Papers by Katerina Harvati
Scientific Reports, 2024
The Balkans are considered a major glacial refugium where flora and fauna survived glacial period... more The Balkans are considered a major glacial refugium where flora and fauna survived glacial periods and repopulated the rest of Europe during interglacials. While it is also thought to have harboured Pleistocene human populations, evidence linking human activity, paleoenvironmental indicators and a secure temporal placement to glacial periods is scant. Here, we present the first intra-tooth multi-isotope analysis for the European straight-tusked elephant Palaeoloxodon antiquus, on an adult male individual excavated in association with lithic artefacts at the MIS 12 site Marathousa 1 (Megalopolis basin, Greece). The studied find also exhibits anthropogenic modifications, providing direct evidence of hominin presence. We employed strontium, carbon and oxygen isotope analysis on enamel bioapatite to investigate its foraging and mobility behaviour, using a sequential sampling strategy along the tooth growth axis of the third upper molar, to assess ecological changes during the last decade of life. We found a geographically restricted range, in a C 3-dominated open woodland environment, and relatively stable conditions over the examined timeframe. Our results show that, despite the severity of the MIS 12 glacial, the Megalopolis basin sustained a mesic habitat, sufficient plant cover and limited seasonal fluctuations in resource availability, pointing to its role as a glacial refugium for both fauna and hominins.
Quaternary, 2024
This article is an open access article distributed under the terms and conditions of the Creative... more This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY
Journal of Human Evolution, 2024
The Kocabaş specimen comes from a travertine quarry near the homonymous village in the Denizli ba... more The Kocabaş specimen comes from a travertine quarry near the homonymous village in the Denizli basin (Turkey). The specimen comprises three main fragments: portions of the right and left parietal and left and right parts of the frontal bone. The fossil was assumed to belong to the Homo erectus s.l. hypodigm by some authors, whereas others see similarities with Middle Pleistocene fossils (Broken Hill 1/Kabwe, Bodo, or Ceprano). Here, we present the first attempt to make a complete reconstruction of the missing medial portion of the frontal bone and a comprehensive geometric morphometric analysis of this bone. We restored the calotte by aligning and mirroring the three preserved fragments. Afterward, we restored the missing portion by applying the thin-plate spline interpolation algorithm of target fossils onto the reconstructed Kocabaş specimen. For the geometric morphometric analyses, we collected 80 landmarks on the frontal bone (11 osteometric points, 14 bilateral curve semilandmarks, and 41 surface semilandmarks). The comparative sample includes 21 fossils from different chronological periods and geographical areas and 30 adult modern humans from different populations. Shape analyses highlighted the presence in Kocabas¸of features usually related to Middle Pleistocene Homo, such as a developed supraorbital torus associated with a relatively short frontal squama and reduced post-toral sulcus. Cluster analysis and linear discriminant analysis classification procedure suggest Kocabaş being part of the same taxonomic unit of Eurasian and African Middle Pleistocene Homo. In light of our results, we consider that attributing the Kocabaş hominin to H. erectus s.l. may be unwarranted. Results of our analyses are compatible with different evolutionary scenarios, but a more precise chronological framework is needed for a thorough discussion of the evolutionary significance of this specimen. Future work should clarify its geological age, given uncertainties regarding its stratigraphic provenance.
Journal of Human Evolution, 2024
The evolution of the human hand is a topic of great interest in paleoanthropology. As the hand ca... more The evolution of the human hand is a topic of great interest in paleoanthropology. As the hand can be involved in a vast array of activities, knowledge regarding how it was used by early hominins can yield crucial information on the factors driving biocultural evolution. Previous research on early hominin hands focused on the overall bone shape. However, while such approaches can inform on mechanical abilities and the evolved efficiency of manipulation, they cannot be used as a definite proxy for individual habitual activity. Accordingly, it is crucial to examine bone structures more responsive to lifetime biomechanical loading, such as muscle attachment sites or internal bone architecture. In this study, we investigate the manual entheseal patterns of Australopithecus afarensis, Australopithecus africanus, and Australopithecus sediba through the application of the validated entheses-based reconstruction of activity method. Using a comparative sample of later Homo and three great ape genera, we analyze the muscle attachment site proportions on the thumb, fifth ray, and third intermediate phalanx to gain insight into the habitual hand use of Australopithecus. We use a novel statistical procedure to account for the effects of interspecies variation in overall size and ray proportions. Our results highlight the importance of certain muscles of the first and fifth digits for humanlike hand use. In humans, these muscles are required for variable in-hand manipulation and are activated during stone-tool production. The entheses of A. sediba suggest muscle activation patterns consistent with a similar suite of habitual manual activities as in later Homo. In contrast, A. africanus and A. afarensis display a mosaic entheseal pattern that combines indications of both humanlike and apelike manipulation. Overall, these findings provide new evidence that some australopith species were already habitually engaging in humanlike manipulation, even if their manual dexterity was likely not as high as in later Homo.
PaleoAnthropology, 2024
The Iwo Eleru skeleton is the only Pleistocene human fossil currently known from western Africa. ... more The Iwo Eleru skeleton is the only Pleistocene human fossil currently known from western Africa. Previously, we showed morphological affinities of the Iwo Eleru cranial remains with Pleistocene archaic African specimens, consistent with former interpretations of this specimen. Those results suggested deep population substructure in Africa and a complex evolutionary process for the origin of modern humans, potentially involving hybridization between Later Stone Age modern human populations and late surviving archaic lineages. Here we conduct a geometric morphometric comparative analysis of the Iwo Eleru mandible to shed further light on the specimen's morphology and evolutionary relationships. We used twenty-five three-dimensional landmark coordinates, collected from a comparative sample comprising Pleistocene and Holocene Homo sapiens, as well as Homo neanderthalensis and Eurasian Middle Pleistocene Homo samples. Results show that the Iwo Eleru mandible is most consistent with the shape variation found in North African Homo sapiens from Epipaleolithic contexts, both in overall shape and in its large size. These findings are discussed in the temporal and geographic framework of the Iwo Eleru skeleton.
Scientific Reports, 2024
Neanderthals' lives were historically portrayed as highly stressful, shaped by constant pressures... more Neanderthals' lives were historically portrayed as highly stressful, shaped by constant pressures to survive in harsh ecological conditions, thus potentially contributing to their extinction. Recent work has challenged this interpretation, leaving the issue of stress among Paleolithic populations highly contested and warranting in-depth examination. Here, we analyze the frequency of dental enamel hypoplasia, a growth disruption indicator of early life stress, in the largest sample of Neanderthal and Upper Paleolithic dentitions investigated to date for these features. To track potential species-specific patterns in the ontogenetic distribution of childhood stress, we present the first comprehensive Bayesian modelling of the likelihood of occurrence of individual and matched enamel growth disruptions throughout ontogeny. Our findings support similar overall stress levels in both groups but reveal species-specific patterns in its ontogenetic distribution. While Neanderthal children faced increasing likelihoods of growth disruptions starting with the weaning process and culminating in intensity post-weaning, growth disruptions in Upper Paleolithic children were found to be limited around the period of weaning and substantially dropping after its expected completion. These results might, at least in part, reflect differences in childcare or other behavioral strategies between the two taxa, including those that were advantageous for modern humans' long-term survival.
Global and Planetary Change, 2024
Southern Europe is hypothesized to have acted as a glacial refugium for hominin populations durin... more Southern Europe is hypothesized to have acted as a glacial refugium for hominin populations during the Pleistocene. Of particular importance is SouthEast Europe, which most likely played a dual role, both as refugium and dispersal corridor, especially during the Middle Pleistocene glaciations, when drastic climatic conditions led to major sea level drops in the Aegean. However, little is known about the palaeoenvironmental conditions at the time of hominin presence in this region, making these hypotheses difficult to test. Here we analyze biomarker data and leaf wax stable isotopic compositions of the MIS 12 Lower Palaeolithic site Marathousa 1 (Megalopolis Basin, Greece) to assess the climatic conditions accompanying the time of hominin presence in the area. Our data indicate a major cooling affecting the north Mediterranean/Aegean domain during this time interval, with lowest temperatures recorded between~440-432 ka. The glacial peak is associated with changes in vegetation (i.e., from more forested to more open landscape), reduction of humidity and water availability (i.e., moisture depletion, increased evaporation). Hominins are present at the Marathousa 1 location at the end of this interval (434-432 ka), confirming that the Megalopolis Basin served as a refugium for hunter-gatherer groups during periods of harsh climatic conditions. Additionally, the progressive cooling is associated with an important sedimentary hiatus between~465-440 ka reflected in all circum-Mediterranean records (both marine and continental), indicating a regional impact of the MIS 12 glaciation over surface processes.
HAL (Le Centre pour la Communication Scientifique Directe), 2012
Historical Biology, Nov 13, 2022
The Early Pleistocene mammal communities of Europe are characterized by a great diversity of larg... more The Early Pleistocene mammal communities of Europe are characterized by a great diversity of large carnivorans. Among them, the largest ever hyaenid, Pachycrocuta brevirostris, a fierce predator with great bone-cracking adaptations that has left its taphonomic signature on several fossiliferous sites. Here, we perform a rigorous taphonomic analysis focusing on bone surface modifications and damage patterns on large-sized ungulate bones from the site Tsiotra Vryssi (1.78 to ~1.5 Ma; Mygdonia Basin, Greece), aiming to identify the main biotic agent responsible for the modifications. Results reveal significant carnivore ravaging of the assemblage, and selective consumption of bones/bone portions related to nutrient value. Comparisons with modifications on similar-sized ungulate carcasses produced by extant and extinct predators, and the similarities with Pachycrocuta-modified assemblages, indicate that Pachycrocuta was the principal agent of modification. Overall, this study not only provides taphonomic evidence for the interpretation of Tsiotra Vryssi, but also offers insights into the palaeobiology, and particularly carcass consumption behaviour of the giant hyaena. Hence, it advances our knowledge on carnivoran guild dynamics and prey-predator relationships during this epoch and has important implications for the investigations of the subsistence behaviour of the meat-eating hominins, who entered Eurasia at ~1.8 Ma, roughly synchronously with Pachycrocuta.
Quaternary International, Dec 1, 2018
Exposures of Middle Pleistocene lacustrine sediments at the margins of an open-cast lignite mine ... more Exposures of Middle Pleistocene lacustrine sediments at the margins of an open-cast lignite mine at Marathousa near Megalopolis, western Arcadia, Greece yielded the partial remains of a Palaeoloxodon antiquus skeleton which exhibited signs of being butchered. Sedimentation occurred between ca. 400 and 480 ka. Lithic artefacts were found in close spatial and stratigraphic association with the elephant remains. A palaeobotanical investigation (involving carpological, phytolith and wood remains) as well as diatom analysis led to a detailed reconstruction of the local environment at the time of sediment deposition. The results of this study enabled the environmental context of the butchering of the Palaeoloxodon antiquus carcass to be established. Palaeobotanical data show that sediment deposition at the Palaeoloxodon antiquus site occurred in shallow water in front of a reed swamp with trees in the immediate surroundings (particularly Alnus and Salix) on a flat plain where the water table was at or just below the ground surface. Warm conditions prevailed at the time of sediment deposition allowing aquatic plants typically recorded in the climatic optima of European interglacial assemblages to occur (e.g. Brasenia schreberi) and palms (Palmae) to live close-by. This suggests that the organic-rich sediments that contain the archaeological finds represent a transition between the underlying clastic sediments deposited during colder conditions and the overlying lignite that was deposited during warm climatic conditions as reported in previous studies.
American Journal of Physical Anthropology, 2012
ABSTRACT Greece lies on the hypothesized migration route of archaic humans into Europe, and its p... more ABSTRACT Greece lies on the hypothesized migration route of archaic humans into Europe, and its paleolithic record is critical inaddressing issues of timing and routes of dispersal of the earliest European colonization. The Aliakmon Paleolithic survey, conducted on the Aliakmon river terraces in 2004-05, is the first systematic survey for this time period in the region. The area preserves Plio-Pleistocene fluvial fossiliferous sediments and has yielded paleolithic artifacts. We confirmed the existence of theterrace systems at previously proposed levels but with more complex dating than originally proposed. Faunal specimens collected include equids, rhinocerotids, suids, bovids, cervids, rodents, canids, proboscideans, and cf. Hippopotamus. Two possible Lower Paleolithic sites were located. Given the scarcity of such sites in Greece, these findings are pivotal in documenting early human presence in the area.
Nature
Modern humans have populated Europe for more than 45,000 years1,2. Our knowledge of the genetic r... more Modern humans have populated Europe for more than 45,000 years1,2. Our knowledge of the genetic relatedness and structure of ancient hunter-gatherers is however limited, owing to the scarceness and poor molecular preservation of human remains from that period3. Here we analyse 356 ancient hunter-gatherer genomes, including new genomic data for 116 individuals from 14 countries in western and central Eurasia, spanning between 35,000 and 5,000 years ago. We identify a genetic ancestry profile in individuals associated with Upper Palaeolithic Gravettian assemblages from western Europe that is distinct from contemporaneous groups related to this archaeological culture in central and southern Europe4, but resembles that of preceding individuals associated with the Aurignacian culture. This ancestry profile survived during the Last Glacial Maximum (25,000 to 19,000 years ago) in human populations from southwestern Europe associated with the Solutrean culture, and with the following Magdal...
HAL (Le Centre pour la Communication Scientifique Directe), Jan 14, 2023
Science Advances
The frontal sinuses are cavities inside the frontal bone located at the junction between the face... more The frontal sinuses are cavities inside the frontal bone located at the junction between the face and the cranial vault and close to the brain. Despite a long history of study, understanding of their origin and variation through evolution is limited. This work compares most hominin species’ holotypes and other key individuals with extant hominids. It provides a unique and valuable perspective of the variation in sinuses position, shape, and dimensions based on a simple and reproducible methodology. We also observed a covariation between the size and shape of the sinuses and the underlying frontal lobes in hominin species from at least the appearance of Homo erectus . Our results additionally undermine hypotheses stating that hominin frontal sinuses were directly affected by biomechanical constraints resulting from either chewing or adaptation to climate. Last, we demonstrate their substantial potential for discussions of the evolutionary relationships between hominin species.
Words, Bones, Genes, Tools: DFG Center for Advanced Studies
The site of Apidima, in southern Greece, is one of the most important Paleolithic sites in Greece... more The site of Apidima, in southern Greece, is one of the most important Paleolithic sites in Greece and southeast Europe. One of the caves belonging to this cave complex, Cave A, has yielded human fossil crania Apidima 1 and 2, showing the presence of an early Homo sapiens population followed by a Neanderthal one in the Middle Pleistocene. Less known are the human remains reportedly recovered from Cave C at Apidima. These include a number of isolated elements, but also a partial skeleton interpreted as a female burial, Apidima 3, proposed by Pitsios (e.g., Pitsios 1999) to be associated with Aurignacian lithics and to date to ca. 30 ka. In light of the rarity of the Upper Paleolithic in Greece, and the general scarcity of human remains associated with the Aurignacian, the remains from Apidima Cave C are potentially very significant in elucidating the arrival of the early Upper Paleolithic populations in Europe. Here we undertake direct Uranium-series dating of three human samples from...
Journal of Archaeological Science, 2021
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Papers by Katerina Harvati
In this study we provide a synthesis of the Proboscidea-Homo record in Early and Middle Pleistocene open-air localities of western Eurasia, by documenting any direct (e.g., presence of cut marks, elephant bone tools, fractures for marrow extraction) and indirect (e.g., association and refitting of lithic artefacts, use-wear analysis) evidence of proboscidean carcass exploitation. Sex and ontogenetic age of butchered proboscideans are recorded, so as to assess possible preferences by humans. Furthermore, we investigate the role of the large carnivores (hyaenids, felids, large canids). We focus on important renewals in the carnivore guilds, and their significance in terms of availability of carrion for scavenging and the hominin-carnivore competition for food resources. The ecological adaptations of the two Middle Pleistocene elephantids in Europe, the straight-tusked elephant Palaeoloxodon antiquus and the steppe mammoth Mammuthus trogontherii, are also evaluated. Finally, we discuss various aspects of the evolution of Pleistocene Homo, including technological advances in material culture, important developments in cognition and relevant inferences about human social behavior. The objective of this study is to re-address the key issues in the Homo-Proboscidea research agenda, assess emerging patterns between ecological, ethological, environmental and cultural parameters, and identify potential biases that obstruct nuanced interpretations of the record.
Of particular interest are an elephant cranium and several postcranial elements, which were found in close anatomical association and are attributed to a single individual of the straight-tusked elephant Elephas (P.) antiquus. The good state of preservation of the MAR bones allows to identify taphonomic modifications. Cut marks and percussion damage indicate hominin exploitation of elephant and other mammal bones by means of butchering activities. Carnivore gnawing is also evident on some specimens, suggesting a certain degree of carnivore competition with humans for early access to the animal carcasses.