Katerina Harvati

University of Tuebingen | EKU Tübingen · Paleoanthropology, Senckenberg Center for Human Evolution and Paleoenvironment, Institute for Archaeological Sciences

Neanderthals are commonly depicted as leading dangerous lives and permanently struggling for survival. This view largely relies on the high incidences of trauma that have been reported1,2 and have variously been attributed to violent social behaviour3,4, highly mobile hunter-gatherer lifestyles2 or attacks by carnivores5. The described Neanderthal...

Two fossilized human crania (Apidima 1 and Apidima 2) from Apidima Cave, southern Greece, were discovered in the late 1970s but have remained enigmatic owing to their incomplete nature, taphonomic distortion and lack of archaeological context and chronology. Here we virtually reconstruct both crania, provide detailed comparative descriptions and an...

Previous scientific consensus saw human evolution as defined by adaptive differences (behavioural and/or biological) and the emergence of Homo sapiens as the ultimate replacement of non-modern groups by a modern, adaptively more competitive group. However, recent research has shown that the process underlying our origins was considerably more compl...

The Middle and Late Pleistocene is arguably the most interesting period in human evolution. This broad period witnessed the evolution of our own lineage, as well as that of our sister taxon, the Neanderthals, and related Denisovans. It is exceptionally rich in both fossil and archaeological remains, and uniquely benefits from insights gained throug...

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 sca...

The early Iron Age (800 to 450 BCE) in France, Germany and Switzerland, known as the ‘West-Hallstattkreis’, stands out as featuring the earliest evidence for supra-regional organization north of the Alps. Often referred to as ‘early Celtic’, suggesting tentative connections to later cultural phenomena, its societal and population structure remain e...

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...

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 simil...

An intensive, target-oriented surface survey conducted in the Megalopolis basin during 2012-2013 led to the discovery of several Palaeolithic sites and findspots with lithics and faunal remains, including Marathousa-1, a Lower Palaeolithic open-air elephant-butchering site, dated to ca. 400-500 ka BP. This study presents the results from the techno...

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 com...

Reconstructing habitual physical activities in the past constitutes a fundamental objective of anthropological sciences. The morphology of muscle attachment sites (entheses) is widely utilized for this purpose, but their reliability has been previously questioned due to important methodological downsides of traditional methodological approaches. Re...

Neanderthals’ lives 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. Her...

The use of reindeer has been a crucial element in the subsistence strategies of past Arctic and Subarctic populations. However, the spatiotemporal occurrence of systematic herding practices has been difficult to identify in the bioarchaeological record. To address this research gap, this study proposes a new virtual anthropological approach for rec...

The Early Pleistocene mammal communities of Europe are characterised 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...

There is a growing consensus that global patterns of modern human cranial and dental variation are shaped largely by neutral evolutionary processes, suggesting that craniodental features can be used as reliable proxies for inferring population structure and history in bioarchaeological, forensic, and paleoanthropological contexts. However, there is...

Unlabelled: Birch tar is the oldest synthetic substance made by early humans. The earliest such artefacts are associated with Neanderthals. According to traditional interpretations, their study allows understanding Neanderthal tool behaviours, skills and cultural evolution. However, recent work has found that birch tar can also be produced with si...

In this article we present the new open-air Middle Pleistocene locality Marathousa 2, which was discovered during a double intensive and targeted field survey in the lignite mines of the Megalopolis Basin (Greece). The locality is situated just below the Lignite Seam III of the Marathousa Member (Choremi Formation), and its similar stratigraphic po...

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...

The use of non-destructive approaches for digital acquisition (e.g. computerised tomography-CT) allows detailed qualitative and quantitative study of internal structures of skeletal material. Here, we present a new R-based software tool, Icex, applicable to the study of the sizes and shapes of skeletal cavities and fossae in 3D digital images. Trad...

Objectives This study uses a virtual framework to examine the left maxillary fragment of the juvenile fossil from Mugharet el'Aliya, Morocco, found in association with an Aterian lithic industry. Previously, this fossil had been ascribed to modern humans or the Neanderthal lineage based on its “archaic”/“Neanderthal‐like” features and apparent larg...

Small flake industries are a commonly identified component of Lower Paleolithic archaeological assemblages in Eurasia. Utilized as blanks for tools, at many sites, their functions are often poorly understood. Here we present a preliminary traceological analysis of lithics from Marathousa 1 (MAR-1; Megalopolis, Greece). MAR-1 dates to ca. 400-500 ka...

Systematic tool use is a central component of the human niche. However, the timing and mode of its evolution remain poorly understood. A newly developed method for the analysis of muscle recruitment patterns (Validated Entheses-based Reconstruction of Activity-V.E.R.A.) has recently been experimentally shown to provide clear and reliable evidence o...

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....

A partial proboscidean skeleton has been recovered during the systematic excavations at the Middle Pleistocene open-air site Marathousa 1 (MAR-1) in the Megalopolis basin (Peloponnese, Greece). The remains of the individual, an adult male of the European straight-tusked elephant Palaeoloxodon antiquus, display human-induced cut marks, which, togeth...

Population affinity identification is important for reconstructing the biological profile of human skeletal remains. Most anthropological methods for predicting population affinity rely on complete crania or cranial parts. However, complete parts are frequently not found in forensic and bioarchaeological contexts. In contrast, the petrous portion o...

The Megalopolis basin (Peloponnese, Greece) is known for its Pleistocene fossiliferous deposits. The basin’s stratigraphic sequence comprises fluviolacustrine deposits containing lignite seams and spans from ca. 900 ka to ca. 150 ka, thus covering part of the Early and the entire Middle Pleistocene. Since 2012 the basin has been investigated for st...

This is an exciting time for our understanding of the origin of our species. Previous scientific consensus saw human evolution as defined by adaptive differences (behavioural and/or biological) and the emergence of Homo sapiens as the ultimate replacement of non-modern groups by a modern, adaptively more competitive one. However, recent research ha...

In the published manuscript, the study presents diverse geochronological and biochronological data providing age constraints on the site of Tsiotra Vryssi (Mygdonia basin, Greece). One of the methods presented is based on burial ages from cosmogenic radionuclides. Table 2 of this study reports cosmogenic simple burial ages of 1.88 ± 0.16 Myr, 2.10...

In this article, we describe an almost complete macaque mandible from the Middle Pleistocene locality Marathousa 1 in the Megalopolis Basin of southern Greece. The mandible belonged to a male individual of advanced ontogenetic age and of estimated body mass~13 kg. Comparative metric analysis of its teeth permits its attribution to the Barbary macaq...

Human paleontology is a relatively limited field, given that human fossil remains tend to be extremely rare. Furthermore, it differs from paleontology in that it is intrinsically bound with Paleolithic archaeology, at least for the later parts of human evolution, and cannot be considered independently from human early prehistory. Modern paleoanthro...

This paper presents the earliest evidence for the exploitation of lignite (brown coal) in Europe and sheds new light on the use of combustion fuel sources in the 2nd millennium BCE Eastern Mediterranean. We applied Thermal Desorption/Pyrolysis–Gas Chromatography-Mass Spectrometry and Polarizing Microscopy to the dental calculus of 67 individuals an...

The sedimentary basins of Greece contain an important record of fossil vertebrates that has been known and studied for nearly two centuries. Here, we present our collective effort to review and summarize this fossil record. A combination of our original research and previously published records permits the complete reassessment of the identified ve...

Objectives: To explore mandibular shape differences between Ouranopithecus macedoniensis and a comparative sample of extant great apes using three-dimensional (3D) geometrics morphometrics. Other objectives are to assess mandibular shape variation and homogeneity within Ouranopithecus, explore the effects of size on mandibular shape, and explore th...

Commingled remains describes the situation of intermixed skeletal elements, an extremely common occurrence in contemporary forensic cases, archaeological mass graves, as well as fossil hominin assemblages. Given that reliable identification is typically impossible for commingled contexts, a plethora of previous studies has focused on the developmen...

A recent protocol of digital restoration is applied to the Middle Pleistocene human cranium from Steinheim (Germany). The retrodeformation of the specimen sheds new light on the taphonomic origin of some peculiar features observed on the cranium and returned a morphology consistent with its attribution to the Neanderthal lineage.

Marathousa 1 (MAR-1), located in the Megalopolis palaeolake basin, southern Greece, preserves an exceptionally well-preserved archaeological and palaeontological assemblage. Radiometric dating, magnetostratigraphy, and geological and biochronological analyses indicate an age of 500-400 ka, and place the locality within the Marine Isotope Stage (MIS...

The lithic industries of Lower Palaeolithic Europe are characterized by a wide range of diversity. Biface assemblages stand alongside small tool assemblages and core-and-flake industries. This variability is still poorly understood, hindering the assessment of hominin techno-economical choices. Recent studies on lithic variability of the Middle Ple...

Traces of Neandertal cultural and skeletal remains are found in most European countries, covering a time-span of more than 250,000 years. Since the recognition of Neandertals as a specific human group, following discoveries in the Neander valley in 1856, much works has been taken on many aspects of their heritage (cultural, anthropological, genetic...

A number of different approaches are currently available to digitally restore the symmetry of a specimen deformed by taphonomic processes. These tools include mirroring and retrodeformation to approximate the original shape of an object by symmetrisation. Retrodeformation has the potential to return a rather faithful representation of the original...

The series aims to further the foundations for a cross-disciplinary field of bio- cultural co-evolution by addressing topics on linguistic, cultural, and biological trajectories of the human past. It aims to push the limits of cooperation between traditional disciplines, bringing together reviews, original research, and perspectives from scholars i...

The series aims to further the foundations for a cross-disciplinary field of bio- cultural co-evolution by addressing topics on linguistic, cultural, and biological trajectories of the human past. It aims to push the limits of cooperation between traditional disciplines, bringing together reviews, original research, and perspectives from scholars i...

In recent decades, a significant number of Pleistocene (ca. 2.6 million years–10,000 years ago) open-air and cave sites yielding elephant or mammoth bones in direct association with hominin remains and/or lithic artifacts have been discovered in Eurasia, Africa and America. Many of them show strong evidence of acquisition and processing of probosci...

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...

Lithics and cut-marked mammal bones, excavated from the paleo-lake Marathousa 1 (MAR-1) sediments in the Megalopolis Basin, southern Greece, indicate traces of hominin activity occurring along a paleo-shoreline ca. 444,000 years (444 ka) ago. However, the local environment and climatic conditions promoting hominin activity in the area during the MI...

The sedimentary basins of Greece contain an important record of fossil vertebrates that has been known and studied for nearly two centuries. Here, we present our collective effort to review and summarize this fossil record. A combination of our original research and previously published records permits the complete reassessment of the identified ve...

Until the early 5th century BC, Phaleron Bay was the main port of ancient Athens (Greece). On its shore, archaeologists have discovered one of the largest known cemeteries in ancient Greece, including a range of burial forms, simple pits, cremations, larnaces (clay tubs), and series of burials of male individuals who appear to have died violent dea...

Significance The microbiome plays key roles in human health, but little is known about its evolution. We investigate the evolutionary history of the African hominid oral microbiome by analyzing dental biofilms of humans and Neanderthals spanning the past 100,000 years and comparing them with those of chimpanzees, gorillas, and howler monkeys. We id...

Interview with paleoanthropologist Katerina Harvati, who studies Neanderthal evolution and modern human origins at the Eberhard Karls University of Tübingen.

Material belonging to the late Miocene hominoid Ouranopithecus macedoniensis has been poorly analyzed using advanced techniques. This study aims to explore mandibular shape variation between Ouranopithecus macedoniensis and a comparative sample of extant great apes, using three-dimensional (3D) geometric morphometrics. Other objectives are to asses...

An accurate reconstruction of habitual activities in past populations and extinct hominin species is a paramount goal of paleoanthropological research, as it can elucidate the evolution of human behavior and the relationship between culture and biology. Variation in muscle attachment (entheseal) morphology has been considered an indicator of habitu...

Systematic tool production and use is one of humanity’s defining characteristics, possibly originating as early as >3 million years ago. Although heightened manual dexterity is considered to be intrinsically intertwined with tool use and manufacture, and critical for human evolution, its role in the emergence of early culture remains unclear. Most...

Background and scope: The late Villafranchian large mammal age (~2.0–1.2 Ma) of the Early Pleistocene is a crucial interval of time for mammal/hominin migrations and faunal turnovers in western Eurasia. However, an accurate chronological framework for the Balkans and adjacent territories is still missing, preventing pan-European biogeographic corre...

KNM-OG 45500 is a hominin fossil composed of parts of a frontal bone, left temporal bone, and cranial vault pieces. Since its discovery along the Olorgesailie Formation (Kenya) in 2003, it has been associated with the Homo erectus hypodigm. The specimen, derived from a geological context dated to ca. 900 Ka BP, has been described as a very small in...

LETTER TO THE EDITOR Cioclovina fractures: Reply to Soficaru and Trinkaus: Perimortem versus postmortem damage: The recent case of Cioclovina 1, Am J Phys Anthropol 2020 172, 135–139 In our recent article (Kranioti et al., 2019), we conducted an exhaustive investigation of the fracture patterns presented by the Upper Paleolithic calvaria from Ciocl...

Report on the 2014 excavation campaign at the Lower Palaeolithic site Marathousa 1 (Megalopolis Basin, Greece).

Objectives Cuncaicha, a rockshelter site in the southern Peruvian Andes, has yielded archaeological evidence for human occupation at high elevation (4,480 masl) during the Terminal Pleistocene (12,500–11,200 cal BP), Early Holocene (9,500–9,000 cal BP), and later periods. One of the excavated human burials (Feature 15‐06), corresponding to a middle...

Objectives: This study characterizes patterns of cranial trauma prevalence in a large sample of Upper Paleolithic (UP) fossil specimens (40,000-10,000 BP). Materials and Methods: Our sample comprised 234 individual crania (specimens), representing 1,285 cranial bones (skeletal elements), from 101 Eurasian UP sites. We used generalized linear mixed...

Background: Recent advances in sequencing have facilitated large-scale analyses of the metagenomic composition of different samples, including the environmental microbiome of air, water, and soil, as well as the microbiome of living humans and other animals. Analyses of the microbiome of ancient human samples may provide insights into human health...

Despite a steady increase in our understanding of the phenotypic variation of Pleistocene Homo, debate continues over phylogenetically informative features. One such trait is the suprainiac fossa, a depression on the occipital bone above inion that is commonly considered an autapomorphy of the Neanderthal lineage. Challenging this convention, depre...

Previous research found sexual dimorphism in the bony labyrinth of a Greek population sample (Osipov et al. 2013). This study intends to investigate the nature of this structure’s sexual dimorphism across populations of diverse geographic origin and to identify the effect of inter-population variation on the accuracy of determining sex using the bo...

Many authors have hypothesized an association between rates of morphological evolution and rates of species diversification, however, this association has yet to be empirically tested in the primate cranium. In this investigation, we used phylogeny-based approaches to examine the relationship between rates of species diversification, rates of crani...

The aim of this paper is to evaluate the craniometric affinities of the only Cuncaicha cranial specimen with other early, middle, and late Holocene South American samples. To do so, the skull was first reconstructed by using computer-aided techniques applied to several μ-CT-scanned fragments. Linear measurements were calculated in the facial skelet...

The morphology of entheses (muscle/tendon attachment sites) on bones is routinely used in paleontological and bioarcheological studies to infer the physical activity patterns of ancient vertebrate species including hominins. However, such inferences have often been disputed due to limitations of the quantitative methods commonly employed and a lack...

Reconstructions of habitual activity in past populations and extinct human groups is a primary goal of paleoanthropological research. Muscle attachment scars (entheses) are widely considered as indicators of habitual activity and many attempts have been made to use them for this purpose. However, their interpretation remains equivocal due to method...

The morphology of entheses (muscle/tendon attachment sites) on bones is routinely used in paleontological and bioarcheological studies to infer the physical activity patterns of ancient vertebrate species including hominins. However, such inferences have often been disputed due to limitations of the quantitative methods commonly employed and a lack...

Objectives: We are testing competing scenarios regarding the population history of the ancient Greek colonization of southern Italy using dental phenotypic evidence. Materials and methods: We collected dental metric and nonmetric trait data for 481 human skeletons from six archaeological sites along the Gulf of Taranto, dating to pre-colonial (9...

Objective: This study aims to develop a comparative basis for assessing the developmental stage of KNM-ER 42700 based on the ontogenetic pattern of the ectocranial surface of the basicranium in modern humans and chimpanzees. Materials and methods: A total of 33 landmarks were collected from an ontogenetic sample of modern humans (80), chimpanzee...

In recent years, a significant number of Pleistocene localities with evidence of proboscidean exploitation by humans have been discovered, substantially enriching our knowledge on Homo subsistence strategies and interactions with megaherbivores. However, the human engagement in proboscidean assemblages, as well as the degree of interaction is not a...

Following the discovery and excavation of the Lower Palaeolithic butchering locality Marathousa 1 (MAR-1; Megalopolis Basin, Peloponnesus, Greece), conducted by the Ephoreia of Paleoanthropology and Speleology, Greek Ministry of Culture in collaboration with the American School of Classical Studies at Athens, led to the discovery of a new open-air...

Following the discovery and excavation of the Lower Palaeolithic butchering locality Marathousa 1 (MAR-1; Megalopolis Basin, Peloponnesus, Greece), conducted by the Ephoreia of Paleoanthropology and Speleology, Greek Ministry of Culture in collaboration with the American School of Classical Studies at Athens, led to the discovery of a new open-air...

Objectives: This study aims to virtually reconstruct the deformed face (XIR-1) and maxilla (RPl-128) of the Late Miocene hominoid Ouranopithecus macedoniensis from Greece, through the application of mirror-imaging and segmentation. Additionally, analysis was conducted through 3D geometric morphometrics, utilizing a comparative sample of fossil hom...

The Cioclovina (Romania) calvaria, dated to ca. 33 cal ka BP and thought to be associated with the Aurignacian lithic industry, is one of the few relatively well preserved representatives of the earliest modern Europeans. Two large fractures on this specimen have been described as taphonomic modifications. Here we used gross and virtual forensic cr...

The fossil record of early Homo sapiens in the African Pleistocene remains sparse. In contrast to its prominent position regarding the cultural evolution of our species, southern Africa plays a secondary role in narratives regarding human biological origins. Reasons for this are a limited and fragmentary fossil record from the Middle Stone Age (MSA...

The calvaria from Cioclovina (Romania) has been argued to possess some traits commonly ascribed to individuals belonging to the Neanderthal lineage, including a suprainiac fossa. However, its supranuchal morphology has only been evaluated with a qualitative analysis of the ectocranial surface. We evaluate whether the morphology of the supranuchal a...

Our current knowledge of the emergence of anatomically modern humans, and the human lineage in general, is limited, in large part because of the lack of a well preserved and well dated fossil record from Pleistocene Africa. Thus, the primary aim of our research is to partly relieve this problem by virtually reconstructing and analyzing the hominin...

The Cioclovina (Romania) calvaria, dated to 33.212 ± 6993 cal BP, has been argued to possess a horizontal rugose concavity directly above inion, known as a suprainiac fossa. This feature has been proposed as a phenotypic marker for introgression of substantial levels of Neanderthal genetic material in modern human populations. A quantitative assess...

The face is the most distinctive feature used to identify others. Modern humans have a short, retracted face beneath a large globular braincase that is distinctively different from that of our closest living relatives. The face is a skeletal complex formed by 14 individual bones that houses parts of the digestive, respiratory, visual and olfactory...

By applying advanced spatial statistical methods, spatial taphonomy complements the traditional taphonomic approach and enhances our understanding of biostratinomic and diagenetic processes. In this study, we elaborate on a specific aspect – spatial anisotropy – of taphonomic processes. We aim to unravel the taphonomic history of the Early Pleistoc...

The Villafranchian ursids of Greece are scanty, represented by a few isolated teeth and bones. During our last field campaigns in the Mygdonia Basin (Macedonia, Greece) we discovered an almost complete cranium, as well as some cranial, dental and postcranial remains, which are presented in this article. The new material originates from the locality...

In this article, we present the first results on the large mammal fauna from the new open-air Lower Palaeolithic locality Marathousa 1 (MAR-1) (Megalopolis Basin, Peloponnesus, Greece). MAR-1 belongs to the Marathousa Member of the Choremi Formation and its large mammal faunal list (collection 2013–2016) includes the castorid Castor fiber, the must...