Mietje Germonpré | Royal Belgian Institute of Natural Sciences (original) (raw)

Papers by Mietje Germonpré

Bulletin-Institut royal des sciences naturelles de Belgique. Sciences de la terre, 1993

Large quantities of fossil bones were recovered in the Flemish Valley at the sites of Zemst, Hofs... more Large quantities of fossil bones were recovered in the Flemish Valley at the sites of Zemst, Hofstade, Overmere and Dendermonde. Most of these Late Pleistocene assemblages date from the Weichselian and were recovered from fluvial deposits formed by braided river Systems. Twenty-three mammalian taxa were collected. The fossils accumulated mainly through graduai, long-term processes as indicated by the scattered and dispersed spatial distribution of the bones, the abundance of scavenged and subaerially weathered bones, the low numbers of carnivores, the low CSI (corrected number of specimens per individual) values and the prépondérance of Voorhies (1969) Group 3. Age profiles of mammoth support this interprétation. However, the age profile of rhinocéros of assemblage Hofstade I suggests episodic mass-mortality events affecting this large herbivore.

Frontiers in Psychology, 2023

The historically known relationship of interspecies companionship between Aboriginal foraging com... more The historically known relationship of interspecies companionship between Aboriginal foraging communities in Australia and free-ranging dingoes provides a model for understanding the human-canid relations that gave rise to the first domesticated dogs. Here, we propose that a broadly similar relationship might have developed early in time between wild-living wolves and mobile groups of foragers in Late Pleistocene Eurasia, with hunter-gatherers routinely raiding wild wolf dens for pre-weaned pups, which were socialized to humans and kept in camp as tamed companions ("pets"). We outline a model in which captive wolf pups that reverted to the wild to breed when they were sexually mature established their territories in the vicinity of foraging communities-in a "liminal" ecological zone between humans and truly wild-living wolves. Many (or most) of the wolf pups humans took from the wilderness to rear in camp may have derived from these liminal dens where the breeding pairs had been under indirect human selection for tameness over many generations. This highlights the importance of the large seasonal hunting/aggregation camps associated with mammoth kill-sites in Gravettian/Epigravettian central Europe. Large numbers of foragers gathered regularly at these locations during the wild wolf birthing season. We infer that if a pattern of this kind occurred over long periods of time then there might have been a pronounced effect on genetic variation in free-ranging wolves that denned and whelped in the liminal zones in the vicinity of these human seasonal aggregation sites. The argument is not that wolves were domesticated in central Europe. Rather, it is this pattern of hunter-gatherers who caught and reared wild wolf pups gathering seasonally in large numbers that might have been the catalyst for the early changes leading to the first domesticated dogs-whether in western Eurasia or further afield.

The Anatomical Record, 2022

Janssens et al. (2021, doi: 10.1002/ar.24624) recently commented on our article (Galeta et al., 2... more Janssens et al. (2021, doi: 10.1002/ar.24624) recently commented on our article (Galeta et al., 2021, doi: 10.1002/ar.24500) regarding the morphological differences between putative Paleolithic dog and Pleistocene wolf crania. The authors argued that these differences reflect the normal population variation of wolves, that some of the cranial measurements used do not reflect morphological changes during domestication, and that our canid dataset was small because we
inexplicably omitted several specimens we analyzed in our previous publications. In this commentary, we briefly address the issue of within and between morpho-population variability.
The results based on our canid sample suggest that the magnitude of morphological differences between distinct morpho-populations (i.e., recent northern dogs and wolves) is at least twice as
large as that observed within morpho-populations (between two groups of recent northern wolves segregated by cluster analysis). The morphological differences between putative Paleolithic dogs
and Pleistocene wolves are relatively large, which may indicate that they did not likely represent a single Late Pleistocene morpho-population. Finally, we clarified the rationale behind the composition of our 2021 dataset to show that we did not adjust the list of the analyzed specimens. Although the sample size was small, the randomization analysis published in 2021 confirmed that the unbalanced composition of the reference sample did not affect the reliability of the morphological segregation of putative Paleolithic dogs and Pleistocene wolves.

Monographien des RGZM, 2021

Most researchers accept that by the end of the Pleistocene dogs were part of the daily life of pr... more Most researchers accept that by the end of the Pleistocene dogs were part of the daily life of prehistoric hunter-gatherers. Recent analyses of the mammal assemblages from the third cave of Goyet (Belgium) reveal that a large component of the material from bone level A1 postdates the Last Glacial Maximum. The biometric study of the large canid remains from this level shows that an ulna can be described as from a medium-sized Palaeolithic dog with an estimated body mass of ~ 20 kg. A direct AMS 14 C date of the bone demonstrates that the dog lived during the Bølling / Allerød interstadial. Human and carnivore modifications of the bone indicate that the animal was dismembered by a contemporaneous human individual, likely to obtain its meat, and then gnawed by a canid-sized carnivore. Presumably, Palaeolithic dogs fulfilled diverse roles in Late Palaeolithic societies including as a source of food.

Human-elephant interactions: from past to present, 2021

Middle and Upper Palaeolithic sites, where mammoths dominate the faunal assemblages, are mainly f... more Middle and Upper Palaeolithic sites, where mammoths dominate the faunal assemblages, are mainly found in Central and Eastern Europe. At these sites concentrations of skulls, tusks and long bones, interpreted as deliberate constructions, often occur. Rare instances of weapon tip fragments embedded in mammoth bones provide direct archaeological evidence of human hunting. Indirect evidence, such as the accumulation of mammoth bones from multiple individuals with specific ontogenetic ages, occurs more frequently. Based on the eruption sequence and wear of deciduous premolars from mammoth calves, we examined whether a season of death could be deduced from the characteristics of the dentition. Our results suggest that the mammoth hunt was not restricted to the cold half of the year.

Umschlagbild: Ein europäischer Waldelefant (Palaeoloxodon antiquus) und ein Hominin aus dem Mitte... more Umschlagbild: Ein europäischer Waldelefant (Palaeoloxodon antiquus) und ein Hominin aus dem Mittelpleistozän (Vorder-und Rückseitenabbildungen von Gleiver Prieto; Copyright Katerina Harvati, Universität Tübingen) Druck und Bindung: readbox unipress in der readbox publishing GmbH Printed in Germany Bibliogra sche Information der Deutschen Nationalbibliothek Die Deutsche Nationalbibliothek verzeichnet diese Publikation in der Deutschen Nationalbibliogra e, detaillierte bibliogra sche Daten sind im Internet über http://dnb.d-nb.de abrufbar. e symposium and the volume "Human-elephant interactions: from past to present" were funded by the Volkswagen Foundation.

Journal of Anthropological Archaeology, 2021

Two canid morphotypes have been proposed for the middle Upper Palaeolithic site of Pˇredmostí (Mo... more Two canid morphotypes have been proposed for the middle Upper Palaeolithic site of Pˇredmostí (Moravia, Czech Republic): Pleistocene wolf and Palaeolithic dog (Germonpr ́e et al., 2012, 2015; Galeta et al., 2020). In Wilc-zy ́nski et al. (2020), faunal assemblages from other Upper Palaeolithic Moravian sites were analyzed and those results used to project a similar, which we interpret as flawed, result for Pˇredmostí: that only wild canids were present. We address issues with their methodology and argue that their conclusion, that dogs were not present at any of those studied sites, is based on cursory taphonomic analysis. Further, their projection of these results to Pˇredmostí, an assemblage they did not study, is unsubstantiated, but does highlight the current dearth of taphonomic information that could aid the study of dog domestication. We also point out an important error in their Fig. 1, where the hemimandibles intended to show the natural variability in large canids from Dolní Vĕstonice II and Pavlov I SE, are in fact canid specimens from Pˇredmostí. Since several of these canid specimens have been published by Germonpr ́e et al. (2015), we find it imperative to address this error to ensure these data are correctly referenced in future works.

Human Ecology, 2021

Several hypotheses have been proposed to explain the initial steps in the domestication process o... more Several hypotheses have been proposed to explain the initial steps in the domestication process of the wolf. We discuss the human-initiated model in which wolf pups were brought to camp sites by male hunters and cared for by nursing women. A good relation between the more sociable and playful pups and the women and their children likely formed affiliative bonds and led to the survival of such pups into maturity. Some of these animals could have reproduced and delivered at least one litter. A selection on the behaviour of subsequent generations could ultimately have led to Palaeolithic dogs.

Journal of Archaeological Science, 2021

Canids from the Upper Paleolithic site of Predmostí are central to debates concerning the domesti... more Canids from the Upper Paleolithic site of Predmostí are central to debates concerning the domestication of dogs as two morphotypes were identified: Pleistocene wolf and Paleolithic dog (Germonpr´e et al., 2015a). In Prassack et al. (2020), we set out to determine whether specimens previously parsed into these two groups differed
significantly in dental microwear textures, a proxy for diet. We did not assume that one group was comprised of dogs, but hypothesized that if they were, they would likely have consumed more bone, leading to microwear surface textures dominated by pitting. We indeed found significantly higher scales of maximum complexity on second lower molar crushing surfaces of the sample identified as Paleolithic dogs by Germonpr´e et al. (2015a),
consistent with larger pits on average and more bone consumption. These results suggest that the two morphotypes
identified by Germonpr´e et al. (2015a) represent ecologically distinct populations. This is in accord
with the interpretation of domestication, but, as we noted, the groups could also represent two distinct wild canid
populations with differing diets. Janssens et al. (2021) recently criticized our study, questioning our methods of
analysis and claiming bias in our interpretation of results. We reply to their issues here, focusing only on those
relevant to Prassack et al. (2020).

Social inequality before farming? Multidisciplinary approaches to the study of social organization in prehistoric and ethnographic hunter-gatherer-fisher societies Edited by Luc Moreau, 2020

The dog is the only domesticated species that dates from before the origin of agriculture when hu... more The dog is the only domesticated species that dates from
before the origin of agriculture when human populations
were living as hunter-gatherers (e.g. Germonpré
et al. 2009, 2015, 2018; Thalmann et al. 2013; Freedman
and Wayne 2017). Morphological and genetic analyses
have shown that dogs descent from an extinct Eurasian
Pleistocene wolf population or possibly several populations
(e.g. Germonpré et al. 2009; Thalmann et al. 2013;
Skoglund et al. 2015; Frantz et al. 2016). Although the
dogs’ ancestor is now known, many questions remain,
such as how the first dogs could have participated in
the daily life of their owners (Losey et al. 2018).
In this contribution, we first summarize two
models on the origin of the dog; then we detail two
canid morphotypes from the Upper Palaeolithic; next,
we look to the Upper Palaeolithic sites that have evidence
for the presence of incipient dogs. After that, we
review ethnographic sources for dog-related practices
among Northern societies and whether and how these
could enhance differential access to resources and
influence social status distinctions. Subsequently, we
discuss what could have been the roles of early dogs
in some Upper Palaeolithic societies. Finally, we propose
a tentatively narrative on how the contributions
of Palaeolithic dogs could have affected differential
wealth and influenced social distinction among past
men and women.

Science, 2020

Dogs were the first domestic animal, but little is known about their population history and to wh... more Dogs were the first domestic animal, but little is known about their population history and to what extent it was linked to humans. We sequenced 27 ancient dog genomes and found that all dogs share a common ancestry distinct from present-day wolves, with limited gene flow from wolves since domestication but substantial dog-to-wolf gene flow. By 11,000 years ago, at least five major ancestry lineages had diversified, demonstrating a deep genetic history of dogs during the Paleolithic. Coanalysis with human genomes reveals aspects of dog population history that mirror humans, including Levant-related ancestry in Africa and early agricultural Europe. Other aspects differ, including the impacts of steppe pastoralist expansions in West and East Eurasia and a near-complete turnover of Neolithic European dog ancestry.

Current Biology, 2020

Extant Canis lupus genetic diversity can be grouped into three phylogenetically distinct clades: ... more Extant Canis lupus genetic diversity can be grouped into three phylogenetically distinct clades: Eurasian and American wolves and domestic dogs.1
Genetic studies have suggested these groups trace their origins to a wolf population that expanded during the last glacial maximum (LGM)1
, 2
, 3
and replaced local wolf populations.4
Moreover, ancient genomes from the Yana basin and the Taimyr peninsula provided evidence of at least one extinct wolf lineage that dwelled in Siberia during the Pleistocene.3
5
Previous studies have suggested that Pleistocene Siberian canids can be classified into two groups based on cranial morphology. Wolves in the first group are most similar to present-day populations, although those in the second group possess intermediate features between dogs and wolves.6
7
However, whether this morphological classification represents distinct genetic groups remains unknown. To investigate this question and the relationships between Pleistocene canids, present-day wolves, and dogs, we resequenced the genomes of four Pleistocene canids from Northeast Siberia dated between >50 and 14 ka old, including samples from the two morphological categories. We found these specimens cluster with the two previously sequenced Pleistocene wolves, which are genetically more similar to Eurasian wolves. Our results show that, though the four specimens represent extinct wolf lineages, they do not form a monophyletic group. Instead, each Pleistocene Siberian canid branched off the lineage that gave rise to present-day wolves and dogs. Finally, our results suggest the two previously described morphological groups could represent independent lineages similarly related to present-day wolves and dogs.

The Anatomical Record, 2020

The antiquity of the wolf/dog domestication has been recently pushed back in time from the Late U... more The antiquity of the wolf/dog domestication has been recently pushed back in time from the Late Upper Paleolithic (~14,000 years ago) to the Early Upper Paleolithic (EUP; ~36,000 years ago). Some authors questioned this early dog domestication claiming that the putative (EUP) Paleolithic dogs fall within the morphological range of recent wolves. In this study, we reanalyzed a data set of large canid skulls using unbalanced-and balanced-randomized discriminant analyses to assess whether the putative Paleolithic dogs are morphologically unique or whether they represent a subsample of the wolf morpho-population. We evaluated morphological differences between 96 specimens of the 4 a priori reference groups (8 putative Paleolithic dogs, 41 recent northern dogs, 7 Pleistocene wolves, and 40 recent northern wolves) using discriminant analysis based on 5 ln-transformed raw and allometrically size-adjusted cranial measurements. Putative Paleolithic dogs are classified with high accuracies (87.5 and 100.0%, cross-validated) and randomization experiment suggests that these classification rates cannot be exclusively explained by the small and uneven sample sizes of
reference groups. It indicates that putative Upper Paleolithic dogs may represent a discrete canid group with morphological signs of domestication (a relatively shorter skull and wider palate and braincase) that distinguish them from sympatric Pleistocene wolves. The present results add evidence to the view that these specimens could represent incipient Paleolithic dogs that were involved in daily
activities of European Upper Paleolithic forager groups.

Journal of Archaeological Science, 2020

Morphological and genetic evidence put dog domestication during the Paleolithic, sometime between... more Morphological and genetic evidence put dog domestication during the Paleolithic, sometime between 40,000 and 15,000 years ago, with identification of the earliest dogs debated. We predict that these earliest dogs (referred to herein as protodogs), while potentially difficult to distinguish morphologically from wolves, experienced behavioral shifts, including changes in diet. Specifically, protodogs may have consumed more bone and other less desirable scraps within human settlement areas. Here we apply Dental Microwear Texture Analysis (DMTA) to canids from the Gravettian site of Předmostí (approx. 28,500 BP), which were previously assigned to the Paleolithic dog or Pleistocene wolf morphotypes. We test whether these groups separate out significantly by diet-related variation in microwear patterning. Results are consistent with differences in dietary breadth, with the Paleolithic dog morphotype showing evidence of greater durophagy than those assigned to the wolf morphotype. This supports the presence of two morphologically and behaviorally distinct canid types at this middle Upper Paleolithic site. Our primary goal here was to test whether these two morphotypes expressed notable differences in dietary behavior. However, in the context of a major Gravettian settlement, this may also support evidence of early stage dog domestication. Dental microwear is a behavioral signal that may appear generations before morphological changes are established in a population. It shows promise for distinguishing protodogs from wolves in the Pleistocene and domesticated dogs from wolves elsewhere in the archaeological record.

Lengths, widths, and size proportions (length to width) of the lower carnassial were measured in ... more Lengths, widths, and size proportions (length to width) of the lower carnassial were measured in 45 teeth of the arctic fox and 35 teeth of the red fox from Belgium radiocarbon dated to 46 640-14 120 ka BP. Data the Late Pleistocene foxes from Belgium were compared to 20 ancient and extant populations form Europe, Asia, and North America. The Pleistocene arctic fox from Belgium showed larger carnassial than in all recent samples of this species, whereas the Belgian fossil red foxes were characterized by the carnassial size comparable to that of the recent Siberian red foxes. Both fox species from the Pleistocene of Belgium showed the highest index of the carnassials length to width, which means increase in carnivorous adaptation. We conclude that the higher level of carnivorous specialization reached by the Belgian arctic and red foxes at the end of the Late Pleistocene reflected their scavenging on kills of large carnivores and human hunters (remains of megafauna). Harsh environmental conditions of that period and specific composition of ecosystems led to adapting to a more carnivorous food niche in both foxes.

Journal of Archaeological Science: Reports, 2020

The Trou de Chaleux is a cave site located in Belgium. It delivered a rich late Magdalenian mater... more The Trou de Chaleux is a cave site located in Belgium. It delivered a rich late Magdalenian material culture constituted mainly of lithic artefacts but also including bone industries and figurative art. This paper presents the results of the analysis of the large collection of bird remains recovered by E. Dupont in 1865, which was yet unstudied from taphonomical and archaeozoological perspectives. In addition to the taxonomic identification, surface alterations were investigated based on a macro-and microscopic analysis, including an analysis of wear traces and elementary composition. Special attention is devoted to the presence of human modifications such as disarticulation or butchering marks, traces of heating, presence of colourants and traces of bone working. The taphonomic history of the bird assemblage is reconstructed and the use of birds by humans characterized, as well as their importance in past human activities. We also discuss evidence for seasonal exploitation and for reconstructing the local environment and integrate our results with evidence from other Magdalenian assemblages from northwestern Europe. At Trou de Chaleux, birds were used for food, as raw material for bone working and for symbolic purposes. The exploitation of avian products was intense, and species have been used for several purposes such as the raven and snowy owl having been exploited both for food and for symbolic reasons. Large bird bones were used as raw material to produce artefacts, but the use-wear analysis did not evidence un-ambiguous traces related to the use of the objects produced. Despite several limiting factors, the bird material from Trou de Chaleux considerably increases the knowledge of past human exploitation of birds during the late Magdalenian in northwestern Europe.

Plos Biology, 2019

While sequencing ancient DNA (aDNA) from archaeological material is now commonplace, very few att... more While sequencing ancient DNA (aDNA) from archaeological material is now commonplace, very few attempts to sequence ancient transcriptomes have been made, even from typically stable deposition environments such as permafrost. This is presumably due to assumptions that RNA completely degrades relatively quickly, particularly when dealing with autolytic, nuclease-rich mammalian tissues. However, given the recent successes in sequencing ancient RNA (aRNA) from various sources including plants and animals, we suspect that these assumptions may be incorrect or exaggerated. To challenge the underlying dogma, we generated shotgun RNA data from sources that might normally be dismissed for such study. Here, we present aRNA data generated from two historical wolf skins, and perma-frost-preserved liver tissue of a 14,300-year-old Pleistocene canid. Not only is the latter the oldest RNA ever to be sequenced, but it also shows evidence of biologically relevant tissue specificity and close similarity to equivalent data derived from modern-day control tissue. Other hallmarks of RNA sequencing (RNA-seq) data such as exon-exon junction presence and high endogenous ribosomal RNA (rRNA) content confirms our data's authenticity. By performing independent technical library replicates using two high-throughput sequencing platforms, we show not only that aRNA can survive for extended periods in mammalian tissues but also that it has potential for tissue identification. aRNA also has possible further potential, such as identifying in vivo genome activity and adaptation, when sequenced using this technology.

Cell, 2019

Horse domestication revolutionized warfare and accelerated travel, trade, and the geographic expa... more Horse domestication revolutionized warfare and accelerated travel, trade, and the geographic expansion of languages. Here, we present the largest DNA time series for a non-human organism to date, including genome-scale data from 149 ancient animals and 129 ancient genomes (≥1-fold coverage), 87 of which are new. This extensive dataset allows us to assess the modern legacy of past equestrian civilizations. We find that two extinct horse lineages existed during early domestication, one at the far western (Iberia) and the other at the far eastern range (Siberia) of Eurasia. None of these contributed significantly to modern diversity. We show that the influence of Persian-related horse lineages increased following the Islamic conquests in Europe and Asia. Multiple alleles associated with elite-racing, including at the MSTN “speed gene,” only rose in popularity within the last millennium. Finally, the development of modern breeding impacted genetic diversity more dramatically than the previous millennia of human management.

Ecology and Evolution, 2019

The current phylogeographic pattern of European brown bears (Ursus arctos) has commonly been expl... more The current phylogeographic pattern of European brown bears (Ursus arctos) has commonly been explained by postglacial recolonization out of geographically distinct refugia in southern Europe, a pattern well in accordance with the expansion/contraction model. Studies of ancient DNA from brown bear remains have questioned this pattern, but have failed to explain the glacial distribution of mitochondrial brown bear
clades and their subsequent expansion across the European continent. We here present 136 new mitochondrial sequences generated from 346 remains from Europe, ranging in age between the Late Pleistocene and historical times. The genetic data show a high Late Pleistocene diversity across the continent and challenge the strict confinement of bears to traditional southern refugia during the last glacial maximum (LGM). The mitochondrial data further suggest a genetic turnover just before this
time, as well as a steep demographic decline starting in the mid‐Holocene. Levels of stable nitrogen isotopes from the remains confirm a previously proposed shift toward increasing herbivory around the LGM in Europe. Overall, these results suggest that in addition to climate, anthropogenic impact and inter‐specific competition may have had more important effects on the brown bear's ecology, demography, and genetic structure than previously thought.

Bulletin-Institut royal des sciences naturelles de Belgique. Sciences de la terre, 1993

Large quantities of fossil bones were recovered in the Flemish Valley at the sites of Zemst, Hofs... more Large quantities of fossil bones were recovered in the Flemish Valley at the sites of Zemst, Hofstade, Overmere and Dendermonde. Most of these Late Pleistocene assemblages date from the Weichselian and were recovered from fluvial deposits formed by braided river Systems. Twenty-three mammalian taxa were collected. The fossils accumulated mainly through graduai, long-term processes as indicated by the scattered and dispersed spatial distribution of the bones, the abundance of scavenged and subaerially weathered bones, the low numbers of carnivores, the low CSI (corrected number of specimens per individual) values and the prépondérance of Voorhies (1969) Group 3. Age profiles of mammoth support this interprétation. However, the age profile of rhinocéros of assemblage Hofstade I suggests episodic mass-mortality events affecting this large herbivore.

Frontiers in Psychology, 2023

The historically known relationship of interspecies companionship between Aboriginal foraging com... more The historically known relationship of interspecies companionship between Aboriginal foraging communities in Australia and free-ranging dingoes provides a model for understanding the human-canid relations that gave rise to the first domesticated dogs. Here, we propose that a broadly similar relationship might have developed early in time between wild-living wolves and mobile groups of foragers in Late Pleistocene Eurasia, with hunter-gatherers routinely raiding wild wolf dens for pre-weaned pups, which were socialized to humans and kept in camp as tamed companions ("pets"). We outline a model in which captive wolf pups that reverted to the wild to breed when they were sexually mature established their territories in the vicinity of foraging communities-in a "liminal" ecological zone between humans and truly wild-living wolves. Many (or most) of the wolf pups humans took from the wilderness to rear in camp may have derived from these liminal dens where the breeding pairs had been under indirect human selection for tameness over many generations. This highlights the importance of the large seasonal hunting/aggregation camps associated with mammoth kill-sites in Gravettian/Epigravettian central Europe. Large numbers of foragers gathered regularly at these locations during the wild wolf birthing season. We infer that if a pattern of this kind occurred over long periods of time then there might have been a pronounced effect on genetic variation in free-ranging wolves that denned and whelped in the liminal zones in the vicinity of these human seasonal aggregation sites. The argument is not that wolves were domesticated in central Europe. Rather, it is this pattern of hunter-gatherers who caught and reared wild wolf pups gathering seasonally in large numbers that might have been the catalyst for the early changes leading to the first domesticated dogs-whether in western Eurasia or further afield.

The Anatomical Record, 2022

Janssens et al. (2021, doi: 10.1002/ar.24624) recently commented on our article (Galeta et al., 2... more Janssens et al. (2021, doi: 10.1002/ar.24624) recently commented on our article (Galeta et al., 2021, doi: 10.1002/ar.24500) regarding the morphological differences between putative Paleolithic dog and Pleistocene wolf crania. The authors argued that these differences reflect the normal population variation of wolves, that some of the cranial measurements used do not reflect morphological changes during domestication, and that our canid dataset was small because we
inexplicably omitted several specimens we analyzed in our previous publications. In this commentary, we briefly address the issue of within and between morpho-population variability.
The results based on our canid sample suggest that the magnitude of morphological differences between distinct morpho-populations (i.e., recent northern dogs and wolves) is at least twice as
large as that observed within morpho-populations (between two groups of recent northern wolves segregated by cluster analysis). The morphological differences between putative Paleolithic dogs
and Pleistocene wolves are relatively large, which may indicate that they did not likely represent a single Late Pleistocene morpho-population. Finally, we clarified the rationale behind the composition of our 2021 dataset to show that we did not adjust the list of the analyzed specimens. Although the sample size was small, the randomization analysis published in 2021 confirmed that the unbalanced composition of the reference sample did not affect the reliability of the morphological segregation of putative Paleolithic dogs and Pleistocene wolves.

Monographien des RGZM, 2021

Most researchers accept that by the end of the Pleistocene dogs were part of the daily life of pr... more Most researchers accept that by the end of the Pleistocene dogs were part of the daily life of prehistoric hunter-gatherers. Recent analyses of the mammal assemblages from the third cave of Goyet (Belgium) reveal that a large component of the material from bone level A1 postdates the Last Glacial Maximum. The biometric study of the large canid remains from this level shows that an ulna can be described as from a medium-sized Palaeolithic dog with an estimated body mass of ~ 20 kg. A direct AMS 14 C date of the bone demonstrates that the dog lived during the Bølling / Allerød interstadial. Human and carnivore modifications of the bone indicate that the animal was dismembered by a contemporaneous human individual, likely to obtain its meat, and then gnawed by a canid-sized carnivore. Presumably, Palaeolithic dogs fulfilled diverse roles in Late Palaeolithic societies including as a source of food.

Human-elephant interactions: from past to present, 2021

Middle and Upper Palaeolithic sites, where mammoths dominate the faunal assemblages, are mainly f... more Middle and Upper Palaeolithic sites, where mammoths dominate the faunal assemblages, are mainly found in Central and Eastern Europe. At these sites concentrations of skulls, tusks and long bones, interpreted as deliberate constructions, often occur. Rare instances of weapon tip fragments embedded in mammoth bones provide direct archaeological evidence of human hunting. Indirect evidence, such as the accumulation of mammoth bones from multiple individuals with specific ontogenetic ages, occurs more frequently. Based on the eruption sequence and wear of deciduous premolars from mammoth calves, we examined whether a season of death could be deduced from the characteristics of the dentition. Our results suggest that the mammoth hunt was not restricted to the cold half of the year.

Umschlagbild: Ein europäischer Waldelefant (Palaeoloxodon antiquus) und ein Hominin aus dem Mitte... more Umschlagbild: Ein europäischer Waldelefant (Palaeoloxodon antiquus) und ein Hominin aus dem Mittelpleistozän (Vorder-und Rückseitenabbildungen von Gleiver Prieto; Copyright Katerina Harvati, Universität Tübingen) Druck und Bindung: readbox unipress in der readbox publishing GmbH Printed in Germany Bibliogra sche Information der Deutschen Nationalbibliothek Die Deutsche Nationalbibliothek verzeichnet diese Publikation in der Deutschen Nationalbibliogra e, detaillierte bibliogra sche Daten sind im Internet über http://dnb.d-nb.de abrufbar. e symposium and the volume "Human-elephant interactions: from past to present" were funded by the Volkswagen Foundation.

Journal of Anthropological Archaeology, 2021

Two canid morphotypes have been proposed for the middle Upper Palaeolithic site of Pˇredmostí (Mo... more Two canid morphotypes have been proposed for the middle Upper Palaeolithic site of Pˇredmostí (Moravia, Czech Republic): Pleistocene wolf and Palaeolithic dog (Germonpr ́e et al., 2012, 2015; Galeta et al., 2020). In Wilc-zy ́nski et al. (2020), faunal assemblages from other Upper Palaeolithic Moravian sites were analyzed and those results used to project a similar, which we interpret as flawed, result for Pˇredmostí: that only wild canids were present. We address issues with their methodology and argue that their conclusion, that dogs were not present at any of those studied sites, is based on cursory taphonomic analysis. Further, their projection of these results to Pˇredmostí, an assemblage they did not study, is unsubstantiated, but does highlight the current dearth of taphonomic information that could aid the study of dog domestication. We also point out an important error in their Fig. 1, where the hemimandibles intended to show the natural variability in large canids from Dolní Vĕstonice II and Pavlov I SE, are in fact canid specimens from Pˇredmostí. Since several of these canid specimens have been published by Germonpr ́e et al. (2015), we find it imperative to address this error to ensure these data are correctly referenced in future works.

Human Ecology, 2021

Several hypotheses have been proposed to explain the initial steps in the domestication process o... more Several hypotheses have been proposed to explain the initial steps in the domestication process of the wolf. We discuss the human-initiated model in which wolf pups were brought to camp sites by male hunters and cared for by nursing women. A good relation between the more sociable and playful pups and the women and their children likely formed affiliative bonds and led to the survival of such pups into maturity. Some of these animals could have reproduced and delivered at least one litter. A selection on the behaviour of subsequent generations could ultimately have led to Palaeolithic dogs.

Journal of Archaeological Science, 2021

Canids from the Upper Paleolithic site of Predmostí are central to debates concerning the domesti... more Canids from the Upper Paleolithic site of Predmostí are central to debates concerning the domestication of dogs as two morphotypes were identified: Pleistocene wolf and Paleolithic dog (Germonpr´e et al., 2015a). In Prassack et al. (2020), we set out to determine whether specimens previously parsed into these two groups differed
significantly in dental microwear textures, a proxy for diet. We did not assume that one group was comprised of dogs, but hypothesized that if they were, they would likely have consumed more bone, leading to microwear surface textures dominated by pitting. We indeed found significantly higher scales of maximum complexity on second lower molar crushing surfaces of the sample identified as Paleolithic dogs by Germonpr´e et al. (2015a),
consistent with larger pits on average and more bone consumption. These results suggest that the two morphotypes
identified by Germonpr´e et al. (2015a) represent ecologically distinct populations. This is in accord
with the interpretation of domestication, but, as we noted, the groups could also represent two distinct wild canid
populations with differing diets. Janssens et al. (2021) recently criticized our study, questioning our methods of
analysis and claiming bias in our interpretation of results. We reply to their issues here, focusing only on those
relevant to Prassack et al. (2020).

Social inequality before farming? Multidisciplinary approaches to the study of social organization in prehistoric and ethnographic hunter-gatherer-fisher societies Edited by Luc Moreau, 2020

The dog is the only domesticated species that dates from before the origin of agriculture when hu... more The dog is the only domesticated species that dates from
before the origin of agriculture when human populations
were living as hunter-gatherers (e.g. Germonpré
et al. 2009, 2015, 2018; Thalmann et al. 2013; Freedman
and Wayne 2017). Morphological and genetic analyses
have shown that dogs descent from an extinct Eurasian
Pleistocene wolf population or possibly several populations
(e.g. Germonpré et al. 2009; Thalmann et al. 2013;
Skoglund et al. 2015; Frantz et al. 2016). Although the
dogs’ ancestor is now known, many questions remain,
such as how the first dogs could have participated in
the daily life of their owners (Losey et al. 2018).
In this contribution, we first summarize two
models on the origin of the dog; then we detail two
canid morphotypes from the Upper Palaeolithic; next,
we look to the Upper Palaeolithic sites that have evidence
for the presence of incipient dogs. After that, we
review ethnographic sources for dog-related practices
among Northern societies and whether and how these
could enhance differential access to resources and
influence social status distinctions. Subsequently, we
discuss what could have been the roles of early dogs
in some Upper Palaeolithic societies. Finally, we propose
a tentatively narrative on how the contributions
of Palaeolithic dogs could have affected differential
wealth and influenced social distinction among past
men and women.

Science, 2020

Dogs were the first domestic animal, but little is known about their population history and to wh... more Dogs were the first domestic animal, but little is known about their population history and to what extent it was linked to humans. We sequenced 27 ancient dog genomes and found that all dogs share a common ancestry distinct from present-day wolves, with limited gene flow from wolves since domestication but substantial dog-to-wolf gene flow. By 11,000 years ago, at least five major ancestry lineages had diversified, demonstrating a deep genetic history of dogs during the Paleolithic. Coanalysis with human genomes reveals aspects of dog population history that mirror humans, including Levant-related ancestry in Africa and early agricultural Europe. Other aspects differ, including the impacts of steppe pastoralist expansions in West and East Eurasia and a near-complete turnover of Neolithic European dog ancestry.

Current Biology, 2020

Extant Canis lupus genetic diversity can be grouped into three phylogenetically distinct clades: ... more Extant Canis lupus genetic diversity can be grouped into three phylogenetically distinct clades: Eurasian and American wolves and domestic dogs.1
Genetic studies have suggested these groups trace their origins to a wolf population that expanded during the last glacial maximum (LGM)1
, 2
, 3
and replaced local wolf populations.4
Moreover, ancient genomes from the Yana basin and the Taimyr peninsula provided evidence of at least one extinct wolf lineage that dwelled in Siberia during the Pleistocene.3
5
Previous studies have suggested that Pleistocene Siberian canids can be classified into two groups based on cranial morphology. Wolves in the first group are most similar to present-day populations, although those in the second group possess intermediate features between dogs and wolves.6
7
However, whether this morphological classification represents distinct genetic groups remains unknown. To investigate this question and the relationships between Pleistocene canids, present-day wolves, and dogs, we resequenced the genomes of four Pleistocene canids from Northeast Siberia dated between >50 and 14 ka old, including samples from the two morphological categories. We found these specimens cluster with the two previously sequenced Pleistocene wolves, which are genetically more similar to Eurasian wolves. Our results show that, though the four specimens represent extinct wolf lineages, they do not form a monophyletic group. Instead, each Pleistocene Siberian canid branched off the lineage that gave rise to present-day wolves and dogs. Finally, our results suggest the two previously described morphological groups could represent independent lineages similarly related to present-day wolves and dogs.

The Anatomical Record, 2020

The antiquity of the wolf/dog domestication has been recently pushed back in time from the Late U... more The antiquity of the wolf/dog domestication has been recently pushed back in time from the Late Upper Paleolithic (~14,000 years ago) to the Early Upper Paleolithic (EUP; ~36,000 years ago). Some authors questioned this early dog domestication claiming that the putative (EUP) Paleolithic dogs fall within the morphological range of recent wolves. In this study, we reanalyzed a data set of large canid skulls using unbalanced-and balanced-randomized discriminant analyses to assess whether the putative Paleolithic dogs are morphologically unique or whether they represent a subsample of the wolf morpho-population. We evaluated morphological differences between 96 specimens of the 4 a priori reference groups (8 putative Paleolithic dogs, 41 recent northern dogs, 7 Pleistocene wolves, and 40 recent northern wolves) using discriminant analysis based on 5 ln-transformed raw and allometrically size-adjusted cranial measurements. Putative Paleolithic dogs are classified with high accuracies (87.5 and 100.0%, cross-validated) and randomization experiment suggests that these classification rates cannot be exclusively explained by the small and uneven sample sizes of
reference groups. It indicates that putative Upper Paleolithic dogs may represent a discrete canid group with morphological signs of domestication (a relatively shorter skull and wider palate and braincase) that distinguish them from sympatric Pleistocene wolves. The present results add evidence to the view that these specimens could represent incipient Paleolithic dogs that were involved in daily
activities of European Upper Paleolithic forager groups.

Journal of Archaeological Science, 2020

Morphological and genetic evidence put dog domestication during the Paleolithic, sometime between... more Morphological and genetic evidence put dog domestication during the Paleolithic, sometime between 40,000 and 15,000 years ago, with identification of the earliest dogs debated. We predict that these earliest dogs (referred to herein as protodogs), while potentially difficult to distinguish morphologically from wolves, experienced behavioral shifts, including changes in diet. Specifically, protodogs may have consumed more bone and other less desirable scraps within human settlement areas. Here we apply Dental Microwear Texture Analysis (DMTA) to canids from the Gravettian site of Předmostí (approx. 28,500 BP), which were previously assigned to the Paleolithic dog or Pleistocene wolf morphotypes. We test whether these groups separate out significantly by diet-related variation in microwear patterning. Results are consistent with differences in dietary breadth, with the Paleolithic dog morphotype showing evidence of greater durophagy than those assigned to the wolf morphotype. This supports the presence of two morphologically and behaviorally distinct canid types at this middle Upper Paleolithic site. Our primary goal here was to test whether these two morphotypes expressed notable differences in dietary behavior. However, in the context of a major Gravettian settlement, this may also support evidence of early stage dog domestication. Dental microwear is a behavioral signal that may appear generations before morphological changes are established in a population. It shows promise for distinguishing protodogs from wolves in the Pleistocene and domesticated dogs from wolves elsewhere in the archaeological record.

Lengths, widths, and size proportions (length to width) of the lower carnassial were measured in ... more Lengths, widths, and size proportions (length to width) of the lower carnassial were measured in 45 teeth of the arctic fox and 35 teeth of the red fox from Belgium radiocarbon dated to 46 640-14 120 ka BP. Data the Late Pleistocene foxes from Belgium were compared to 20 ancient and extant populations form Europe, Asia, and North America. The Pleistocene arctic fox from Belgium showed larger carnassial than in all recent samples of this species, whereas the Belgian fossil red foxes were characterized by the carnassial size comparable to that of the recent Siberian red foxes. Both fox species from the Pleistocene of Belgium showed the highest index of the carnassials length to width, which means increase in carnivorous adaptation. We conclude that the higher level of carnivorous specialization reached by the Belgian arctic and red foxes at the end of the Late Pleistocene reflected their scavenging on kills of large carnivores and human hunters (remains of megafauna). Harsh environmental conditions of that period and specific composition of ecosystems led to adapting to a more carnivorous food niche in both foxes.

Journal of Archaeological Science: Reports, 2020

The Trou de Chaleux is a cave site located in Belgium. It delivered a rich late Magdalenian mater... more The Trou de Chaleux is a cave site located in Belgium. It delivered a rich late Magdalenian material culture constituted mainly of lithic artefacts but also including bone industries and figurative art. This paper presents the results of the analysis of the large collection of bird remains recovered by E. Dupont in 1865, which was yet unstudied from taphonomical and archaeozoological perspectives. In addition to the taxonomic identification, surface alterations were investigated based on a macro-and microscopic analysis, including an analysis of wear traces and elementary composition. Special attention is devoted to the presence of human modifications such as disarticulation or butchering marks, traces of heating, presence of colourants and traces of bone working. The taphonomic history of the bird assemblage is reconstructed and the use of birds by humans characterized, as well as their importance in past human activities. We also discuss evidence for seasonal exploitation and for reconstructing the local environment and integrate our results with evidence from other Magdalenian assemblages from northwestern Europe. At Trou de Chaleux, birds were used for food, as raw material for bone working and for symbolic purposes. The exploitation of avian products was intense, and species have been used for several purposes such as the raven and snowy owl having been exploited both for food and for symbolic reasons. Large bird bones were used as raw material to produce artefacts, but the use-wear analysis did not evidence un-ambiguous traces related to the use of the objects produced. Despite several limiting factors, the bird material from Trou de Chaleux considerably increases the knowledge of past human exploitation of birds during the late Magdalenian in northwestern Europe.

Plos Biology, 2019

While sequencing ancient DNA (aDNA) from archaeological material is now commonplace, very few att... more While sequencing ancient DNA (aDNA) from archaeological material is now commonplace, very few attempts to sequence ancient transcriptomes have been made, even from typically stable deposition environments such as permafrost. This is presumably due to assumptions that RNA completely degrades relatively quickly, particularly when dealing with autolytic, nuclease-rich mammalian tissues. However, given the recent successes in sequencing ancient RNA (aRNA) from various sources including plants and animals, we suspect that these assumptions may be incorrect or exaggerated. To challenge the underlying dogma, we generated shotgun RNA data from sources that might normally be dismissed for such study. Here, we present aRNA data generated from two historical wolf skins, and perma-frost-preserved liver tissue of a 14,300-year-old Pleistocene canid. Not only is the latter the oldest RNA ever to be sequenced, but it also shows evidence of biologically relevant tissue specificity and close similarity to equivalent data derived from modern-day control tissue. Other hallmarks of RNA sequencing (RNA-seq) data such as exon-exon junction presence and high endogenous ribosomal RNA (rRNA) content confirms our data's authenticity. By performing independent technical library replicates using two high-throughput sequencing platforms, we show not only that aRNA can survive for extended periods in mammalian tissues but also that it has potential for tissue identification. aRNA also has possible further potential, such as identifying in vivo genome activity and adaptation, when sequenced using this technology.

Cell, 2019

Horse domestication revolutionized warfare and accelerated travel, trade, and the geographic expa... more Horse domestication revolutionized warfare and accelerated travel, trade, and the geographic expansion of languages. Here, we present the largest DNA time series for a non-human organism to date, including genome-scale data from 149 ancient animals and 129 ancient genomes (≥1-fold coverage), 87 of which are new. This extensive dataset allows us to assess the modern legacy of past equestrian civilizations. We find that two extinct horse lineages existed during early domestication, one at the far western (Iberia) and the other at the far eastern range (Siberia) of Eurasia. None of these contributed significantly to modern diversity. We show that the influence of Persian-related horse lineages increased following the Islamic conquests in Europe and Asia. Multiple alleles associated with elite-racing, including at the MSTN “speed gene,” only rose in popularity within the last millennium. Finally, the development of modern breeding impacted genetic diversity more dramatically than the previous millennia of human management.

Ecology and Evolution, 2019

The current phylogeographic pattern of European brown bears (Ursus arctos) has commonly been expl... more The current phylogeographic pattern of European brown bears (Ursus arctos) has commonly been explained by postglacial recolonization out of geographically distinct refugia in southern Europe, a pattern well in accordance with the expansion/contraction model. Studies of ancient DNA from brown bear remains have questioned this pattern, but have failed to explain the glacial distribution of mitochondrial brown bear
clades and their subsequent expansion across the European continent. We here present 136 new mitochondrial sequences generated from 346 remains from Europe, ranging in age between the Late Pleistocene and historical times. The genetic data show a high Late Pleistocene diversity across the continent and challenge the strict confinement of bears to traditional southern refugia during the last glacial maximum (LGM). The mitochondrial data further suggest a genetic turnover just before this
time, as well as a steep demographic decline starting in the mid‐Holocene. Levels of stable nitrogen isotopes from the remains confirm a previously proposed shift toward increasing herbivory around the LGM in Europe. Overall, these results suggest that in addition to climate, anthropogenic impact and inter‐specific competition may have had more important effects on the brown bear's ecology, demography, and genetic structure than previously thought.

SOCIAL INEQUALITY BEFORE FARMING

The dog is the oldest domesticated species and the only animal that was domesticated during the P... more The dog is the oldest domesticated species and the only animal that was domesticated during the Pleistocene - before the emergence of agriculture - when human populations were living as hunter-gatherers. Today, owned dogs can assist their owners in various ways. They can function as watchdog, facilitate transport as beasts of burden, aid in hunting, play a ritual role or provide company. In some cultures, they are consumed and their skin or hair can be used for the tailoring of cloths. We have shown previously that in several Upper Palaeolithic sites two morphotypes of fossil large canids can be distinguished: Palaeolithic dogs and Pleistocene wolves. The remains of Palaeolithic dogs occur in certain early and mid Upper Palaeolithic sites located above 45° latitude; their geographic distribution in post-Last Glacial Maximum sites is more widespread. We adapt here a table proposed by Sigaut (1980) and compare canid products and roles, from living and dead animals, that could have been of possible use in Upper Palaeolithic societies. These roles and products are based on data from the ethnographic literature and confronted with the possible registration of uses in the archaeological record. It is in a framework of an animated worldview of Upper Palaeolithic peoples drawn upon a range of archaeozoological, archaeological, and ethnographic data that we review whether some of these uses and products could have led to differential access to resources and could possibly have enhanced inequality among Upper Palaeolithic hunter-gatherers.

Grey wolves (Canis lupus) are one of the few large terrestrial carnivores that maintained a wide ... more Grey wolves (Canis lupus) are one of the few large terrestrial carnivores that maintained a wide geographic distribution across the Northern Hemisphere throughout the Pleistocene and Holocene. Recent genetic studies have suggested that, despite this continuous presence, major demographic changes occurred in wolf populations between the late Pleistocene and early Holocene, and that extant wolves trace their ancestry to a single late Pleistocene population. Both the geographic origin of this ancestral population and how it became widespread remain a mystery. Here we analyzed a large dataset of novel modern and ancient mitochondrial wolf genomes, spanning the last 50,000 years, using a spatially and temporally explicit modeling framework to show that contemporary wolf populations across the globe trace their ancestry to an expansion from Beringia at the end of the Last Glacial Maximum - a process most likely driven by the significant ecological changes that occurred across the Northern Hemisphere during this period. This study provides direct ancient genetic evidence that long-range migration has played an important role in the population history of a large carnivore and provides an insight into how wolves survived the wave of megafaunal extinctions at the end of the last glaciation. Moreover, because late Pleistocene grey wolves were the likely source from which all modern dogs trace their origins, the demographic history described in this study has fundamental implications for understanding the geographical origin of the dog.