptations to a highly specialized predatory behaviour. Given that the elongated and laterally compressed canines of sabretooths were more vulnerable to fracture than the shorter canines of conical-tooth cats, it has been long hypothesized that the heavily muscled forelimbs of sabretooths were used for immobilizing prey before developing a quick and precise killing bite. However, the effect of this unique adaptation on the covariation between the fore- and the hindlimb has not been explored in a quantitative fashion. In this paper, we investigate if the specialization of sabretooth predators decoupled the morphological variation of their forelimb with respect to their hindlimb or, in contrast, both limbs vary in the same fashion as in conical-tooth cats, which do not show such extreme adaptations in their forelimb. We use 3D geometric morphometrics and different morphological indices to compare the fore- and hindlimb of conical- and sabretooth predators. Our results indicate that the limb bones of sabretooth predators covary following the same trend of conical-tooth cats. Therefore, we show that the predatory specialization of sabretooth predators did not result in a decoupling of the morphological evolution of their fore- and hindlimbs. The role of developmental constraints and natural selection on this coordinate variation between the fore- and the hindlimb is discussed in the light of this new evidence.

Neurocranium versus Face: A Morphometric Approach with Classical Anthropometric Variables for Characterizing Patterns of Cranial Integration in Extant Hominoids and Extinct Hominins.

The relative importance of the two main cranial complexes, the neurocranium and the splanchnocranium, has been examined in the five species of extant hominoids and in a huge sample of extinct hominins using six standard craniometric variables that measure the length, width and height of each cranial module. Factor analysis and two-block partial least squares were used for establishing the major patterns of developmental and evolutionary integration between both cranial modules. The results obtained show that all extant hominoids (including the anatomically modern humans) share a conserved pattern of developmental integration, a result that agrees with previous studies. The pattern of evolutionary integration between both cranial modules in australopiths runs in parallel to developmental integration. In contrast, the pattern of evolutionary and developmental integration of the species of the genus Homo is the opposite, which is probably the consequence of distinctive selective regimes for both hominin groups.

Patterns of morphological integration in the appendicular skeleton of mammalian carnivores.

We investigated patterns of evolutionary integration in the appendicular skeleton of mammalian carnivores. The findings are discussed in relation to performance selection in terms of organismal function as a potential mechanism underlying integration. Interspecific shape covariation was quantified by two-block partial least-squares (2B-PLS) analysis of 3D landmark data within a phylogenetic context. Specifically, we compared pairs of anatomically connected bones (within-limbs) and pairs of both serially homologous and functional equivalent bones (between-limbs). The statistical results of all the comparisons suggest that the carnivoran appendicular skeleton is highly integrated. Strikingly, the main shape covariation relates to bone robustness in all cases. A bootstrap test was used to compare the degree of integration between specialized cursorial taxa (i.e., those whose forelimbs are primarily involved in locomotion) and noncursorial species (i.e., those whose forelimbs are involved in more functions than their hindlimb) showed that cursors have a more integrated appendicular skeleton than noncursors. The findings demonstrate that natural selection can influence the pattern and degree of morphological integration by increasing the degree of bone shape covariation in parallel to ecological specialization.

A three-dimensional analysis of the morphological evolution and locomotor behaviour of the carnivoran hind limb.

BACKGROUND: The shape of the appendicular bones in mammals usually reflects adaptations towards different locomotor abilities. However, other aspects such as body size and phylogeny also play an important role in shaping bone design.We used 3D landmark-based geometric morphometrics to analyse the shape of the hind limb bones (i.e., femur, tibia, and pelvic girdle bones) of living and extinct terrestrial carnivorans (Mammalia, Carnivora) to quantitatively investigate the influence of body size, phylogeny, and locomotor behaviour in shaping the morphology of these bones. We also investigated the main patterns of morphological variation within a phylogenetic context. RESULTS: Size and phylogeny strongly influence the shape of the hind limb bones. In contrast, adaptations towards different modes of locomotion seem to have little influence. Principal Components Analysis and the study of phylomorphospaces suggest that the main source of variation in bone shape is a gradient of slenderness-robustness. CONCLUSION: The shape of the hind limb bones is strongly influenced by body size and phylogeny, but not to a similar degree by locomotor behaviour. The slender-robust "morphological bipolarity" found in bone shape variability is probably related to a trade-off between maintaining energetic efficiency and withstanding resistance to stresses. The balance involved in this trade-off impedes the evolution of high phenotypic variability. In fact, both morphological extremes (slender/robust) are adaptive in different selective contexts and lead to a convergence in shape among taxa with extremely different ecologies but with similar biomechanical demands. Strikingly, this "one-to-many mapping" pattern of evolution between morphology and ecology in hind limb bones is in complete contrast to the "many-to-one mapping" pattern found in the evolution of carnivoran skull shape. The results suggest that there are more constraints in the evolution of the shape of the appendicular skeleton than in that of skull shape because of the strong biomechanical constraints imposed by terrestrial locomotion.

On the relationships of postcanine tooth size with dietary quality and brain volume in primates: implications for hominin evolution.

Brain volume and cheek-tooth size have traditionally been considered as two traits that show opposite evolutionary trends during the evolution of Homo. As a result, differences in encephalization and molarization among hominins tend to be interpreted in paleobiological grounds, because both traits were presumably linked to the dietary quality of extinct species. Here we show that there is an essential difference between the genus Homo and the living primate species, because postcanine tooth size and brain volume are related to negative allometry in primates and show an inverse relationship in Homo. However, when size effects are removed, the negative relationship between encephalization and molarization holds only for platyrrhines and the genus Homo. In addition, there is no general trend for the relationship between postcanine tooth size and dietary quality among the living primates. If size and phylogeny effects are both removed, this relationship vanishes in many taxonomic groups. As a result, the suggestion that the presence of well-developed postcanine teeth in extinct hominins should be indicative of a poor-quality diet cannot be generalized to all extant and extinct primates.

A three-dimensional analysis of morphological evolution and locomotor performance of the carnivoran forelimb.

In this study, three-dimensional landmark-based methods of geometric morphometrics are used for estimating the influence of phylogeny, allometry and locomotor performance on forelimb shape in living and extinct carnivorans (Mammalia, Carnivora). The main objective is to investigate morphological convergences towards similar locomotor strategies in the shape of the major forelimb bones. Results indicate that both size and phylogeny have strong effects on the anatomy of all forelimb bones. In contrast, bone shape does not correlate in the living taxa with maximum running speed or daily movement distance, two proxies closely related to locomotor performance. A phylomorphospace approach showed that shape variation in forelimb bones mainly relates to changes in bone robustness. This indicates the presence of biomechanical constraints resulting from opposite demands for energetic efficiency in locomotion -which would require a slender forelimb- and resistance to stress -which would be satisfied by a robust forelimb-. Thus, we interpret that the need of maintaining a trade-off between both functional demands would limit shape variability in forelimb bones. Given that different situations can lead to one or another morphological solution, depending on the specific ecology of taxa, the evolution of forelimb morphology represents a remarkable "one-to-many mapping" case between anatomy and ecology.

The oldest human fossil in Europe, from Orce (Spain).

The Orce region has one of the best late Pliocene and early Pleistocene continental paleobiological records of Europe. It is situated in the northeastern sector of the intramontane Guadix-Baza Basin (Granada, Andalusia, southern Spain). Here we describe a new fossil hominin tooth from the site of Barranco León, dated between 1.02 and 1.73 Ma (millions of years ago) by Electron Spin Resonance (ESR), which, in combination with paleomagnetic and biochronologic data, is estimated to be close to 1.4 Ma. While the range of dates obtained from these various methods overlaps with those published for the Sima del Elefante hominin locality (1.2 Ma), the overwhelming majority of evidence points to an older age. Thus, at the moment, the Barranco León hominin is the oldest from Western Europe.

Cenozoic climate change influences mammalian evolutionary dynamics.

Global climate change is having profound impacts on the natural world. However, climate influence on faunal dynamics at macroevolutionary scales remains poorly understood. In this paper we investigate the influence of climate over deep time on the diversity patterns of Cenozoic North American mammals. We use factor analysis to identify temporally correlated assemblages of taxa, or major evolutionary faunas that we can then study in relation to climatic change over the past 65 million years. These taxa can be grouped into six consecutive faunal associations that show some correspondence with the qualitative mammalian chronofaunas of previous workers. We also show that the diversity pattern of most of these chronofaunas can be correlated with the stacked deep-sea benthic foraminiferal oxygen isotope (δ(18)O) curve, which strongly suggests climatic forcing of faunal dynamics over a large macroevolutionary timescale. This study demonstrates the profound influence of climate on the diversity patterns of North American terrestrial mammals over the Cenozoic.

Cranial shape transformation in the evolution of the giant panda (Ailuropoda melanoleuca).

In this study, landmark-based methods of geometric morphometrics are used for investigating the main aspects of cranial shape transformation in the evolution of the giant panda, Ailuropoda melanoleuca. Specifically, we explore if the highly derived cranial adaptations for bamboo feeding of the living panda were developed early in the panda's lineage. Results obtained show that the overall cranial morphologies of the oldest known panda, the "pygmy" Ailuropoda microta, and the late Pleistocene Ailuropoda baconi are both very similar to that of their closest living relative, A. melanoleuca, which agrees with a previous proposal based on qualitative criteria. However, we also describe several differences between the crania of A. microta, A. baconi, and A. melanoleuca, including the development of the postorbital process, the orientation of the occipital region, and the expansion of the braincase. As a result, the cranial morphology of A. microta shows a less specialized morphology toward a fibrous and durophagous diet compared to the giant panda. These results are confirmed by a comparative analysis of the dimensions of the upper teeth in bears, which has revealed differences in relative tooth size between A. microta and A. melanoleuca, most probably as a result of mosaic evolution. Therefore, we conclude that cranial shape did not remain essentially uniform in the Ailuropoda lineage, as previously thought, but underwent a number of changes during more than 2 Myr.