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In this letter, we revisit a study we published in 2017, following comment in a paper by Marchesini et al. published in this volume. We provide some further analyses that help us to reinforce the original conclusions of our earlier paper, and to address the points raised by Marchesini et al. We conclude that the concerns raised in their review do not alter the inference we presented earlier, and we identify issues with analyses presented by Marchesini et al. that limit their utility. The key points of inference remain that this species in Europe shows remarkably low levels of diversity within populations and strong structure among populations which can be explained by a combination of natural and anthropogenic processes.

SUMMARY: The chance discovery of an 18th-century knuckle-bone floor at the National Trust property of Belton House in Lincolnshire prompted a review of all known post-medieval knuckle-bone floors i...

Species that evolved in temperate regions during the Pleistocene experienced periods of extreme climatic transitions. Consequent population fragmentation and dynamics had the potential to generate small, isolated populations where the influence of genetic drift would be expected to be strong. We use comparative genomics to assess the evolutionary influence of historical demographics and natural selection through a series of transitions associated with the formation of the genus Capreolus, speciation within this genus during the Quaternary and during divergence among European roe deer (C. capreolus) populations. Our analyses were facilitated by the generation of a new high‐coverage reference genome for the Siberian roe deer (Capreolus pygargus). We find progressive reductions in effective population size (Ne), despite very large census sizes in modern C. capreolus populations and show that low Ne has impacted the C. capreolus genome, reducing diversity and increasing linkage disequilibrium. Even so, we find evidence for natural selection shared among C. capreolus populations, including a historically documented founder population that has been through a severe bottleneck. During each phylogenetic transition there is evidence for selection (e.g. using evidence from non‐synonymous change or population data), including at loci associated with diapause (delayed embryonic development), a phenotype restricted to this genus among the even‐toed ungulates. Together these data allow us to assess expectations for the origin and diversification of a mammalian genus during a period of extreme environmental change.

International audience; AbstractSusceptibility to prion diseases is largely determined by the sequence of the prion protein gene (PRNP), which encodes the prion protein (PrP). The recent emergence of chronic wasting disease (CWD) in Europe has highlighted the need to investigate PRNP gene diversity in European deer species, to better predict their susceptibility to CWD. Here we report a large genetic survey of six British deer species, including red (Cervus elaphus), sika (Cervus nippon), roe (Capreolus capreolus), fallow (Dama dama), muntjac (Muntiacus reevesii), and Chinese water deer (Hydropotes inermis), which establishes PRNP haplotype and genotype frequencies. Two smaller data sets from red deer in Norway and the Czech Republic are also included for comparison. Overall red deer show the most PRNP variation, with non-synonymous/coding polymorphisms at codons 98, 168, 226 and 247, which vary markedly in frequency between different regions. Polymorphisms P168S and I247L were only found in Scottish and Czech populations, respectively. T98A was found in all populations except Norway and the south of England. Significant regional differences in genotype frequencies were observed within both British and European red deer populations. Other deer species showed less variation, particularly roe and fallow deer, in which identical PRNP gene sequences were found in all individuals analysed. Based on comparison with PRNP sequences of North American cervids affected by CWD and limited experimental challenge data, these results suggest that a high proportion of wild deer in Great Britain may be susceptible to CWD.

The archaeological record concerning the distribution and timing of fallow deer translocation across the Mediterranean has been growing in the last years. This knowledge has provided important insights into the movement, trade patterns and ideology of past societies. Unfortunately, the dispersal of fallow deer to the western part of the Mediterranean is insufficiently understood. To fill this gap, this article presents the results of a multidisciplinary investigation (combining the zooarchaeological evidence with AMS radiocarbon dating, isotope analyses and preliminary aDNA results) from a set of remains recovered from archaeological sites on Mallorca (Balearic Islands, Western Mediterranean). The purpose is to explore the timing, circumstances and meaning of the fallow deer's introduction to this island, as well as their subsequent management and extirpation. The results of these proxies confirm the arrival of the fallow deer during the Roman period and, after a short expansion with the establishment of vivaria, its disappearance after the Byzantine period., This research was conducted as part of the Dama International Project, funded by the Arts and Humanities Research Council (Standard Grant No. AH/I026456/1), supported by research leave funded by the University of Nottingham. It is also a contribution of the Research Project “Cambios holocénicos en la biodiversidad animal de las islas de la Macaronesia y de Baleares”/(CGL2012- 38087) of the Spanish Ministerio de Ciencia e Innovación.

Barbuda and Antigua's national animal is the fallow deer, Dama dama dama, a species native to the eastern Mediterranean that has been transported around the world by people during the last 8,000 years. The timing and circumstances by which fallow deer came to be established on Barbuda are currently uncertain but, by examining documentary, osteological and genetic evidence, this paper will consider the validity of existing theories. It will review the dynamics of human-Dama relationships from the 1500s AD to the present day and consider how the meaning attached to this species has changed through time: from a symbol of colonial authority and dominance, to a 'walking larder' after the slave emancipation of 1834, and now an important part of the island's economy and cultural heritage that requires careful management.

Species that have been translocated and otherwise manipulated by humans may show patterns of population structure that reflect those interactions. At the same time, natural processes shape populations, including behavioural characteristics like dispersal potential and breeding system. In Europe, a key factor is the geography and history of climate change through the Pleistocene. During glacial maxima throughout that period, species in Europe with temperate distributions were forced south, becoming distributed among the isolated peninsulas represented by Anatolia, Italy and Iberia. Understanding modern patterns of diversity depends on understanding these historical population dynamics. Traditionally, European fallow deer (Dama dama dama) are thought to have been restricted to refugia in Anatolia and possibly Sicily and the Balkans. However, the distribution of this species was also greatly influenced by human-mediated translocations. We focus on fallow deer to better understand the relative influence of these natural and anthropogenic processes. We compared modern fallow deer putative populations across a broad geographic range using microsatellite and mitochondrial DNA loci. The results revealed highly insular populations, depauperate of genetic variation and significantly differentiated from each other. This is consistent with the expectations of drift acting on populations founded by small numbers of individuals, and reflects known founder populations in the north. However, there was also evidence for differentiation among (but not within) physically isolated regions in the south, including Iberia. In those regions we find evidence for a stronger influence from natural processes than may be expected for a species with such strong, known anthropogenic influence.

The potential impact of population bottlenecks and founder events on genetic diversity and indirect measures of fitness (such as fluctuating asymmetry; FA) has important conservation implications. Here we take advantage of historical events that generated a remnant roe deer (Capreolus capreolus) population in the north of the British Isles that retained diversity, while populations in the south were apparently extirpated during the early mediaeval era. The southern population was later re-established from small founder populations of introduced European roe deer starting in the 19th century. We assess the impact of these events, using the northern remnant population as a reference, based on measures of FA at 16 bilateral cranial traits. Comparing the northern and southern populations we find evidence of differential impact on both the level of FA and the relationship between FA and levels of genetic diversity.

Evidence from numerous Pan-African savannah mammals indicates that open-habitat refugia existed in Africa during the Pleistocene, isolated by expanding tropical forests during warm and humid interglacial periods. However, comparative data from other taxonomic groups are currently lacking. We present a phylogeographic investigation of the African puff adder (Bitis arietans), a snake that occurs in open-habitat formations throughout sub-Saharan Africa. Multiple parapatric mitochondrial clades occur across the current distribution of B. arietans, including a widespread southern African clade that is subdivided into four separate clades. We investigated the historical processes responsible for generating these phylogeographic patterns in southern Africa using species distribution modelling and genetic approaches. Our results show that interior regions of South Africa became largely inhospitable for B. arietans during glacial maxima, whereas coastal and more northerly areas remained habitable. This corresponds well with the locations of refugia inferred from mitochondrial data using a continuous phylogeographic diffusion model. Analysis of data from five anonymous nuclear loci revealed broadly similar patterns to mtDNA. Secondary admixture was detected between previously isolated refugial populations. In some cases, this is limited to individuals occurring near mitochondrial clade contact zones, but in other cases, more extensive admixture is evident. Overall, our study reveals a complex history of refugial isolation and secondary expansion for puff adders and a mosaic of isolated refugia in southern Africa. We also identify key differences between the processes that drove isolation in B. arietans and those hypothesized for sympatric savannah mammals.

Human influence typically impacts on natural populations of conservation interest. These interactions are varied and sometimes complex, and may be negative and unintended or associated with conservation and management strategy. Understanding the details of how these interactions influence and are influenced by natural evolutionary processes is essential to the development of effective conservation strategies. In this study, we investigate a species in Britain that has experienced both negative impact through overhunting in historical times and management efforts through culls and translocations. At the same time, there are regional populations that have been less affected by human influence. We use mtDNA and nuclear microsatellite DNA markers to investigate patterns of connectivity and diversity and find multiple insular populations in Britain that probably evolved within the Holocene (when the habitat was free of ice). We identify three concurrent processes. First, surviving indigenous populations show highly provincial patterns of philopatry, maintaining and generating population structure on a small geographic scale. Second, founder populations into habitat extirpated of native populations have expanded, but remained largely insular. Third, introductions into established populations generate some admixture. We discuss the implications for the evolution of diversity of the integration of natural processes with anthropogenic influences on population size and distribution.

The semi-domestic status of the European fallow deer (Dama dama dama,) renders its ancient biogeography a reflection of human activity with the potential to provide important insights into the movement, trade patterns and ideology of past societies. Given this potential, fallow deer have received surprisingly little attention from scientists within the fields of archaeology, biology and zoology. Here we present new AMS radiocarbon dates, stable carbon and nitrogen isotope data and genetic evidence (the first ancient DNA sequences for the species) resulting from the analysis of a set of remains recovered from the Roman settlement at Monkton on the Isle of Thanet, Kent, UK. By viewing our results against the very limited available comparative data, this paper provides new information for the establishment and management of fallow deer in Britain. We argue that much more could be achieved with even a slight increase in sample sizes and a plea is made for greater research into this culturally significant species.

The Anglo-Normans first introduced fallow deer (Dama dama) to Ireland in the thirteenth century, however no biomolecular research has previously been undertaken to examine the timing, circumstances and impact of the arrival of this species. This study combines historical, zooarchaeological, genetic and isotopic data from both medieval and post-medieval samples to address this lack of research. The paper identifies a peak in the presence of fallow deer in Ireland between the thirteenth and fourteenth centuries, with a corresponding peak in documentary evidence for their presence in the thirteenth century. The deer are predominantly male, and from castle sites, supporting the historical evidence for their link with elite hunting. The English origin of the source populations shows correspondence between the documentary evidence, suggesting a western bias-and genetic evidence-with a similarity to southern and western England. Furthermore a stable isotope study identifies two possible first-generation imports, one dating from the medieval period and one from the post-medieval period. British Arts and Humanities Research Council

Extant patterns of population structure and levels of diversity are a consequence of factors that vary in both space and time. Our objective in this study is to investigate a species that has responded to both natural and anthropogenic changes in ways that have shaped modern populations and provide insight into the key processes. The roe deer (Capreolus capreolus) is one of the two species of deer native to Britain. During the last glacial maximum (LGM), the British habitat was largely under ice and there was a land bridge to mainland Europe. As the Earth warmed during the early Holocene, the land bridge was lost. Subsequent hunting on the British mainland left the southern region extirpated of roe deer, whereas a refugial population remained in the north. Later reintroductions from Europe led to population expansion, especially in southern United Kingdom. Here, we combine data from ancient and modern DNA to track population dynamics and patterns of connectivity, and test hypotheses about the influence of natural and anthropogenic environmental change. We find that past expansion and divergence events coincided with a warming environment and the subsequent closure of the land bridge between Europe and the United Kingdom. We also find turnover in British roe deer haplotypes between the late-Holocene and modern day that have likely resulted from recent human disturbance activities such as habitat perturbation, overhunting and restocking.