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Plan of Neolithic site at Skara Brae, Orkney, showing locations of excavated trenches (I–IV on figure) and dwelling structures (shaded grey).
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Rodents have important effects on contemporary human societies, sometimes providing a source of food but more often as agricultural pests, or as vectors and reservoirs of disease. Skeletal remains of rodents are commonly found in archaeological assemblages from around the world, highlighting their potential importance to ancient human populations....
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... In this study, we investigated the genomic interplay between selection and genetic drift leading to the island syndrome in a system with an exceptionally long and clear history of population isolation. Common voles (Microtus arvalis) were introduced to the Orkney archipelago in the north of Scotland by Neolithic farmers >5000 years ago, likely as a food resource (Haynes et al. 2003;Martínková et al. 2013;Romaniuk et al. 2016). Common voles are very widespread in continental Europe with multiple distinct evolutionary lineages (Heckel et al. 2005;Beysard and Heckel 2014), but Orkney vole ancestry can be traced back specifically to populations in the Western evolutionary lineage from Belgium or northern France (Martínková et al. 2013;Wang et al. 2023). ...
... Such low levels of locally reduced diversity after selective sweeps would take long periods to be restored (Nei et al. 1975). It is possible that these signals of selection relate to the introduction history of Orkney voles, when they were kept by Neolithic farmers during the sea journey and afterward probably as a food item (Romaniuk et al. 2016). This history has probably involved voles originating from different local populations (Wang et al. 2023) and thus might have challenged the immune system through the transmission of pathogens from different geographic origins. ...
Island populations often experience different ecological and demographic conditions than their counterparts on the continent, resulting in divergent evolutionary forces affecting their genomes. Random genetic drift and selection both may leave their imprints on island populations, although the relative impact depends strongly on the specific conditions. Here we address their contributions to the island syndrome in a rodent with an unusually clear history of isolation. Common voles ( Microtus arvalis ) were introduced by humans on the Orkney archipelago north of Scotland >5000 years ago and rapidly evolved to exceptionally large size. Our analyses show that the genomes of Orkney voles were dominated by genetic drift, with extremely low diversity, variable Tajima's D , and very high divergence from continental conspecifics. Increased d N / d S ratios over a wide range of genes in Orkney voles indicated genome-wide relaxation of purifying selection. We found evidence of hard sweeps on key genes of the lipid metabolism pathway only in continental voles. The marked increase of body size in Orkney—a typical phenomenon of the island syndrome—may thus be associated to the relaxation of positive selection on genes related to this pathway. On the other hand, a hard sweep on immune genes of Orkney voles likely reflects the divergent ecological conditions and possibly the history of human introduction. The long-term isolated Orkney voles show that adaptive changes may still impact the evolutionary trajectories of such populations despite the pervasive consequences of genetic drift at the genome level.
... Malgré cette carence, en combinant plusieurs critères, la détermination de l'origine anthropogène d'un ensemble mésofaunique (proies entre 15 et 0.5 kg) est relativement fiable surtout si le rôle de l'Homme dans sa formation est prédominant (Cochard 2007). En revanche, pour la microfaune (<500g), l'identification est beaucoup plus problématique en raison généralement de la discrétion des marques de boucherie sur ces proies (e.g.Romaniuk et al. 2016 ; Andrade 2019 ;Fernandez et al. 2019 ;Lopez et Chiavazza 2020 ;Badenhorst et al. 2023). ...
En biologie évolutive, l'approche fonctionnelle vise à comprendre comment des traits, qu'ils soient morphologiques ou comportementaux, évoluent pour s’ajuster aux exigences environnementales, afin d’optimiser la survie et la reproduction d'un organisme. L’écologie comportementale humaine propose d’utiliser cette approche pour appréhender les comportements de subsistance des chasseurs-cueilleurs actuels et passés. Les archéozoologues ont examiné plus spécifiquement les prédictions des modèles économétriques issus de la théorie de l’approvisionnement optimal (optimal foraging theory) pour appréhender les décisions et les stratégies liées à l’alimentation et plus globalement à l’économie de la matière première alimentaire. Bien que la dimension évolutionniste de cette démarche ait séduit plusieurs générations d’archéozoologues en Amérique du Nord, elle peine à obtenir une reconnaissance académique en Europe. Elle y est souvent perçue comme une approche réductrice et déshumanisante en raison de l’accent qu’elle met sur des données quantifiables et économiques des comportements de prédation, au détriment de dimensions plus sociales et culturelles. Ce jugement idéologique occulte malheureusement la puissance heuristique des modèles d’optimisations pour l’archéologie et j’ai souhaité, à travers ce manuscrit, démystifier et clarifier cette approche en déconstruisant avec pédagogie ses soubassements conceptuels et méthodologiques. Cet objectif m’a conduit à remonter l’histoire des grands piliers conceptuels de l’écologie comportementale pour mettre en évidence les spécificités de cette discipline vis-à-vis des autres sciences du comportement, actuelles et passées, en particulier l’éthologie et la sociobiologie. Par la suite, je me suis penché sur l’écologie comportementale humaine en examinant son évolution, les controverses qu’elle a suscitées aux États-Unis et en France, ainsi que la réception de cette discipline par l'archéologie. Les deux derniers chapitres se focalisent sur le principal modèle de l’optimal foraging theory utilisé en archéozoologie : le modèle de la sélection optimale des proies (prey choice model). Après avoir examiné en détail les concepts et le fonctionnement de ce modèle, je m’attarde sur la manière de l’opérationnaliser en archéologie et sur les limites méthodologiques associées à son application. Je termine par une mise en perspective de ses résultats, en mettant l'accent sur les variations de l'efficacité du foraging.
... Abbreviation: PW, Papa Westray. al. 1997, Flowerdew andTattersall 2008), was established in Mainland Orkney as early as the fourth-third millennia BC (Romaniuk et al. 2016). All the major Orkney islands seem currently to be inhabited by both house and field mice (Berry 2000: 142-3 Rackham 1996). ...
Archaeological material adds a temporal dimension to evolutionary studies that is valuable for elucidating long-term population stability and evolutionary shifts for species closely associated with humans. Here, a two-dimensional geometric morphometrics approach on first upper molars was applied to modern and archaeological samples to assess the evolution of house mice in Orkney, an archipelago north of Scotland. Modern populations included localities in Orkney, north Scotland, and France. Two archaeological sites in Orkney represented the Norse period: Birsay Beachview (Mainland) and Tuquoy (Westray) (10 th-14 th /15 th centuries AD). The archaeological specimens were larger than modern specimens from similar settings, suggesting processes leading to a recent decline in the size of Orkney house mice. Molar morphology and associated non-metric traits distinct to the Orkney lineage were already established in the Norse period, as indicated by morphological similarity of the Birsay samples to modern Orkney and north Scotland populations. Stability of human settlement is likely to influence morphological evolution in house mice. The Birsay site, located in Birsay village, which has been inhabited since Norse times, might represent the ancestral house mouse population in Orkney. Tuquoy, a settlement abandoned by the end of the medieval period, provided samples different from modern house mouse populations in Westray and neighbouring isles.
... The post-glacial land connection between the Orkney archipelago and mainland Britain disappeared too early for natural species migration to take place [1]. All terrestrial mammals, and many other faunal and floral species, were most likely introduced by humans since the time of the Neolithic intensification of maritime contacts [2][3][4][5][6][7][8][9]. One of the most studied Orcadian mammals is the 'Orkney vole', an introduced population of the common vole Microtus arvalis, which is widespread in mainland Europe but otherwise absent from the British Isles [10]. ...
... Molecular phylogenetic studies pointed towards its introduction from the western European mainland around the fourth millennium BC, possibly as a single human-mediated event [5,7,[11][12][13]. Previous research on archaeological material from Skara Brae, both micromammal remains [9,14] and coprolites [15], revealed large quantities of vole remains in refuse deposits as well as individual vole bones within the faeces of dogs, perhaps showing the outcome of pest control methods or even intentional collecting of those animals by Neolithic Orcadians. ...
... Methods for quantitative and taphonomic analysis were based on those of Andrews [28], together with more recent studies [9,14,34,35] and considering their applicability to the archaeological setting [36]. For quantitative analysis, the number of identifiable specimens (NISP) was established for each sample, counted individually for key anatomical elements (mandibles, maxillae, scapulae, humeri, radii, ulnae, pelves, femora, tibiae, vertebrae, metacarpals, phalanges, calcanei, tali, ribs as well as loose incisors and molars) and jointly for other bones. ...
Micromammals, like rodents and shrews, adapt rapidly to take advantage of new food sources, habitats and ecological niches, frequently thriving in anthropogenic environments. Their remains, often retrieved during archaeological investigations, can be a valuable source of information about the past environmental conditions as well as interspecies interactions and human activity. However, the research on such finds rarely covers multiple approaches, often relying on single species or data type (e.g. identification/information for proxy studies). Here we investigate micromammal remains from the Norse and medieval (AD tenth–fourteenth centuries) archaeological site at Tuquoy, Orkney, to elucidate the relationships between micromammals, humans and other species present using a variety of data. Four micromammal species were identified, and their species dynamics as well as relationships with humans could be inferred by tracking changes in spatial and temporal location of remains, from their taphonomic history and by age estimation for individual animals. A larger, predatory assemblage was also identified, with species composition differing from that in the rest of the archaeological assemblage, and possibly therefore representing small mammal species composition in the wild. The assemblage was probably deposited by a diurnal raptor, though identification to species is not certain due to post-depositional processes.
... This could be supported by the presence of burnt elements in units III and IV (Table 3), where human occupation -either short term or long-term -is documented (Darlas and Psathi 2016;Kolendrianou et al. 2020). Burning of small vertebrates could indicate some purpose, possibly for feeding (Romaniuk et al. 2016); however, microvertebrate burning in Kalamakia is believed to have been unintentional (bones finding themselves near hearths in the interior of the cave and not being tossed deliberately into the fire) because of the absence of cutmarks or any sign of rodenthuman interaction on any of the specimens. Unfortunately, no linear marks indicative of trampling (Fernández-Jalvo and Andrews 2016) were observed in order to fully support this hypothesis, either during the microstereoscopical examination of the specimens from the site or under the electron microscope, where only a handful of them were examined. ...
The present study focuses on the microvertebrate fossil material retrieved from the three fossiliferous units of Kalamakia cave dating from ca. 90-25 kya BP, to identify the accumulation agents of the assemblage and the processes that affected it post-depositionally. This was done through the quantification of specific indices described in the literature concerning breakage, digestion and other types of modifications observed upon the surface of the specimens. Furthermore, the Taxonomic Habitat Index was calculated to reconstruct the possible habitat types in the surrounding area. Regarding the results on taphonomy, significant amounts of post-depositional breakage were observed, possibly resulting from trampling and, secondarily, modifications from the occasionally naturally occurring high humidity within a cave. The main accumulators of the microvertebrate assemblage seem to have been barn owls either independently or jointly with long-eared owls and/or owlets. Concerning the habitats possibly present in the area, the existence of mixed habitats varying between relative expansions of shrublands, grasslands, rocky areas and (limited) deciduous forests was identified, while the occasional presence of water bodies was also revealed. Shifts between the percentages of different habitats were not great but could be correlated with climatic events within a geochronological context. ARTICLE HISTORY
... Although small mammal remains are regularly recorded at Neolithic sites in Europe, these types of finds have served, for the most part, as sources of information on environmental and climatic conditions [37][38][39][40]. Certain small mammals also were a source of food for ancient populations [41][42][43]. Studies on the potential of rodents as pests in praehistoric and historic times mainly refer to species introduced in southern, western, and northern Europe, such as the house mouse (Mus musculus), the house rat (Rattus rattus), and the brown rat (Rattus norvegicus). ...
... This raises the question of whether other Pre-Neolithic rodents in Europe, prior to the arrival of the exogenous species, already played a role as commensals or pests. The analyses of rodents from the Neolithic site Skara Brae (Orkney) suggests that endemic voles were subjected to pest control while likewise serving as a source of food [42]. Otherwise, rodents are hardly cited as field pests in prehistoric times although finds of voles appear regularly at archaeological sites throughout Europe [40,[48][49][50]. ...
... That this was not always successful can be gleaned from the wood mouse finds in the granary of the Neolithic lakeshore settlement Chalain 3 in eastern France [48]. It is also conceivable that dogs were trained to ward off rodents, an assumption advanced for the Neolithic settlement of Orkney [42,43]. The use of herbal repellents [34], for example, dwarf elder (Sambucus ebulus), which was found in all wells but in higher numbers in wells 250 and 990 [24,25], is not proven so far. ...
Pests appear to have accompanied humans and their crops since the beginning of farming. Nevertheless, their study is only rarely integrated into research on farming in prehistory. An assemblage of invertebrates and small mammals was recovered from the waterlogged layers of three wells at the Middle Neolithic site (4250–3700 cal B.C.) of Les Bagnoles (SE France). The microfaunal remains were retrieved from sediment samples by wet sieving (wash-over technique). The most common among the rodents is the wood mouse. The assemblage also consists of insect remains of grain weevil, seed beetle, and corn ground beetle. The different finds not only offer data on the role of insect and rodent pests in the Neolithic but on the possible strategies adopted by the early farming communities in the western Mediterranean in response to pest infestation. The findings appear to confirm the hypothesis that the wood mouse was a commensal and storage pest in settlements long before the arrival of the invasive house mouse during the Bronze Age. The presence of the main storage pest, the grain weevil, suggests a long-term grain storage issue at Les Bagnoles. The combination of the results of the site’s archaeobotanical findings with those of other sites in the western Mediterranean suggests that the shift from naked to glume wheat around 4000 B.C. may also stem from a reaction to the problem of grain weevil infestation.
... Orkney vole remains from Skara Brae coprolites are much smaller than the North American pocket gophers (Pappogeomys/ Cratogeomys) in canid coprolites studied by Bravo-Cuevas et al. (2017), but show a similar level of preservation, with fragments being easily identifiable anatomically and in many cases also taxonomically. Previous research on the Skara Brae micromammal assemblages found significant concentrations of rodents within the site centre (Trench I) and periphery (Trench II), possibly related to human occupation (Romaniuk et al. 2016). Two likely explanations were that these are the remains produced as a by-product of pest control, or processing of micromammals as a food source. ...
... The micromammal inclusions show severe taphonomic alterations of bone and tooth surfaces characteristic of carnivore digestion, especially the loss of enamel or dentine and the thinning or cracking of bone. Previous analysis of the micromammal skeletal assemblage from Skara Brae (Romaniuk et al. 2016) did not provide such finds. Signs of digestion on the micromammal remains were scarce and did not follow a pattern that has previously been identified as indicative of any mammalian species. ...
Coprolites (fossilized faeces) can provide valuable insights into species' diet and related habits. In archaeozoological contexts, they are a potential source of information on human-animal interactions as well as human and animal subsistence. However, despite a broad discussion on coprolites in archaeology, such finds are rarely subject to detailed examination by researchers, perhaps due to the destructive nature of traditional analytical methods. Here, we have examined coprolitic remains from the Neolithic (third millennium BCE) settlement at Skara Brae, Orkney, using a range of modern methods: X-ray computed tomography, scanning electron microscopy, lipid and protein analysis (shotgun proteomics of the coprolite matrix as well as collagen peptide mass fingerprinting of isolated bone fragments). This combined approach minimised destructiveness of sampling, leaving sufficient material for subsequent study, while providing more information than traditional morphological examination alone. Based on gross visual examination, coprolites were predominantly attributed to domestic dogs (Canis familiaris), with morphologically identified bone inclusions derived from domestic sheep (Ovis aries) and common voles (Microtus arvalis). Partial dissection of a coprolite provided bone samples containing protein markers akin to those of domestic sheep. Considering the predominance of vertebral and distal limb bone fragments, Skara Brae dogs were probably consuming human butchery or meal refuse, either routinely fed to them or scavenged. The presumably opportunistic consumption of rodents may also have played a role in pest control.
... That such voyages occurred, as part of a strategy to enhance power and create a new social order, is hard to deny; indeed, more distant voyages between Orkney and the Continent are attested to by the appearance of the Orkney vole (Microtus arvalis orcadensis), between 3455 BC and 3100 BC (95% probability; 3315-3135 BC at 64% probability: Bayliss et al. 2017, figs 4 and 5 and table S.3). DNA analysis has demonstrated that this species must have come by boat from the Continent -although, as explained by Sheridan and Pétrequin (2014), not from Belgium as others have claimed -and it may well have been used as a self-replenishing food source on the journey; it is known to have been cooked and eaten at Skara Brae (Romaniuk et al. 2016). ...
This contribution offers an overview of the appearance, spread and regionally specific developmental trajectories of funerary monuments in Neolithic Scotland, setting these within the broader context of the arrival of farming groups from Brittany and northern France in the early centuries of the 4th millennium, and the subsequent expansion of farming to the Northern and Western Isles and the northernmost part of Scotland a few generations later. The monuments are also considered within the broader picture of Scottish Neolithic funerary practices more generally. Possible reasons for the monuments' specific developmental trajectories are offered, with competitive conspicuous consumption argued to be a major driver of developments in northern Scotland and Orkney during the second half of the 4th millennium. Bayesian models are presented for the first time for Clyde cairns and for mainland Scottish Orkney-Cromarty type passage tombs, and a chronological conundrum concerning Maeshowe-type passage tombs is explored.
... By this period, ca. 5000 years BP, the longtailed field mouse (Apodemus sylvaticus) and the common vole (Microtus arvalis) had been introduced to the archipelago (Romaniuk et al. 2016). The Orkney vole is Associate editor: Bastiaan Star * Pascale Chevret pascale.chevret@univ-lyon1.fr recognisable by its large size and characteristic tooth shape (Cucchi et al. 2014). ...
... This could contribute to their restricted overland dispersal in Mainland, given that wood mice were introduced there several millennia before the arrival of house mice (e.g. (Romaniuk et al. 2016)). Indeed, competition with the wood mouse appeared to be an important factor influencing tooth-shape diversification of house mice from the islands of Orkney (Ledevin et al. 2016). ...
Following human occupation, the house mouse has colonised numerous islands, exposing the species to a wide variety of environments. Such a colonisation process, involving successive founder events and bottlenecks, may either promote random evolution or facilitate adaptation, making the relative importance of adaptive and stochastic processes in insular evolution difficult to assess. Here, we jointly analyse genetic and morphometric variation in the house mice (Mus musculus domesticus) from the Orkney archipelago. Genetic analyses, based on mitochondrial DNA and microsatellites, revealed considerable genetic structure within the archipelago, suggestive of a high degree of isolation and long-lasting stability of the insular populations. Morphometric analyses, based on a quantification of the shape of the first upper molar, revealed considerable differentiation compared to Western European populations, and significant geographic structure in Orkney, largely congruent with the pattern of genetic divergence. Morphological diversification in Orkney followed a Brownian motion model of evolution, suggesting a primary role for random drift over adaptation to local environments. Substantial structuring of human populations in Orkney has recently been demonstrated, mirroring the situation found here in house mice. This synanthropic species may thus constitute a bioproxy of human structure and practices even at a very local scale.
... Studies on micromammals in archeology contribute to our knowledge about the faunal environments of human settlements and their interactions (Jenkins 2012, Romaniuk et al. 2016. Also, micromammals are considered better proxies, compared to macromammals, for assumptions on paleoclimate and paleoenvironment as they are more sensitive to small variations in climate and have greater capacity to adapt to environmental changes thanks to their success in adaptive radiation (Renfrew and Bahn 2011). ...
The primary aim of this study was to establish for the very first time the results on the rodent fauna gathered from the ongoing Neolithic excavations at the Tepecik- Çiftlik site in southern Cappadocia (Niğde, Turkey). So far, the fauna being study is represented by species of arvicolines (Microtus cf. arvalis and Arvicola cf. amphibius), murines (Mus cf. musculus), sousliks (Spermophilus xanthoprymnus), hamsters (Mesocricetus brandti) and molerats (Spalax xanthodon), and their remains are described in detail. Paleoenvironmental assumption based on common preferences of the fauna elements indicates, at first sight, a rather dry steppe environment with sparse plant cover or perennial short grasses. On the other hand, the presence of Arvicola cf. amphibius points out streams and marsh-like vegetation cover with bodies of water . The predation or burn marks observed on some specimens and the presence of subterraneous rodents raise questions concerning their taphonomy