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Phylogeographic, ancient DNA, fossil and morphometric analyses reveal ancient and modern introductions of a large mammal: The complex case of red deer (Cervus elaphus) in Ireland

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... As mentioned above, we favour red deer as the likely candidate species for the Category 1 footprint tracks, but do not exclude ovicaprids. At present, the available evidence suggests that red deer were introduced to Ireland after the Early Neolithic: ~5800 cal BP, with the earliest faunal remains dated by association to ~5400 cal BP and by radiocarbon to ~4900-4500 cal BP (Carden et al. 2012). Ovicaprids also appear in the Irish record at roughly the same time: ~5400 and 4500 cal BP for sheep and goat respectively (Montgomery et al. 2014). ...
... If the Trawgar prints are contemporary with obtained radiocarbon dates (~7300-6300 cal BP), this would make them ~1500-500 years older than the presumed earliest of these introductions: that of red deer at ~5800 cal BP (Carden et al. 2012). If this is the case, then it implies the early introduction of red deer (or even sheep/goat) during the Mesolithic. ...
... However, we consider this an unlikely scenario for two main reasons. Firstly, it is at odds with the total absence of red deer or ovicaprids from Irish Mesolithic archaeological sites (Carden et al. 2012;Warren 2022). Secondly, the nature of the depositional environment, in particular the fact that deposition of a relatively thick clay layer implies standing water. ...
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During the winter storms of 2019, a deposit of organic-rich clay was fortuitously exposed in the intertidal zone of a beach near Streedagh (Co. Sligo). Impressed in the clay surface were a series of small (<10cm) paired indentations arranged in rough alignments. Their size and shape suggest that they are animal tracks; possibly red deer but not excluding sheep/goat. Animal tracks in Holocene sediments are well-documented from intertidal contexts in Great Britain but, to date, none have been reported from Ireland. This paper describes these tracks and discusses their chronological and paleoenvironmental context. Radiocarbon dating of the clay surface places it between ~7300–6300 cal BP, though for taphonomic reasons the tracks are argued to have been made later in the Holocene: probably after 5800 and before 4000 cal BP As such, the deposit provides an example of palaeoenvironmental evidence, that can be found even under the high-energy conditions characteristic of the Irish coast, and hints at the possibility that Holocene ichnological evidence can also be found here.
... Three species of deer are well distributed throughout Ireland: red deer, sika deer and fallow deer. Red deer are native to Ireland (but see Carden et al. 2012), whereas fallow deer were introduced by the Anglo-Normans in the 12th century (Beglane et al. 2018) and sika deer were initially introduced for ornamental purposes in the 1860s to the Powercourt Estate, Co. Wicklow, which is situated close to the capital city of Dublin (Powerscourt 1884). ...
... One of the population predicted hotspots was centred around the Killarney National Park, a herd under conservation measures such as a hunting ban in the area (Carden et al. 2012). ...
... Table 3. Kendall correlation coefficients between the relative abundances of red, sika and fallow deer as predicted by our integrated species distribution models (ISDMs) and 1) culling returns data at the Irish county level (n = 26) and 2) deer densities from faecal pellet counts at the Coillte property level (n = the prediction values extracted at the Coillte property centroids and the deer densities estimated in them by faecal pellet counts (n = 417). Page 11 of 14 three species have been introduced (or re-introduced in the case of the red deer; Powerscourt 1884, Carden et al. 2012, Beglane et al. 2018). ...
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Using geospatial data of wildlife presence to predict a species distribution across a geographic area is among the most common tools in management and conservation. The collection of high‐quality presence–absence (PA) data through structured surveys is, however, expensive, and managers usually have access to larger amounts of low‐quality presence‐only (PO) data collected by citizen scientists, opportunistic observations and culling returns for game species. Integrated species distribution models (ISDMs) have been developed to make the most of the data available by combining the higher‐quality, but usually scarcer and more spatially restricted, PA data with the lower‐quality, unstructured, but usually more extensive PO datasets. Joint‐likelihood ISDMs can be run in a Bayesian context using integrated nested laplace approximation methods that allow the addition of a spatially structured random effect to account for data spatial autocorrelation. Here, we apply this innovative approach to fit ISDMs to empirical data, using PA and PO data for the three prevalent deer species in Ireland: red, fallow and sika deer. We collated all deer data available for the past 15 years and fitted models predicting distribution and relative abundance at a 25 km² resolution across the island. Model predictions were associated to spatial estimate of uncertainty, allowing us to assess the quality of the model and the effect that data scarcity has on the certainty of predictions. Furthermore, we checked the performance of the three species‐specific models using two datasets, independent deer hunting returns and deer densities based on faecal pellet counts. Our work clearly demonstrates the applicability of spatially explicit ISDMs to empirical data in a Bayesian context, providing a blueprint for managers to exploit unexplored and seemingly unusable data that can, when modelled with the proper tools, serve to inform management and conservation policies.
... The genetic structure of large mammal species is affected by natural and anthropogenic factors. Anthropogenic impacts like selective hunting, translocations, and habitat destruction/fragmentation can blur natural patterns of genetic diversity and relationships (Frantz et al. 2006;Haanes et al. 2010;Dellicour et al. 2011;Carden et al. 2012;Stanton et al. 2016;Zachos et al. 2016;Galarza et al. 2017;Queirós et al. 2020). Such impacts, particularly in game species, can cause the disintegration of populations into several subpopulations with a more or less pronounced genetic exchange Willems et al. 2016;Iacolina et al. 2019;Mihalik et al. 2020). ...
... The red deer (Cervus elaphus L. 1758) is one of the most widespread and valuable European game species (Ludt et al. 2004;Milner et al. 2006). Consequently, its populations have been extensively managed, introduced, restocked, and selectively hunted for centuries or even millennia (Martínez et al. 2002;Haanes et al. 2010;Carden et al. 2012;Rivrud et al. 2013;Queirós et al. 2014;Hoffmann et al. 2016;Stanton et al. 2016;Frantz et al. 2017;Galarza et al. 2017). Furthermore, during the last decades, the keeping of deer in enclosures has spread all over the world, and the species is farmed for venison and antler products (Milner et al. 2006;Wada et al. 2010;Zachos and Hartl 2011;Bana et al. 2018). ...
... The blurred structuring could also be explained by humaninduced translocations, as such activities are thought to have concerned important game species for centuries or even millennia (Scandura et al. 2011;Zachos and Hartl 2011;Steinbach et al. 2018;Queirós et al. 2020). It is believed that the present gene pool of many European red deer populations is affected by human-induced translocations (Frantz et al. 2006;Carden et al. 2012;Krojerová-Prokešová et al. 2015;Stanton et al. 2016;Galarza et al. 2017;Iacolina et al. 2019;Queirós et al. 2020). However, the results of the assignment test, namely, the low number of misassigned deer, indicate that the transportation of deer from greater distances was not common in this region. ...
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After the last glacial, the Carpathian Basin was repopulated from either eastward or northward colonisation routes for various species; one of these was the emblematic member of the European megafauna, the red deer, Cervus elaphus. We analysed 303 red deer individuals from the middle of the region, in seven Hungarian game reserves, at ten microsatellite loci (C01, C229, T26, T108, T123, T156, T172, T193, T501, T507), to investigate the genetic diversity of these subpopulations. We discovered high levels of genetic diversity of red deer subpopulations; allelic richness values ranging 4.99-7.01, observed heterozygosity 0.729-0.800, polymorphic information content 0.722-0.806, and Shannon's information index 1.668-2.064. Multi-locus analyses indicated population admixtures of various degrees that corresponded to geographical location, and complex genetic structures were shown by clustering. Populations in the southwestern and the northeastern parts of the region formed two highly separated groups, and the red deer from populations in between them were highly admixed (in western Pannonia/Transdanubia, where the Danube flows into the Carpathian Basin). This pattern corresponds to the distribution of mitochondrial as well as Y-chromosome lineages. Assignment tests showed that a large fraction of individuals (29.4%) are found outside of their population of origin, indicating that the dispersal of red deer is rather common, which could be expected considering the life course of the species.
... Phylogeographic studies of the most widely studied group, the terrestrial mammals, have shown distinct mitochondrial DNA (mtDNA) lineages in small mammals (Searle et al., 2009;Stojak et al., 2016;Vega et al., 2020) and ungulates (Carden et al., 2012;Sommer et al., 2008) that are consistent with contraction and re-expansion from these refugial regions (Hewitt, 1999;Taberlet et al., 1998). ...
... This uncertainty and lack of resolution in phylogeographic structure outside the "traditional" (and well-established) refugia means that inferring post-glacial colonization patterns of previously glaciated regions during the LGM is challenging with more limited genetic marker sets. This is particularly evident in the British Isles and Scandinavia where these regions present differing problems in terms of how and when contemporary populations of terrestrial species reached these areas in post-glacial periods (Carden et al., 2012(Carden et al., , 2020Herman et al., 2014). ...
... The island of Ireland has long presented a biogeographical quandary in terms of how and when terrestrial species colonized it (Carden et al., 2012(Carden et al., , 2020McDevitt et al., 2011). The latitude of the area means that it was covered almost entirely by the ice sheet during the LGM and it did not become an island until approximately 15,000 years ago (almost twice as long as Britain; Edwards & Brooks, 2008). ...
Article
Carnivores tend to exhibit a lack of (or less pronounced) genetic structure at continental scales in both a geographic and temporal sense and this can confound the identification of post‐glacial colonization patterns in this group. In this study we used genome‐wide data (using Genotyping‐by‐Sequencing (GBS)) to reconstruct the phylogeographic history of a widespread carnivore, the red fox (Vulpes vulpes), by investigating broad‐scale patterns of genomic variation, differentiation and admixture amongst contemporary populations in Europe. Using 15,003 single nucleotide polymorphisms (SNPs) from 524 individuals allowed us to identify the importance of refugial regions for the red fox in terms of endemism (e.g. Iberia). In addition, we tested multiple post‐glacial re‐colonization scenarios of previously glaciated regions during the Last Glacial Maximum using an Approximate Bayesian Computation (ABC) approach that were unresolved from previous studies. This allowed us to identify the role of admixture from multiple source population post‐Younger Dryas in the case of Scandinavia and ancient land‐bridges in the colonization of the British Isles. A natural colonization of Ireland was deemed more likely than an ancient human‐mediated introduction as has previously been proposed and potentially points to an increased mammalian fauna on the island in the early post‐glacial period. Using genome‐wide data has allowed us to tease apart broad‐scale patterns of structure and diversity in a widespread carnivore in Europe that was not evident from using more limited marker sets and provides a foundation for next‐generation phylogeographic studies in other non‐model species.
... There is some debate, however, as to whether or not giant deer survived the Younger Dryas in Ireland. As Carden et al. (2012) have demonstrated, misidentification of deer species has been a reoccurring issue in reconstructing faunal biogeography. Additionally, as Monaghan (2017) points out, redating of some giant deer specimens indicate they are older than initially thought and pre-date the Younger Dryas. ...
... Isotopic data consistently reflect terrestrially-focused diets in Britain before 10,000 cal BP, with subsistence patterns becoming more variable throughout the Mesolithic (Pickard and Bonsall, 2020;Barton and Roberts, 2004). Terrestrial components of Mesolithic diets in Scotland are notably dominated by red deer and wild boar (Sus scrofa) (Kitchener et al., 2004;Mithen et al., 2020) e both of which were absent in Ireland following deglaciation (Carden et al., 2012;Warren et al., 2014). The Late Mesolithic site of Cnoc Coig in western Scotland is the only location where diets appear to have been heavily focused on marine resources (Pickard and Bonsall, 2020), and temporally corresponds to the exploitation of coastal resources and the spread of shell middens after 7000 cal BP (Mithen et al., 2020). ...
... Mesolithic hunters were well-adapted to hunting red deer throughout northern Europe. However, while red deer inhabited Ireland prior to the LGM, they were scarcely present across the island from the Younger Dryas until the Neolithic (Carden et al., 2012). Red deer not only served as a vital subsistence resource and held particular spiritual role in Mesolithic peoples' lives, but they were also critical in the construction of microlith composite tools e the hallmark of the European Mesolithic. ...
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Human migration throughout northern Europe following the Last Glacial Maximum is an ideal situation to investigate human colonization and adaptation in new landscapes. This is particularly so in Ireland, which possesses a distinctly compressed archaeological record compared to the rest of Europe. While various periods of Paleolithic occupations are well-documented throughout Europe, including Britain, the initial colonization of Ireland appears to be delayed until the Early Holocene. An assessment of archaeological and paleoenvironmental data suggests that inhospitable environmental conditions, specifically the absence of mature woodland ecosystems, substantially delayed the human colonization of Ireland. Once Mesolithic peoples reached Ireland, the absence of familiar fauna led them to quickly modify existing technologies. These local adaptions are reflected in the discontinuation of composite microlith technologies that characterize the rest of the European Mesolithic record. Within 1000 years of colonization, Mesolithic hunter-gatherers developed a uniquely Irish macrolith-based technology.
... Ireland's fauna is considered as impoverished relative to, and despite its proximity to, Britain, its nearest landmass (Yalden 1999). Most of our current knowledge regarding Irish pre-and post-Last Glacial Maximum (LGM) colonization processes and origins comes from studies of terrestrial mammals using skeletal remains and phylogeographical studies involving ancient and/or modern DNA, such as brown bear, pygmy shrew, red deer, stoat, red fox and badger (Mart ınkov a et al. 2007;Edwards et al. 2011;McDevitt et al. 2011McDevitt et al. , 2020Carden et al. 2012;O'Meara et al. 2012;Vega et al. 2020). These studies indicate post-LGM introductions or early colonization event(s) of these species to Ireland. ...
... These studies indicate post-LGM introductions or early colonization event(s) of these species to Ireland. The mammalian skeletal remains analysed in two previous studies, brown bear (Edwards et al. 2011) and red deer (Carden et al. 2012), were derived primarily from numerous limestone cave systems within Ireland, since there are few MIS 3 deposits containing faunal remains dated to before the LGM, for example the Aghnadarragh Interstadial (McCabe et al. 1987). For this reason, there have been difficulties in the identification of any substantial fauna from MIS 5 or older ( O Drisceoil & Jennings 2006;Coxon & McGarron 2009). ...
... Pretreatment methods denoted by AI are ion-exchanged gelatine dates and AF is ultrafiltered collagen. An asterisk denotes previous identifications of red deer; recent re-examination and aDNA extractions of this bone confirmed species identification as reindeer (see Carden et al. 2012). See Table 1 It is also probable on the basis of the dates from Castlepook Cave and their abundant presence that reindeer and brown bear were present throughout this 20 000-year period. ...
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With the improvements in the techniques of radiocarbon dating since the 1990s and the pretreatment of bone samples, it has become apparent that some of the radiocarbon dates from Irish caves such as Castlepook Cave may not be as accurate as previously determined. The faunal remains and sediments within Castlepook Cave are important as this cave is one of only two caves in Ireland in which both pre‐ and post‐Last Glacial Maximum sediments containing faunal remains are found. In this study, samples of identified bones were re‐dated from this cave as well as two others (Foley and Shandon caves) from the same region in the south‐southeast of Ireland. The new dates from Castlepook Cave disclosed results that were significantly older by between c . 5000 and 12 000 radiocarbon years. This was not as apparent in dates obtained from Foley and Shandon caves, where the new dates were older by c . 4000 radiocarbon years on average. The results imply a much older presence of certain mammals during MIS 3 in Ireland and suggest that certain species, notably giant deer and spotted hyaena, may have become locally extirpated before the Last Glacial Maximum, around 40 cal. ka BP.
... Although red foxes from central Italy are identified as a distinct cluster in fastSTRUCTURE, admixture was identified with neighbouring populations north of the Alps in central Europe and the Balkans (Figs. 1B and 1C). (48). It has existed as an island for approximately 15,000 years (almost twice as long as Britain; 49) and humans have been proposed as the primary mechanism of transport for its mammalian fauna in ancient and modern times (10,48). ...
... (48). It has existed as an island for approximately 15,000 years (almost twice as long as Britain; 49) and humans have been proposed as the primary mechanism of transport for its mammalian fauna in ancient and modern times (10,48). An estimate of 10.2 kyrs BP was estimated for a split between Irish and British red foxes using mtDNA data, but with a 95% CI range that incorporated the possibility of natural colonization before Ireland became an island (18). ...
... mountain hare Lepus timidus and arctic lemming Dicrostonyx torquatus) were also present in the early post-glacial period. While humans were an important factor in determining later faunal assemblages on Ireland (10,48), the early post-glacial period clearly warrants further investigation on the island (41,48). ...
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Carnivores tend to exhibit a lack of (or less pronounced) genetic structure at continental scales in both a geographic and temporal sense using various mitochondrial DNA markers on modern and/or ancient specimens. This tends to confound the identification of refugial areas and post-glacial colonization patterns in this group. In this study we used Genotyping-by-Sequencing (GBS) to reconstruct the phylogeographic history of a widespread carnivore, the red fox (Vulpes vulpes), in Europe by investigating broad-scale patterns of genomic variation, differentiation and admixture amongst contemporary populations. Using 15,003 single nucleotide polymorphisms (SNPs) from 524 individuals allowed us to identify the importance of refugial regions for the red fox in terms of endemism (e.g. Iberia) and sources of post-glacial re-expansion (e.g. Carpathians and Balkans) across northern regions of the continent. In addition, we tested multiple post-glacial re-colonization scenarios of previously glaciated regions during the Last Glacial Maximum using an Approximate Bayesian Computation (ABC) approach. We identified the role of ancient and temporary land-bridges in the colonization of Scandinavia and the British Isles, with a natural colonization of Ireland deemed more likely than an ancient human-mediated introduction as has previously been proposed. Using genome-wide data has allowed us to tease apart broad-scale patterns of structure and diversity in a widespread carnivore in Europe that was not always evident from using more limited marker sets.
... Primate populations that have experienced recent habitat loss and fragmentation show low levels of genetic diversity, due to elevated genetic drift (Mbora and McPeek, 2010) and in those species characterised by female philopatry, a mtDNA pattern which is relatively homogenised within populations, as is found in forest patches for example, but heterogeneous between populations (Shimada, 2000;Mbora and McPeek, 2010) is typically seen. These discrete habitats act as a mechanism of geographic isolation and it is probable that recent phylogeographic history has shaped the pattern of genetic differentiation (Carden et al. 2012) in many of the African monkeys. Although there is often a level of discordance between mitochondrial phylogeny and morphology, many of the African non-human primate taxa such as the guenons and papionins can be broken down into several well-supported major haplogroups, reflecting distinct geographic populations ). ...
... The use of ancient DNA (aDNA) over the last two decades has become an invaluable analytical tool to investigate extinct species and populations (Callaway, 2011;Brace et al. 2014;Brace et al. 2015) and temporal genetic changes (Gilbert et al. 2005;Helgen et al. 2008;Carden et al. 2012;Langille et al. 2014). The techniques were originally designed to successfully extract the residual amounts of DNA remaining in samples that are hundreds or thousands of years old (Rohland and Hofreiter, 2007) and it was first used to extract DNA from the extinct quagga (Equus quagga) (Higuchi et al. 1984). ...
... Compared with contemporary DNA preparations from fresh tissue, aged material subjected to aDNA techniques are usually much shorter in length. However, more recently-aged samples that may not even exceed 100 years old can also be subjected to aDNA techniques (Helgen et al. 2008;Carden et al. 2012), in instances where sample preservation may be poor such as in historic museum collections where environmental conditions are not controlled and where these conditions are often characterised as being warm and damp. In instances where fresh and 'old' material are directly compared, then because of variation in sample preservation and quality, all samples (both historic and modern) can be subjected to aDNA techniques in order to standardise the methods. ...
Conference Paper
Across the world, islands were and still are inhabited by unique species, often restricted to their own island and found nowhere else. After their ancestors managed to reach an island from a mainland population and become isolated from this mainland and its ecological restrictions, they often evolved spectacular adaptations. The more extrinsic barriers to gene flow there are and the more distant the populations, the greater the probability of a profound genetic and morphological change. Whereas many other insular mammalian taxa such as proboscideans, rodents and cervids react in readily identifiable trends, primates respond in varied and unpredictable ways. In order to better understand the underlying evolutionary principles behind island speciation, this thesis focuses on the small cercopithecine monkeys taken from western Africa to the Caribbean during the transatlantic slave trade. These Chlorocebus monkeys inhabit Nevis, St Kitts and Barbados but have long been assumed to originate solely from the Senegambia region. This thesis investigates the very early phase associated with island separation, using mitochondrial analysis and 3D geometric morphometric techniques to thoroughly assess whether any changes are present in these populations. An additional assessment is made of the three western African species of Chlorocebus, which is still largely subject to taxonomic discord. The results here show that the existing taxonomic status of African Chlorocebus does not fully describe the whole situation and that changes should be made to resolve this. The molecular results from this thesis show that rather than originating from one Senegambian location, Caribbean Chlorocebus monkeys instead originate from three different African species, across the entire western African coast, meaning their current designation as ‘African green monkeys’ is inaccurate. Additionally, morphological adaptations within these three insular populations are also already apparent, both across the three island groups and between Caribbean and African Chlorocebus.
... As in the case of most temperate European mammals, the current phylogenetic status, diversity and distribution of red deer (Cervus elaphus L. 1758) may be the consequence of multiple climatic fluctuations of the Pleistocene epoch (Hewitt, 2004;Sommer et al., 2008;Meiri et al., 2013;Stanton et al., 2016). It is assumed that the species was widely distributed in the southern part of Central Europe 60 000–25 000 years ago, but was subsequently restricted to refugia south of the permafrost in southern Europe during the last glacial maximum, 25 000–14 700 years ago, according to fossil evidence (Hewitt, 2004;Sommer et al., 2008;Sommer and Zachos, 2009;Carden et al., 2012;Stanton et al., 2016). In general, five major European refugia have been identified: the Balkans, the Iberian Peninsula with the southern part of France, the Italian Peninsula, the Carpathians and the Caspian/Caucasus region (Hewitt, 2004;Sommer and Zachos, 2009;Karaiskou et al., 2014;Carranza et al., 2016). ...
... These suture or admixture zones can provide useful insights into evolutionary mechanisms of reproductive isolation, selection, and the genetic basis of local adaptation.The red deer is one of the most widespread, most abundant and most important game species in Europe (Ludt et al., 2004;Milner et al., 2006;Zachos and Hartl, 2011). As a result, red deer have been extensively managed, introduced, restocked, and selectively hunted throughout its history and distribution area (Frantz et al., 2006;Milner et al., 2006;Csányi and Lehoczki, 2010;Haanes et al., 2010;Zachos and Hartl, 2011;Carden et al., 2012;Stanton et al., 2016), and are farmed for meat and for antler products (Csányi and Lehoczki, 2010;Wada et al., 2010;Zachos and Hartl, 2011;Frank et al., 2016). The species has been the subject of several studies with a phylogeographic focus at local (Zachos et al., 2003;Feulner et al., 2004;Haanes et al., 2010;Carden et al., 2012;Fickel et al., 2012;Karaiskou et al., 2014;Krojerová-Prokešová et al., 2015;Markov et al., 2015;Borowski et al., 2016;Carranza et al., 2016;Stanton et al., 2016) as well as larger geographical scales (Ludt et al., 2004;Skog et al., 2009;Niedziałkowska et al., 2011;Meiri et al., 2013). ...
... As a result, red deer have been extensively managed, introduced, restocked, and selectively hunted throughout its history and distribution area (Frantz et al., 2006;Milner et al., 2006;Csányi and Lehoczki, 2010;Haanes et al., 2010;Zachos and Hartl, 2011;Carden et al., 2012;Stanton et al., 2016), and are farmed for meat and for antler products (Csányi and Lehoczki, 2010;Wada et al., 2010;Zachos and Hartl, 2011;Frank et al., 2016). The species has been the subject of several studies with a phylogeographic focus at local (Zachos et al., 2003;Feulner et al., 2004;Haanes et al., 2010;Carden et al., 2012;Fickel et al., 2012;Karaiskou et al., 2014;Krojerová-Prokešová et al., 2015;Markov et al., 2015;Borowski et al., 2016;Carranza et al., 2016;Stanton et al., 2016) as well as larger geographical scales (Ludt et al., 2004;Skog et al., 2009;Niedziałkowska et al., 2011;Meiri et al., 2013). Based on previous studies of mitochondrial DNA (mtDNA) sequences, three deeply divergent lineages of red deer are known within Europe, enabling classification of individuals into Iberian (termed haplogroup A sensuSkog et al., 2009), Balkan (haplogroup C) and Mediterranean (haplogroup B) groups, suggesting retraction into three separate refugia during the last glaciation, the Iberian Peninsula, the Balkans and the Italian Peninsula (Ludt et al., 2004;Skog et al., 2009;Meiri et al., 2013;Karaiskou et al., 2014;Carranza et al., 2016). ...
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Changes in the distributional range of European red deer (Cervus elaphus) during the Quaternary and the recolonization of Europe from different refugia, created a clear phylogeographical pattern. In Central Europe, two distinct − Iberian and Balkan − lineages of European red deer are present; however, this suture zone has not yet been studied in detail. The sequences of the complete mitochondrial control region were used to investigate the genetic structure of the red deer in the Carpathian Basin, a region where both lineages are expected to be present. We sequenced the complete control region of 96 red deer samples, discovered 39 haplotypes, and found high levels of haplotype diversity (H = 0.806 for the southwestern and H = 0.954 for the northeastern population). We discovered a high number of haplotypes belonging to both lineages, as well as unique haplotypes, which do not fit into described haplotypes. This is in accordance with previous assumptions that the Carpathian Basin is a suture zone between the Iberian and the Balkan red deer haplogroups.
... There is evidence for the maritime colonization of Europe by Neolithic people of the Near East via the Mediterranean [5], with isolated long-distance sea travel around the Mediterranean as early as the Mesolithic [6]. There are fewer such examples of ancient sea travel in northern Europe, although it is thought that humans introduced fur-bearing species to the Scottish Isles from the Neolithic onwards [7] and transported red deer (Cervus elaphus) from Britain to Ireland during the Irish Bronze Age [8]. There is also evidence of long-distance movement of wildlife (Orkney vole) by Neolithic people between continental Europe and Orkney [4]. ...
... Ancient DNA (aDNA) can give insights into the dynamics of populations that would not be possible to determine from archaeological information or DNA analysis of modern populations alone [25,26]. This is particularly relevant in the case of red deer, due to the complex history of human-mediated translocations [8]. This study uses DNA from opportunistically collected deer bone to provide a unique, serially sampled dataset from a restricted geographical range. ...
... We included all available and comparable aDNA sequences from previously published studies, including Norway [20], Ireland [8] and Italy [21]. When an age of a sample was estimated in one of these studies, we used the midpoint of this estimate to assign to a time period. ...
Article
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Red deer (Cervus elaphus) have played a key role in human societies throughout history, with important cultural significance and as a source of food and materials. This relationship can be traced back to the earliest human cultures and continues to the present day. Humans are thought to be responsible for the movement of a considerable number of deer throughout history, although the majority of these movements are poorly described or understood. Studying such translocations allows us to better understand ancient human–wildlife interactions, and in the case of island colonizations, informs us about ancient human maritime practices. This study uses DNA sequences to characterise red deer genetic diversity across the Scottish islands (Inner and Outer Hebrides and Orkney) and mainland using ancient deer samples, and attempts to infer historical colonization events. We show that deer from the Outer Hebrides and Orkney are unlikely to have originated from mainland Scotland, implying that humans introduced red deer from a greater distance. Our results are also inconsistent with an origin from Ireland or Norway, suggesting long-distance maritime travel by Neolithic people to the outer Scottish Isles from an unknown source. Common haplotypes and low genetic differentiation between the Outer Hebrides and Orkney imply common ancestry and/or gene flow across these islands. Close genetic proximity between the Inner Hebrides and Ireland, however, corroborates previous studies identifying mainland Britain as a source for red deer introductions into Ireland. This study provides important information on the processes that led to the current distribution of the largest surviving indigenous land mammal in the British Isles.
... Ancient DNA is also more likely to survive in bones recovered from caves compared to bones from other contexts. For instance, in a recent Irish project nine red deer bones from five different caves all yielded aDNA whereas 11 red deer bones from ten aboveground sites and one cave did not yield any aDNA (Carden et al. 2012). At present aDNA has only been successfully extracted from one prehistoric human skeleton in Ireland -that of a 2-3 year old child of Bronze Age date from Glencurran Cave (Dowd and Bunce 2009, 9). ...
... The horse tooth from Plunkett Cave is not an isolated occurrence. A small but significant number of horse, red deer and wolf bones from five caves have returned Iron Age dates (Table 7.1) as part of two different radiocarbon dating projects Carden et al. 2012). The number of Iron Age dates is disproportionately larger than those from any other prehistoric period, particularly considering that the Iron Age is the shortest of all periods. ...
... The current distribution of red deer is assumed to be strongly influenced primarily by colonization history during the Late Pleistocene and the early Holocene (Sommer et al., 2008) and also by human activities (e.g. Carden et al., 2012;Doan et al., 2017;Fernandez-Garcia et al., 2014). The red deer is also one of the most abundant large mammal species in archaeological sites dated to the Late Pleistocene across Europe (Sommer & Nadachowski, 2006). ...
... In addition to natural environmental change, human wildlife management over recent centuries has affected red deer populations via selective hunting, translocations, isolation of deer in enclosures and through the creation of barriers resulting in habitat fragmentation (Carden et al., 2012;Hartl et al., 2003;. ...
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Aim: The Expansion-Contraction model has been used to explain the responses of species to climatic changes. During periods of unfavourable climatic conditions, species retreat to refugia from where they may later expand. This paper focuses on the palaeoecology of red deer over the past 54 ka across Europe and the Urals, to reveal patterns of change in their range and explore the role of environmental conditions in determining their distribution. Location: Europe and western Asia to 63°E. Taxon: Red deer (Cervus elaphus). Methods: We collected 984 records of radiocarbon-dated red deer subfossils from the Late Pleistocene and the Holocene, including 93 original dates. For each deer sample we compiled climatic and biome type data for the corresponding time intervals. Results: During the last 54 ka changes in red deer range in Europe and the Urals were asynchronous and differed between western and eastern Europe and western Asia due to different environmental conditions in those regions. The range of suitable areas for deer during the Last Glacial Maximum (LGM) was larger than previously thought and covered vast regions not only in southern but also in western and eastern Europe. Throughout the period investigated the majority of specimens inhabited forests in the temperate climatic zone. The contribution of forests in deer localities significantly decreased during the last 4 ka, due to deforestation of Europe caused by humans. Mean January temperature was the main limiting factor for species distribution. Over 90% of the samples were found in areas where mean January temperature was above −10°C. Main conclusions: Red deer response to climatic oscillations are in agreement with the Expansion-Contraction model but in contradiction to the statement of only the southernmost LGM refugia of the species. During the last 54 ka red deer occurred mostly in forests of the temperate climatic zone.
... The new D-loop sequences (670 bp) were aligned together with 624 sequences retrieved from GenBank, comprising the representative haplotypes from the majority of red deer studies published throughout Europe and North Africa up to 2017 [13,17,24,30,31,34,[64][65][66][67][68][69][70][71][72][73][74][75][76][77][78][79][80][81][82]. Phylogeographic analyses were performed at two geographical scales: the central and northern European level and the Iberian level. ...
... However, the absence of fossil records from the LGM in the British Isles (Fig 9; see also S8 Fig) makes this hypothesis unlikely, even though the surrounded areas of continental self remain unexplored for fossil remains. Furthermore, the high levels of mitochondrial diversity in red deer in the British Isles may also reflect the intensity of sampling for mitochondrial studies [67,68,71] and/or massive re-introductions from the rest of western Europe [80]. Moreover, a demographic expansion of red deer populations in the British Isles was only estimated around 7-4 ky BP, which is most simply explained by the impact on red deer populations of the transition from pre-Neolithic hunter-gatherer societies to Neolithic farming, known as the Neolithic revolution, which was one of the most pronounced cultural changes in European prehistory [119]. ...
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The red deer (Cervus elaphus) is a widespread wild ungulate in Europe that has suffered strong anthropogenic impacts over their distribution during the last centuries, but also at the present time, due its economic importance as a game species. Here we focus on the evolutionary history of the red deer in Iberia, one of the three main southern refugial areas for temperate species in Europe, and addressed the hypothesis of a cryptic refugia at higher latitudes during the Last Glacial Maximum (LGM). A total of 911 individuals were sampled, genotyped for 34 microsatellites specifically developed for red deer and sequenced for a fragment of 670 bp of the mitochondrial (mtDNA) D-loop. The results were combined with published mtDNA sequences, and integrated with species distribution models and historical European paleo-distribution data, in order to further examine the alternative glacial refugial models and the influence of cryptic refugia on European postglacial colonization history. Clear genetic differentiation between Iberian and European contemporary populations was observed at nuclear and mtDNA levels, despite the mtDNA haplotypes central to the phylogenetic network are present across western Europe (including Iberia) suggesting a panmictic population in the past. Species distribution models, fossil records and genetic data support a timing of divergence between Iberian and European populations that overlap with the LGM. A notable population structure was also found within the Iberian Peninsula, although several populations displayed high levels of admixture as a consequence of recent red deer translocations. Five D-loop sub-lineages were found in Iberia that belong to the Western European mtDNA lineage, while there were four main clusters based on analysis of nuclear markers. Regarding glacial refugial models, our findings provide detailed support for the hypothesis that red deer may have persisted in cryptic northern refugia in western Europe during the LGM, most likely in southern France, southern Ireland, or in a region between them (continental shelf), and these regions were the source of individuals during the European re-colonization. This evidence heightens the importance of conserving the high mitochondrial and nuclear diversity currently observed in Iberian populations.
... A similar mixture of mtDNA lineages has been found in populations from Bulgaria, Hungary and the Czech Republic (Markov et al., 2012(Markov et al., , 2015Krojerová-Prokešová, Barančeková & Koubek, 2015). Red deer translocations have also been suggested for populations in Germany (Ludt et al., 2004), Scotland (Carden et al., 2012) and Spain (Fernández-García et al., 2013). Moreover, Kuznetsova et al. (2007) found the mtDNA lineages of both Western and Eastern red deer in the Ukrainian population, which probably resulted from the introduction of Cervus elaphus maral to Europe. ...
... This suggests that Crimea was probably colonized naturally by immigrant red deer from Western Europe. However, translocations by humans cannot be excluded, as it is well known that this happened in Sardinia, Corsica, Crete and Ireland during the Neolithic period (Hajji et al., 2008;Carden et al., 2012;Harris, 2014). The reasons for such management are not well known, but studies by Sykes et al. (2006Sykes et al. ( , 2011 and Davis & MacKinnon (2009) have showed that fallow deer (Dama dama) was introduced to Britain and Portugal by the Romans, probably for keeping them in special parks (vivaria), which enhanced their owner's status. ...
Article
The present distribution of many species is a result of climatic changes during the Pleistocene and human activity. The impact of climate has been accompanied by restrictions of populations into refugia during glacial periods, and subsequent expansions during more favourable conditions, whereas human influence has been associated with hunting practices and translocations. One mammalian species that has been subject to such transformations is the red deer, Cervus elaphus, but the exact nature of these changes has been difficult to determine using only modern DNA. In this study, we obtained new cytochrome b sequences from subfossil remains of deer found in the Crimean Peninsula. A comparison of these sequences with the available recent and ancient sequences allowed to us to reconstruct phylogeographic relationships between Cervus lineages and to determine their potential migration routes at both local and Eurasian scales. Our analyses showed that the Crimean Peninsula was not a glacial refugium for red deer, but rather that red deer colonized Crimea in three independent waves from both Western and Eastern red deer populations. The immigrations were related to local extinctions and replacements of native populations.
... Humans may also have altered Ireland's plant communities and woodland cover through reintroductions of species such as red deer (C. elaphus), which had disappeared from the island during the last Ice Age and were reintroduced in the Neolithic (28), and introductions of species such as fallow deer (Dama dama) during the medieval (29). These events also do not correspond temporally with changes observed in later Holocene Irish fauna d 15 N values and therefore would not have been a primary cause of landscape changes that influenced the N cycle at a regional scale. ...
... Faunal samples (n = 712) from at least 90 (this number is the minimum estimate as several extinct Irish Giant Deer from natural history collections do not have associated provenance information) archaeological and natural history sites in 16 Irish counties were collected from private-sector archaeology companies, universities, and museums. Data for 45 additional animals were sourced from the literature (table S2) (28,(36)(37)(38). Species identifications were reconfirmed by an archaeozoologist (F.B.). ...
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Humans have always affected their ecosystems, but finding evidence for significant and lasting changes to preindustrial landscapes is rare. We report on human-caused changes to the nitrogen cycle in Ireland in the Bronze Age, associated with intensification of agriculture and animal husbandry that resulted in long-term changes to the nitrogen isotope values of animals (wild and domesticates) during the Holocene. Major changes to inputs and cycling of soil nitrogen occurred through deforestation, land clearance and management, and more intensive animal husbandry and cereal crop cultivation in the later Bronze Age; after this time, the Irish landscape took on its current form. Within the debate concerning the onset of the Anthropocene, our data suggest that human activity in Ireland was significant enough in the Bronze Age to have long-term impact, thereby marking a profound shift in the relationship between humans and their environment.
... A similar mixture of mtDNA lineages has been found in populations from Bulgaria, Hungary and the Czech Republic (Markov et al., 2012(Markov et al., , 2015Krojerová-Prokešová, Barančeková & Koubek, 2015). Red deer translocations have also been suggested for populations in Germany (Ludt et al., 2004), Scotland (Carden et al., 2012) and Spain (Fernández-García et al., 2013). Moreover, Kuznetsova et al. (2007) found the mtDNA lineages of both Western and Eastern red deer in the Ukrainian population, which probably resulted from the introduction of Cervus elaphus maral to Europe. ...
... This suggests that Crimea was probably colonized naturally by immigrant red deer from Western Europe. However, translocations by humans cannot be excluded, as it is well known that this happened in Sardinia, Corsica, Crete and Ireland during the Neolithic period (Hajji et al., 2008;Carden et al., 2012;Harris, 2014). The reasons for such management are not well known, but studies by Sykes et al. (2006Sykes et al. ( , 2011 and Davis & MacKinnon (2009) have showed that fallow deer (Dama dama) was introduced to Britain and Portugal by the Romans, probably for keeping them in special parks (vivaria), which enhanced their owner's status. ...
Article
ity. The impact of climate has been accompanied by restrictions of populations into refugia during glacial periods, and subsequent expansions during more favourable conditions, whereas human influence has been associated with hunting practices and translocations. One mammalian species that has been subject to such transformations is the red deer, Cervus elaphus, but the exact nature of these changes has been difficult to determine using only modern DNA. In this study, we obtained new cytochrome b sequences from subfossil remains of deer found in the Crimean Peninsula. A comparison of these sequences with the available recent and ancient sequences allowed to us to recon- struct phylogeographic relationships between Cervus lineages and to determine their potential migration routes at both local and Eurasian scales. Our analyses showed that the Crimean Peninsula was not a glacial refugium for red deer, but rather that red deer colonized Crimea in three independent waves from both Western and Eastern red deer populations. The immigrations were related to local extinctions and replacements of native populations.
... Ancient DNA studies have revealed that the present distribution of phylogeographic lineages may be more limited than before the anthropogenic impacts on large mammal populations, including those of red deer 27 (for an example in brown bears see refs 28 and 29). They have also been performed to elucidate the recolonization of Scandinavia 30 and specifically to address the colonization of islands: Carden et al. 31 were able to distinguish ancient and modern introductions of red deer to Ireland, and Stanton et al. 32 provided aDNA evidence that red deer populations on Orkney and the Outer Hebrides off Scotland, but not the Inner Hebrides, were probably established through long-distance translocation in the Neolithic. ...
... However, our study has also confirmed that there were two genetically distinct autochthonous populations of red deer in mainland Italy: one inhabiting the northern and the second occurring in the central and perhaps southern Italian Peninsula as it was hypothesized by the morphometric studies of red deer remains dated to the older Holocene 59 . Like previous studies 27,31,32 our analyses have shown that aDNA data derived from subfossil material can shed light on otherwise unanswerable questions pertaining to the historical biogeography of red deer (and other taxa), in particular when it comes to the origin of long-established island populations. ...
Article
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We present ancient mitochondrial DNA analyses of 31 complete cytochrome b gene sequences from subfossil red deer remains from the Tyrrhenian islands (Corsica and Sardinia) and mainland Italy in a European-wide phylogeographic framework. Tyrrhenian and North African red deer, both going back to human introductions, were previously the only red deer to harbour the mitochondrial B lineage whose origin, however, remained unknown. Our ancient Italian samples from the central part of the peninsula that were radiocarbon-dated to an age of ca. 6300 to 15 600 cal BP all showed B haplotypes, closely related or even identical to those found on Sardinia. Genetic diversity in the mainland population was considerably higher than on the islands. Together with palaeontological evidence our genetic results identify the Italian Peninsula as the ultimate origin of the B lineage and thus the Tyrrhenian and North African red deer. This is in line with previous biogeographic findings that uncovered distinct intraspecific phylogeographic lineages in Italian mammals, underlining Italy’s status as a hotspot of European mammalian diversity.
... A misidentified 'hyena' bone fragment from France was established as that of a herbivore using this technique (Stuart and Lister 2014). Irish errors in identification of bones based on morphology include a seal bone from Dalkey Island, off Co. Dublin, published as a bear [corrected in Woodman et al. (1997)], and a reindeer from Shandon cave published in error as a red deer [corrected in Carden et al. (2012)]. ...
... 4500 cal. BP) at which point Carden et al. (2012) have argued for its introduction by people (Table 10). (Woodman et al. 1997). ...
Chapter
The study of Irish Quaternary vertebrates has a long history, but with significant work in recent decades that allows for clear sequencing of faunas and notable advances in understanding for particular species. The primary resource for study is the museum collections of bones excavated from Irish caves in the late nineteenth and early twentieth centuries. Known faunas are much more limited in diversity and antiquity than those of Britain or continental Europe. The oldest vertebrate fossils were discovered in sediments from 109 to 74 ky BP in the form of woolly mammoth (Mammuthus primigenius) and musk ox (Ovibos moschatus) late in MIS 5, or early in MIS 4. The pre-Last Glacial Maximum fauna also included spotted hyena (Crocuta crocuta), the giant deer (Megaloceros giganteus), horse (Equus ferus), brown bear (Ursus arctos), reindeer (Rangifer tarandus), red deer (Cervus elaphus), Arctic fox (Alopex lagopus), stoat (Mustela erminea), collared lemming (Dicrostonyx torquatus), Norwegian lemming (Lemmus lemmus) and mountain hare (Lepus timidus). Faunas of the Woodgrange Interstadial include remains from open sites as well as caves and are dominated by discoveries of giant deer, but red deer, reindeer, brown bear, wolf, stoat and hare are also present. By the Holocene, Ireland was an island with its first human settlements, and faunas show their impacts through introductions and extirpations, but there are also indications of some species surviving through the Younger Dryas into the Holocene.
... Indeed, consideration of (any) megafaunal population dynamics and extirpations within Ireland must now consider potential anthropogenic related factors and impacts on such events that heretofore have not been considered. For example, giant deer, reindeer and red deer all disappeared from Ireland during or shortly following the Younger Dryas, presumably as the result of unfavourable climatic and subsequent ecosystem conditions (Woodman et al., 1997;Carden et al., 2012). ...
... Previous studies indicate a link between Irish biota and those in south-west Europe in terms of species assemblages (Corbet, 1961) and genetic affiliations (Grindon and Davison, 2013;Beatty and Provan, 2014). This was proposed to have been associated with Early Mesolithic human mobility, though the model was criticised as too simplistic and unrealistic (Carden et al., 2012). Certainly the new evidence for a human presence in Ireland around 12,800e12,600 cal BP will influence future interpretations of colonisation and megafaunal extirpation theories, or at the very least will stimulate further debates regarding human-mediated (deliberate or accidental) introductions of species to Ireland. ...
... The timing and mechanisms that have shaped the fauna of Ireland remain pervasive questions in studies of Quaternary north-western Europe (Woodman et al., 1997;Searle, 2008;Carden et al., 2012;Montgomery et al., 2014;Sleeman, 2014;Monaghan, 2017;Stimpson, 2024). While Ireland's fauna is unique and distinct, it continues to pose many questions for zoologists, archaeologists and biogeographers, not least due to a relatively limited taxonomic breadth and sparse chronological records (e.g. ...
Article
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The nature of the mechanisms that have shaped the animal communities of the island of Ireland remains a pervasive question in the study of the Quaternary of north-western Europe. Archived Quaternary faunal bone assemblages from antiquarian excavations of cave sites are a direct line of evidence with demonstrable potential to shed light on this issue, but are currently constrained by limited publication, understanding of early excavation protocols, and a lack of chronological reference. Alice and Gwendoline Cave in the west of Ireland was the subject of extensive excavations in 1902, which yielded a substantial faunal assemblage and the first evidence of an Upper Palaeolithic human presence on the island during the terminal Pleistocene. Here, we report further results from this important site. Archaeological excavations and a radiocarbon dating programme in 2019-2020 have shed light on the environmental context of the faunal assemblage, the cave taphonomy and site chronology. Nineteen radiocarbon dates are now available, including new direct dates for brown bear (Ursus arctos), giant deer (Megaloceros giganteus) and reindeer (Rangifer tarandus). The archaeological excavations and the analysis of sedimentary lipids – the first of its kind for an Irish cave site, both suggest that the sediments are primarily of exogenous origin, likely carried into the cave through fissures and openings by runoff or colluvial movements. Our analyses indicate that there were at least three main ‘pulses’ of bone-bearing sediment accumulation: at the beginning of the Younger Dryas (c. 12,700 cal. BP), in the aftermath of the Younger Dryas (11,300 cal. BP) and in the Early Holocene (c. 10,000 cal. BP), with a hiatus in sediment deposition after 9700 cal. BP.
... The Iberian Clade bears appear in the Holocene after 6,000 BP (Fig. 14), raising an intriguing possibility that the arrival of this population may have been due to human influence. Red Deer occur in Ireland for the first time as a human introduction at about 5,000 BP (Carden et al. 2012), showing a willingness of Neolithic settlers to bring large mammals with them, if they had a perceived value as a wild resource. The most recent bears in the Irish landscape are dated to 3,100 BP (calibrated to 1,271-1,048 BCE). ...
Article
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The Brown Bear (Ursus arctos) is known in the fossil state from over thirty localities in Ireland. All localities are summarised here, as is the current state of knowledge regarding this species on the island. All Irish fossil bears belong to this species, which inhabited the island at three discrete periods during the late Pleistocene and through the Holocene up to 3,000 years ago. The species Ursus planafrons Denny, 1864, based on Irish type material is here assigned to Ursus arctos Linnaeus, 1758.
... As neither an accepted domesticated nor a "pristine" wild species, both Persian and European fallow deer have been under-researched relative to other cervids such as reindeer (Rangifer tarandus) and red deer (Cervus elaphus) which have been the subject of numerous studies concerning their ancient range and management (19)(20)(21)(22)(23). By contrast, there is no consensus regarding the European fallow deer's glacial refugia or natural post-glacial distribution. ...
Article
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Over the last 10,000 y, humans have manipulated fallow deer populations with varying outcomes. Persian fallow deer ( Dama mesopotamica ) are now endangered. European fallow deer ( Dama dama ) are globally widespread and are simultaneously considered wild, domestic, endangered, invasive and are even the national animal of Barbuda and Antigua. Despite their close association with people, there is no consensus regarding their natural ranges or the timing and circumstances of their human-mediated translocations and extirpations. Our mitochondrial analyses of modern and archaeological specimens revealed two distinct clades of European fallow deer present in Anatolia and the Balkans. Zooarchaeological evidence suggests these regions were their sole glacial refugia. By combining biomolecular analyses with archaeological and textual evidence, we chart the declining distribution of Persian fallow deer and demonstrate that humans repeatedly translocated European fallow deer, sourced from the most geographically distant populations. Deer taken to Neolithic Chios and Rhodes derived not from nearby Anatolia, but from the Balkans. Though fallow deer were translocated throughout the Mediterranean as part of their association with the Greco-Roman goddesses Artemis and Diana, deer taken to Roman Mallorca were not locally available Dama dama , but Dama mesopotamica . Romans also initially introduced fallow deer to Northern Europe but the species became extinct and was reintroduced in the medieval period, this time from Anatolia. European colonial powers then transported deer populations across the globe. The biocultural histories of fallow deer challenge preconceptions about the divisions between wild and domestic species and provide information that should underpin modern management strategies.
... These connections, possibly composed of gravel or ice connecting Ireland to Scotland or Wales, would have allowed fauna recolonisation during the postglacial period (Yalden 1981). Even taking into account the challenges of recolonisation via shifting low-lying landbridges that might restrict some species more than others, this landbridge model fails to explain the unusual composition of fauna found in early Holocene Ireland (Carden et al. 2012;Woodman, McCarthy, and Monaghan 1997). ...
... They now comprise Ireland's largest extant terrestrial wildlife, yet there is no coordinated national management plan for deer in Ireland. Red deer are typically considered native to Ireland and stemming from their ancient Neolithic introduction is a single population of conservation importance, yet the total population has seen numerous modern introductions from continental European and British stock (Carden et al. 2012;McDevitt et al. 2009). Subsequently, fallow deer became established after their introduction during the post-medieval period (Beglane et al. 2018). ...
Article
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Systematic camera trap surveys are important for gathering information on terrestrial wildlife. Such surveys reveal distributions, abundances, and behaviours that can inform conservation and wildlife management by providing evidence of animal presence at known locations and times. However, in Ireland, international-standard camera trap surveys have not been undertaken to inform the management of large terrestrial wildlife. Through participation in a continent-wide initiative (Snapshot Europe) with a shared methodology, we undertook Ireland’s first systematic camera trap survey for large mammals over a two-month period in 2021 in a known deer “hotspot” - central Wicklow, Ireland. We captured eight wild mammal species from 19 forest camera trap sites. Sika (Cervus nippon) or sika-red deer hybrids (Cervus nippon × Cervuselaphus) were detected at all sites and comprised 92% of all independent camera trap events of wild mammals. Sika (and hybrid) events occurred at an average rate of 1.1 events per site per day. Females were more often detected than males (male:female = 1:1.42). We also noted that most females were not accompanied by juveniles (female:juvenile = 1:0.31). Overall, we demonstrate the efficacy of even this exploratory survey for revealing wildlife dynamics and generating data for science-informed management and conservation. We recommend that further surveys should be carried out across Ireland to achieve minimum international standards for monitoring terrestrial wildlife.
... They now comprise Ireland's largest extant terrestrial wildlife, yet there is no coordinated national management plan for deer in Ireland. Red deer are typically considered native to Ireland and stemming from their ancient Neolithic introduction is a single population of conservation importance, yet the total population has seen numerous modern introductions from continental European and British stock (Carden et al. 2012;McDevitt et al. 2009). Subsequently, fallow deer became established after their introduction during the post-medieval period (Beglane et al. 2018). ...
Article
Full-text available
Systematic camera trap surveys are important for gathering information on terrestrial wildlife. Such surveys reveal distributions, abundances, and behaviours that can inform conservation and wildlife management by providing evidence of animal presence at known locations and times. However, in Ireland, international-standard camera trap surveys have not been undertaken to inform the management of large terrestrial wildlife. Through participation in a continent-wide initiative (Snapshot Europe) with a shared methodology, we undertook Ireland’s first systematic camera trap survey for large mammals over a two-month period in 2021 in a known deer “hotspot” - central Wicklow, Ireland. We captured eight wild mammal species from 19 forest camera trap sites. Sika (Cervus nippon) or sika-red deer hybrids (Cervus nippon × Cervus elaphus) were detected at all sites and comprised 92% of all independent camera trap events of wild mammals. Sika (and hybrid) events occurred at an average rate of 1.1 events per site per day. Females were more often detected than males (male:female = 1:1.42). We also noted that most females were not accompanied by juveniles (female:juvenile = 1:0.31). Overall, we demonstrate the efficacy of even this exploratory survey for revealing wildlife dynamics and generating data for science-informed management and conservation. We recommend that further surveys should be carried out across Ireland to achieve minimum international standards for monitoring terrestrial wildlife.
... In the wake of their migrations, Neolithic farmers brought deer to new regions as a source of venison, skin, bones, and antlers. For example, red deer were translocated from peninsular Italy to Sardinia at least 4000-5000 years ago (Vigne 1992;Doan et al. 2017Doan et al. , 2022, from the Scottish mainland to the Hebrides and Orkney around 5000 ya (Stanton et al. 2016), and from Britain to Ireland in the same period (Carden et al. 2012). From the Middle Ages to the end of the eighteenth century, red deer were considered "high game" (Hochwild) in all of Europe, i.e., they were the exclusive quarry of kings, princes, high aristocracy, and high clergy. ...
... A current increase in the use of cave deposits was fuelled by the finding that cave sediments can provide data on past hominin presence (Slon et al. 2017, Zhang et al. 2020, Vernot et al. 2021. Cave sediments from different climatic regions have been targeted for non-hominin mammals, revealing, for example 25000-year-old DNA of wolves and bison from a cave in the Caucasus, Georgia (Gelabert et al. 2021), black bears Ursus americanus and giant short-faced bears Arctodus simus in northern Mexico (Pedersen et al. 2021), extirpated red deer Cervus elpahus from a cave in Ireland (Carden et al. 2012), and various mammals from a cave in Australia (Haouchar et al. 2014). Moreover, stalagmites in a cave in Georgia were shown to contain aDNA of various mammals such as bears Ursus sp., roe deer Capreolus sp., and horseshoe bats Rhinolophus sp. ...
Article
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1. Terrestrial mammals shape their ecosystems, and mammalian community assemblages can be important indicators of ecosystem functioning and ecosystem changes over time. Numerous taxa of terrestrial mammals are currently threatened by habitat loss and face displacement to new geographical areas or systems to which they are less suited and where they may affect the original communities. 2. Understanding past ecosystem changes is important for predicting future responses of species assemblages to changes in their environments. Thus, ecological and evolutionary history, as well as adaptive capacity, are important predictors of future population viability. Genomic and metagenomic approaches using environmental or ancient DNA offer a wealth of information regarding genome-wide variation of changing communities or of taxonomic groups over time, which may help explain past changes and predict future responses of communities to changes in their environment; however, to date, such studies are relatively scarce. 3. We review studies on environmental DNA and environmental genomics of terrestrial mammals to assess the potential of such approaches regarding past, contemporary, and future terrestrial ecosystems, identify inherent challenges, and discuss potential applications. We elaborate on lessons to be learned from mammal genomics of past ecosystems and compare metabarcoding with general metagenetic and metagenomic techniques. We provide a comprehensive overview of current applications, challenges, and future potential of environmental DNA with regards to terrestrial mammals. 4. As current major challenges regarding mammalian eDNA we identify its scarcity and patchy distribution, along with the persistent necessity of genomic reference data. While the latter are steadily increasing, the former can only be tackled by explicitly mapping the environment to gain understanding of spatial eDNA distribution. Such understanding may facilitate informed choices of sample sites and substrates and, together with new sequencing techniques, this can allow mammalian eDNA to be maximally exploited as a source of biodiversity data.
... In the wake of their migrations, Neolithic farmers brought deer to new regions as a source of venison, skin, bones, and antlers. For example, red deer were translocated from peninsular Italy to Sardinia at least 4000-5000 years ago (Vigne 1992;Doan et al. 2017Doan et al. , 2022, from the Scottish mainland to the Hebrides and Orkney around 5000 ya (Stanton et al. 2016), and from Britain to Ireland in the same period (Carden et al. 2012). From the Middle Ages to the end of the eighteenth century, red deer were considered "high game" (Hochwild) in all of Europe, i.e., they were the exclusive quarry of kings, princes, high aristocracy, and high clergy. ...
Chapter
An up-to-date synthesis of the biology, ecology, behaviour and conservation status of the red deer. After introducing the taxonomic status and the the systematic of the species, we provide an account of its current distribution. We then describe the main morphological, physiological and genetic features; the main life history traits (growth, survival and reproduction); the relationships with the environment (space use, diet) and how internal and external variables impact on population dynamics, including competition with other species; the social behaviour throughout the year and the mating system; the most relevant diseases and their demographic impacts; the issues surrounding management and conservation. This chapter will provide researchers and people interested in red deer with the opportunity to access the most relevant advances on the biology of this species.
... Currently, little is known about many aspects of wild-deer ecology in Ireland. We do know the natural history, origins of all three species along with previous distributions (Ní Lamhna 1979;McDevitt et al, 2009;Carden et al, 2011;Carden et al, 2012;Belgane et al, 2018) and the degree of hybridisation between red and sika deer and their distribution (McDevitt et al, 2009;Smith et al, 2014). However, detailed census data, spatial deer-habitat relationships and up-to-date deer distribution research, which has been lacking until recently, is forthcoming by members of the team in 2021, in conjunction with the Irish Deer commission (Carden et al, in prep.). ...
... Deer have cultural, ecological and an increasing economic importance. Being among the most important game animals for trophies, their populations have been managed, translocated and selectively hunted throughout their history and distribution area [26][27][28][29][30]. Recently a worldwide "deer industry" has been developed, whereby animals are farmed for venison and to some extent for antler products [31][32][33]. ...
Article
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Microsatellites are widely applied in population and forensic genetics, wildlife studies and parentage testing in animal breeding, among others, and recently, high-throughput sequencing technologies have greatly facilitated the identification of microsatellite markers. In this study the genomic data of Cervus elaphus (CerEla1.0) was exploited, in order to identify microsatellite loci along the red deer genome and for designing the cognate primers. The bioinformatics pipeline identified 982,433 microsatellite motifs genome-wide, assorted along the chromosomes, from which 45,711 loci mapped to the X- and 1096 to the Y-chromosome. Primers were successfully designed for 170,873 loci, and validated with an independently developed autosomal tetranucleotide STR set. Ten X- and five Y-chromosome-linked microsatellites were selected and tested by two multiplex PCR setups on genomic DNA samples of 123 red deer stags. The average number of alleles per locus was 3.3, and the average gene diversity value of the markers was 0.270. The overall observed and expected heterozygosities were 0.755 and 0.832, respectively. Polymorphic Information Content (PIC) ranged between 0.469 and 0.909 per locus with a mean value of 0.813. Using the X- and Y-chromosome linked markers 19 different Y-chromosome and 72 X-chromosome lines were identified. Both the X- and the Y-haplotypes split to two distinct clades each. The Y-chromosome clades correlated strongly with the geographic origin of the haplotypes of the samples. Segregation and admixture of subpopulations were demonstrated by the use of the combination of nine autosomal and 16 sex chromosomal STRs concerning southwestern and northeastern Hungary. In conclusion, the approach demonstrated here is a very efficient method for developing microsatellite markers for species with available genomic sequence data, as well as for their use in individual identifications and in population genetics studies.
... Wild animal (red deer and hare) and dog were recovered from a small number of EN II sites. Although red deer is considered a wild animal, it appears to have been introduced to Ireland during the Neolithic period (Carden et al. 2012). Birds were present at three EN II sites, but fish were not recorded at any sites. ...
Article
When compared with earlier periods, the Neolithic in Ireland (4000–2500 cal BC) witnessed enormous changes in the foods being produced, and the work involved in their production and processing. Several crops were introduced – archaeobotanical studies indicate that emmer wheat became the dominant crop, with evidence also for barley (hulled and naked) and flax. Gathered resources were not abandoned; on the contrary, there is substantial evidence for a variety of nuts, fruits and leafy greens. Zooarchaeological studies indicate that new animals also arrived, including domesticated cattle, pig and sheep. Recent studies have provided substantial information on the timing and nature of these new ways of farming and living, but the focus is often on ingredients rather than food products. There are many challenges in determining which foods were being made with these new crops and animals, and in assessing their dietary and social importance. While cereals have been found at many sites, for example, it is not clear if they are being ground, boiled or other techniques are used for their processing. In this paper we explore aspects of food production, processing and foodways in Neolithic Ireland, drawing upon evidence from archaeobotany, zooarchaeology, isotopes, human skeletal remains and artefacts.
... Another potentially significant discovery is the identification of possible red deer bone from Cairn K. If this can be further substantiated, it would support the presence of red deer in Neolithic Ireland, rather than the antlers being imported, thus contributing to a long-running debate regarding this question (Woodman & McCarthy 2003;Carden et al. 2012). ...
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The first detailed investigation of the human remains from the Carrowkeel passage tomb complex since their excavation in 1911 has revealed several new and important insights about life, death, and mortuary practice in Neolithic Ireland. Osteological analysis provides the first conclusive proof for the occurrence of dismemberment of the dead at Irish passage tombs, practised contemporarily with cremation as one of a suite of funerary treatments. The research also highlights changes in burial tradition at the complex over the course of the Neolithic. Providing a chronology for these changes allows them to be linked to wider trends in monument construction, which may relate to changes in both land use and climate during the period. Multi-isotope analysis hints at the presence of non-local individuals among the interred and the possible existence of different food sourcing areas at the onset of the later Neolithic period. Preliminary results from ancient DNA sequencing of six individuals from Carrowkeel provide evidence for the genetic ancestry of Irish Neolithic populations, demonstrating their Anatolian origins and links along the Atlantic façade.
... The arguments made here about numbers are, however, translatable to these other species, and since they are considerably smaller, only a single boatload of sheep or pigs need ever have set sail from Continental Europe to be the source of all British and Irish domesticates. It is of considerable interest, however, that there is now evidence for the movement of wild deer populations at the start of the Neolithic, as suggested for Ireland (Woodman and McCarthy 2003;Carden et al. 2012) but also apparently taking place in Scotland (Stanton, Mulville, and Bruford 2016). It seems that the importation of new domesticates inspired people to move other species around as well. ...
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Set against the new chronological framework for the introduction and spread of the Neolithic into Britain this paper considers the logistics of introducing domestic cattle from continental Europe. Cattle were the most extensively utilised domestic animal at the start of the Neolithic and understanding the processes behind their introduction is critical for exploring early Neolithic farming systems. We begin by exploring the realities of creating a viable new animal population and then moving that population to Britain across the water from Continental Europe. Using a series of simulated models we demonstrate that only a very small number of cattle need to have been introduced into Britain in order to enable the uptake and spread of domestic cattle across the whole island. This, in turn, may effect how we consider the mechanisms involved in the adoption of the Neolithic, and can better inform our understanding of the kinds of processes involved in the transition to the Neolithic.
... This is most often associated with the consumption of contaminated water plants. Given the lack of domestic livestock or wild ruminants in Mesolithic Irish archaeological sites (Carden et al., 2012;Woodman, 2015), the presence of Fasciola sp. is unlikely. Echinostoma sp., on the other hand, parasitizes a variety of animals, including humans (as definitive hosts). ...
... This was complemented by the use of domesticates-cattle, sheep/ goat and pig-whose remains dominate the limited available evidence for the Early Neolithic faunal record (F McCormick 2007;Schulting 2013). Alongside these introduced resources, which probably included red deer (Carden et al. 2012;Bergh & Hensey 2013), a range of other wild mammals and gathered plant foods, including hazelnuts and fruits, were also used (F McCormick 2007;McClatchie, Bogaard et al. 2014). Smyth and Evershed (2015a; have established, using organic residue analysis of Carinated Bowls, that the consumption of milk and dairying was practised, as was the use of pottery for the processing of meat products from the 38th century BC onwards. ...
... The reliance on domesticates precludes characterization of these Neolithic colonizers as hunter-gatherers, yet genetic information on red deer (Cervus elaphus) populations in the Orkneys (Stanton et al., 2016) and Ireland (Carden and Edwards, 2012) demonstrate that they were brought from the continental mainland by Neolithic colonizers. Without this genetic evidence, recognition of wild faunal materials could easily support conclusions of hunter-gatherer modes of subsistence in use in the region. ...
... We consider it likely that the late-surviving Hainan population may therefore have been the wild source population for the nineteenth-century Beijing herd, although we note that in the absence of genetic data from extinct mainland Chinese populations, we cannot rule out the possibility of generally low intraspecific variation across allopatric milu populations, so that the Hainan population might not be the only one that was phylogenetically close to the Beijing herd. There is evidence of ancient human transportation of other continental deer species to several island systems [42][43][44][45], raising the possibility that the Hainan milu population could itself have been originally introduced from mainland China. However, although this possibility cannot be discounted, ancient deer introductions were typically made to islands that lacked native large deer, and the local presence on Hainan of other surviving deer species that are considered native [17], together with the low cultural and economic importance of the island during most of China's history, makes it more likely that milu were native to Hainan. ...
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Determining the ‘dynamic biogeography’ of range collapse in threatened species is essential for effective conservation, but reconstruction of spatio-temporal patterns of population vulnerability and resilience can require use of non-standard ecological data such as historical archives. Père David’s deer or milu, one of the few living mammal species that has become extinct in the wild, is historically known only from a small captive herd of unknown provenance that survived until 1900 in the Imperial Hunting Park near Beijing, from which all living individuals are descended. Using ancient DNA analysis, we demonstrate that two fawns collected in 1868 from Hainan Island, off the southern Chinese mainland, represent the only known wild milu specimens and were sampled from probably the last wild population. The Hainan milu population shows extremely low genetic differentiation from descendants of the Beijing herd, suggesting that this now-extinct population may have been the source of the captive herd. This revised extinction model refutes the supposed long-term survival of a captive milu herd for centuries or millennia after final extinction of wild populations, highlighting the vulnerability of remnant mammal populations in the absence of proactive management and the importance of historical museum collections for providing unique new insights on evolution, biogeography and conservation. Milu experienced a pattern of final population persistence on an island at the periphery of their former range, consistent with the ‘range eclipse’ or ‘contagion’ model of range collapse, and matching the spatial extinction dynamics of other extinct mammals such as the thylacine and woolly mammoth.
... Anthropogenic activities have led to the introduction of new species to local ecosystems both intentionally and unintentionally. For example, Mus musculus domesticus was shown genetically to have been taken unintentionally with the Vikings from Norway to the British Isles [23], while Cervus elaphus was introduced intentionally to Ireland from Britain during the Irish Bronze Age [24]. Our analyses of A. agrarius suggest that their distribution also have been influenced by human unintentionally induced introductions. ...
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Species from the steppe region of Eastern Europe likely colonized northwestern Europe in connection with agriculture after 6500 BP. The striped field mouse (Apodemus agrarius Pallas, 1783), is a steppe-derived species often found in human crops. It is common on the southern Danish islands of Lolland and Falster, which have been isolated from mainland Europe since approximately 10 300-8000 BP. Thus, this species could have been brought in with humans in connection with agriculture, or it could be an earlier natural invader. We sequenced 86 full mitochondrial genomes from the northwestern range of the striped field mouse, analysed phylogenetic relationships and estimated divergence time. The results supported human-induced colonization of Denmark in the Subatlantic or Subboreal period. A newly discovered population from Central Jutland in Denmark diverged from Falster approximately 100-670 years ago, again favouring human introduction. One individual from Sweden turned out to be a recent introduction from Central Jutland.
... The Mesolithic red deer (Cervus elaphus) was found to be one of the most representative species on the southern slope of the Alps (Hohenstein et al. 2016). Other studies (Carden et al. 2012) indicate that the red deer was introduced into Ireland during the Neolithic period. On the territory of Lithuania, among terrestrial mammals the red deer has been known since the Late Neolithic (Stančikaitė et al. 2009). ...
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The aim of the study was to compare characteristics of longissimus dorsi (LD) and semimembranosus (SM) muscles from free-living and farmed red deer (Cervus elaphus) in Lithuania. No significant differences were found either between LD and SM muscles of free-living and farmed red deer or within these muscles of red deer from different environments regarding meat pH and colour parameters. The assessment of meat toughness employing the Warner-Bratzler (WB) method showed the longissimus muscle of free-living deer to be tenderer than the semimembranosus muscle. The TPA (texture profile analysis) method, in contrast to the WB test, showed higher tenderness of the semimembranosus muscle in farmed red deer. The present study revealed only small differences in properties of free-living and farmed red deer meat, i.e. intramuscular fat in the longissimus muscle of free-living red deer had a lower and a more favourable n-6/n-3 PUFA ratio and the semimembranosus muscle a lower content of cholesterol than the respective muscles of farmed deer. Therefore, the meat of free-living red deer could be considered to be more acceptable in terms of healthy nutrition.
... Deer are the largest terrestrial wild mammal and an important wildlife species on the island of Ireland, with species including fallow (Dama dama), sika (Cervus nippon) and red deer (Cervus elaphus) present. While the population of red deer in Killarney, Co. Kerry are descendants of a c. 5,000 year old early introduction to the island [1], other populations of red and sika deer date back for only approximately 150 years, whilst muntjac deer and roe deer are relatively recent introductions although the abundance and distribution range of these two species is relatively unknown [2]. The presence of muntjac deer (Muntiacus sp) was confirmed in Northern Ireland in 2009 and 2015. ...
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Background Deer are an important wildlife species in both the Republic of Ireland and Northern Ireland having colonised most regions across the island of Ireland. In comparison to cattle and sheep which represent the main farmed ruminant species on the island, there is a lack of data concerning their exposure, as measured by the presence of antibodies, to important viral pathogens of ruminants. A study was therefore undertaken to investigate the seroprevalence of wild deer to four viruses, namely bovine viral diarrhoea virus (BVDV), bovine herpesvirus-1 (BoHV-1), Schmallenberg virus (SBV) and bluetongue virus (BTV). Results Two panels of sera were assembled; Panel 1 comprised 259 samples (202 collected in the Republic of Ireland and 57 in Northern Ireland) between 2013 and 2015, while Panel 2 comprised 131 samples collected in the Republic of Ireland between 2014 and 2015. Overall sika deer (Cervus nippon) were sampled most commonly (54.8%), followed by fallow deer (Dama dama) (35.3%), with red deer (Cervus elaphus) (4.3%) and hybrid species (0.3%) sampled less frequently, with the species not being recorded for the remaining 5.3% of deer sampled. Age was not recorded for 96 of the 390 deer sampled. 196 of the remainder were adults, while 68 and 30 were yearlings and calves, respectively. Using commercially available enzyme-linked immunosorbent assays, true prevalence and 95% confidence intervals were calculated as 9.9%, (6.8-13.0% CI), SBV; 1.5% (0.1-3.0% CI), BoHV-1; 0.0%, 0-1.7% CI), BVDV; and 0.0%, (0.01-0.10% CI), BTV. Conclusions The results indicate a very low seroprevalence for both BVDV and BoHV-1 in the wild deer tested within the study and, are consistent with a very low prevalence in Ireland. While serological cross-reaction with cervid herpesviruses cannot be excluded, the results in both cases suggest that the presence of these viruses in deer is not a significant risk to their control and eradication from the cattle population. This is important given the ongoing programme to eradicate BVDV in Ireland and deliberations on a national eradication programme for BoHV-1. The SBV results show consistency with those reported from cattle and sheep on the island of Ireland, while the BTV results are consistent with this virus remaining exotic to Ireland. The results provide a baseline against which future surveys of either wild or farmed/captive deer populations can be compared.
... The same is true for many wild animal species: they are given short shrift in archaeological discussions of food even though in the past they made a significant contribution to human diet and were often translocated and managed alongside animals that became the true 'domesticates' (e.g. Vigne, Daujat and Monchot 2015;Carden et al. 2012;Carden 2012;Stanton, Mulville and Bruford 2016). ...
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This paper brings together zooarchaeological data from Neolithic to post-medieval sites in England to explore the plasticity of cultural attitudes to the consumption of wild animals. It shows how, through time, game has been considered variously as ‘tabooed’ and ‘edible’, each having implications for patterns of biodiversity and wildlife management. The essential points being made are that deeper-time studies can reveal how human perceptions of ‘surplus foods’ have the potential to both create and remedy problems of environmental sustainability and food security. Perhaps more significantly, this paper argues that understanding the bio-cultural past of edible wild animal species has the potential to transform human attitudes to game in the present. This is important at a time when food security and the production of surplus are pressing national and global concerns.
... The situation also differs in the apparent lack of two major wild-game species found on Mesolithic sites throughout most of Britain, red deer and aurochs, as well as the smaller roe deer (McCormick 1999(McCormick , 2007Woodman et al. 1997). Recent aDNA work has strongly supported the Neolithic re-introduction of red deer to Ireland (Carden et al. 2012). This leaves wild pig and bear as the only important terrestrial land mammals prior to the arrival of Neolithic domesticates. ...
... The population genetic structure of the red deer as an important game animal could have been blurred by numerous human-mediated introductions and translocations [20][21][22]. Thus, the biogeographic structure of this animal could have been affected, or alternatively, long-term processes (density blocking and/or selection) still play a major role. ...
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The trajectories of postglacial range expansions, the occurrence of lineage patches and the formation and maintenance of secondary contact between lineages may mostly reflect neutral demographic processes, including density blocking, that may leave long-lasting genetic signatures. However, a few studies have recently shown that climate may also play a role. We used red deer, a large, mobile herbivore that is assumed to be sensitive to climate change, to test hypotheses of possible selection on the mitochondrial DNA cytochrome b gene (mtDNA cytb) and competitive and/or density-blocking (using mtDNA control region). We searched for a possible link between the phylogeographic structure and abiotic climatic variables. Finally, we tested for isolation by distance and isolation by environment and assessed the impact of human-mediated translocations on the genetic structure of red deer. Our analysis of 30 red deer populations in Poland using the mtDNA control region (N = 357) and cytochrome b (N = 50) markers not only confirmed the presence of the Western and SouthEastern lineages of the species but also indicated the presence of a previously unnoticed, rare relic haplotype that grouped together C. e. italicus from Italy (the Mesola deer). No significant signs of positive selection were detected for the mtDNA cytb gene in the studied red deer. However, a significant signal for purifying selection was found in our study that may explain the narrowness of the contact zone because gene flow between the Western and SouthEastern lineages should drive relatively strong mito-nuclear incompatibilities. MtDNA control region differentiation among red deer populations in Poland correlated with different abiotic climatic variables. Strikingly, the southernmost ice sheet limits during the Elsterian was the most important factor, and it explained the largest amount of variation. However, neither isolation by distance (IBD) nor isolation by environment (IBE) were recorded, and a very limited impact of human translocations was evident. The above-mentioned results suggest that in contemporary red deer populations in Poland, the phylo-geographic pattern is well preserved, and long-term processes (density and/or competitive blocking) still play a major role.
Article
Climate change is allowing fire to expand into previously unburnt ecosystems and regions. While management policies such as fire suppression have significantly altered their frequency and intensity. To prevent future biodiversity/ecosystem services loss, and the large financial burden of wildfires, management plans will be required to adapt to future climate and land use changes. Long-term ecological data offer a unique perspective to assess fire variability under different climate and land-use conditions. In this study, we focus on Killarney National Park, Ireland. An area which today is under threat from an increase in fire activity. Comparing palaeoecological and archaeological records, we reconstruct the past fire dynamic and its impact on the landscape, and evaluate the role of climate vs humans in influencing the natural fire regime over the millennial time-scale. Our results indicate that fire has been present in the landscape since the beginning of the Holocene, with fire in the early Holocene being largely controlled by climate and microsite conditions, and in the late Holocene being increasingly influenced by human activity. The knowledge of past fire regimes can help inform future management in order to protect the semi-natural native woodland. The park's present landscape mosaic, could be preserved by limiting forest encroachment through moderate grazing and burning, while also protecting any fragmented forest from excessive grazing and large/intense fires, via traditional fire management strategies such as fuel load management. However, a fire management strategy should only be implemented following careful consideration of all ecosystem factors and controls.
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Many species of carnivorans exhibit a relationship between skeletal phenotype and habitat and/or climate as a result of behavioural or phenotypic adaptations to their environment. Mustelidae contains several species with shortened limbs and elongated bodies compared to other carnivorans. As a result of this unique bauplan, these species may not respond to changes in habitat/climate in similar ways to other carnivorans. I examined how limb skeletal morphology relates to biome and climate in North American pine martens, Martes americana and Martes caurina, which span coniferous, mixed broadleaf and boreal forest biomes. I found Martes limb proportions correlate with temperature and non-snow precipitation. These species do not follow Allen's Rule and instead have proportionally longer limbs in colder regions. There is a significant difference in morphology between specimens from the three biomes. Differences were found in the robusticity of the proximal ends of each bone, with specimens from coniferous forests being the most robust. Previous studies have shown that differences in limb proportions and robusticity correlate with vegetation density and degrees of arboreality vs. fossoriality. This suggests that specimens from coniferous forests may be less arboreal and occupy less dense forests than those from broadleaf/boreal biomes.
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Research into deer population dynamics has mostly focused on community or landscape scales in the Republic of Ireland (ROI). However, in order to propose national deer management strategies to confront the increasing deer populations in recent years, a larger scale analysis of deer populations is necessary. In this inter-county, national study, generalised linear mixed models (GLMM) were utilized to examine the relationship between the harvest rate (defined as the numbers harvested per 100 ha in this study) data and factors identified in past research as contributing to deer population dynamics on a county basis, where annual county-level deer harvest records were used to reflect deer populations by species. The procedures were repeated for each species and significant factors and their impacts were identified for each county. The results suggested that multiple factors affect deer population dynamics; for instance, increases in forest cover tended to result in increases in deer populations of all species, and the presence of red and sika deer populations in a county had a positive influence on hybrid deer populations. National migration maps for each deer species were produced, as well as predictions of future population trends. The outcome of this study can be used to guide the development of a national deer management strategy and the prioritisation, on a regional basis, of deer control measures.
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Aim We documented how the similarity of mammal assemblages on continental and oceanic islands has changed since initial human colonization, since European arrival and overall. We investigated how levels of similarity might change in the future. Location Continental and oceanic islands worldwide. Time period Human settlement of islands to the present, as well as projections for the future. Major taxa studied Mammals. Methods We used mammal occurrence data on islands to calculate the change in similarity using a pairwise approach based on Jaccard's index and a multisite approach based on Jaccard's and Sørensen's measures. We divided the mammal assemblages into two time periods, before and after island colonization or trade began with Europeans. We unpacked the mechanisms driving changes in similarity, exploring how initial similarity interacts with seven types of species turnover events to determine overall change. Finally, we calculated how future similarity levels will change if past trends in introductions and extinctions continue. Results Mammals, on both continental and oceanic islands, show one of the most pronounced cases of homogenization ever observed, and on oceanic islands mammals show the largest increase in homogenization ever observed for a terrestrial group. Most of the homogenization observed to date has been driven by recent historical changes, not by changes that occurred before European arrival. If current patterns of species introductions and extinctions continue, then oceanic islands will experience little additional homogenization, whereas continental islands will homogenize greatly beyond current levels. Main conclusions Mammal assemblages on oceanic islands show nearly an order of magnitude greater change in similarity than plant and bird assemblages. Projections of future similarity indicate that continental and oceanic islands are on different trajectories of change. These trajectories could be altered by management actions, but in some cases those actions that would be impactful run counter to current conservation norms.
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Caves have been an important source of vertebrate fossils for much of Southeast Asia, particularly for the Quaternary. Despite this importance, the mechanisms by which vertebrate remains accumulate and preserve in Southeast Asian caves has never been systematically reviewed or examined. Here, we present the results of three years of cave surveys in Indonesia and Timor-Leste, describing cave systems and their attendant vertebrate accumulations in diverse geological, biogeographical, and environmental settings. While each cave system is unique, we find that the accumulation and preservation of vertebrate remains are highly dependent on local geology and environment. These factors notwithstanding, we find the dominant factor responsible for faunal deposition is the presence or absence of biological accumulating agents, a factor directly dictated by biogeographical history. In small, isolated, volcanic islands, the only significant accumulation occurs in archaeological settings, thereby limiting our understanding of the palaeontology of those islands prior to human arrival. In karstic landscapes on both oceanic and continental islands, our understanding of the long-term preservation of vertebrates is still in its infancy. The formation processes of vertebrate-bearing breccias, their taphonomic histories, and the criteria used to determine whether these represent syngenetic or multiple deposits remain critically understudied. The latter in particular has important implications for arguments on how breccia deposits from the region should be analysed and interpreted when reconstructing palaeoenvironments.
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Biological invasions are one of the great threats to Earth’s ecosystems and biodiversity in the Anthropocene. However, species introductions and invasions extend deep into the human past, with the translocation of both wild and domestic species around the world. Here, we review the human translocation of wild plants and animals to the world’s islands. We focus on establishing criteria used to differentiate natural from human-assisted dispersals and the differences between non-native and invasive species. Our study demonstrates that, along with a suite of domesticates, ancient people transported numerous wild plants and animals to islands and helped shape ecosystems in ways that have important ramifications for modern conservation, restoration, and management.
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This paper reviews the human colonisation of Ireland. Archaeological evidence suggests humans have been discontinuously present in Ireland from c. 12.8 to 12.5 ka cal BP. This date seems anomalously recent in comparison to other areas of northwest Europe, including Britain, where hominins were present from c. 0.78 to 0.99 mya. Explanations for this apparently delayed colonisation include taphonomic factors and research biases as well as human activity in the past. These factors are outlined in a comparative context. The paper examines pre-Late Glacial Maximum evidence and the (re) colonisation of the British-Irish Isles following the retreat of the Ice, synthesising archaeological and genetic data as appropriate. An important emphasis is placed on the need to consider the process of colonisation, how hunter-gatherers encounter new worlds and to examine why things happen at the times that they do.
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Founder analysis is a method for analysis of nonrecombining DNA sequence data, with the aim of identification and dating of migrations into new territory. The method picks out founder sequence types in potential source populations and dates lineage clusters deriving from them in the settlement zone of interest. Here, using mtDNA, we apply the approach to the colonization of Europe, to estimate the proportion of modern lineages whose ancestors arrived during each major phase of settlement. To estimate the Palaeolithic and Neolithic contributions to European mtDNA diversity more accurately than was previously achievable, we have now extended the Near Eastern, European, and northern-Caucasus databases to 1,234, 2, 804, and 208 samples, respectively. Both back-migration into the source population and recurrent mutation in the source and derived populations represent major obstacles to this approach. We have developed phylogenetic criteria to take account of both these factors, and we suggest a way to account for multiple dispersals of common sequence types. We conclude that (i) there has been substantial back-migration into the Near East, (ii) the majority of extant mtDNA lineages entered Europe in several waves during the Upper Palaeolithic, (iii) there was a founder effect or bottleneck associated with the Last Glacial Maximum, 20,000 years ago, from which derives the largest fraction of surviving lineages, and (iv) the immigrant Neolithic component is likely to comprise less than one-quarter of the mtDNA pool of modern Europeans.
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Mount Sandel has long been an iconic site for the Irish Mesolithic, having produced evidence for a sequence of occupation huts and pits and the earliest radiocarbon dates for the Mesolithic on the island. This paper presents details of a recent programme of redating whereby the application of Bayesian modelling has confirmed the early date for the site but also helped to refine its internal chronology. The major phase of hut building at Mount Sandel took place within a much shorter period of time than had previously been thought, perhaps only a generation or two. The dating of pits of differing sizes suggests that many of them were created during other slightly later visits to the area. The implications of the dating programme for the place of Mount Sandel in the Irish Mesolithic, and for the chronology of the period and its relations with that on the British mainland, are discussed.
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1. Throughout Europe, the range of many deer species is expanding. We provide current distribution maps for red deer Cervus elaphus, sika Cervus nippon, fallow deer Dama dama and muntjac deer Muntiacus sp. in Ireland, and estimates of range expansion rates for red deer, sika and fallow deer. 2. There was a considerable expansion in the ranges of red deer, sika and fallow deer between 1978 and 2008. The compound annual rate of expansion was 7% for red deer, 5% for sika and 3% for fallow deer. The total range increase was 565% for red deer, 353% for sika and 174% for fallow deer. The potential implications of these expansions are discussed. 3. There are unknown numbers of red-sika hybrid deer in some parts of Ireland. Range expansion is likely to lead to further hybridizations with implications for the genetic integrity of deer stocks. 4. Sightings of free-roaming muntjac deer were first recorded in 2007. The distribu-tion of confirmed sightings of single and multiple animals in the eastern region of Ireland suggests multiple releases. 5. Deer are already impacting on both the economic and biodiversity values of habitats in Ireland, where, at present, no sustainable deer management policy exists.
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Red deer (n=149) from eight geographical locations, including the endangered endemic populations from the Tyrrhenian islands (Sardinia and Corsica), were analysed at eight polymorphic microsatellite loci. Two questions were addressed: (1) Is there a founder effect in the Corsican population, which was reintroduced to the island using Sardinian deer after the species’ extinction on Corsica? (2) What is the origin of the Tyrrhenian or Corsican red deer (Cervus elaphus corsicanus)? Our results showed signs of a founder effect for the red deer on Corsica in that these deer showed differentiation from the Sardinian population as measured by FST values, assignment tests (with and without a priori definition of populations) and individual-based dendrograms. Genetic variability, however, did not differ significantly between the two populations. With respect to the phylogeography of C. e. corsicanus we found that both deer from North-Africa and Mesola on the Italian mainland were genetically close to the Corsican red deer, but phylogenetic trees based on genetic distances were only poorly supported statistically. Among all populations studied the Mesola red deer showed the lowest distance values from Corsican red deer and yielded allele frequencies that were more similar to those of C. e. corsicanus than were those of North-African red deer. These results are in line with recent palaeontological and archaeozoological findings which suggest that the Corsican red deer is derived from small Italian red deer introduced from the mainland to Sardinia and Corsica during the Late Neolithic and just before the beginning of Classical Antiquity, respectively. They also suggest a possible recent introduction of Tyrrhenian red deer to North-Africa (rather than the other way around), thus accounting for the close genetic relationship (especially based on mitochondrial DNA) that has repeatedly been found between C. e. corsicanus and C. e. barbarus.
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The North Atlantic INTIMATE group of the INQUA Palaeoclimate Commission has previously recommended an Event Stratigraphy approach for the synchronisation of records of the Last Termination using the Greenland GRIP isotopic record as the regional stratotype and INTCAL98 for the calibration of radiocarbon dates [Lowe, J.J., Hoek, W., INTIMATE Group, 2001. Inter-regional correlation of palaeoclimatic records for the Last Glacial-Interglacial Transition: a protocol for improved precision recommended by the INTIMATE project group. Quaternary Science Reviews 20, 1175–1187]. Here, we present a revised protocol for time-stratigraphic correlation in the North Atlantic region over a more extended time period (30–8 ka). This employs the new NGRIP isotopic record and associated Greenland Ice Core Chronology 2005 (GICC05) as the regional stratotype, INTCAL04 for the calibration of radiocarbon dates, Bayesian-based statistical procedures for the construction of age models, and tephrochronology to validate correlations between regional site records.
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Much of Ireland's Pleistocene and Early Holocene mammalian faunas are derived from a series of late 19th/early 20th century cave excavations. In many instances it would appear that the deposits containing these faunal remains were disturbed. This project assessed the chronological range of the mammalian species present in the caves using 14C dating, in particular accelerator mass spectrometry (AMS). The research has shown that (1) a wide range of mammals colonised Ireland in the period between at least 45 ka and 20 ka, with some elements surviving until close to the Last Glacial Maximum; (2) a more restricted range of species re-colonised Ireland during the Lateglacial period, with evidence for a slightly more temperature fauna being replaced by an Arctic fauna at about 11 ka; (3) certain elements of Ireland's Holocene fauna may have survived through from the Lateglacial into the Holocene; (4) there is a lack of evidence for red deer, Cervus elaphus, being present in the Early Holocene in Ireland; and (5) horse is only documented in the Irish Holocene from 4 ka. The paper also discusses the implications of the Quaternary Fauna Project for the Late Pleistocene of Ireland, the mechanism and period of colonisation of Ireland as well as the introduction of domesticates in the Mid Holocene.
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