The utility of body size as a functional trait to link the past and present in a diverse reptile clade.

Understanding the relationships between functional traits and environment is increasingly important for assessing ecosystem health and forecasting biotic responses to future environmental change. Taxon-free analyses of functional traits (ecometrics) allow for testing the performance of such traits through time, utilizing both the fossil record and paleoenvironmental proxies. Here, we test the role of body size as a functional trait with respect to climate, using turtles as a model system. We examine the influence of mass-specific metabolic rate as a functional factor in the sorting of body size with environmental temperature and investigate the utility of community body size composition as an ecometric correlated to climate variables. We then apply our results to the fossil record of the Plio-Pleistocene Shungura Formation in Ethiopia. Results show that turtle body sizes scale with mass-specific metabolic rate for larger taxa, but not for the majority of species, indicating that metabolism is not a primary driver of size. Body size ecometrics have stronger predictive power at continental than at global scales, but without a single, dominant predictive functional relationship. Application of ecometrics to the Shungura fossil record suggests that turtle paleocommunity ecometrics coarsely track independent paleoclimate estimates at local scales. We hypothesize that both human disruption and biotic interactions limit the ecometric fit of size to climate in this clade. Nonetheless, examination of the consistency of trait-environment relationships through deep and shallow time provides a means for testing anthropogenic influences on ecosystems.

Improving the relevance of paleontology to climate change policy.

Paleontology has provided invaluable basic knowledge on the history of life on Earth. The discipline can also provide substantial knowledge to societal challenges such as climate change. The long-term perspective of climate change impacts on natural systems is both a unique selling point and a major obstacle to becoming more pertinent for policy-relevant bodies like the Intergovernmental Panel on Climate Change (IPCC). Repeated experiments on the impacts of climate change without anthropogenic disturbance facilitate the extraction of climate triggers in biodiversity changes. At the same time, the long timescales over which paleontological changes are usually assessed are beyond the scope of policymakers. Based on first-hand experience with the IPCC and a quantitative analysis of its cited literature, we argue that the differences in temporal scope are less of an issue than inappropriate framing and reporting of most paleontological publications. Accepting that some obstacles will remain, paleontology can quickly improve its relevance by targeting climate change impacts more directly and focusing on effect sizes and relevance for projections, particularly on higher-end climate change scenarios.

Insect herbivory within modern forests is greater than fossil localities.

Fossilized leaves provide the longest running record of hyperdiverse plant-insect herbivore associations. Reconstructions of these relationships over deep time indicate strong links between environmental conditions, herbivore diversity, and feeding damage on leaves. However, herbivory has not been compared between the past and the modern era, which is characterized by intense anthropogenic environmental change. Here, we present estimates for damage frequencies and diversities on fossil leaves from the Late Cretaceous (66.8 Ma) through the Pleistocene (2.06 Ma) and compare these estimates with Recent (post-1955) leaves collected via paleobotanical methods from modern ecosystems: Harvard Forest, United States; the Smithsonian Environmental Research Center, United States; and La Selva, Costa Rica. Total damage frequency, measured as the percentage of leaves with any herbivore damage, within modern ecosystems is greater than any fossil locality within this record. This pattern is driven by increased frequencies across nearly all functional feeding groups within the Recent. Diversities of total, specialized, and mining damage types are elevated within the Recent compared with fossil floras. Our results demonstrate that plants in the modern era are experiencing unprecedented levels of insect damage, despite widespread insect declines. Human influence, such as the rate of global climate warming, influencing insect feeding and timing of life cycle processes along with urbanization and the introduction of invasive plant and insect species may drive elevated herbivory. This research suggests that the strength of human influence on plant-insect interactions is not controlled by climate change alone but rather, the way in which humans interact with terrestrial landscape.

Functional traits of fossil plants.

A minuscule fraction of the Earth's paleobiological diversity is preserved in the geological record as fossils. What plant remnants have withstood taphonomic filtering, fragmentation, and alteration in their journey to become part of the fossil record provide unique information on how plants functioned in paleo-ecosystems through their traits. Plant traits are measurable morphological, anatomical, physiological, biochemical, or phenological characteristics that potentially affect their environment and fitness. Here, we review the rich literature of paleobotany, through the lens of contemporary trait-based ecology, to evaluate which well-established extant plant traits hold the greatest promise for application to fossils. In particular, we focus on fossil plant functional traits, those measurable properties of leaf, stem, reproductive, or whole plant fossils that offer insights into the functioning of the plant when alive. The limitations of a trait-based approach in paleobotany are considerable. However, in our critical assessment of over 30 extant traits we present an initial, semi-quantitative ranking of 26 paleo-functional traits based on taphonomic and methodological criteria on the potential of those traits to impact Earth system processes, and for that impact to be quantifiable. We demonstrate how valuable inferences on paleo-ecosystem processes (pollination biology, herbivory), past nutrient cycles, paleobiogeography, paleo-demography (life history), and Earth system history can be derived through the application of paleo-functional traits to fossil plants.

Craniomandibular variation in the endemic Hispaniolan primate, Antillothrix bernensis.

Here we describe new fossil material of Antillothrix bernensis, a Pleistocene-Holocene primate taxon from Hispaniola. It is now represented by seven crania, five mandibles, and dozens of postcranial elements from several paleontologically rich cave systems. The five adult crania included here share a similar overall profile as well as specific features such as a deep depression at the glabella. The complete anterior dentition of Antillothrix can now be described for the first time; short canine crowns, in the apicobasal dimension, compare well with titi monkeys, but the new crania and mandibles lack the specialized tall-crowned incisors of the extant pitheciids. They do, however, have a diastema between the lateral maxillary incisors and canines, a feature not present in the previously known crania. The new mandibles deepen posteriorly and have a medial inflection of the mandibular ramus, as in some pitheciids, but also share with Xenothrix a significant vertical narrowing of the corpus under P4/M1 not observed among extant taxa. Two of the specimens, a cranium and a mandible that do not fit together, exhibit congenitally absent third molars-a rarity among extant, noncallitrichine taxa. There is an approximately 1-kg range in the estimated body mass among the full Antillothrix sample (from 2.4 to 3.4 kg), as well as a range of approximately 5 cm3 of endocranial volume (from 40 to 45 cm3). With these extended ranges from the new specimens, Antillothrix can no longer be described as a taxon with a brain size smaller than that expected for its body size. Neither of these ranges in the brain size or body size is large enough to indicate a substantial level of sexual dimorphism or to necessitate separating the sample into male and female individuals. Given this, and the similar canine sizes for all specimens where they are present, the sample is consistent with a morphologically variable but monomorphic species.

Selection by differential survival among marine animals in the Phanerozoic.

The Gaia hypothesis posits that the Earth and its biosphere function as a single self-stabilizing system, but a key challenge is explaining how this could have arisen through Darwinian evolution. One theory is that of "selection by differential survival," in which a clade's extinction probability decreases with age as it accumulates adaptations resisting environmental disturbances. While this is hard to assess during early Earth history, we can assess whether this process operated among marine animal genera throughout the Phanerozoic. To that end, we analyzed time ranges of 36,117 extinct animal genera using fossil occurrence data from the Paleobiology Database in order to calculate marine metazoan extinction age selectivity, extinction rates, and speciation rates over the Phanerozoic. We identify four signatures of selection by differential survival: lower extinction rates among older lineages, heritability and taxonomically nested propagation of extinction resistance, reduced age selectivity during rare environmental perturbations, and differential extinction rather than speciation as the primary driver of the phenomenon. Evidence for this process at lower taxonomic levels also implies its possibility for life as a whole - indeed, the possibility of Gaia.

Fossil dermal denticles reveal the preexploitation baseline of a Caribbean coral reef shark community.

Preexploitation shark baselines and the history of human impact on coral reef-associated shark communities in the Caribbean are tpoorly understood. We recovered shark dermal denticles from mid-Holocene (∼7 ky ago) and modern reef sediments in Bocas del Toro, Caribbean Panama, to reconstruct an empirical shark baseline before major human impact and to quantify how much the modern shark community in the region had shifted from this historical reference point. We found that denticle accumulation rates, a proxy for shark abundance, declined by 71% since the mid-Holocene. All denticle morphotypes, which reflect shark community composition, experienced significant losses, but those morphotypes found on fast-swimming, pelagic sharks (e.g., families Carcharhinidae and Sphyrnidae) declined the most. An analysis of historical records suggested that the steepest decline in shark abundance occurred in the late 20th century, coinciding with the advent of a targeted shark fishery in Panama. Although the disproportionate loss of denticles characterizing pelagic sharks was consistent with overfishing, the large reduction in denticles characterizing demersal species with low commercial value (i.e., the nurse shark Ginglymostoma cirratum) indicated that other stressors could have exacerbated these declines. We demonstrate that the denticle record can reveal changes in shark communities over long ecological timescales, helping to contextualize contemporary abundances and inform shark management and ecology.

Hominin locomotion and evolution in the Late Miocene to Late Pliocene.

In this review, we present on the evolution of the locomotor adaptation of hominins in the Late Miocene to Late Pliocene, with emphasis on some of the prominent advances and debates that have occurred over the past fifty years. We start with the challenging issue of defining hominin locomotor grades that are currently used liberally and offer our own working definitions of facultative, habitual, and obligate bipedalism. We then discuss the nature of the Pan-Homo last common ancestor and characterize the locomotor adaptation of Sahelanthropus, Orrorin, and Ardipithecus-often referred to as facultative bipeds-and examine the debates on the extent of bipedality and arboreality in these taxa. Moreover, the question of Middle Pliocene hominin locomotor diversity is addressed based on information derived from the 'Little Foot' specimen from Sterkfontein, footprints from Laetoli, and the Burtele Foot in Ethiopia. Our review suggests that the most convincing evidence for locomotor diversity comes from Burtele, whereas the evidence from Sterkfontein and Laetoli is unconvincing and equivocal, respectively. Finally, we address the decades old issue of the significance of arboreality in the otherwise habitual biped, Australopithecus, with emphasis on Australopithecus afarensis and its implications for the paleobiology of these creatures. We conclude that many of the apelike features encountered, mostly in the upper part of the Australopithecus skeleton, were retained for their significance in climbing. Approaches that have investigated character plasticity and those exploring internal bone structure have shown that the shoulder and limbs in Au. afarensis and Australopithecus africanus were involved in arboreal activities that are thought to be key for feeding, nesting, and predator avoidance. We conclude that many of the so-called retained ape-like features persisted due to stabilizing selection, that early hominins engaged in a considerable amount of arboreality even after Australopithecus had become a habitual biped, and arboreality only ceased to be an important component of hominin locomotor behavior after the emergence of Homo erectus.

Biased, Spasmodic, and Ridiculously Incomplete: Sequence Stratigraphy and the Emergence of a New Approach to Stratigraphic Complexity in Paleobiology, 1973-1995.

This paper examines the emergence of a new approach to stratigraphic complexity, first in geology and then, following its creative appropriation, in paleobiology. The approach was associated with a set of models that together transformed stratigraphic geology in the decades following 1970. These included the influential models of depositional sequences developed by Peter Vail and others at Exxon. Transposed into paleobiology, they gave researchers new resources for studying the incompleteness of the fossil record and for removing biases imposed by the processes of sedimentary accumulation. In addition, they helped reconfigure the cultural landscape of paleobiology, consolidating a growing emphasis on fieldwork and eroding the barrier that had been erected in the 1970s between "paleontology" and "paleobiology." This paper traces these developments, paying special attention to the simulation models of stratigraphic paleobiologist Steven Holland. It also considers how the integration of sequence and event stratigraphy and paleobiology has begun to influence long-running discussions of incompleteness and bias in the fossil record.

Resilient biotic response to long-term climate change in the Adriatic Sea.

Preserving adaptive capacities of coastal ecosystems, which are currently facing the ongoing climate warming and a multitude of other anthropogenic impacts, requires an understanding of long-term biotic dynamics in the context of major environmental shifts prior to human disturbances. We quantified responses of nearshore mollusk assemblages to long-term climate and sea-level changes using 223 samples (~71,300 specimens) retrieved from latest Quaternary sediment cores of the Adriatic coastal systems. These cores provide a rare chance to study coastal systems that existed during glacial lowstands. The fossil mollusk record indicates that nearshore assemblages of the penultimate interglacial (Late Pleistocene) shifted in their faunal composition during the subsequent ice age, and then reassembled again with the return of interglacial climate in the Holocene. These shifts point to a climate-driven habitat filtering modulated by dispersal processes. The resilient, rather than persistent or stochastic, response of the mollusk assemblages to long-term environmental changes over at least 125 thousand years highlights the historically unprecedented nature of the ongoing anthropogenic stressors (e.g., pollution, eutrophication, bottom trawling, and invasive species) that are currently shifting coastal regions into novel system states far outside the range of natural variability archived in the fossil record.

Global warming generates predictable extinctions of warm- and cold-water marine benthic invertebrates via thermal habitat loss.

Anthropogenic global warming is redistributing marine life and may threaten tropical benthic invertebrates with several potential extinction mechanisms. The net impact of climate change on geographical extinction risk nevertheless remains uncertain. Evidence of widespread climate-driven extinctions and of potentially unidentified mechanisms exists in the fossil record. We quantify organism extinction risk across thermal habitats, estimated by paleoclimate reconstructions, over the past 300 million years. Extinction patterns at seven known events of rapid global warming (hyperthermals) differ significantly from typical patterns, resembling those driven by global geometry under simulated global warming. As isotherms move poleward with warming, the interaction between the geometry of the globe and the temperature-latitude relationship causes an uneven loss of thermal habitat and a bimodal latitudinal distribution of extinctions. Genera with thermal optima warmer than ~21°C show raised extinction odds, while extinction odds continually increase for genera with optima below ~11°C. Genera preferring intermediate temperatures generally have no additional extinction risk during hyperthermals, except under extreme conditions as the end-Permian mass extinction. Widespread present-day climate-driven range shifts indicate that occupancy loss is already underway. Given the most-likely projections of modern warming, our model, validated by seven past hyperthermal events, indicates that sustained warming has the potential to annihilate cold-water habitat and its endemic species completely within centuries.

Evolution of static allometry and constraint on evolutionary allometry in a fossil stickleback.

Allometric scaling describes the relationship of trait size to body size within and among taxa. The slope of the population-level regression of trait size against body size (i.e. static allometry) is typically invariant among closely related populations and species. Such invariance is commonly interpreted to reflect a combination of developmental and selective constraints that delimit a phenotypic space into which evolution could proceed most easily. Thus, understanding how allometric relationships do eventually evolve is important to understanding phenotypic diversification. In a lineage of fossil Threespine Stickleback (Gasterosteus doryssus), we investigated the evolvability of static allometric slopes for nine traits (five armour and four non-armour) that evolved significant trait differences across 10 samples over 8500 years. The armour traits showed weak static allometric relationships and a mismatch between those slopes and observed evolution. This suggests that observed evolution in these traits was not constrained by relationships with body size, perhaps because prior, repeated adaptation to freshwater habitats by Threespine Stickleback had generated strong selection to break constraint. In contrast, for non-armour traits, we found stronger allometric relationships. Those allometric slopes did evolve on short time scales. However, those changes were small and fluctuating and the slopes remained strong predictors of the evolutionary trajectory of trait means over time (i.e. evolutionary allometry), supporting the hypothesis of allometry as constraint.

Macroevolutionary diversification rates show time dependency.

For centuries, biologists have been captivated by the vast disparity in species richness between different groups of organisms. Variation in diversity is widely attributed to differences between groups in how fast they speciate or go extinct. Such macroevolutionary rates have been estimated for thousands of groups and have been correlated with an incredible variety of organismal traits. Here we analyze a large collection of phylogenetic trees and fossil time series and describe a hidden generality among these seemingly idiosyncratic results: speciation and extinction rates follow a scaling law in which both depend on the age of the group in which they are measured, with the fastest rates in the youngest clades. Using a series of simulations and sensitivity analyses, we demonstrate that the time dependency is unlikely to be a result of simple statistical artifacts. As such, this time scaling is likely a genuine feature of the tree of life, hinting that the dynamics of biodiversity over deep time may be driven in part by surprisingly simple and general principles.

Evolutionary trend in dental size in fossil orangutans from the Pleistocene of Chongzuo, Guangxi, southern China.

More than 800 isolated teeth of fossil Pongo have been recovered from cave sites in the vicinity of Chongzuo in Guangxi, southern China, ranging from the Early to Late Pleistocene (2.0-0.1 Ma). These collections provide a unique regional window into the evolutionary history of orangutans over a two-million-year period at the northernmost extent of their former geographic range. Here we investigate the nature and timing of the evolutionary change in the dental size of fossil orangutans from Chongzuo. Fossil tooth size (mesiodistal length∗buccolingual breadth) was compared against an extant Pongo pygmaeus standard (n = 106 individuals). During the course of the Pleistocene, orangutans from southern China exhibited a progressive reduction in overall dental size. Early Pleistocene Pongo has cheek teeth with occlusal areas that are 38.1% larger than those of extant P. pygmaeus. Those from the Middle and Late Pleistocene are 25.2% and 18.9% larger, respectively. Previously, the size difference in dentition between the Early to Middle Pleistocene and Middle to Late Pleistocene samples was used to differentiate time-successive species of Pongo, namely Pongo weidenreichi and Pongo devosi. However, with access to larger samples and better representation of populations through time, the evidence in support of this taxonomic arrangement requires reconsideration. Diminution of the teeth now appears to be a gradual evolutionary transformation rather than a punctuated event. Moreover, the morphological features that distinguish the Chongzuo fossil orangutans from extant Pongo spp. remain uniform throughout the Pleistocene. Retaining P. weidenreichi and P. devosi as anagenetic species remains an option, but, given the current evidence, we consider it preferable to assign all of the fossil orangutans from Chongzuo to P. weidenreichi. Beyond resolving questions of alpha taxonomy, the study of fossil orangutan dental size provides a basis for estimating body mass, which has implications for interpreting the paleobiology of Pleistocene Pongo in southern China.

Reconstructing the ecology of a Cretaceous cockroach: destructive and high-resolution imaging of its micro sensory organs.

Animals highly depend on their sensory organs to detect information about their surrounding environment. Among animal sensory organs, those of insects have a notable ability to detect information despite their small size, which might be, therefore, one of the reasons for the evolutionary success of insects. However, insect sensory organs are seldom fossilized in sediments due to their small size and fragility. A potential solution for this problem is the study of exceptionally well-preserved fossil material from amber. Unfortunately, the resolution of existing non-destructive analysis is insufficient to observe details of these micro sensory organs even with amber preservation. Here, we focus on the analysis of the micro sensory organs of an extinct male cockroach (Huablattula hui Qiu et al., 2019) in Cretaceous amber by combining destructive and non-destructive methods. Compared to extant species inhabiting dark environments, H. hui has relatively large compound eyes, and all the antennal sensilla for detecting multimodal information observed here are fewer or smaller. The characteristics of these sensory organs support the diurnality of the bright habitats of H. hui in contrast to many extant cockroaches. Like extant male mantises, grooved basiconic type sensilla exist abundantly on the antenna of the fossilized specimen. The abundance of grooved basiconic sensilla in mantid males results from using sex pheromones, and therefore, H. hui may have likewise used mantis-like intersexual communication. These lines of evidence suggest that the ecology and behavior of Cretaceous cockroaches were more diverse than those of related extant species.

The evolution of the human trophic level during the Pleistocene.

The human trophic level (HTL) during the Pleistocene and its degree of variability serve, explicitly or tacitly, as the basis of many explanations for human evolution, behavior, and culture. Previous attempts to reconstruct the HTL have relied heavily on an analogy with recent hunter-gatherer groups' diets. In addition to technological differences, recent findings of substantial ecological differences between the Pleistocene and the Anthropocene cast doubt regarding that analogy's validity. Surprisingly little systematic evolution-guided evidence served to reconstruct HTL. Here, we reconstruct the HTL during the Pleistocene by reviewing evidence for the impact of the HTL on the biological, ecological, and behavioral systems derived from various existing studies. We adapt a paleobiological and paleoecological approach, including evidence from human physiology and genetics, archaeology, paleontology, and zoology, and identified 25 sources of evidence in total. The evidence shows that the trophic level of the Homo lineage that most probably led to modern humans evolved from a low base to a high, carnivorous position during the Pleistocene, beginning with Homo habilis and peaking in Homo erectus. A reversal of that trend appears in the Upper Paleolithic, strengthening in the Mesolithic/Epipaleolithic and Neolithic, and culminating with the advent of agriculture. We conclude that it is possible to reach a credible reconstruction of the HTL without relying on a simple analogy with recent hunter-gatherers' diets. The memory of an adaptation to a trophic level that is embedded in modern humans' biology in the form of genetics, metabolism, and morphology is a fruitful line of investigation of past HTLs, whose potential we have only started to explore.

Developmental bias, macroevolution, and the fossil record.

A fuller understanding of the role of developmental bias in shaping large-scale evolutionary patterns requires integrating bias (the probability distribution of variation accessible to an ancestral phenotype) with clade dynamics (the differential survival and production of species and evolutionary lineages). This synthesis could proceed as a two-way exchange between the developmental data available to neontologists and the strictly phenotypic but richly historical and dynamic data available to paleontologists. Analyses starting in extant populations could aim to predict macroevolution in the fossil record from observed developmental bias, while analyses starting in the fossil record, particularly the record of extant species and lineages, could aim to predict developmental bias from macroevolutionary patterns, including the broad range of extinct phenotypes. Analyses in multivariate morphospaces are especially effective when coupled with phylogeny, theoretical and developmental models, and diversity-disparity plots. This research program will also require assessing the "heritability" of an ancestral bias across phylogeny, and the tendency for bias change in strength and orientation over evolutionary time. Such analyses will help find a set of general rules for the macroevolutionary effects of developmental bias, including its impact on and interactions with the other intrinsic and extrinsic factors governing the movement, expansion, and contraction of clades in morphospace.

Endocranial morphology of the Brazilian Permian dicynodont Rastodon procurvidens (Therapsida: Anomodontia).

Dicynodontia is a major clade of terrestrial tetrapods that greatly diversified during the Permian and Triassic periods, reaching a worldwide distribution. In this study, the endocranial cavity of the Brazilian Permian dicynodont Rastodon procurvidens is described based on a digital endocast obtained using digital imaging (X-ray computed tomography) and 3D modeling. It was possible to reconstruct the brain, olfactory bulbs, inner ear, some neurovascular canals, cranial nerves, the nasal cavity, and the maxillary recesses. The endocast of R. procurvidens preserves a typical plesiomorphic morphology of non-mammaliaform therapsids, being predominantly tubular and displaying a relatively short and robust hindbrain. Encephalization quotients (EQs) were calculated for R. procurvidens, resulting in EQs of 0.09 ± 0.03 and 0.13 ± 0.05 (Jerison's EQ and Manger's EQ, respectively). Finally, some biological implications of the endocast morphology were inferred for R. procurvidens. Its inner ear is especially small, and its orientation implies a slightly downturned head posture in life. Furthermore, the presence of uncompressed maxillary recesses in R. procurvidens indicates a correlation between the enlargement of the recesses and the reduction of the tusks, also seen in other dicynodonts with reduced tusks.

Fossil Cercopithecidae from the Early Pliocene Sagantole Formation at Gona, Ethiopia.

The Early Pliocene Sagantole Fm. in the Gona Project area, Afar State, Ethiopia, is noted for discoveries of the early hominin Ardipithecus ramidus. A large series of fossil cercopithecid primates dated to between 4.8 and 4.3 Ma has also been collected from these sediments. In this paper, we use qualitative analysis and standard dental and postcranial measures to systematically describe the craniodental remains and tentatively allocate postcrania to taxa where we are able to. We then use these data to compare these specimens to fossil assemblages from contemporary sites, interpret their paleobiology, and discuss implications for the paleoecology of the Gona Sagantole Fm. We recognize three cercopithecid species in the Gona Sagantole Fm. Pliopapio alemui makes up approximately two-thirds of the identifiable specimens; nearly all of the rest are allocated to Kuseracolobus aramisi, and a single molar indicates the presence of a second, somewhat larger but morphologically distinct papionin. Among the Early Pliocene cercopithecids from Gona are also a number of postcranial elements. None of the postcranial remains are directly associated with any of the cranial material. Nonetheless, some of the distal humeri and proximal femora can be tentatively allocated to either Pl. alemui or K. aramisi based on a combination of size, as the latter is approximately 50% larger than the former, and morphology. If these assignments are correct, they suggest K. aramisi was primarily arboreal and similar to most extant colobines, whereas Pl. alemui was more mixed in its substrate use, being more terrestrially adapted than K. aramisi, but less so than extant Papio or Theropithecus. Thus, we interpret the predominance of Pl. alemui over K. aramisi is consistent with a somewhat more open environment at Gona than at Aramis.

The study of the lower limb entheses in the Neanderthal sample from El Sidrón (Asturias, Spain): How much musculoskeletal variability did Neanderthals accumulate?

Entheses have rarely been systematically studied in the field of human evolution. However, the investigation of their morphological variability (e.g., robusticity) could provide new insight into their evolutionary significance in the European Neanderthal populations. The aim of this work is to study the entheses and joint features of the lower limbs of El Sidrón Neanderthals (Spain; 49 ka), using standardized scoring methods developed on modern samples. Paleobiology, growth, and development of both juveniles and adults from El Sidrón are studied and compared with those of Krapina Neanderthals (Croatia, 130 ka) and extant humans. The morphological patterns of the gluteus maximus and vastus intermedius entheses in El Sidrón, Krapina, and modern humans differ from one another. Both Neanderthal groups show a definite enthesis design for the gluteus maximus, with little intrapopulation variability with respect to modern humans, who are characterized by a wider range of morphological variability. The gluteus maximus enthesis in the El Sidrón sample shows the osseous features of fibrous entheses, as in modern humans, whereas the Krapina sample shows the aspects of fibrocartilaginous ones. The morphology and anatomical pattern of this enthesis has already been established during growth in all three human groups. One of two and three of five adult femurs from El Sidrón and from Krapina, respectively, show the imprint of the vastus intermedius, which is absent among juveniles from those Neanderthal samples and in modern samples. The scant intrapopulation and the high interpopulation variability in the two Neanderthal samples is likely due to a long-term history of small, isolated populations with high levels of inbreeding, who also lived in different ecological conditions. The comparison of different anatomical entheseal patterns (fibrous vs. fibrocartilaginous) in the Neanderthals and modern humans provides additional elements in the discussion of their functional and genetic origin.