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Highlights The oldest human evidence in western Europe is currently documented at 1.4 Ma. Understanding the climatic variability that these hominins encountered is relevant. Paleoclimatic data are provided for an archaeopaleontological sequence. The Early Pleistocene hominins were able to cope with changing climatic conditions. Abstract The oldest evidence of a human presence in western Europe is currently documented at the late Early Pleistocene sites of Barranco León and Fuente Nueva 3 (Guadix-Baza Basin, SE Spain), dated respectively to 1.4 and 1.2 Ma. Understanding the climatic variability that these hominins encountered and coped with is of outstanding importance for placing their activities in an ecological context and understanding their capacity to adapt to changing conditions. Glacial-interglacial variability during this period was considerably less intense than during later phases of the Pleistocene. To date, however, no quantitative estimates are available for this climate variability in Early Pleistocene hominin sites, and no ‘cold’ reconstructions directly associated with the early hominin occupations of Europe have been performed either. Here stratigraphically constrained quantitative climatic reconstructions are provided for the sites of Barranco León and Fuente Nueva 3, using an improvement on the Mutual Ecogeographic Range method by projecting the niche envelope of the extant representatives ofOphisaurussp. We ascertain differences in temperature and rainfall between the different layers of the two sites, in line with previous reconstructions that revealed warm, humid periods (‘interglacial') as well as more temperate but drier periods (‘glacial'), consistent with Early Pleistocene climate cyclicity. Our new estimates suggest that late Early Pleistocene hominins, though conditioned to some degree by climatic factors, were able to deal with changing climatic and environmental conditions (‘interglacial' and ‘glacial') in the southwestern extremity of the European continent.

Venta Micena (Orce, Guadix-Baza Basin, Spain) is an Early Pleistocene locality renowned for the richness and quality of its palaeontological record. VM is spread over an area of 2.5 km2, where several exposed fossil outcrops are visible amidst its gorges and ravines. The best known of these sites, VM3, has been interpreted as a hyaena den. In addition, a new site, named VM4, has recently been the focus of fieldwork and taphonomic studies. The publication by Luz on et al. (2021) pointed out that VM4 presents a more complex history than VM3. First, two different sub-levels were identified: VM4-I and VM4-II. Secondly, the preliminary taphonomic analysis showed conspicuous differences with regard to VM3. Nevertheless, such interpretation has been challenged by Palmqvist et al. (2022) who proposed that VM3 and VM4 are both the result of a single depositional process, entailing the selective transport of skeletal parts by the giant extinct hyaena Pachycrocuta brevirostris back to its den. Using well-preserved faunal elements whose depositional context and provenance are reliable, in this paper we show that: 1) there are two clearly defined sub-levels in VM4 with some shared taphonomic characteristics as well as some notable differences; 2) VM3 and VM4 exhibit enough divergence to support differences in site formation processes; 3) The interpretation of both VM4-I and VM4-II is more consistent with their characterisation as open-air sites in which multiple agents and depositional processes contributed to its formation, rather than with hyaena dens. Nevertheless, excavations are still in progress at VM4 and therefore any results and interpretations ought to be considered as provisional., Consejeía de Cultura de la Junta de Andalucia through the General Research Project Primeras ocupaciones humanas y contexto paleoecologico a partir de los depositos Plio-pleistocenos de la cuenca Guadix-Baza BC.03.032/17, Inside the Artefacts & Ecofacts" Excellence Unit (University of Granada) HUM-607 PID 2021.125098NB.I00 MCIN/AEI/10.13039/501100011033/FEDER, Una manera de hacer Europa ProyExcel_00274 2021 SGR 01238, Agencia de Gestio D'Ajuts Universitaris de Recerca Agaur (AGAUR) Generalitat de Catalunya, National Health and Medical Research Council (NHMRC) of Australia PRE2020-094482 CEX2019-000945-M-20-1, Spanish Government, Archaeometrical Studies. Inside the Artefacts & Ecofacts" Excellence Unit (University of Granada

We know that the fossil record is incomplete. But how incomplete? Here we very coarsely estimate the completeness of the mammalian record in the Miocene, assuming that the duration of a mammalian species is about 1 Myr and the species diversity has stayed constant and is structurally comparable to the taxonomic diversity today. The overall completeness under these assumptions appears to be around 4%, but there are large differences across taxonomic groups. We find that the fossil record of proboscideans and perissodactyls as we know it for the Miocene must be close to complete, while we might know less than 15% of the species of artiodactyl or carnivore fossil species and only about 1% of primate species of the Miocene. The record of small mammals appears much less complete than that of large mammals.

Venta Micena is an area containing several palaeontological sites marking the beginning of the Calabrian stage (Early Pleistocene). The richness of the fossil accumulation including species of Asian, African and European origin, makes Venta Micena a key site for the the palaeoecological and palaeoenvironmental study of southern Europe during the Early Pleistocene. Thus, research has been focused on Venta Micena 3, which was originally interpreted as a single palaeosurface associated with a marshy context, in which most of the fauna was accumulated by Pachycrocuta brevirostris. Recent excavations have unearthed a new site, Venta Micena 4, located in the same stratigraphic unit (Unit C) and in close proximity to Venta Micena 3. Here we show the frst analyses regarding the taphonomic and spatial nature of this new site, defning two stratigraphic boundaries corresponding to two diferent depositional events. Furthermore, the taphonomic analyses of fossil remains seem to indicate a diferent accumulative agent than Pachycrocuta, thus adding more complexity to the palaeobiological interpretation of the Venta Micena area. These results contribute to the discussion of traditional interpretations made from Venta Micena 3., Regional Government of Andalusia, with both contracts associated to General Research Project BC.03.032/17, Spanish Public Employment Service (Spanish Ministry of Labour and Social Economy), Spanish Ministry of Science, Innovation and Universities, with an FPI Predoctoral Grant (Reg. PRE2019-089411) associated to project RTI2018-099850-B-IOO and the University of Salamanca, Institut Català de Paleoecologia Humana i Evolució Social (IPHES-CERCA) has received fnancial support from the Spanish Ministry of Science and Innovation through the “María de Maeztu” program for Units of Excellence (CEX2019-000945-M), Projects 2017SGR-859 (Gov. of Catalonia, AGAUR), and CGL2016-80000-P (Spanish Ministry of Economy and Competitiveness), Research Group HUM-607 (Junta de Andalucía).

A species factory refers to the source that gives rise to an exceptionally large number of species. However, what is it exactly: a place, a time or a combination of places, times and environmental conditions, remains unclear. Here we search for species factories computationally, for which we develop statistical approaches to detect origination, extinction and sorting hotspots in space and time in the fossil record. Using data on European Late Cenozoic mammals, we analyse where, how and how often species factories occur, and how they potentially relate to the dynamics of environmental conditions. We find that in the Early Miocene origination hotspots tend to be located in areas with relatively low estimated net primary productivity. Our pilot study shows that species first occurring in origination hotspots tend to have a longer average longevity and a larger geographical range than other species, thus emphasizing the evolutionary importance of the species factories.

The Red Queen's hypothesis portrays evolution as a never-ending competition for expansive energy, where one species’ gain is another species’ loss. The Red Queen is neutral with respect to body size, implying that neither small nor large species have a universal competitive advantage. Here we ask whether, and if so how, the Red Queen's hypothesis really can accommodate differences in body size. The maximum population growth in ecology clearly depends on body size—the smaller the species, the shorter the generation length, and the faster it can expand given sufficient opportunity. On the other hand, large species are more efficient in energy use due to metabolic scaling and can maintain more biomass with the same energy. The advantage of shorter generation makes a wide range of body sizes competitive, yet large species do not take over. We analytically show that individuals consume energy and reproduce in physiological time, but need to compete for energy in real time. The Red Queen, through adaptive evolution of populations, balances the pressures of real and physiological time. Modeling competition for energy as a proportional prize contest from economics, we further show that Red Queen's zero-sum game can generate unimodal hat-like patterns of species rise and decline that can be neutral in relation to body size.

The Early Pleistocene dispersal of Homo out of Africa is a highly studied and debated topic. One of the controversies centres on the question of what type of environments hominin species expanded out of Africa into. We conducted a literature review of 163 papers published since 2000 studying the environmental settings of the first Out of Africa expansion. We found that the literature is polarised between two types of hypotheses. On one hand there are papers which describe Homo in the Early Pleistocene as inflexible (compared to Homo sapiens) and incapable of persisting in non-savannah environments, e.g. the ‘savannahstan’ hypothesis. On the other hand there are papers which describe Homo as flexible and able to persist in various environment types, e.g. the variability selection hypothesis. By investigating these hypotheses we are able to move closer to answering the question - as Homo dispersed out of Africa, did they diversify to exploit new environments, or remain within the ranges of their African niche? We analysed the reconstructions of early Homo environments included in these papers. We found that the qualitative language used to describe hominin environments is problematic and impedes the formation of clear conclusions about the environments occupied by early Homo species. However, by forcibly quantifying the descriptions used in 69 (of the original 163) papers we found that the research does not strongly support the savannahstan hypothesis. Instead the environments inhabited by Homo are consistently reconstructed as a mix of environment types (grassland, forest, savannah etc.), with a slight skew towards open habitats. Based on these results, we tentatively suggest that Homo exhibited a preference for heterogeneous “edge” environments during the Pleistocene and as they dispersed out of Africa. However, in order to further investigate the potential preference of Homo for heterogeneous environments and to build confidence in reconstructions of early human environments in general, quantified reconstructions of the vegetation composition and distribution at early Homo sites are needed.

New mammal species have always been a source of major zoological intrigue, and whilst discoveries of large mammals are becoming less frequent, species like the saola and dingiso continue to enter the zoological literature. Modern communities are often assumed to be complete and are used for constructing community-level models. One example is ecometric modelling. Here, we estimate the number of large herbivorous mammals not yet described based on description curves. We also investigate the effect of species description on the stability and accuracy of previously established relationships between traits and climate. The contemporary description record of large herbivorous mammals is incomplete, with at least 83 species undescribed. Primates, artiodactyls and marsupials are estimated to contain the greatest undescribed diversity, with particular gaps in the Neotropic, Afrotropic and Indomalayan realms. We find that beyond ~40% completeness, there is a limited impact on trait–environment relationships of increased species description, but that there is a high mismatch between true and predicted climatic values for published models. Consequently, mammalian species that have not yet been discovered are unlikely to have a large impact on the accuracy of trait-environment models, but we suggest possible alterations to previous approaches that might improve the accuracy of future models.

Little is known about the subsistence practices of the first European settlers, mainly due to the shortage of archaeological sites in Europe older than a million years. This article contributes to the knowledge of the subsistence of the first Europeans with new zooarchaeology and taphonomic data from the Palaeolithic site of Barranco León (Orce, Granada, Spain). We present the results of the analysis of the faunal assemblages retrieved in the context of new excavations undertaken between 2016 and 2020. We have followed a standard methodology for the identification and quantification of species, mortality profiles, skeletal representation and taphonomic analysis. With regard to the taphonomic evidence, we have documented the extent of rounding, abrasion and other alterations. Finally, we examined traces from the activities of carnivores and hominins that led to the accumulation and alteration of the bone assemblages. Results indicate that the archaeo-paleontological deposits from Barranco León present a dual-patterned mixed taphonomic origin. The first phase primarily involved waterborne processes (BL-D1), which led to the accumulation of lithic raw materials, a few archaeological stone tools, and some faunal remains with percussion and cutmarks. The second phase (BL-D2) contains several stone tools associated with faunal remains with more anthropogenic alterations, such as cutmarks and percussion marks. After analysing the Barranco León zooarchaeological assemblage, the present study concludes that hominins had access to the meat and within-bone nutrients of animals of diverse sizes. However, the specific carcass acquisition mechanisms that hominins followed are less certain because the presence of tooth marks suggests that carnivores also played a role in the accumulation and modification of the Barranco León faunal assemblage., CRUE-CSIC agreement Springer Nature, Junta de Andalucia, Consejeria de Educacion, Cultura y Deporte: Orce Research Project `Primeras ocupaciones humanas y contexto paleoecologico a partir de los depositos pliopleistocenos de la cuenca Guadix-Baza: zona arqueologica de la Cuenca de Orce (Grana BC.03.032/17, Spanish Government PRE2019-089411 RTI2018-099850-B-I00, University of Salamanca, Arts and Humanities Research Council through a Baillie Gifford AHRC Scholarship (OOC-DTP program) at the University of Oxford AH/R012709/1, National Health and Medical Research Council (NHMRC) of Australia PRE2020-094482, European Union-NextGenerationEU 2021URV-MS-03, Ministry of Universities and Recovery, Transformation and Resilience Plan, Generalitat de Catalunya 2017SGR-859 2017SGR-1666, Spanish Ministry of Science and Innovation through the `Maria de Maeztu' program for Units of Excellence CEX2019-000945-M, Spanish Government CEX2019-000945-M-20-1

Nearly five decades ago Berggren and Van Couvering proposed an Old World “Hipparion Datum” wherein a North American Hipparion extended its range across Eurasia and Africa as an “instantaneous prochoresis” populating the Old World. Four decades ago Woodburne and Bernor examined European and North African hipparion assemblages and proposed a number of distinct hipparion lineages, sharply departing from the mono-generic paradigm of previous work. Through the 1980s until now, hipparion systematic studies have delineated multiple superspecific groups of hipparions. Herein, we define 10 recognizable genus-rank Eurasian and African taxa delineating their chronologic occurrences, geographic extent and where data exists, their body mass and paleodietary preferences. Our study supports the current interpretation that a species of North American Cormohipparion extended its range into the Old World in the early late Miocene. Regional first occurrences of Cormohipparion are recognized in the Potwar Plateau, Pakistan and Sinap Tepe, Turkey 10.8 Ma. The slightly derived lineage Hippotherium is recorded earlier in the Pannonian C of the Vienna Basin, 11.4–11.0 Ma marking the chronologic “Hipparion” Datum at the lower boundary of Mammal Neogene (MN) Unit 9. Within MN 9, 11.2–9.9 Ma, Cormohipparion underwent a minor diversification whereas Hippotherium diversified in Central and Western Europe and China and Sivalhippus (S. nagriensis) originated in the Indian Subcontinent. Whereas Cormohipparion did not survive into the late Vallesian, MN10 (9.9–8.9 Ma), Hippotherium and Sivalhippus did and the Cremohipparion and Hipparion s.s. lineages originated. During the early and middle Turolian (MN11–12, 8.9–6.8 Ma) Hippotherium, Sivalhippus, Cremohipparion and Hipparion persisted and new lineages, Eurygnathohippus, Plesiohipparion, Baryhipparion and Shanxihippus originated. An initial extinction interval occurred at the end of the Miocene, MN13 (6.8–5.3 Ma) wherein all but one endemic species of Hippotherium, H. malpassi (Italy), Hipparion and several species of Cremohipparion became extinct. Lineage and species reduction continued across the Mio-Pliocene boundary so that by the beginning of the Pliocene (MN14, 5.3 Ma) only African species of Eurygnathohippus, Chinese Plesiohipparion houfenense and Proboscidipparion sinense remained. The later Pliocene (MN15–16, ca. 5.0–2.5 Ma) documents the persistence of endemic Chinese Baryhipparion insperatum, modest diversification of African Eurygnatohippus spp. and Chinese Plesiohipparion and Proboscidipparion spp. Eurygnatohippus made a limited geographic extension into the Indian subcontinent during MN16, whereas Pleisohipparion and Proboscidipparion extended their ranges into Eurasia during MN15 and MN16. The latest occurring hipparions are Proboscidipparion sinense at 1.0 Ma in China and Eurygnatohippus cornelianus in Africa 300 kg), with the smaller forms being predominately grass feeders and larger ones being mixed feeders. Decreased hipparion lineage and species diversity in the Pliocene was accompanied by increased average body size and hypsodonty probably in response to more seasonal Eurasian and African environments. There is no evidence that hipparions ever adapted to cold and dry Old World Pleistocene environments.

The Guadix-Baza Basin (GBB) in Andalucía, Spain, comprises palaeontological and archaeological sites dating from the Early Pliocene to the Middle Pleistocene, including some of the earliest sites with evidence for the presence of early humans (Homo sp.) in Europe. Thus, the history of climate and environments in this basin contributes significantly to our understanding of the conditions under which early humans spread into Europe during the Early Pleistocene. Here we present estimates of precipitation and primary productivity in the GBB from the Pliocene to the Middle Pleistocene based on dental ecometrics in fossil communities of large herbivorous mammals, and perform an ecometrics-based distribution modelling to analyse the environmental conditions of Early and Middle Pleistocene human sites in Europe. Our results show that Early Pleistocene humans generally occupied on average relatively diverse habitats with ecotones, such as woodlands and savannas, but avoided very open and harsh (cool or dry) environments. During the Middle Pleistocene in Europe, humans occupied a comparatively much broader range of environments than during the Early Pleistocene, but were on average more concentrated in environments where the dental ecometric of mammals indicate wooded palaeoenvironments. In the earliest human occupation sites of the GBB, Barranco Leon and Fuente Nueva 3, the mean annual precipitation and net primary production estimates indicate climatic conditions close to modern Mediterranean sclerophyllous woodland environments, but with slightly higher primary productivity,indicating some similarity with East African woodlands. On the other hand, the environments did not resemble African grassland savannas. The browse-dominated diets of ungulates from Barranco Leon and Fuente Nueva 3 further suggest palaeoenvironments where grasses were a minor component of the vegetation. In the slightly older site of Venta Micena that has no evidence for the presence of hominins, dental ecometric estimates indicate climate and environments similar to Mediterranean “forest steppe” environments existing in the surroundings of Baza today. Grasses were prevalent in the diet of some taxa, especially equids, in Venta Micena, but most of the species show browse-dominated diets even there.

The writings of Gottfried Wilhelm Leibniz (1646–1716) provide a window on early evolutionary thinking of a kind interestingly different from the roots of modern evolutionary theory as it emerged in the years following the French Revolution. Here we relate aspects of Leibniz’s thinking to methods of modern palaeoecology and show that, despite a different terminology and a different hierarchic focus, Leibniz emerges as a strikingly modern theoretician, who viewed the living world as dynamic and capable of adaptive change. The coexistence approach of palaeoecological reconstruction, developed by Volker Mosbrugger and collaborators, with its core assumption of harmoniously co-adapted communities with strong historical legacy, represents, in a positive sense, a more Leibnizian view than functionally based and theoretically history-free approaches, such as ecometrics. Recalling Leibniz’s thinking helps to highlight how palaeoecological reconstruction is about much more than reliably establishing the ecological and climatic situation of a given fossil locality. While reliable reconstructions of past conditions are certainly of great value in research, it is arguably the need to think deeply about how the living world really works that keeps palaeoecological reconstruction such a long-running and central aspect of evolutionary science. And while we struggle to understand the coexistence and dynamic interaction of endless levels of living agents of the living world, simultaneously large and small, global and local, the coexistence approach of palaeoecological reconstruction remains both an outstandingly operational method and part of a philosophical tradition reaching back to the very earliest evolutionary thinking.

Open Access funding provided thanks to the CRUE-CSIC agreement with Springer Nature. This research was funded by the Junta de Andalucia, Consejeria de Educacion, Cultura y Deporte: Orce Research Project "Primeras ocupaciones humanas y contexto paleoecologico a partir de los depositos pliopleistocenos de la cuenca Guadix-Baza: zona arqueologica de la Cuenca de Orce (Granada, Espana), 2017-2020 (Ref. BC.03.032/17)". We also received support from the PALARQ Foundation with the convocatory of Analitics 2019: "Identificando Carnivoros a partir de analisis Tafonomicos de ultima generacion aplicando Fotogrametria y Morfometria Geometrica de las Marcas de Diente. Aplicacion a Yacimientos del Pleistoceno Inferior Iberico: FN3, Venta Micena 3 y 4 (Granada), Ponton de la Oliva (Patones, Madrid)". Lloyd Austin Courtenay is also funded by the Spanish Ministry of Science, Innovation and Universities with an FPI Predoctoral Grant (Ref. PRE2019-089411) associated to project RTI2018-099850-B-I00 and the University of Salamanca. The Institut Catala de Paleoecologia Humana I Evolucio Social (IPHES-CERCA) has received financial support from the Spanish Ministry of Science and Innovation through the "Maria de Maeztu" program for Units of Excellence (CEX2019-000945-M)., Over the last few decades, several types of evidence such as presence of hominin remains, lithic assemblages, and bones with anthropogenic surface modifications have demonstrated that early human communities inhabited the European subcontinent prior to the Jaramillo Subchron (1.07–0.98 Ma). While most studies have focused primarily on early European lithic technologies and raw material management, relatively little is known about food procurement strategies. While there is some evidence showing access to meat and other animal-based food resources, their mode of acquisition and associated butchery processes are still poorly understood. This paper presents a taphonomic and zooarchaeological analysis of the Fuente Nueva-3 (FN3) (Guadix-Baza, Spain) faunal assemblage, providing a more in-depth understanding of early hominin subsistence strategies in Europe. The present results show that hominins had access to the meat and marrow of a wide range of animal taxa, including elephants, hippopotami, and small- and medium-sized animals. At the same time, evidence of carnivore activity at the site suggests that these communities likely faced some degree of competition from large predators when acquiring and processing carcasses., CRUE-CSIC agreement, Springer Nature, Junta de Andalucia BC.03.032/17, PALARQ Foundation, Spanish Ministry of Science, Innovation and Universities, National Health and Medical Research Council of Australia PRE2019-089411 RTI2018-099850-B-I00, University of Salamanca, Spanish Government CEX2019-000945-M

Pastoralism is globally significant in social, environmental, and economic terms. However, it experiences crises rooted in misconceptions and poor interdisciplinary understanding, while being largely overlooked in international sustainability forums and agendas. Here, we propose a transdisciplinary research approach to understand pastoralist transitions using (1) social, economic, and environmental dimensions, (2) diverse geographic contexts and scales to capture emerging properties, allowing for cross-system comparisons, and (3) timescales from the distant past to the present. We provide specific guidelines to develop indicators for this approach, within a social-ecological resilience analytical framework to understand change. Distinct systems undergo similar transitions over time, crossing critical thresholds and then either collapsing or recovering. Such an integrated view of multidimensional interactions improves understanding of possible tipping points, thereby supporting better-informed decision making. The need for a paradigm shift in pastoralism science and policy is pressing. This research approach, including participatory methods, can provide the solutions urgently needed. peerReviewed

The Early Pleistocene sites of Barranco Leon and Fuente Nueva 3 (Guadix-Baza Basin, SE Spain) have yielded abundant Oldowan lithic artifacts and one hominin tooth (Homo sp. in level D1 or D2 of Barranco Leon), today considered to be among the earliest evidence for a hominin presence in Western Europe, at ca. 1.4-1.2 Ma. Here, for the first time, the stratigraphic succession of these two sites are studied more precisely from a palaeoenvironmental point of view, taking into account the different levels of the depositional sequences to analyze the successive fossil assemblages of amphibians and reptiles. Palaeoenvironmental reconstructions are carried out by applying the "habitat weighting" method, which uses the modern distribution by habitat of amphibian and reptile species in order to interpret past landscapes. The successive herpetofaunal assemblages from Barranco Leon show a certain tendency towards more arid conditions from level D1 to level E, whereas in Fuente Nueva 3 environmental reconstructions reveal oscillating conditions, with a tendency towards more arid conditions in the basal part of the sequence, up to level 5, where the tendency shifts back to more humid conditions. Our results show that the layers included in this study with the highest density of anthropic evidence (such as level 5 in FN 3 and levels D1 and D2 in BL) are situated within the late Early Pleistocene climatic and environmental cyclicity, yielding different environmental conditions: a humid, wooded biotope for BL, and a more open and drier biotope in FN 3. This suggests that the hominins of the late Early Pleistocene, although conditioned to some extent by climatic factors, were able to cope with changing environmental conditions, both "interglacial" and "glacial", in the southwestern extremity of the European continent. (C) 2020 Elsevier Ltd. All rights reserved.

The Guadix-Baza Basin (Granada Province, Southern Spain) is the richest area in Western Europe for the study of early hominin dispersal and evolution, having yielded the earliest localities with evidence of hominin occupation (a deciduous human molar, lithic industries and cutmarks) together with a rich large and small vertebrate assemblage dated to around 1.4 Ma. A key question is whether environmental changes were involved in the arrival of hominins in this region at this time. To answer this question, possible environmental differences between one older site lacking evidence for hominin occurrence (Venta Micena VM, ~1.6 Ma) and younger sites with undisputable evidence (Barranco León BL and Fuente Nueva-3 FN3, ~1.2-1.5 Ma) were investigated using various approaches, including carbon and oxygen isotopes in tooth enamel, tooth wear analysis, ecometrics and microvertebrates (amphibians, reptiles, mammals) as proxies for palaeoclimate.Tooth enamel powders were collected from herbivorous mammal specimens from the three sites. For several specimens, enamel was sampled serially to document intra-annual dietary and/or habitat changes for the studied individuals. A large diversity of herbivorous taxa was sampled, including cervids, bovids, equids, rhinoceros, hippopotamus and mammoths. The analyses were conducted at the University of Tübingen (Senckenberg Centre for Human Evolution and Palaeoenvironment).Carbon isotopic results from the three sites showed that the plants foraged by herbivores were essentially of C3 photosynthetic pathway (trees, shrubs and C3-grass adapted to mild growth season), which is consistent with the results of tooth wear analysis indicating browsing or mixed feeding with browsing preference for most taxa. The consumption of some C4 plants for some herbivores has been detected only in Barranco León, which is consistent with the results of palaeoclimatic investigations based on ecometrics and microvertebrate fauna, indicating a particularly wet and warm climate for this site compared to both others and colder conditions in Venta Micena, the site devoid of hominins. Oxygen isotopic results seem to be essentially related to browsing (high values) versus grazing (lower values) and to different habitats (lowest oxygen isotopic values for semi-aquatic hippos). The differences in isotopic results among taxa are in agreement with those of dietary preferences from mesowear tooth analysis. In addition, in a context of still Mediterranean climate with 4-months aridity during summer, isotopic variations within teeth suggest in some cases significant changes in foraging through a year, which could be related to local seasonal changes or mobility across areas with different vegetation types.The first results of this multidisciplinary research project financed by the Leakey Foundation and a General Research Project from the Andalusian Regional Government help us to refine the palaeoenvironmental reconstruction around the time of the earliest arrival of hominins in Southern Spain. In combination with the climatic data provided by ecometrics and microvertebrate investigations, it allows us to develop a more detailed framework for the interpretation of the carbon and oxygen isotopic data from tooth enamel in a Mediterranean climate context, which corresponds to the type of climatic conditions where the earliest hominins occur in Europe in the Early Pleistocene.

A new study by Fraser et al (2018) urges the use of phylogenetic comparative methods, whenever possible, in analyses of mammalian tooth wear. We are concerned about this for two reasons. First, this recommendation may mislead the research community into thinking that phylogenetic signal is an artifact of some sort rather than a fundamental outcome of the evolutionary process. Secondly, this recommendation may set a precedent for editors and reviewers to enforce phylogenetic adjustment where it may unnecessarily weaken or even directionally alter the results, shifting the emphasis of analysis from common patterns manifested by large clades to rare cases.

How food intake and mastication scale to satisfy the metabolic needs of mammals has been the subject of considerable scientific debate. Existing theory suggests that the negative allometric scaling of metabolic rate with body mass is compensated by a matching allometric scaling of the chewing rate. Why empirical studies have found that the scaling coefficients of the chewing rate seem to be systematically smaller than expected from theory remains unknown. Here we explain this imparity by decoupling the functional surface area of teeth from overall surface area. The functional surface area is relatively reduced in forms emphasizing linear edges (e.g., lophodont) compared with forms lacking linear structures (e.g., bunodont). In forms with reduced relative functional surface, the deficit in food processed per chew appears to be compensated for by increased chewing rate, such that the metabolic requirements are met. This compensation accounts for the apparent difference between theoretically predicted and observed scaling of chewing rates. We suggest that this reflects adaptive functional evolution to plant foods with different fracture properties and extend the theory to incorporate differences in functional morphology. (C) 2017 Elsevier Ltd. All rights reserved.

Over the past five decades, fossil deposits within the Upper Burgi, KBS and Okote members at East Turkana in northern Kenya have provided many important insights into hominin behavior and ecology during a critical period in hominin evolution between 2.0 and 1.4 Ma. In this study, we use a large compilation of faunal abundance data from paleontological and archaeological collections at East Turkana dating to this time interval to investigate temporal patterns in large mammal taxa, ecosystem evolution and hominin ecology. Our analyses indicate that although portions of the ecosystem were dominated by mesic grasslands, the relative proportion of mesic and arid grassland environments varied though time. We document a major transition in the family Suidae with an increase in the abundance of fossils attributed to the Metridiochoerus lineage coeval with the local extinction of the Notochoerus lineage and decline in abundance of the Kolpochoerus lineage. Finally, by comparing the proportional representation of mammalian taxa found in paleontological collections versus those found in archaeological collections, our data suggest that archaeological sites at East Turkana, particularly those c.1.5 Ma, contain disproportionately large numbers of alcelaphin bovid remains. This could reflect 1) hominin prey choice, 2) hominin hunting/scavenging habitat choice, or 3) a combination the two.

More than half a century ago, Percy Butler touted the importance of analyzing teeth to understand their function in an evolutionary context. There have been many advances in the study of dental functional morphology since that time. Here we review the various approaches to characterizing and comparing occlusal form that have been developed, especially dental topographic analysis. We also report on a new study of dental topography of platyrrhine primates (n = 341 individuals representing 16 species) with known differences in both dietary preferences and other food items eaten. Results indicate frugivores, gummivores, folivores, and seed eaters each have a unique combination of slope, relief, angularity, sharpness, and occlusal orientation patch size and count values. Likewise, among frugivores, those that supplement their diets with hard objects, insects, leaves, and seeds, also each have a distinctive suite of topographic features. We conclude that both primary and secondary diet choices select for occlusal form, and that functional morphology more reflects the types of foods and mechanical challenges they pose rather than the frequencies in which they are eaten.

After a long period of inactivity, recent excavations at the late Miocene Maragheh Formation unexpectedly resulted in the discovery of the first fossil hominoid and second Mesopithecus remains from this area. The discovery motivated a new international initiative to conduct research in these rich fossil sites. These studies focused on the fossil hominoid and its locality, aiming to reveal more about the context of this fossil discovery. Detailed stratigraphy, sedimentology and magnetostratigraphy studies were conducted. New samples from volcaniclastic key horizons (pumice beds) in Dareh Gorg, where the hominoid fossil site is located, were dated by radiometric methods. The radiometric age determinations provide a firm tie-point for the geochronology. The polarity pattern in the palaeomagnetically investigated section corroborates the K-Ar results. The preliminary magnetostratigraphic results suggest that the hominoid locality can be correlated to the normal polarity chron C4n.2n (8.108–7.695 Ma), C4n.1n (7.642–7.528 Ma) or C3Br.1n (7.285–7.251 Ma), placing it at intervals corresponding to the mammal units MN11 or possibly early MN12. The study of fossil hominoid indicates broad affinities with a number of contemporaneous taxa from the Balkan-Iranian palaeoprovince, as well as Siwaliks and southeast Asia. A preliminary analysis of the accompanying (in situ) fauna at the hominoid site indicates the highest similarity of this level to Turolian hominoid- and Mesopithecus-bearing localities in Turkey, Greece and Bulgaria. However, some environmental differences are observed among these localities, based on their faunal structure and taxon properties, as well as in the different masticatory adaptations of their hominoids.

The Maragheh Formation is an important deposit, which yields the savanna-type large mammal assemblage known as the “Pikermian Fauna.” Our high-resolution facies analysis of the interval between the Lower Pumice and the White Tuff demonstrated that debris-flow deposits and paleosols are dominant in the studied sequence. Fluvial channel-fill and small pond facies are the subordinate components in this interval. Most of the channel-fill deposits are interpreted as having been accumulated from ephemeral streams. The wide distribution of the Middle Pumice, the fact that it contains grains with older ages than those of the Lower Pumice, and the presence of “traction carpet” deposits allow the interpretation of the pumice interval as having been deposited from a hyperconcentrated flow probably caused by crater-lake destruction around the peak of Mt. Sahand, which supplied older rocks to the flow. The internal architecture of fluvial channel-fill deposits and the structures of paleosols (rhizoliths, cracks and slickensides: probable Vertic Inceptisols) imply a seasonal climate during deposition of the studied interval. This is consistent with previous environmental reconstructions based on mammal fossils (woodland-dominated savannah) as well as the results of phytolith analysis and δ18O data obtained from northern Iran. However, flood deposits covering the paleosols, showing sheet-like geometry, may not have been affected by large trees on the flood plain: the extent of woodland around the study site appears to have been limited.

Maragheh in northwestern of Iran is a world famous Miocene fossil-bearing area. The area has yielded classical late Miocene Turolian age fauna that has been collected and studied sporadically over the last 150 years. However, the precise correlation of these sediments to the Global Time Scale (GTS) has remained ambiguous. To address this, 115 levels along an approximately 27-m-thick interval were collected from the middle Maragheh Formation at Dareh Gorg (Gort Daresi) section. Characteristic remanent magnetization directions obtained by alternating field demagnetization produce a polarity pattern that is supported by thermal demagnetization on a set of sister specimens. Three polarity intervals were recognised, the middle part of the section at around 15-21 m showing reversed polarity, bounded by normal polarities above and below. Based on the palaeontological constraints and recent K-Ar age determinations from the Maragheh Fm, three correlations to the geomagnetic polarity time scale appear likely. According to these correlations, recently discovered hominoid locality is correlated to C3Br.1n, C4n.1n, or to C4n.2n. For a unique correlation, however, additional palaeomagnetic data is required from the upper and lower parts of the section.

Pumice beds of the Maragheh Formation, which is distributed on the plains at the foot of the late Cenozoic Sahand volcano, NW Iran, consist of the Mordaq tuff bed, lower pumice beds (A and B), middle pumice bed and upper pumice bed, in ascending order. New hornblende and plagioclase K–Ar ages of the pumices from these are: Mordaq tuff bed: 8.14 ± 0.27 Ma, lower pumice beds (A): 7.54 ± 0.22 Ma, lower pumice beds (B): 6.95 ± 0.28 Ma, middle pumice bed: 7.87 ± 0.29 Ma, upper pumice bed: 6.96 ± 0.31 Ma. The middle pumice bed is a flood deposit, and the chemical compositions of glass, hornblende and biotite in pumices overlap those from the lower pumice beds (A). These features indicate that the pumices of the middle pumice bed were derived from a similar source to the lower pumice beds (A). Chemical compositions of pumices from the Maragheh Formation fall in the dacite field of the total alkali-silica (TAS) classification diagram and slightly differ from those of the Plio–Pleistocene rocks from the Sahand volcano. Chemical compositions of late Miocene Maragheh Formation pumices show adakitic features with high Sr contents (698–545 ppm), and low Y (12–8 ppm). It is inferred that the source materials of the adakitic dacite magma were amphibole eclogite or garnet amphibolite originating from the Neo-Thetys plate subducted beneath the Eurasia plate.

Miocene hominoid fossils are known from Africa and Eurasia, in the latter ranging widely from western Europe to Anatolia and from South Asia to Southeast/East Asia. Iran is located between the known western and eastern Eurasian hominoid distributions and is potentially important in understanding Miocene hominoid dispersal patterns. Maragheh is a late Miocene fossil locality in northwestern Iran, well known since the nineteenth century for its abundant mammalian fossils. However, until now, the only primate fossils reported from Maragheh or Iran were the Old World monkey Mesopithecus pentelicus. Recent field research at Maragheh has changed this situation by the discovery of the first hominoid fossil from Iran, a maxillary fragment with well-preserved second and third molars. Here, we provide a detailed description of this new specimen, comparing it with other similarly large-sized Eurasian late Miocene hominoids, Ouranopithecus, Ankarapithecus, Sivapithecus, and Indopithecus. Molar morphology of the Maragheh hominoid is similar to that of these Eurasian Miocene genera, with only minor differences in morphology and wear pattern. Based on the presently available materials, we tentatively prefer the interpretation that the Maragheh hominoid may be related more closely to either Ankarapithecus or Sivapithecus rather than to Ouranopithecus, but the fragmentary nature of the fossil makes evaluations difficult. Future discoveries of this Iranian hominoid are needed to determine its phylogenetic position with more certainty.

International audience; As organisms are adapted to their environments, assemblages of taxa can be used to describe environments in the present and in the past. Here, we use a data mining method, namely redescription mining, to discover and analyze patterns of association between large herbivorous mammals and their environments via their functional traits. We focus on functional properties of animal teeth, characterized using a recently developed dental trait scoring scheme. The teeth of herbivorous mammals serve as an interface to obtain energy from food, and are therefore expected to match the types of plant food available in their environment. Hence, dental traits are expected to carry a signal of environmental conditions. We analyze a global compilation of occurrences of large herbivorous mammals and of bioclimatic conditions. We identify common patterns of association between dental traits distributions and bioclimatic conditions and discuss their implications. Each pattern can be considered as a computational biome. Our analysis distinguishes three global zones, which we refer to as the boreal-temperate moist zone, the tropical moist zone and the tropical-subtropical dry zone. The boreal-temperate moist zone is mainly characterized by seasonal cold temperatures, a lack of hypsodonty and a high share of species with obtuse lophs. The tropical moist zone is mainly characterized by high temperatures, high isothermality, abundant precipitation and a high share of species with acute rather than obtuse lophs. Finally, the tropical dry zone is mainly characterized by a high seasonality of temperatures and precipitation, as well as high hypsodonty and horizodonty. We find that the dental traits signature of African rain forests is quite different from the signature of climatically similar sites in North America and Asia, where hypsodont species and species with obtuse lophs are mostly absent. In terms of climate and dental signatures, the African seasonal tropics share many similarities with Central-South Asian sites. Interestingly, the Tibetan plateau is covered both by redescriptions from the tropical-subtropical dry group and by redescriptions from the boreal-temperate moist group, suggesting a combination of features from both zones in its dental traits and climate.

The spatiotemporal pattern of the late Cenozoic spread of C 4 vegetation is an important indicator of environmental change that is intertwined with the uplift of the Himalaya and Tibetan Plateau, and the development of the East Asian monsoons. To explore the spread of C 4 vegetation in China and shed new light on regional climatic evolution, we measured δ 13 C values of more than 200 small mammal teeth (primarily rodents and lagomorphs) using a laser ablation isotope ratio mass spectrometry approach. Small mammals are highly sensitive indicators of their environment because they have limited spatial ranges and because they have minimal time-averaging of carbon isotope signatures of dietary components. The specimens originate from four classic Late Miocene fossil localities, Lufeng, Yuanmou, Lingtai, and Ertemte, along a southwest-northeast transect from Yunnan Province to Inner Mongolia. In Yunnan (Lufeng, Yuanmou) and on the Loess Plateau (Lingtai), the small mammal δ 13 C values record nearly pure C 3 ecosystems, and mixed but C 3 -based ecosystems, respectively, in agreement with previous studies based on carbon isotopes of large herbivores and soil carbonates. In Inner Mongolia, the micromammalian tooth enamel δ 13 C record picks up the presence of C 4 vegetation where large mammal samples do not, indicating a mixed yet C 3 -dominated ecosystem at ~ 6 Ma. As a whole, the results support a scenario of northward increasing C 4 grass abundance in a pattern that mirrors northward decreasing precipitation of the summer monsoon system. The results highlight differences between large and small mammals as indicators of C 4 vegetation in ancient ecosystems, particularly the ability of small mammal δ 13 C values to detect the presence of minor components of the vegetation structure.

The standard adaptive dynamics framework assumes two timescales, i.e. fast population dynamics and slow evolutionary dynamics. We further assume a third timescale, which is even slower than the evolutionary timescale. We call this the geological timescale and we assume that slow climatic change occurs within this timescale. We study the evolution of our model population over this very slow geological timescale with bifurcation plots of the standard adaptive dynamics framework. The bifurcation parameter being varied describes the abiotic environment that changes over the geological timescale. We construct evolutionary trees over the geological timescale and observe both gradual phenotypic evolution and punctuated branching events. We concur with the established notion that branching of a monomorphic population on an environmental gradient only happens when the gradient is not too shallow and not too steep. However, we show that evolution within the habitat can produce polymorphic populations that inhabit steep gradients. What is necessary is that the environmental gradient at some point in time is such that the initial branching of the monomorphic population can occur. We also find that phenotypes adapted to environments in the middle of the existing environmental range are more likely to branch than phenotypes adapted to extreme environments.

Most suids (Mammalia: Suidae, pigs) worldwide are omnivores living in closed environments, but the African warthog (Phacochoerus) has special adaptations for grazing in open environments. Similar specializations have been recorded from Plio-Pleistocene African suids. Four genera, Nyanzachoerus, Notochoerus, Kolpochoerus, and Metridiochoerus, have been discovered in late Miocene to middle Pleistocene locations around the Turkana Basin. We analyse the relative abundances of these four suid genera compared to other mammals, from approximately 8-0.7 Ma. The dataset includes most of the mammal specimens collected from locations around the Kenyan side of the Turkana Basin. Species of genus Nyanzachoerus were dominant before 4 Ma, but their relative abundance decreases through time thereafter. At the same time, Notochoerus increases in relative abundance, followed by Kolpochoerus, and finally Metridiochoerus. Their peak relative abundances do not overlap: Notochoerus peaks at 3.44-2.53 Ma, Kolpochoerus at 2.53-1.87 Ma, and Metridiochoerus at 1.38-0.7 Ma. We interpret the palaeoecology of these suids based on their relative abundances over time and on published isotope and pollen data. We find that Nyanzachoerus was replaced by its abrasive-diet-specialized successor Notochoerus, possibly in response to a rapid decrease in forest cover. Notochoerus adapted at first to the expanding wood- and grasslands, and then to more arid shrublands. After a period of severe aridity around 2.7-2.5 Ma, more variable environments allowed Kolpochoerus and Metridiochoerus to disperse, while Notochoerus disappeared, perhaps having lost its competitive edge. Further changes in the environment encouraged the expansion of grasslands over shrublands, favouring Metridiochoerus. Kolpochoerus persisted in the more closed environments near water sources.

Damiao, Inner Mongolia, has three main fossil horizons representing the early, middle, and late Miocene. The middle Miocene locality DM01 is the only primate locality from the region and also represents the latest occurrence of pliopithecoids in northern China. The presence of pliopithecoid primates in central Asia after the middle Miocene climatic optimum seems to contradict the general trend of strengthening climatic zonality and increasing aridity. To investigate this enigma, we employ faunal similarity, ecometrics, and stable isotope analysis. Our results support previous inferences concerning the presence of locally humid environments within the increasingly arid surroundings that characterized central Asia. Hypsodonty, estimated mean annual precipitation (MAP), local sedimentology, and large mammal fossils suggest more humid and possibly more forested and wooded environments for the DM01 locality. We compared our results with the adjacent fossil-rich middle Miocene Tunggur localities. However, the small mammal fauna and isotope data are consistent with a mosaic of forest and grassland environment for all Damiao localities. Based on our results, Tunggur may have been too seasonal or not sufficiently humid for pliopithecids. This is supported by the higher mean hypsodonty and lower estimated MAP estimates, as well as slightly higher d13C values. We suggest that DM01, the driest known Asian pliopithecid locality, may have been a more humid refugium within a generally drier regional context.

The relative weights of physical forcing and biotic interaction as drivers of evolutionary change have been debated in evolutionary theory. The recent finding that species, genera, clades, and chronofaunas all appear to exhibit a symmetrical pattern of waxing and waning lends support to the view that biotic interactions shape the history of life. Yet, there is similarly abundant evidence that these primary units of biological evolution arise and wane in coincidence with major climatic change. We review these patterns and the process-level explanations offered for them. We also propose a tentative synthesis, characterized by interdependence between physical forcing and biotic interactions. We suggest that species with evolutionary novelties arise predominantly in “species factories” that develop under harsh environmental conditions, under dominant physical forcing, whereas exceptionally mild environments give rise to “oases in the desert,” characterized by strong competition and survival of relics.

Although patterns of tooth wear are crucial in palaeo-reconstructions, and dental wear abnormalities are important in veterinary medicine, experimental investigations on the relationship between diet abrasiveness and tooth wear are rare. Here, we investigated the effect of four different pelleted diets of increasing abrasiveness (due to both internal [phytoliths] and external abrasives [sand]) or whole grass hay fed for two weeks each in random order to 16 rabbits (Oryctolagus cuniculus) on incisor and premolar growth and wear, and incisor and cheek tooth length. Wear and tooth length differed between diets, with significant effects of both internal and external abrasives. While diet abrasiveness was linked to tooth length for all tooth positions, whole forage had an additional effect on upper incisor length only. Tooth growth was strongly related to tooth wear and differed correspondingly between diets and tooth positions. At 1.4-3.2 mm per week, the growth of cheek teeth measured in this study was higher than previously reported for rabbits. Dental abnormalities were most distinct on the diet with sand. This study demonstrates that concepts of constant tooth growth in rabbits requiring consistent wear are inappropriate, and that diet form (whole vs. pelleted) does not necessarily affect cheek teeth. Irrespective of the strong effect of external abrasives, internal abrasives have the potential to induce wear and hence exert selective pressure in evolution. Detailed differences in wear effects between tooth positions allow inferences about the mastication process. Elucidating feedback mechanisms that link growth to tooth-specific wear represents a promising area of future research.

The Anthropocene is recognized (though not yet formally defined) as the time when human impacts are widespread on Earth. While some of the impacts are essential to supporting large human populations and can be sustainable in the long run, others can irretrievably damage the life support systems upon which the global society has come to depend, or spark rapid changes to which societies cannot adapt fast enough. Among these dangerous trends are increasing climate disruption, extinctions, loss of non-human-dominated ecosystems, pollution, and population overgrowth. Interactions between these five trends exacerbate their potential to trigger harmful global change. Reducing the resultant risks requires effective cooperation between scientists and policy makers to develop strategies that guide for environmental health over the next few decades. To that end, the Scientific Consensus on Maintaining Humanity’s Life Support Systems in the 21st Century was written to make accessible to policy makers and others the basic scientific underpinnings and widespread agreement about both the dangers of and the solutions to climate disruption, extinctions, ecosystem loss, pollution and population overgrowth. When it was released in May 2013, the document included endorsements by 522 global change scientists, including dozens of members of various nations’ most highly recognized scientific bodies, from 41 countries around the world. Since then, endorsements have grown to more than 1300 scientists plus more than 1700 others – business people, NGO representatives, students, and the general public – spanning more than 60 countries. Now also available in Spanish and Chinese, the document has proven useful in helping to stimulate national and international agreements. Further information about the genesis, uses, the signatories, and how to endorse it can be found at http://consensusforaction.stanford.edu/ . Such communication between scientists, policy makers, and the public at large will be essential for effective guidance to address global change as the Anthropocene progresses.

The evolution of a particular sensory organ is often discussed with no consideration of the roles played by other senses. Here, we treat mammalian vision, olfaction and hearing as an interconnected whole, a three-dimensional sensory space, evolving in response to ecological challenges. Until now, there has been no quantitative method for estimating how much a particular animal invests in its different senses. We propose an anatomical measure based on sensory organ sizes. Dimensions of functional importance are defined and measured, and normalized in relation to animal mass. For 119 taxonomically and ecologically diverse species, we can define the position of the species in a three-dimensional sensory space. Thus, we can ask questions related to possible trade-off vs. co-operation among senses. More generally, our method allows morphologists to identify sensory organ combinations that are characteristic of particular ecological niches. After normalization for animal size, we note that arboreal mammals tend to have larger eyes and smaller noses than terrestrial mammals. On the other hand, we observe a strong correlation between eyes and ears, indicating that co-operation between vision and hearing is a general mammalian feature. For some groups of mammals we note a correlation, and possible co-operation between olfaction and whiskers.

Data, whether images, measurements, counts, occurrences, or character codings, are a cornerstone of vertebrate paleontology. Every published paper, master's thesis, and doctoral dissertation relies on these data to document patterns and processes in evolution, ecology, taphonomy, geography, geologic time, and functional morphology, to name just a few. In turn, the vertebrate paleontology community relies on published data in order to reproduce and verify others' work, as well as to expand upon published analyses in new ways without having to reconstitute data sets that have been used by earlier authors and to accurately preserve data for future generations of researchers. Here, we review several databases that are of interest to vertebrate paleontologists and strongly advocate for more deposition of basic research data in publicly accessible databases by vertebrate paleontologists.

A major focus in evolutionary biology is to understand how the evolution of organisms relates to changes in their physical environment. In the terrestrial realm, the interrelationships among climate,vegetation, and herbivores lie at the heart of this question. Here we introduce and test a scoring scheme for functional traits present on theworn surfaces of large mammalian herbivore teeth to capture their relationship to environmental conditions. We modeled local precipitation, temperature, primary productivity, and vegetation index as functions of dental traits of large mammal species in 13 national parks in Kenya over the past 60 y. We found that these dental traits can accurately estimate local climate and environment, even at small spatial scales within areas of relatively uniform climate (within two ecoregions), and that they predict limiting conditions better than average conditions. These findings demonstrate that the evolution of key functional properties of organisms may be more reflective of demands during recurring adverse episodes than under average conditions or during isolated severe events. This article contains supporting information online at www.pnas.org/lookup/suppl/doi:10.1073/pnas.1609409113/-/DCSupplemental

Author correction: vol 2, pg, 402, 2018 DOI:10.1038/s41559-018-0468-8 Despite much interest in the ecology and origins of the extensive grassland ecosystems of the modern world, the biogeographic relationships of savannah palaeobiomes of Africa, India and mainland Eurasia have remained unclear. Here we assemble the most recent data from the Neogene mammal fossil record in order to map the biogeographic development of Old World mammalian faunas in relation to palaeoenvironmental conditions. Using genus-level faunal similarity and mean ordinated hypsodonty in combination with palaeoclimate modelling, we show that savannah faunas developed as a spatially and temporally connected entity that we term the Old World savannah palaeobiome. The Old World savannah palaeobiome flourished under the influence of middle and late Miocene global cooling and aridification, which resulted in the spread of open habitats across vast continental areas. This extensive biome fragmented into Eurasian and African branches due to increased aridification in North Africa and Arabia during the late Miocene. Its Eurasian branches had mostly disappeared by the end of the Miocene, but the African branch survived and eventually contributed to the development of Plio-Pleistocene African savannah faunas, including their early hominins. The modern African savannah fauna is thus a continuation of the extensive Old World savannah palaeobiome.

Animal clades tend to follow a predictable path of waxing and waning during their existence, regardless of their total species richness or geographic coverage. Clades begin small and undifferentiated, then expand to a peak in diversity and range, only to shift into a rarely broken decline towards extinction. While this trajectory is now well documented and broadly recognised, the reasons underlying it remain obscure. In particular, it is unknown why clade extinction is universal and occurs with such surprising regularity. Current explanations for paleontological extinctions call on the growing costs of biological interactions, geological accidents, evolutionary traps, and mass extinctions. While these are effective causes of extinction, they mainly apply to species, not clades. Although mass extinctions is the undeniable cause for the demise of a sizeable number of major taxa, we show here that clades escaping them go extinct because of the widespread tendency of evolution to produce increasingly specialised, sympatric, and geographically restricted species over time.

Although ecometric methods have been used to analyse fossil mammal faunas and environments of Eurasia and North America, such methods have not yet been applied to the rich fossil mammal record of eastern Africa. Here we report results from analysis of a combined dataset spanning east and west Turkana from Kenya between 7 and 1 million years ago (Ma). We provide temporally and spatially resolved estimates of temperature and precipitation and discuss their relationship to patterns of faunal change, and propose a new hypothesis to explain the lack of a temperature trend. We suggest that the regionally arid Turkana Basin may between 4 and 2 Ma have acted as a ‘species factory’, generating ecological adaptations in advance of the global trend. We show a persistent difference between the eastern and western sides of the Turkana Basin and suggest that the wetlands of the shallow eastern side could have provided additional humidity to the terrestrial ecosystems. Pending further research, a transient episode of faunal change centred at the time of the KBS Member (1.87–1.53 Ma), may be equally plausibly attributed to climate change or to a top-down ecological cascade initiated by the entry of technologically sophisticated humans.This article is part of the themed issue ‘Major transitions in human evolution’.

We discuss a proboscidean bone fragment discovered in southern Finland, including the morphological analysis of the bone, as well as pollen and diatom analyses from sediment contained in the marrow cavity. Preliminary analysis of the bone suggested petrification and thus an apparently old age, while the microfossil assemblages include numerous unequivocally pre-Quaternary pollen, spore, and diatom types. A Miocene age for the bone is determined based on the presence of the diatom genus Alveolophora , indicating a minimum age of 5 Ma, and based on the earliest appearance of proboscideans outside Africa, setting a maximum age of 19 Ma. Based on morphology, the bone is determined as a partial humerus of the left foreleg of a large proboscidean. The bone is tentatively assigned to cf. Deinotherium sp., which is consistent with the diatom-based minimum age. The pollen assemblage is rich in spores of shoreline pteridophytes, while the diatom assemblage is also consistent with a shoreline freshwater environment, suggesting that the bone was deposited post-mortem near the shore of a lake or a stream. Miocene sediments do not currently exist in southern Finland or in the near vicinity. This implies that the bone has been transported over a considerable distance. Due to the discovery of the bone in early-Holocene Baltic Sea clay, the final transport phase and deposition must have taken place via iceberg rafting. This was likely preceded by one or more phases of glacial and/or glacio-fluvial transport. While we are unable to conclusive ascertain the region of origin, the alkaline composition of the contained sediment and diatoms point towards the Russian Plain region in the east. This specimen represents the oldest mammalian bone discovered in Finland and the northernmost discovery of a Miocene proboscidean bone in Europe.