Several millennia of human-mediated translocation of non-native pig species (genus Sus) to the islands of Wallacea and Oceania have considerably altered local ecosystems. To investigate the timing and trajectory of these introductions, we conducted both genomic analyses of 576 pig nuclear genomes and a geometric morphometric analysis of 708 modern and ancient dental remains. Our analyses demonstrate that free-living and domestic pigs in Wallacea and Oceania have diverse ancestries resulting from the introduction of multiple sequential pig populations followed by gene flow. Despite the variability in their genomic ancestry, these pigs all have a distinct tooth morphology as well as a genetic link to the Chinese domestic pig populations that accompanied the dispersal of Austronesian language speakers ~4000 to 3000 years ago via Taiwan and the Philippines.

Due to the insular location, farming emerged in Cyprus mostly through the introduction of plants and animals from nearby continent, throughout the 9th and 8th millennia BCE. These included wild cultivated einkorn and emmer during the Pre-Pottery Neolithic A (PPNA) and ungulates during the PPNB. However, large osteoarchaeological samples from Klimonas and Shillourokambos sites, revealed an interesting diversity in the processes. Cattle and sheep were introduced as early domesticates, c. 8500 and 7900 BCE respectively, and raised throughout the late 9th and the 8th millennia. However, shortly after their introduction, the domestic goats went feral and became game, before being domesticated again 8-10 centuries later. Even more intriguing is the history of suids. The wild boar was introduced long before the PPN and rapidly evolved into an endemic insular sub-species (Sus scrofa circeus). From 10,200 to 8500 BCE, it was the only large game on the Cyprus island. Then, from 8500 to 7500 BCE, Sus s. circeus was domesticated, giving rise to all subsequent Neolithic pigs, without any visible introduction of continental domestic pigs. This diversity of processes highlights the complexity of dispersal of domesticates at the beginning of the Neolithic period, involving transfer and acclimatisation components as well as possible local domestications. It suggests that local domestication played a more significant role than expected in the Neolithic dispersal on the continents.

Captive environments induce plastic and adaptive phenotypic changes to the inner and outer anatomy of vertebrate bones. However, the respective effects of these changes are not well documented. This study examines the impact of captivity on the microanatomy of the talus, an ankle bone that lacks muscular attachment but is involved in supporting the body weight during locomotion. This study focuses on the wild boar (Sus scrofa) as a wild ungulate model and on an experimental setting through which these wild boars experienced different mobility patterns during their growth: 20 lived in their natural habitat, while 21 were born in the wild but raised in captivity, either in a large enclosure (10) or in a stable (11). We quantitatively investigated the microanatomical structure of their talus using 3D cartographies of compact bone thickness and five microanatomical parameters, while describing the distribution of compact bone and the trabecular network from the cartographies and virtual sections. While the absolute thickness of compact bone remains similar regardless of the mobility context, specimens that have been raised in captivity have tali with greater bone density and a tighter network of bone trabeculae but a lower relative maximum thickness of compact bone. Our results show that captivity induces a plastic response in the bone microanatomy, which could therefore serve as a valuable indicator of the early process of domestication of wild boars and, more generally, wild ungulates in the bioarchaeological records.

Abstract The timing and causes of brain size reduction in domestic dogs remain uncertain. Using endocast volume as a proxy for brain size, this study provides a first insight into long-term brain size evolution in the wolf-dog lineage. We compared endocranial volumes of 185 modern and 22 prehistoric wolves and dogs ranging from Western Europe to Australia, and spanning the Pleniglacial (35 000 yr BP) to the Late Neolithic (5000 yr BP). Our results reveal that Pleistocene so-called ‘protodogs’ show no brain size reduction compared with coeval Pleistocene wolves. Instead, we observed a slightly larger relative endocranial volume in the 35 000-year-old 'protodog' from Goyet, which could suggest increased behavioural flexibility in the presence of humans. In contrast, Late Neolithic dogs show a drastic brain size reduction (46%) with endocranial volumes comparable to modern small terrier and toy breeds. We speculate that the anxious and wary temperaments of these Late Neolithic dogs, induced by the brain tissue reorganization associated with such a size reduction, could have served an alerting purpose, among the many other potential roles dogs could have played within these Late Neolithic socio-ecosystems.

One of the key challenges in the archaeology of sheep domestication is reconstructing the complex history of environmental and anthropogenic transformations undergone by sheep since the beginning of the domestication process of their wild ancestors. In recent years, GMM studies of sheep astragalus bones have contributed to our understanding of morphological differences between wild and domestic caprine species. However, the respective influences of biological and ecological factors on astragalus morphological variations in sheep remain poorly documented. This limitation hinders a comprehensive understanding of its biosystematic resolution and, consequently, its use as a proxy in archaeological contexts to investigate early selective breeding and the emergence of sheep breeds in Southwest Asia. This paper presents the results of a morphological study of 96 astragali using 3D geometric morphometrics, focusing primarily on modern Eurasian and African sheep breeds and landraces. The study is based on a well-documented comparative collection encompassing phenotypical traits (breed, sex, age, presence/absence of horns, coat and tail type, weight, body length); ecological characteristics (climate, geography, environment, elevation, topography); and breeding strategies (mobility). The results demonstrate that the 3D astragalus morphological pattern is a reliable marker for distinguishing one sheep breed from another. They suggest that astragalus morphology is only slightly influenced by phenotypic markers. The study further explores the effects of environmental and climatic factors on phenotypic variation and highlights the potential of the astragalus as an ecomorphological marker. Finally, the current limitations in interpreting the relationship between astragalus morphological variation and mobility strategies in archaeological contexts are discussed.

The domestication process of South American Camelid (SAC) is still poorly understood despite its major role in the history of Andean societies. This study is aimed at documenting the elusive process of intensification in human and SAC interactions at the key site of Telarmachay (Peru), between 9.0-8.0 and 3.7-3.1 Kyr BP. We assessed taxonomic identification and intraspecific morphological modifications using talus bone 3D geometric morphometrics (GM), associated with stable isotope analysis of bone collagen (delta 13C and delta 15N) and dental palaeodemography, to track changes in animal diet, environmental setting and culling practices. The talus bone 3D GM documents more than 5,000 years of morphological changes suggesting intensification of vicuna, and possibly guanaco, population management between 6.4-6.0 and 5.8-4.5 Kyr BP, but does not support phenotypic divergence towards alpaca and llama domestic phenotypes. Comparative stable isotope analysis across the Puna ecozones shows that the changes in human and SAC interactions at Telarmachay were not driven by major modification of the camelid environment and diet. This study highlights the need for multiproxy bioarchaeological approaches to study the domestication process as an elusive, nonlinear multispecies process shaped by trial and error, environmental contingencies, and unintended consequences.

Due to its commensal relationship with humans, facilitating its dispersal from its natural range in East Asia, the brown rat (Rattus norvegicus) is now one of the most cosmopolitan and invasive mammals. Brown rats are also vectors of human pathogens and the most commonly used mammalian model (after mice) for biomedical research. Despite their major role in biodiversity loss and medicine, our understanding of a likely origin for their commensal relationship with humans in China still relies on modern population genetics, without a reliable timeframe of localisation. In this research, we have tried to fill this gap using direct information from the archaeological remains of rats collected by Chinese colleagues in several archaeological sites dated between 9000 and 3500 BP. Thanks to high-resolution morphometrics and direct 14C dating, we have identified the occurrence of the brown rat (Rattus norvegicus) within the domestic area deposits of the early Neolithic site of Jiahu by 7500 cal BP, suggesting a tentative link between rice agriculture and an early commensal behaviour of this species. ### Competing Interest Statement The authors have declared no competing interest. This research was also granted by the Agence National pour la Recherche (ANR, http://www.agence-nationale-recherche.fr/), through the LabEx ANR-10-LABX-0003-BCDiv, in the framework of the Investissements d'avenir programme (ANR-11-IDEX-0004-02) This research was also granted by the Agence National pour la Recherche (ANR, http://www.agence-nationale-recherche.fr/), through the LabEx ANR-10-LABX-0003-BCDiv, in the framework of the Investissements d'avenir programme (ANR-11-IDEX-0004-02) D. Orton’s contribution was funded by UKRI (Frontier Research Guarantee project RATTUS

House mice (Mus musculus) have been associated with humans since the beginning of sedentism, enabling them to become successful global colonisers. Three main subspecies originated approximately 0.5 Ma in a region extending from Southwest Asia to northern India. Molecular data suggest that a complex scenario of secondary admixture occurred thereafter in the Iranian region, leading to the formation of a Central Iranian lineage, but this evidence was overlooked in previous bioarchaeological analysis. The early Neolithic settlement of Ganj Dareh is located in this cradle area. It delivered remains of commensal house mice formerly attributed to M. m. domesticus. A geometric morphometric analysis of the first lower molars is used here to characterize the signature of hybridization between M. m. musculus and M. m. domesticus. The subspecific attribution of the Ganj Dareh mouse remains is re-evaluated through the inclusion of modern specimens from Central Iran as a separate group in the reference dataset. The results indicate that, contrary to what was previously thought, the Ganj Dareh specimens are likely related to the Central Iranian lineage. Their idiosyncrasy compared with modern representatives, however, suggests a complex temporal dynamic of admixture, which may have been influenced by early human settlements and movements.

The Roman conquest of the northern Alpine foreland in 15 BCE introduced larger-sized horses and hybrid mules to the region. To investigate their genetic profiles and influence on local breeding, we analyzed our previously generated shallow shotgun DNA data from 402 Late Iron Age and Roman equids, supplemented with mitochondrial and nuclear capture data from 40 to 31 equids from the same regions and periods. Late Iron Age and Roman horses exhibit high matrilineal diversity, with unique haplogroups in the latter indicating exogenous animals. The preferred use of male mounts in army service mentioned in historical writings is confirmed. The absence of donkey jacks in our study area and the lack of mule matrilineal lineages unique to Late Iron Age horses contradict local hybrid breeding. Alleles associated with speed, endurance, or rare coat colors have not been identified, nor has a genetic basis for the larger body size of Roman horses, although illustrated by archaeozoological and historical data.

Human mediated translocation of non-native pig species (genus Sus ) to the islands of Wallacea and Oceania has significantly altered local ecosystems. To investigate the timing and trajectory of these introductions, we conducted both genomic analyses of 576 pig nuclear genomes and a geometric morphometric analysis of 714 modern and ancient dental remains. Our analyses demonstrate that feral and domestic pigs in Wallacea and Oceania possess diverse ancestries resulting from the introduction of multiple, sequential pig populations followed by gene flow. Despite the variability in their genomic ancestry these pigs all possess a distinct tooth morphology, and a genetic link to the Chinese domestic pig populations that accompanied the dispersal of Austronesian language speakers ∼4,000-3,000 years ago via Taiwan and the Philippines. ### Competing Interest Statement The authors have declared no competing interest.

The timing and causes of brain size reduction in domestic dogs remain uncertain. Using endocasts volume as a proxy for brain size, this study provides a first insight into long-term brain size evolution in the wolf-dog lineage. We compared endocranial volumes of 185 modern and 22 prehistoric wolves and dogs ranging from Western Europe to Australia, and spanning the Pleniglacial (35 Ky BP) to the Late Neolithic (5 Ky BP). Our results reveal that Pleistocene so called protodogs show no brain size reduction compared to coeval Pleistocene wolves. Instead, we observed a slightly larger relative endocranial volume in the 35,000-year-old specimen from Goyet, which could suggest increased behavioural flexibility in the presence of humans. This hypothesis needs to be tested further. In contrast, Late Neolithic dogs show a drastic 46% brain size reduction with an endocranial volumes comparable to modern small terrier and toy breeds. The anxious and wary temperaments of these Late Neolithic dogs, induced by the brain tissue reorganization associated with such a size reduction, could have served an alerting purpose, among the many other potential roles dogs could have played within this Late Neolithic socio-ecosystems. ### Competing Interest Statement The authors have declared no competing interest. This project has received financial support from the CNRS through the MITI interdisciplinary programs DIM PAMIR -, IDF-DIM-PAMIR-2024-4-023 the Australian Research Council Discovery Grant, DP210101960

The mouse remains recovered from the excavations at the Early Bronze Age settlement at Dhaskalio in the Cyclades were identified as Mus musculus domesticus, the western subspecies of the house mouse, using 2D Geometric Morphometrics of the first lower molar. This paper presents the results of the Geometric Morphometrics analysis and discusses the occurrence of the house mouse in Dhaskalio Phases A, B and C, and particularly its earliest presence in Phase A, corresponding to the earlier part of the Early Cycladic II period (c. 2750-2550 BCE). This find suggests the house mouse arrival on Dhaskalio as a boat stowaway, probably hidden in grain cargoes; this was probably part of its broader dispersal around the Aegean, as also indicated by contemporaneous evidence from other sites. It also gives some insight into the subsistence and import strategies employed at the site from the earliest period. Overall, the chronostratigraphic context of this discovery in comparison to other synchronous contexts is discussed in the framework of maritime networks of the Aegean Early Bronze Age.

Dental anomalies are frequent in boars and pigs, and they generally affect the first premolar loci. The prevalence of these dental anomalies was investigated in a large number of populations around the world. These studies mainly focused on the influence of domestication, size, sexual dimorphism or food hardness on these anomalies. However, they rarely considered ontogenetic aspects, while these are crucial for understanding their aetiology during animal growth and how the dental row-jaw complex is affected. Here, we studied the incidence of missing first upper and lower premolars in a French population of captive wild boars to discuss the functional and developmental reasons for missing teeth and to assess the impact of missing teeth on the growth of the dental row-jaw complex. Using the CT-scan data of the cranium and mandible of 24 wild boars investigated six times each during their growth, and presenting a balanced sex ratio, we recorded the number of missing teeth. We then quantified the shape of the upper and lower jaws using 3D geometric morphometrics. We found a similar prevalence of missing first premolar (37.5%) between the upper and the lower jaws, which is higher than the frequencies observed in most continental populations of wild boars. The increasing number of anomalies during ontogeny suggests a relaxed constraint on the dentition associated with a different feeding behaviour in captivity. The absence of first premolars does not appear to be associated with size variation or sexual dimorphism, nor does it affect the place of the dentition within the jaw, the latter being more influenced by the dimorphic shape of the canines and the timing of dental eruption.

The growth of teeth and jaw bones is intimately linked in vertebrates, especially in mammals due to their specialized dentition and limited body growth. However, the relative patterns of growth and level of integration (i.e., co-variation) of these structures are insufficiently known, which hinders our ability to understand how the jaw bones accommodate the diverse dental shapes and eruption patterns observed in mammals. Here, we studied the cranial ontogenetic series of 23 ungulate species among artiodactyls, perissodactyls, and hyracoids having different dental shapes and eruption patterns. We evaluated the variation of the teeth-palate complex, as well as the co-variation of teeth and palate during growth using 3D geometric morphometrics. We found a major ontogenetic component common to all the species studied, corresponding to an elongated dental row relative to the palate in juveniles and vice versa in adults. This pattern agrees with the prolonged growth of the palate as compared to teeth during development but is also reminiscent of an intraspecific allometric pattern previously observed in some dwarf ungulates. Moreover, most artiodactyls, especially ruminants, departed from other ungulates in having a higher co-variation between the dental row and the palate. This stronger integration seen in ruminants might be associated with their inherited rapid growth and relatively fast eruption pattern. This is in contrast to ungulates with late eruption of last molars, whose teeth-palate complex might be less constrained, but further investigation is needed to substantiate these hypotheses and better understand the factors influencing covariations within the upper jaw.

The jaw system in mammals is complex and different muscle morphotypes have been documented. Pigs are an interesting group of animals as they are omnivorous and have a bunodont crushing dentition. Moreover, they have interacted with humans for over 10,000 years and grow nearly two orders of magnitude in size. Despite being a model system for studies on cranial form and function, data on the growth of the jaw adductor muscles are scant. Moreover, whether captivity impacts the growth and architecture of the jaw adductors remains unknown. Based on dissection data of the jaw adductors of 45 animals ranging from less than 1 kg to almost 100 kg, we show that muscle masses, muscle fiber lengths, and cross-sectional areas scale as predicted for geometrically similar systems or with slight negative allometry. Only the fiber length of the lateral pterygoid muscle grew with slight positive allometry. Animals raised in captivity in stalls or in an enclosure were overall very similar to wild animals. However, some muscles were larger in captive animals. Interestingly, variation in bite force in captive animals was well predicted by the variation in the size of the superficial masseter muscle relative to the overall jaw adductor mass. We show that muscle masses, muscle fiber lengths, and cross-sectional areas in wild boar scale as predicted for geometrically similar systems or with slight negative allometry. Animals raised in captivity in stalls or in an enclosure were overall very similar to wild animals.image

This chapter gathers and combines different types of archaeozoological information for documenting the strategies, techniques and practices for the acquisition and material use of wild boar –the only large mammal available to the late PPNA Klimonas villagers. A pilot study of the stable oxygen and carbon isotopes suggests late winter/early spring births and excludes open landscape and dense forest covers. Based on the male and female canine count and the mixture analyses of osteological measurements, the sex ratio appears to slightly favour females (under 24–30 months) and old males (71%). Only one very young foetal bone was found. Age-at-death profiles are homogeneous across the site, except in the Communal building (St 10). Here we notice an extra peak at 6–19 months, likely related to symbolic practices (chap. 31). All the other age profiles point to the same low selective culling strategy, with an apparently deliberate intention to spare individuals under six months. Compared with the middle A2 phase at Shillourokambos (c. 7600–7400 cal BC), where pig rearing is well attested, the wild boar acquisition strategy at Klimonas clearly refers to hunting. This activity focused on groups of animals, either family or subadult groups. Together with modern comparisons, the age-at-death profiles suggest that the hunting season lasted from the end of summer until the beginning of winter. The fact that all the body parts were brought back to the site suggests that the hunting grounds were located close to the village. Hunting appears to have been a well-mastered seasonal activity. Altogether, excepting the presence of only one ungulate species, the meat and fat procurement at Klimonas did not differ from that of all the other Late PPNA sites of Southwest Asia.