Article

Phylogeography, population genetics and conservation of the European red deer Cervus elaphus

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  • Natural History Museum Vienna, Austria
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Abstract

ABSTRACT • During the Last Glacial Maximum, European red deer Cervus elaphus occurred in refugia in Iberia/southern France, Italy, the Balkans and the Carpathians. Most of Europe, including large parts of the east and north-east, is now inhabited by red deer from the western lineage. The eastern lineage is largely confined to south-eastern Europe; a third lineage comprises Sardo-Corsican and Barbary red deer. • Sardo-Corsican, Barbary and Mesola red deer are genetically unique units. They exhibit low levels of genetic diversity and deserve particular protection, since conservation strategies should target genetic information. • Hybridization between sika Cervus nippon and red deer occurs rarely, but may lead to extensive introgression, particularly in parts of the British Isles. Further expansion of both species may lead to increased hybridization in continental Europe. • Although hunting has an impact on red deer gene pools, the main threat today is habitat fragmentation in human-dominated landscapes. The resulting increase in genetic drift and inbreeding reduces variability in isolated populations and may lead to inbreeding depression. To support vital meta-populations, migration corridors should be established.

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... The red deer (Cervus elaphus) is a good model species to address the hypothesis of cryptic refugia and its influence on the European postglacial colonization history, owing to its current and past widespread natural distribution across Europe [15,16], and known phylogeographical structure [17]. The most comprehensive study on a large biogeographical scale, using mitochondrial (mtDNA) cytochrome b data, has revealed that red deer originated in Asia and evolved into two distinct groups: a Western group with four lineages (Western Europe, Balkans, Africa and Middle East); and an Eastern group consisting of three lineages (North Asia/ America, South-Asia and East-Asia) [18]. ...
... 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. ...
... Populations from south-western of France, southern Ireland or a region between them could rapidly expand to central and northern Europe because there was a connection between the British Isles and mainland Europe [126] (Fig 10). The small haplotype diversity observed in the central and northern European countries, excluding the British Isles, is consistent with this rapid expansion after the LGM [17,19], while the high mtDNA diversity currently found in the British Isles could be a result of the sampling bias/reintroductions mentioned above or a local refugium (in or surrounding) of this species during the LGM [26]. Sampling of ancient red deer specimens in the continental shelf and southern France would help to elucidate the precise location of cryptic refugia, since current populations in France mostly derive from restocking events after World War II, as a consequence of almost complete extirpation of populations over the middle of the nineteenth century [127]. ...
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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.
... Међутим, према неким ауторима (Whitehead, 1972) Наведене врсте и подврсте у оквиру породице јелена су описане на основу морфолошких, социјалних и генетских одлика, али и до дан-данас мишљења научника нису јединствена у погледу укупног броја врста и подврста (нпр. Данилкин, 1999, Zachos, Hartl, 2011, Lovari et al., 2016. ...
... Када је реч о обичном јелену, неки аутори (Zachos, Hartl, 2011) наводе четири главна питања у вези његовог очувања, и то: 1) угрожене генетске линије и замагљивање природног генетичког структурирања кроз транслокације и реинтродукције, 2) локална хибридизација са сика јеленом Cervus nippon, 3) селективни лов, и 4) смањену ефективну величину популације услед фрагментације станишта. ...
... Дакле, ови резултати иду у прилог констатацији да је фрагментација шума једна од најважнијих претњи за конзервацију обичног јелена (нпр. Zachos, Hartl, 2011). Насупрот очекиваном, популације у ограђеним просторима нису представљале ни вишу диференцијацију нити нижу генетску разноврсност од оних у слободној природи (тзв. ...
... The red deer, Cervus elaphus, is one of the most widespread wild ungulates across Europe and a species with a great economic, social, and ecosystem value (Milner et al., 2006;Apollonio et al., 2010). Several subspecies have been recognized using phenotypic or biogeographical traits, though there is still an ongoing debate about this subspecific taxonomy since it is not congruent with the evolutionary units that have been described by genetic studies (Zachos and Hartl, 2011;Meiri et al., 2018). Due to its importance and wide distribution, red deer have been largely studied over their geographical range, with numerous studies focused on understanding their evolutionary history (Ludt et al., 2004;Skog et al., 2009;Meiri et al., 2013;Stanton et al., 2016;Queirós et al., 2019). ...
... Due to its importance and wide distribution, red deer have been largely studied over their geographical range, with numerous studies focused on understanding their evolutionary history (Ludt et al., 2004;Skog et al., 2009;Meiri et al., 2013;Stanton et al., 2016;Queirós et al., 2019). Despite being intensively exploited as a natural resource since the Pleistocene (Sommer et al., 2008), studies based on mitochondrial DNA have shown a natural wide-scale phylogeographic pattern of red deer across Europe (Niedziałkowska et al., 2011;Zachos and Hartl, 2011;Meiri et al., 2013;Queirós et al., 2019). Three main mitochondrial D-loop lineages have been described, with a clear spatial pattern: A Western European lineage distributed in Western Europe; an Eastern European lineage distributed in the Balkans (Eastern Europe); and a Mediterranean lineage distributed in Africa and the islands of Sardinia and Corsica. ...
... Anthropogenic hybridization in the red deer between autochthonous European populations and introduced sika deer (Cervus nippon) or wapiti deer (Cervus canadensis) has been often documented (McDevitt et al., 2009;Senn and Pemberton, 2009;Biedrzycka et al., 2012;Smith et al., 2014Smith et al., , 2018. However, anthropogenic admixture between red deer is more challenging to address because of the inconsistent support between subspecies defined using morphological characteristics or biogeography and the genetic evolutionary units (Zachos and Hartl, 2011;Meiri et al., 2018). In addition, such analysis requires large genetic databases to accurately infer non-native or hybrid individuals across Europe. ...
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Anthropogenic hybridization is one of the greatest threats to global biodiversity. It incites human-mediated gene flow between non-native/exotic and native taxa, which can have irreversible effects on native species or locally adapted populations, eventually leading to extinction. The red deer, Cervus elaphus, is a game species that, due to its extraordinary economic value, has been introduced in several regions throughout Europe. However, the consequences of those introductions on native populations, namely on their genetic background, have been poorly addressed. This study is focused on the Iberian Peninsula and aims to: (i) assess the extent of anthropogenic hybridization/introgression of introduced red deer into the native Iberian populations; (ii) evaluate the impact of red deer management regimes on the observed hybridization/introgression patterns; and (iii) assess how hybridization/introgression influence the current genetic diversity of native Iberian populations. A set of 11 microsatellites and a 329 bases pair fragment of the mitochondrial D-loop gene were used to estimate nuclear admixture and mitochondrial introgression in 1,132 individuals sampled across 46 red deer populations throughout Iberia. A Bayesian approach implemented in the STRUCTURE program was employed to investigate the proportion of admixture between native populations and non-native red deer. Results showed that 17% of individuals presented signs of non-native recent ancestors and 10.1% had non-native mitochondrial haplotypes, reaching an overall hybridization/introgression rate of 23%. Non-native or hybrid individuals were found throughout 40 Iberian red deer populations, and the percentages per population varied between 3.3 and 75.0%, independently of the management regime. Mitochondrial introgression was observed across 15 Iberian red deer populations, being more frequent in free-ranging individuals (16.2%) than in fenced populations (9.2%) but was completely absent from public-owned populations. Nuclear genetic diversity correlated positively with the proportion of hybrid individuals in public-owned populations. The genetic footprint of historical and current human-mediated translocations of non-native red deer into the Iberian Peninsula is evidenced in this study, highlighting the need to implement effective measures to avoid such practices both in Portugal and Spain, in order to preserve the endogenous genetic patrimony of the Iberian red deer populations.
... Red deer (Cervus elaphus L.) is currently one of the most widespread European ungulate species and its evolutionary history is relatively well established [1,2,3,4,5]. Molecular studies revealed the existence of at least three distinct mitochondrial DNA (mtDNA) lineages (so called matrilines) in Europe reflecting the main refugia during the Last Glacial Maximum (LGM) and the early Late glacial (25,000-14,700 years ago) in Iberia (lineage A), the Balkan region (lineage C) and the Mediterranean (lineage B) [3,5,6]. ...
... Red deer (Cervus elaphus L.) is currently one of the most widespread European ungulate species and its evolutionary history is relatively well established [1,2,3,4,5]. Molecular studies revealed the existence of at least three distinct mitochondrial DNA (mtDNA) lineages (so called matrilines) in Europe reflecting the main refugia during the Last Glacial Maximum (LGM) and the early Late glacial (25,000-14,700 years ago) in Iberia (lineage A), the Balkan region (lineage C) and the Mediterranean (lineage B) [3,5,6]. Lineage B has originally been attributed to red deer from Sardinia/N-Africa, but has recently been associated, based on ancient DNA material from mainland Italy, to red deer from the Italian refugial lineage [7]. ...
... In addition, northern areas may have acted as cryptic refugia in North-West Europe through the LGM [9].The Bavarian-Bohemian region was identified as part of a suture zone between western and eastern European red deer matrilines [10]. However, a detailed sampling for molecular analyses is missing from other regions of Central Europe, including the region from the North of the Swiss Alps to the East of France [3,4,5,9]. ...
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In north-eastern France, red deer (Cervus elaphus L.) populations were rebuilt from a few hundred individuals, which have subsisted in remote valleys of the Vosges mountains, and to a lesser extent from individuals escaped from private enclosures; at present times, this species occupies large areas, mainly in the Vosges Mountains. In this study, we examined the population dynamics of red deer in the Vosges Mountains using ancient and contemporary mitochondrial DNA (mtDNA) from 140 samples (23 ancient + 117 modern) spanning the last 7’000 years. In addition, we reconstructed the feeding habits and the habitat of red deer since the beginning of agriculture applying isotopic analyses in order to establish a basis for current environmental management strategies. We show that past and present red deer in the Vosges Mountains belong to mtDNA haplogroup A, suggesting that they originated from the Iberian refugium after the last glacial maximum (LGM). Palaeogenetic analysis of ancient bone material revealed the presence of two distinct haplotypes with different temporal distributions. Individuals belonging to the two haplotype groups apparently occupied two different habitats over at least 7’000 years. AM6 correlates with an ecological type that feeds in densely forested mountain landscapes, while AM235 correlates with feeding in lowland landscapes, composed of a mixture of meadows and riverine, herb-rich woodlands. Our results suggest that red deer of north-eastern France was able to adapt, over the long term, to these different habitat types, possibly due to efficient ethological barriers. Modern haplotype patterns support the historical record that red deer has been exposed to strong anthropogenic influences as a major game species.
... The initial transmission of microsatellites from various ungulates to red deer (Kühn et al. 1996;Slate et al. 1998;Roed and Midthjell 1998;Poetsch et al. 2001) had a huge impact on the efficiency of population genetic studies in this species and led to numerous studies on a wide variety of issues. In the foreground were questions about the degrees of relationship and assignments of subspecies and subpopulations (Kuehn 2004;Hajji et al. 2008), the impact of management (Hmwe et al. 2006a;Höglund et al. 2013;Galarza et al. 2015Galarza et al. , 2017, the effect of landscape fragmentation and isolation of populations (Perez-Espona et al. 2008;Dellicour et al. 2011;Frantz et al. 2012), and ways for the conservation of this species (Zachos and Hartl 2011;Zachos et al. 2016). The major advantage of microsatellites is their high variability compared to SNPs and mitochondrial markers, which make them particularly suitable for the investigation of spatially and temporally more closely related populations (Zachos et al. 2016). ...
... Microsatellite studies were done to compare populations of specific areas within countries (Zachos et al. 2007;Niedzialkowska et al. 2012;Willems et al. 2016) or between countries (Fickel et al. 2012;Höglund et al. 2013;Krojerova-Prokesova et al. 2015). Other studies compared populations of entire regions (Zachos and Hartl 2011;Zachos et al. 2016). However, sample numbers, gender relations, microsatellite loci, and microsatellite loci numbers vary considerably between studies. ...
... Twenty-four references were extracted from literature in order to determine the sample size of the populations examined and the microsatellite markers used (Kuehn et al. 2003;Zachos et al. 2003;Kuehn 2004;HMWE et al. 2006aHMWE et al. , 2006bFrantz et al. 2008;Hajji et al. 2008;Kinser and Herzog 2008;Nielsen et al. 2008;Perez-Espona et al. 2008;Dellicour et al. 2011;Zachos and Hartl 2011;Fickel et al. 2012;Frantz et al. 2012;Perez-Gonzales et al. 2012;Niedzialkowska et al. 2012;Höglund et al. 2013;Sprem et al. 2013;Queiros et al. 2014;Galarza et al. 2015;Krojerova et al. 2015;Hoffmann et al. 2016;Willems et al. 2016;Zachos et al. 2016). Only literature making statements on European red deer populations, using microsatellites and being available in the accessible databases (e.g., Medline), was considered. ...
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Population genetic parameters from different studies might be significantly influenced by differences in sample size, fraction of males and females, marker number, and sets of markers used, reducing the comparability between studies. This hypothesis was tested on a red deer population of 205 individuals with an estimated size of 1000 animals. Four tests were performed: (1) the population was subdivided into 10 populations each with 10 to 150 individuals and genotyped with 16 markers, (2) the total population was genotyped 10 times with different panels of microsatellite loci containing 2 to 14 markers, (3) a subset of 8 microsatellite loci was used to genotype the total population; markers of this subset were replaced one by one with a different marker set and genotyping results were compared to the results of the original subset and (4) the effect of sex was estimated. Additionally, 24 references from literature, including 256 European red deer populations, were analyzed. A median of 25 individuals per population was investigated in published studies using 11 microsatellite markers (5 to 22). Sixty-eight percent of possible study comparisons matched with less than 10% of microsatellite loci. Our results show that the factors investigated, except for the factor gender, lead to significant deviations in the population genetic results, especially with sample sizes below 30, with less than 6 to 8 microsatellite markers and with the use of different panels of microsatellite loci. This is also true with respect to population genetic structure and the use of Bayesian methods. Therefore, populations from different studies should be compared with each other with caution.
... ). In the past 10 years, several studies have dealt with the phylogeography of red deer in Europe (Ludt, Schroeder, Rottmann, & Kuehn, 2004;Meiri et al., 2013;Skog et al., 2009;Sommer et al., 2008;Zachos & Hartl, 2011). Based on mitochondrial DNA (mtDNA) from extant specimens, three main lineages can be defined: a western (lineage A), an eastern (lineage C), and a North African and Sardinian lineage (lineage B) (Skog et al., 2009;Zachos & Hartl, 2011). ...
... In the past 10 years, several studies have dealt with the phylogeography of red deer in Europe (Ludt, Schroeder, Rottmann, & Kuehn, 2004;Meiri et al., 2013;Skog et al., 2009;Sommer et al., 2008;Zachos & Hartl, 2011). Based on mitochondrial DNA (mtDNA) from extant specimens, three main lineages can be defined: a western (lineage A), an eastern (lineage C), and a North African and Sardinian lineage (lineage B) (Skog et al., 2009;Zachos & Hartl, 2011). Western and eastern lineages show continuity at least until the Late Pleistocene (Meiri et al., 2013). ...
... The specimens from the cave present a rather diverse haplotype configuration. Half of the haplotypes from Liñares fit in the Western European clade, which is in line with previous red deer studies (Ludt et al., 2004;Meiri et al., 2013;Skog et al., 2009;Sommer et al., 2008;Zachos & Hartl, 2011) and also with data from other species (Hofreiter et al., 2004;Fortes et al., 2016;Niedziałkowska, 2017). For instance, studies on cave bear suggest some gene flow between the Iberian Peninsula, France, and Central Europe during marine isotope stage 3 (MIS 3, 59,000-24,000 BP) (Fortes et al., 2016). ...
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The major climatic oscillations that characterized the Quaternary had a great influence on the evolution and distribution of several species. During cold periods, the distribution of temperate-adapted species became fragmented with many surviving in southern refugia (Iberian, Italian, and Balkan Peninsulas). Red deer was one of the species that contracted its original range to southern refugia. Currently, two main lineages have been described for the species: western and eastern. We have analyzed fossils pre-dating the last glacial maximum (LGM) from Liñares cave (NW Spain) that belongs to the peripheral range of the western clade, and fossils from the Danish Holocene belonging to the central part of the same clade. Phylogenetic analyses place our samples in the western clade. However, some specimens from Liñares represent an early split in the tree along with other pre-LGM western samples from previous studies. Despite low bootstrap values in the Bayesian phylogenies, haplotype networks connect these foreign haplotypes to the eastern clade. We suggest a mixed phylogeographical model to explain this pattern with range expansions from the east during the expansion phase after the cold periods in marine isotope stage 3. We find slight isolation by distance in post-LGM populations that could be a consequence of the recolonization from southern refugia after the LGM.
... If particular areas have been recolonized from a number of different refugia, the different genetic lineages met and could form suture zones of different widths (Sommer and Zachos, 2009;Fickel et al., 2012;Meiri et al., 2013;Krojerová-Prokešová et al., 2015). In effect, the patterns of genetic discontinuities would be visible across the zone, as well documented for species like the European hedgehog, brown bear and roe deer (e.g., Hewitt, 2004;Sommer and Zachos, 2009 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). ...
... In effect, the patterns of genetic discontinuities would be visible across the zone, as well documented for species like the European hedgehog, brown bear and roe deer (e.g., Hewitt, 2004;Sommer and Zachos, 2009 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 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). ...
... In effect, the patterns of genetic discontinuities would be visible across the zone, as well documented for species like the European hedgehog, brown bear and roe deer (e.g., Hewitt, 2004;Sommer and Zachos, 2009 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 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). ...
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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.
... Red deer Cervus elaphus stags produce their rutting calls for attracting potential mates and deterring competitive males (Clutton-Brock and Albon 1979). Studies of acoustic variation of stag rutting calls (Frey et al. 2012;Passilongo et al. 2013;Della Libera et al. 2015;Volodin et al. 2015aVolodin et al. ,b, 2019Golosova et al. 2017) are in agreement with the subdivision of red deer to phylogenetic lineages (Mahmut et al. 2002;Ludt et al. 2004;Skog et al. 2009;Zachos and Hartl 2011;Zachos et al. 2016). The acoustics of stag rutting calls proved to be helpful population markers in red deer (Frey et al. 2012;Passilongo et al. 2013;Volodin et al. 2019) in addition to the genetic markers, such as mtDNA and microsatellites (Feulner et al. 2004;Niedziałkowska et al. 2012;Krojerova-Prokešova et al. 2015;Carranza et al. 2016;Zachos et al. 2016). ...
... After the Late Pleistocene glacial maximum occurred 25-12 thousand years ago (Clark et al. 2009), the species Cervus elaphus recolonized Europe (Ludt et al. 2004;Skog et al. 2009;Zachos and Hartl 2011;Niedziałkowska et al. 2021). Recolonization started from the three main refugia, corresponding to red deer mitochondrial DNA (mtDNA) lineages A, B, and C (Ludt et al. 2004;Skog et al. 2009;Niedziałkowska et al. 2011Niedziałkowska et al. , 2021. ...
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This study investigates a population of red deer Cervus elaphus, founded by 10 individuals introduced in the nineteenth century from Germany to the Voronezh region of the European part of Southern Russia and then developed without further introductions. We characterize for the first time the vocal phenotype of the Voronezh red deer male rutting calls in comparison with similar data on the Pannonian (native Central European) and Iberian (native West European) red deer obtained by the authors during preceding studies. In addition, we provide for the first time the genetic data on Pannonian red deer. In Voronezh stags, the number of roars per bout (2.85 ± 1.79) was lower than in Pannonian (3.18 ± 2.17) but higher than in Iberian (2.11 ± 1.71) stags. In Voronezh stags, the duration of main (the longest within bouts) roars was longer (2.46 ± 1.14 s) than in Pannonian (1.13 ± 0.50 s) or Iberian (1.90 ± 0.50 s) stags. The maximum fundamental frequency of main roars was similar between Voronezh (175 ± 60 Hz) and Pannonian (168 ± 61 Hz) but higher in Iberian stags (223 ± 35 Hz). Mitochondrial cytochrome b gene analysis of red deer from the three study populations partially supports the bioacoustical data, of closer similarity between Voronezh and Pannonian populations. In contrast, microsatellite DNA analysis delineates Voronezh red deer from either Pannonian or Iberian red deer. We discuss that population bottlenecking might affect the acoustics of the rutting roars, in addition to genotype.
... This mechanism results in low f0 values, ranging of 40-380 Hz in male and female red deer (see discussions in Volodin et al., 2015;Golosova et al., 2017). This is in spite of their origin from the three different European isolates of the last Pleistocene glacial maximum: Iberian Peninsula/Southern France, North Italy and the Balkans (Zachos & Hartl, 2011) and their respectively different Cytochrome b haplogroups (Skog et al., 2009;Zachos & Hartl, 2011). ...
... This mechanism results in low f0 values, ranging of 40-380 Hz in male and female red deer (see discussions in Volodin et al., 2015;Golosova et al., 2017). This is in spite of their origin from the three different European isolates of the last Pleistocene glacial maximum: Iberian Peninsula/Southern France, North Italy and the Balkans (Zachos & Hartl, 2011) and their respectively different Cytochrome b haplogroups (Skog et al., 2009;Zachos & Hartl, 2011). ...
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In female terrestrial mammals, vocal aging has only been studied in humans and pandas. In cervids displaying convergent sex dimorphism of vocal apparatus with humans, vocal aging is only investigated in males. This cross-sectional study examined acoustic variables of nasal (closed-mouth) and oral (open-mouth) contact calls of 32 farmed Iberian red deer hinds (Cervus elaphus hispanicus) aged of 4-18 years and their relationships with caller´s age, weight, social discomfort score (bites of other hinds on hind pelt) and body condition score (fat reserves). Decrease of fundamental frequency was associated with age in both oral and nasal calls, but more prominently in the nasal calls. An increase in call duration, peak frequency and power quartiles was associated with a higher degree of bites due to social aggression. Weight and body condition weakly influenced acoustic traits. We discuss that vocal aging of hinds parallels that of vocal aging in human females.
... Allendorf et al. 2001, Wolf et al. 2001, it may be of key importance for the survival of some taxa under rapidly changing environmental conditions. However, in the current management paradigm that aims to minimise interventions to populations, humanmediated hybridisation is often seen as 'genetic pollution' (Zachos & Hartl 2011) and hybrid populations are not given equal protection in many countries, compared to 'pure' populations (Allendorf et al. 2004). Thus, genetic monitoring and the dissemination of relevant empirical data are expected to support conservation actions, including the implementation of officially accepted molecular protocols aimed at identifying the genetic status of populations, and to estimate contemporary vs. historical patterns of hybridisation. ...
... Originally from Japan, the sika deer is strongly genetically differentiated from the red deer; they nevertheless hybridise, especially in some parts of the British Isles (McDevitt et al. 2009, Senn & Pemberton 2009, Senn et al. 2010, Smith et al. 2014, and extensive introgression has occurred, both in enclosures (Bartoš 2009) and in the wild (Senn & Pemberton 2009). Moreover, hybridisation between native red deer hinds and introduced sika deer stags in continental Europe was reported, using both mtDNA and microsatellite, in five regions of Poland, in the Kaliningrad District (Russia), in Lithuania (Biedrzycka et al. 2012), in the Czech Republic (Bartoš 2009), and in Austria (reviewed in Zachos & Hartl 2011). Hybridisation increases the body mass of both sika-like males and females, and increases incisor arcade breadth and jaw length of sika-like females (Senn et al. 2010). ...
Article
Hybridisation and gene introgression are important sources of diversification, the relevance of which in the evolutionary processes is well recognised. Their fitness consequences in animal populations, however, are not sufficiently well understood, despite hybridisation rates becoming increasingly important worldwide following human‐related activities such as domestication, game management and habitat alteration. In Europe, the density and distribution of native ungulates have largely been influenced by humans since pre‐historic times. This, alongside the introduction of non‐native and domesticated species, may bear major consequences at the genetic and population levels. We provide an updated overview of recent hybridisation events in wild European ungulates; we describe their ecological drivers, extent, current distribution, potential consequences and proposed management strategies. We reviewed the scientific literature published between 2000 and 2018 and found that confirmed hybridisation was described in 75 of the 89 references we included, involving nearly all the species that we investigated. Most researchers relied on genetic information for hybrid identification, which often involved a domestic counterpart. However, introductions and translocations also led to crossbreeding between wild ungulate (sub)species. Only 43 papers provided management recommendations, mostly focused on preventing hybridisation and removing hybrids. Hybridisation proved to be relatively common in several ungulate taxa in Europe. Despite reported changes in phenotype and fitness‐related traits in some species, the consequences of hybridisation for adaptation, life history, and evolutionary potential remain largely unknown. The current conservation paradigm aims to prevent the spread of domestic or non‐native genes in native populations; accordingly, conservation plans should: 1) determine the genetic origin of possible source populations; 2) protect native populations from the risk of crossbreeding with non‐native ones, and 3) establish permanent monitoring.
... The vocal features of a certain subspecies are constant across different populations, e.g. in Iberian red deer C.e. hispanicus (Frey et al., 2012;Passilongo et al., 2013;Volodin et al., 2015a) and Siberian wapiti C.e. sibiricus (Volodin et al., 2013b;Golosova et al., 2017). Therefore, bioacoustical analysis represents a powerful tool for defining and validating subspecies of red deer, provided they were not subjected to the introgression of genes from other subspecies as a result of multiple anthropogenic translocations of red deer over historical times (Zachos and Hartl, 2011;Zachos et al., 2016;Frantz et al., 2017). Although subspecies-specific rutting vocalizations do not provide reliable reproductive barriers against hybridization (Long et al., 1998;Nussey et al., 2006;Wyman et al., 2016), they may serve for the affiliation of native populations of Cervus elaphus with a particular subspecies (Frey et al., 2012;Volodin et al., 2015a, b;Golosova et al., 2017). ...
... We also compared stag rutting roars of the Pannonian red deer with stag roars of other European subspecies, regarding fundamental frequency and duration (Fig. 5, Table A1 in Appendix A). Overall, the acoustics of stag rutting roars in the European subspecies of red deer fit well to the respective three haplogroups of mitochondrial DNA (A, B, and C), which, in their order, fit to the Please cite this article in press as: Volodin (Skog et al., 2009;Zachos and Hartl, 2011). The roars of male Corsican red deer C.e. corsicanus (haplogroup B, Skog et al., 2009;Doan et al., 2017) are the lowest in maximum fundamental frequency and are well distinguishable from the roars of both the Mesola red deer stags C.e. italicus (lineage intermediate between A and C haplogroups, Zachos et al., 2014) and from the roars of the Scottish and the Iberian stags (haplogroup A, Skog et al., 2009) (Fig. 5). ...
Article
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The acoustics of male rutting roars, aside from genetic markers, are useful tools for characterization of populations and subspecies of red deer Cervus elaphus. This study of rutting mature male Pannonian red deer from Southern Hungary presents a description of the calling posture, a graphical reconstruction of the oral vocal tract length during rutting roar production and a spectrographic analyses of 1740 bouts containing a total of 5535 rutting roars. In addition, this study provides the first direct comparison of the bouts and main (=longest) rutting roars between Pannonian and Iberian red deer stags, representative of the Western and Eastern lineages of European red deer. The bouts of the Pannonian stags comprised 1-15 roars per bout; 24.37% were single-roar bouts and 23.68% were two-roar bouts. The duration of the main roars within bouts ranged from 0.52 s to 4.60 s, 1.13±0.50 s on average. Main common roars (66.3% of the 1740 main roars), were longer than harsh roars (1.27±0.55 s vs 0.87±0.25 s) and higher in maximum fundamental frequency (179±61 Hz vs 147±54 Hz). In multi-roar bouts, main harsh roars were first roars in 47.4%, intermediate roars in 19.2% and last roars in 18.8% of the bouts. Bout structure and the acoustics of main roars in the Pannonian stags differed from those in the Iberian stags and in stags from other populations of Cervus elaphus. These results support a power of rutting vocalizations for consideration as an additional tool for discriminating populations and subspecies within Cervus elaphus.
... Mitochondrial markers (for a review see Zachos and Hartl 2011) identified in European red deer modern populations three major haplogroups including the several subspecies described for this taxon (Skog et al. 2009 in Italy. C. e. italicus (formerly in haplogroup C) (Ludt et al. 2004;Skog et al. 2009) and haplogroup E occur in Armenia, Crimea, and Southern Russia (Doan et al. 2022). ...
... From Iberia, lineage A colonized the British Isles and parts of Central Europe; in the Balkan region, lineage C migrated to South-East Central Europe and the Carpathians. The Mediterranean (lineage B) could be originated from the Italian refugial lineage, and lineage D has been proposed for the Middle East and South-Eastern Europe (Zachos and Hartl 2011;Meiri et al. 2013;Doan et al. 2017;Schnitzler et al. 2018;Queiros et al. 2019). These hypotheses have been recently reconsidered and new data suggested that the species survived the LGM not only in the well-known southern European refugia, but also in more northern areas of Western and Eastern Europe and in the Urals (Niedziałkowska et al. 2021). ...
Article
The red deer Cervus elephus has been a common species in Italy until the Middle Ages and the Renaissance, when its distribution range started to considerably decrease, due to gradual deforestation and hunting pressure. Afterwards, the red deer has been reintroduced to many regions of the world, including Italy. In the Italian Apennines, the Acquerino-Cantagallo Natural Reserve (ACQUERINO) hosts one of the largest peninsular red deer populations, originated from a series of successful reintroductions. In this study, we meant to detect the level of genetic variability of Acquerino-Cantagallo Natural Reserve deer population and to investigate the genetic relationships with the other Italian and European populations. We identified five mitochondrial DNA control region (D-loop) haplotypes, four falling in lineage A and one falling in lineage C, derived from at least two maternal lineages, confirming that ACQUERINO population should be the result of multiple reintroductions. Haplotype diversity (H = 0.50) and nucleotide (π = 0.004) diversity were low, but included into the deer range values. ACQUERINO population showed low levels of genetic diversity when compared to other European and Mediterranean populations, confirming that this expanding population may have been generated from a low number of founders.
... Jele? szlachetny (Cervus elaphus) jest jednym z najbardziej rozpowszechnionych du?ych ssak?w w Europie wyst?puj?cym, poza p??nocn? Skandynawi? i Irlandi?, praktycznie na ca?ym kontynencie (Zachos i Hartl, 2011). W roku 2010 liczebno?? jeleni w Polsce wynosi?a 22 577 sztuk, przy zag?szczeniu 27,59 sztuk na 1000 ha lasu oraz strukturze p?ci 1:1,67 byk?w do ?a? (Jaworski, 2011). ...
Article
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The aim of the study was to analyze the rates of reproduction and changes in body weight of deer (Cervus elaphus) during the rut. The study was conducted in 2010 on the Bomafar farm, in Dąbrówka, in Lubuskie province. The study included 6 herds with a total of 250 deer hinds and 195 calves. Harem mating system was used. In one of the analyzed herds two bulls were used for mating. Data on age and body weight of females in the 16-26.08.2010 and 17.11-1.12.2010 periods of time were collected. In the same periods of time counting and weighing of calves was carried out. The greatest weight of females was found in a herd in which there were the oldest hinds (p≤0.05). Relative gains reached the highest values (p≤0.01) in herd of two-year old hinds, due to an incomplete somatic growth. The highest fawning rates characterized hinds in 6th, 11th and 12th years, respectively. It was found that male calves achieve higher body weight in mid-August and before weaning, compared with female calves. Sex ratio of emerging deer was close to 1: 1.
... In response to ever-increasing anthropogenic changes to natural ecosystems, genetic monitoring through the usage of different molecular markers is the best estimator of natural populations' sustainability, since genetic variability underpins populations' long-term potential for survival and adaptation (Palsbøll et al., 2007;Schwartz et al., 2007). Mitochondrial DNA is one of the most extensively used molecular markers in determining molecular diversity and phylogeography of many species (e.g., Castor fiber (Durka et al., 2005), Cervus elaphus (Zachos and Hartl, 2011), Sus scrofa (Alexandri et al., 2012;Veličković et al., 2015), Ursus actros (Hirata et al., 2013)), given the high evolutionary rate and lack of recombination (Avise, 2004). Even though analyses based solely on mtDNA have their own limitations due to mtDNA being a single locus marker with an effective population size of one-fourth of nuclear autosomal sequences, it is still a choice in preliminary analyses of genetic variability of wild populations. ...
Article
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Even though the phylogeography of hedgehogs has been well studied, information on the genetic variability of the northern white-breasted hedgehog Erinaceus roumanicus from the Balkans is lacking, since the previous studies were based on very limited sampling across the Balkans. The aim of this study is to estimate the genetic diversity and population structuring of E. roumanicus from the Central Balkans and to complement an already proposed phylogeographic scenario of this species. Tissue samples of 108 road-killed northern white-breasted hedgehogs were collected across the Central Balkan countries of Serbia, Montenegro, Bosnia and Herzegovina, and Macedonia. A partial fragment of the mtDNA control region (CR) was amplified and sequenced. Nine of 13 haplotypes detected in this study have not been previously published. The results indicate a moderate level of haplotype diversity of E. roumanicus from the Central Balkans and differentiation into four spatial groups, which are named after the approximate sampling localities as northwesterncentral, northeastern, southwestern, and southeastern groups. The observed population structure in the Central Balkans remains less pronounced in further phylogenetic and phylogeographic analyses of the dataset comprising E. roumanicus and E. concolor mtDNA CR sequences. The central position of Balkan haplotypes in a median joining network indicated its role as a primary source population for postglacial northward expansion.
... Thus, climate as a single factor driving extensive changes in its population may be less likely than for small mammals. Correspondingly, red deer as an important game species has been hunted, translocated and introduced since ancient times (Hartl, Zachos & Nadlinger, 2003;Zachos & Hartl, 2011). ...
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.
... Various management practices (selective harvesting, translocation, captive-breeding, fencing, introduction of non-native species) lead to changes in the gene pool of ungulates within populations (Randi 2005;Linnell and Zachos 2011;Niedziałkowska et al. 2012;Fernández-García et al. 2014). Consequences of such changes may include a decline in genetic variability, an increase in inbreeding, a decrease in population viability and the loss of local adaptations (Zachos et al. 2007;Zachos and Hartl 2011;Galarza et al. 2015). There is an urgent need for research to quantify these risks and identify solutions, as well as to develop policies and regulations to manage these emerging threats. ...
Article
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Wildlife management systems face growing challenges to cope with increasingly complex interactions between wildlife populations, the environment and human activities. In this position statement, we address the most important issues characterising current ungulate conservation and management in Europe. We present some key points arising from ecological research that may be critical for a reassessment of ungulate management in the future. Wildlife management systems face growing challenges to cope with the increasingly multifaceted interactions between wildlife populations, the environment and human activities. Dense ungulate populations, such as those occurring in many parts of Europe, have impacts on agriculture and forestry production. In addition, landscape fragmentation, due to the development of infrastructure, is leading to an increase in ungulate-traffic collisions and the synurbization of many species, while a few species and subspecies are still endangered (Apollonio et al. 2010). Simultaneously, populations of large carnivores—the ungulates’ natural limiting factor—are returning and stabilising in some areas of Europe (Chapron et al. 2014). In such a context, ungulates should not be managed in isolation, but as important “ecosystem engineers” (Smit and Putman 2011) and to do so means that they need to be considered as an integral part of both ecosystems and ecosystem management. It is thus necessary to develop a holistic approach that enables the widespread presence of ungulates to be considered as an opportunity and a renewable resource rather than a nuisance, and to manage ungulate species and populations in the face of a changing environment and human society. In this position statement, we address some of the most important issues characterising current ungulate management and conservation in Europe. We present some key points arising from ecological research that may be critical for a reassessment of ungulate management in the future and highlight some of the knowledge gaps. The goal of this statement is to provide wildlife managers and policymakers with a modern perspective of the position of ungulates in the ecosystem. We believe it should be used to support the redirection of current management plans and goals to ensure the viability and endurance of sustainable ungulate populations. We propose ten management and conservation measures to help achieve this.
... The Sika deer have been introduced to many countries and the hybridisation with native Red deer population has been demonstrated in British Isles (Malcolm, 2015), Czech Republic (Bartoš et al., 1982;Macháček et al., 2014), Poland (Biedrzycka et al., 2012) or Germany (Herzog et al., 2016). However, in addition to local hybridization with sika three major conservation issues exist: threatened genetic lineages, and blurring of natural genetic structuring through translocations and reintroductions; selective hunting; and reduced effective population sizes due to habitat fragmentation (Zachos and Hartl, 2011). For long-term conservation and development purposes, it therefore appears compulsory to manage Red deer wildlife to maintain both species survival and within species genetic diversity. ...
... Sequence variation of the CR-1 mtDNA is commonly used to develop phylogeographic models in mammals (e.g. Durka et al., 2005 for Castor fiber, Zachos and Hartl, 2011 for Cervus elaphus, Alexandri Table S1. Veličković et al., 2015 for Sus scrofa, Hirata et al., 2013 for Ursus actros; see also Avise, 2004 for general aspects). ...
Article
The contemporary geographical distribution and genetic structure of temperate species have been strongly influenced by the climatic oscillations during the Late Quaternary. As spatial genetic reconstructions are often markedly affected by geographically meaningful sample distributions, we focused in our study on the analyses of mtDNA control region sequences of brown hares from different regions in northern, central and south-central Balkans that have so far not been covered, with the aim to delineate the most likely glacial refugia wherefrom the postglacial northward expansion into central Europe has originated. Three major haplogroups (“Anatolia/Middle East”, “the Balkans”, and “central Europe”) were revealed with apparent south-north gradual decrease in molecular diversity indices. Moreover, phylogenetic and demographic history analyses identified the southeastern central Balkans as the putative origin for most populations from the southern and northern Balkans, while populations from central and northwestern Europe have originated from the northern Balkans.
... It is a problem in Europe in terms of the areas, in which free-living red deer cross-breed with the sika deer. Therefore, this is a debatable issue, which attracts a lot of attention from scientists (Goodman et al. 1999, Diaz et al. 2006, Bartoð 2009, McDevitt et al. 2009, Senn and Pemberton 2009, Senn et al. 2010, Zachos and Hartl 2011. ...
Article
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Genetic diversity of the red deer Cervus elaphus (6 individuals from enclosures and 33 individuals living in the wild) and the sika deer Cervus nippon (30 individuals from enclosures) species was studied by using tissue samples from the legally hunted animals in Lithuania. Genotyping was based on seven microsatellites loci (STR) of nuclear DNA. The STR loci were variable, with 1 -17 alleles and higher than intermediate values of heterozygosity (sika deer: Ho=0.695, He=0.694; wild red deer: Ho=0.626, He=0.624; red deer from enclosures: Ho=0.639, He=0.667). Among 69 individuals phenotypically designated as red deer and sika deer by hunters, based on Bayesian method 31 individuals had Q-values of 0.95–1.0 (“pure red deer”), 3 individuals had Q-values of 0.75–0.95 (hybrid of the “red type”), 2 individuals had values of 0.25 ˂ Q ≤ 0.75 (intermediate hybrid), 6 individuals had Q-values of 0.05–0.25 (hybrid of the “sika type”), and 27 individuals had Q values of 0–0.05 (“pure sika”).
... Thus, climate as a single factor driving extensive changes in its population may be less likely than for small mammals. Correspondingly, red deer as an important game species has been hunted, translocated and introduced since ancient times (Hartl, Zachos & Nadlinger, 2003;Zachos & Hartl, 2011). ...
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.
... Red deer are larger than sika, typically around 30 cm taller at the shoulders and whilst the red stags can grow antlers of 12 points or more, sika antlers rarely exceed eight points ( Whitehead 1964, Harrington, 1973). Hybridisation is a problem in Europe in terms of the areas in which free-living red deer cross-breed with the sika deer ( Goodman et al., 1999;Diaz et al., 2006;Bartoš, 2009;McDevitt et al., 2009;Senn & Pemberton, 2009;Senn et al., 2010;Zachos & Hartl, 2011). Our study also suggests the possibility of hybridization of the two species in the wild, as there were wild living specimens, which are attributed to red deer according to their phenotype, but are closer to sika deer according to their genotype. ...
Article
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The sika deer ( Cervus nippon ) are considered to be an invasive species in Europe. The main problem that the European free-living sika deer pose is damage they do to forests as well as hybridisation with the local red deer. The aim of this study was to analyse the genetic diversity of the sika deer and the red deer in Lithuania, and to determine the hybridization, which might be present in red deer population from the past release of sika deer into the nature. Tissue samples were collected from 30 sika deer individuals from enclosures, and 33 wild-living red deer. Samples were genotyped using seven microsatellite (STR) loci; genetic diversity indices were calculated and individuals were classified using Principal Coordinate Analysis (PCoA); the genetic structure of sika deer and red deer was investigated according to Bayesian clustering method using STRUCTURE software. The STR loci were highly polymorphic with up to 17 alleles per locus, and with an average heterozygosity Ho=0.695 and Ho=0.626 for sika deer and red deer respectively. Overall inbreeding coefficient (F IS ) values are 0.004 and 0.127 in sika and red deer respectively. According to the PCoA sika deer samples differ from those of red deer; however, few red deer individuals mix with sika deer. These animals were attributed to red deer according to their phenotype; however according to their genotype they are closer to sika deer. Thus, it can be concluded that wild red deer in Lithuania may hybridise with sika deer.
... Red deer Cervus elaphus are the most abundant ungulates in central European forests (Zachos and Hartl 2011). In many forest ecosystems, red deer cause severe damage to young trees through browsing and antler rubbing (trophic and nontrophic interactions respectively; Gill 1992). ...
Article
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Trophic plant–animal interactions (e.g. browsing by ungulates, insect attack) are an important and well‐studied source of mortality in many tree populations. Non‐trophic tree–animal interactions (e.g. deer antler rubbing) also frequently lead to tree death, and thus have significant effects on forest ecosystem functioning, but they are much less well studied than trophic interactions are. As deer populations have increased in recent decades in the Northern Hemisphere, their impact on tree populations via browsing and antler rubbing will increase. The aim of the study was to illustrate the potential ability of non‐trophic plant–animal interactions to regulate the dynamics of a natural forest. Specifically, we wanted to determine whether and how density and distance‐dependent processes affect sapling mortality caused by an antler rubbing by red deer Cervus elaphus. We used a spatially explicit approach to examine density and distance‐dependent mortality effects in almost two thousand Picea abies saplings over 20 years, based on a fully mapped permanent 14.4 ha plot in a natural subalpine old‐growth spruce forest. Antler rubbing by deer was the main identified cause of sapling mortality, and it showed a strong spatial pattern: positive density dependence of survival among spruce saplings. Deer selectively killed spruce saplings that were isolated from conspecifics. In consequence, non‐trophic plant–deer interactions were a major driver of the spatial pattern of P. abies sapling survival. The other mortality causes (e.g. breaking, overturning) did not show density‐dependent patterns or their effects were much weaker. In the medium and long term, the density‐dependent pattern of sapling mortality due to antler rubbing can alter the tree stand structure. Our results highlight the ecological relevance of non‐trophic plant–animal interactions for forest ecosystem functioning. This article is protected by copyright. All rights reserved.
... Diversity values are therefore usually not directly comparable, although the relative diversity when comparing different populations often holds across different marker systems. Zachos and Hartl (2011) give an overview of studies (including diversity values) until 2010. The only continent-wide population genetic study yielding directly comparable diversity values (including estimates of effective population sizes N e ) based on highly variable microsatellite data is the one by Zachos et al. (2016). ...
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.
... Die älteste Hirschart Europas ist allerdings nicht der Rothirsch, sondern das Reh. Die gegenwärtige genetische Situation des Menschen) fassten Hartl et al. (2003) und Zachos und Hartl (2011) zusammen. ...
Conference Paper
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Red deer (Cervus elaphus) in Austria. The relationship between man and red deer was and is controversial. Different interests collide in the cultural landscape. Here is an attempt to give an overview of the historical development of red deer and the human-deer interrelation in Austria. From this, guidelines for future deer management are compiled. Dealing with red deer in our cultural landscape is one of the major challenges in wildlife management, as is the handling of wild boar and wolf. Politics and society are also required to support reality-based, sensible solutions instead of merely making ideologically justified demands. An integrated management approach well adapted to the respective regional situation (starting situation, objectives, environment factors) is required, which includes landowners, hunters, foresters, torrent and avalanche control, leisure activities, nature conservation, settlement development, transport infrastructure and land-use planning. In order to integrate red deer in parts of the cultural landscape sustainably and as harmless as possible, the habitat design (esp. rest areas for wildlife, and forests with low susceptibility to damage by ungulates) as well as the regulation of red-deer populations and their distribution on a sufficiently large area must be well coordinated, both spatially, temporally and concerning the concrete measures. Accompanying this, it requires a regular, objective monitoring of success as the basis for adaptive management, and a flexible wildlife-ecological spatial planning with a clear distinction between different land-use priorities. Wildlife-ecological connections are complex. But the problems with red deer are still easily solvable, which can be shown by some positive examples that have emerged with regionally-adapted management cooperation. So we can be optimistic. However, there is the danger of simplifying complex problems too much or even presenting them in a monocausal way. This leads to increased conflicts between the interest groups and prevents sustainable solutions of problems. Main points for the management and hunting of red deer: (1) Making habitat suitable for red deer; (2) critically scrutinizing needs of supplementary feeding, duration of shooting times and methods of hunting, and optimizing them according to location; (3) minimizing game damage by taking appropriate measures, esp. regulation of number and distribution of stocks, agricultural and forestry prevention measures, resting areas for game; (4) (temporary) disturbance-free areas where leisure activities etc. are omitted for efficient game stock regulation; (5) Prevention of diseases and epidemics, and (6) Seeing the “big picture", i.e. wildlife-ecological overview for a sustainable red deer management.
... 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.
... Biological differences between subspecies are based on the genetic differentiation of C. e. hispanicus due to the isolation of the territory by the Pyrenees (Garde et al. 2010). Genetic studies (Skog et al. 2009;Zachos & Hartl, 2011) support the existence of different lineages (Iberian for C. e. hispanicus and Balkan/Carpathian for C. e. hippelaphus) belonging to different glacial refuges. ...
... In absolute terms however, the data of the Hessian population show rather favourable heterozygosity and lower Fis values in comparison with other national (Poetsch et al. 2001;Kuehn et al. 2003;Zachos et al. 2007;Edelhoff et al. 2020) and international studies (Hmwe et al. 2006a, b;Nussey et al. 2007;Nielsen et al. 2008;Sanchez-Fernandez et al. 2008;Zsolnai et al. 2009). The lowest heterozygosity and the highest F values are typically found in small islet populations and populations with longer history of isolation and low population sizes (Hmwe et al. 2006a, b;Hajji et al. 2008;Zachos and Hartl 2011;Zachos et al. 2016;Edelhoff et al. 2020). However, it must be taken into account that both measures are decisively influenced by the markers used, their number, and the sample size (Reiner et al. 2019). ...
Article
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Nineteen red deer areas in a densely populated region with a huge network of fenced motorways and the division into administrative management units (AMUs) with restricted ecological connectivity were investigated. In the season 2018/2019, a total of 1291 red deer samples (on average 68 per area) were collected and genotyped using 16 microsatellite markers. The results show a clear genetic differentiation between most of the AMUs. Fourteen AMUs may be combined into four regions with a considerable internal genetic exchange. Five areas were largely isolated or showed only a limited gene flow with neighbouring areas. Ten of the 19 AMUs had an effective population size below 100. Effective population sizes greater than 500–1000, required to maintain the evolutionary potential and a long-term adaptation potential, were not achieved by any of the studied AMUs, even when AMUs with an appreciable genetic exchange were aggregated. Substantial genetic differentiation between areas can be associated with the presence of landscape barriers hindering gene flow, but also with the maintenance of ‘red deer–free’ areas. Efforts to sustainably preserve the genetic diversity of the entire region should therefore focus on measures ensuring genetic connectivity. Opportunities for this goal arise from the establishment of game bridges over motorways and from the protection of young male stags migrating through the statutory ‘red deer–free’ areas.
... It is known that hybridisation between Cervus nippon and Cervus elaphus (mainly Cervus elaphus females and Cervus nippon males) occurs and that hybrids are fertile. Hybridisation may lead to extensive introgression (Zachos & Hartl, 2011). Studies on population genetics and subspecies of red deer exclusively used mtDNA, which may suggest relationships that are not reproducible when using paternal genes. ...
Article
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Systematic relationships of cervids have been controversial for decades. Despite new input from molecular systematics, consensus could only be partially reached. The initial, gross (sub) classification based on morphology and comparative anatomy was mostly supported by molecular data. The rich fossil record of cervids has never been extensively tested in phylogenetic frameworks concerning potential systematic relationships of fossil cervids to extant cervids. The aim of this work was to investigate the systematic relationships of extant and fossil cervids using molecular and morphological characters and make implications about their evolutionary history based on the phylogenetic reconstructions. To achieve these objectives, molecular data were compiled consisting of five nuclear markers and the complete mitochondrial genome of 50 extant and one fossil cervids. Several analyses using different data partitions, taxon sampling, partitioning schemes, and optimality criteria were undertaken. In addition, the most extensive morphological character matrix for such a broad cervid taxon sampling was compiled including 168 cranial and dental characters of 41 extant and 29 fossil cervids. The morphological and molecular data were analysed in a combined approach and other comprehensive phylogenetic reconstructions. The results showed that most Miocene cervids were more closely related to each other than to any other cervids. They were often positioned between the outgroup and all other cervids or as the sister taxon to Muntiacini. Two Miocene cervids were frequently placed within Muntiacini. Plio-and Pleistocene cervids could often be affiliated to Cervini, Odocoileini or Capreolini. The phylogenetic analyses provide new insights into the evolutionary history of cervids. Several fossil cervids could be successfully related to living representatives, confirming previously assumed affiliations based on comparative morphology and introducing new hypotheses. New systematic relationships were observed, some uncertainties persisted and resolving systematics within certain taxa remained challenging.
... Red deer (Cervus elaphus L.) belongs to the most abundant cervid species in Europe. Wild living and farming populations of red deer have an increasing economic, cultural, and ecological importance [1]. Red deer are a seasonally reproduced ruminant, and females exhibit several estrous cycles in autumn [2,3]. ...
Article
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The aim was to estimate the effective pharmacological method of the estrous cycle synchronization by checking the effects of synchronization by measurement of progesterone (P4) and 17-beta estradiol (E2) concentration by RIA and artificial insemination. The experiment was performed at the red deer farm in Rudzie (North-East Poland; 3 year’s old). The herd (N = 14) was kept away from bulls and was divided in two groups of seven animals. In the Group I, CIDR insert (0.3 g of P4) was applicated intravaginally for 12 days; a second insert replaced the first one for the next 12 days, and next 200 IU of equine chorionic gonadotropin (eCG) was injected intramuscularly (Folligon). Estrus was expected 48 h after eCG injection. In the Group II, Chronogest sponge (20 mg of flugestone acetate) was applicated intravaginally and after 7 days replaced with second chronogest sponge for 7 days. After removing the sponge, on the same day eCG was injected and estrus was expected after 48 h. Artificial insemination was provided with frozen-thawed semen twice: 12 and 24 h after expected estrus. The peripheral blood from the jugular vein was collected each time when the inserts or sponge were applicated and 40 days after insemination. The concentration of P4 and E2 in plasma was measured by RIA. The effectiveness of insemination was monitored by pregnancy-associated glycoproteins determination and observed by the number of calves born. Two pregnancies were confirmed in Group I and five in Group II based on PAG concentration. One newborn was observed in Group I and five in Group II. Both methods of synchronization are effective in hinds based on the received profile of steroids. Although the sponge shape in case of chronogest is better comparing with CIDR, which was not completely deposited in the vagina of hind, potentially leads to bacteria inflammation, and it disturbs the rightful endocrine regulation. Moreover, pregnancy rate and hormone responsiveness were better in Group II.
... The low values of H O observed in our study are rare for red deer and usually only found in populations with longterm low effective population sizes such as the red deer from Sardinia or from Mesola in northern Italy (Hmwe et al. 2006). Both, H O and H E values, in the Hasselbusch AMU are among the lowest ever found in a population of this species (Zachos and Hartl 2011;Zachos et al. 2016). We note that comparisons of genetic diversity estimates across studies should be performed with caution (Reiner et al. 2019). ...
Article
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Red deer (Cervus elaphus) throughout central Europe are influenced by different anthropogenic activities including habitat fragmentation, selective hunting and translocations. This has substantial impacts on genetic diversity and the long-term conservation of local populations of this species. Here we use genetic samples from 480 red deer individuals to assess genetic diversity and differentiation of the 12 administrative management units located in Schleswig Holstein, the northernmost federal state in Germany. We applied multiple analytical approaches and show that the history of local populations (i.e., translocations, culling of individuals outside of designated red deer zones, anthropogenic infrastructures) potentially has led to low levels of genetic diversity. Mean expected heterozygosity was below 0.6 and we observed on average 4.2 alleles across 12 microsatellite loci. Effective population sizes below the recommended level of 50 were estimated for multiple local populations. Our estimates of genetic structure and gene flow show that red deer in northern Germany are best described as a complex network of asymmetrically connected subpopulations, with high genetic exchange among some local populations and reduced connectivity of others. Genetic diversity was also correlated with population densities of neighboring management units. Based on these findings, we suggest that connectivity among existing management units should be considered in the practical management of the species, which means that some administrative management units should be managed together, while the effective isolation of other units needs to be mitigated.
... The average density of red deer (Cervus elaphus) in the Babia Góra forests was estimated at 8.6 ± 2.6 individuals/km 2 in 2010-2016 (mean ± sd; data from the BgNP Directorate). Red deer is the only ungulate in the subalpine spruce forest of the Babia Góra Massif, and the most abundant ungulate in most Central European forests (Zachos and Hartl, 2011). Rowan is the main palatable tree species in the Babia Góra subalpine spruce forest. ...
Article
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Browsing damage by ungulates is among the most decisive factors affecting the establishment and growth of young trees. In recent decades, ungulate populations have been expanding in the Northern Hemisphere; impairment of tree regeneration by their activity is an increasing problem. Herbivore-induced changes in tree regeneration may alter the composition and biodiversity of the future tree stand. In this study we determined where young trees can leave the seedling bank and succeed to higher forest strata in a protected natural forest that is under strong herbivore pressure. We studied rowan (Sorbus aucuparia) regeneration in a subalpine spruce forest growing in a strictly protected area of Babia Góra National Park (Western Carpathians). Rowan is one of the most palatable forest species, so browsing can significantly limit its growth. We predicted that factors, that restrict the movement of red deer (Cervus elaphus), decrease their visits in some forest patches, which will result in a higher share of rowan saplings. We also considered two other factors that can affect the distribution of rowan saplings: light availability and distance to maternal trees. In particular, we tested whether the occurrence of rowan saplings was related to: (i) slope angle, amount of logs lying on the forest floor, and distance to the hiking trails; (ii) distance to fruit-bearing trees; and (iii) canopy openness. The results confirmed our main predictions concerning the relation between the occurrence of palatable tree saplings and the availability of a forest area to ungulates. Factors related to the availability of terrain to red deer significantly influenced the distribution of rowan saplings taller than 1 m. The probability of rowan sapling occurrence increased when the amount of logs was higher and the distance to hiking trails was shorter. Slope had a significant negative impact on sapling occurrence. We found no effect of proximity of fruit-bearing rowan trees or canopy gap area. Thus, in the natural forest, the likelihood that young palatable trees will make the transition from seedlings to taller saplings seems to be determined mainly by factors related to ungulate activity. This suggests that the increase of ungulate populations and their browsing behavior will affect the species composition and spatial structure of future tree stands.
... Phylogenetic and phylogeographic reconstructions of the Sardinian red deer, carried out by different scholars, using different methodologies, have found close relationships with the North African red deer (Hartl et al. 1995;Ludt et al. 2004Ludt et al. , 1075Zachos and Hartl 2011); with the red deer from continental Italy (currently present in Gran Bosco della Mesola) and Spain (Hmwe et al. 2006, 691;Hajji et al. 2008, 669;Zachos et al. 2003) and even Bulgarian red deer, which are phylogenetically very similar to the Eastern subspecies (Hartl et al. 1995), even though the latter hypothesis has not been confirmed by later studies (Hajji et al. 2008, 660). By integrating genetic, palaeontological and historical/archaeological data Hajji et al. (2008, 669) propose an anthropic introduction of the red deer on the island. ...
... The common and scientific species names for our three study species are taken from Burgin et al. (2020). For red deer, we additionally considered Ludt et al. (2004) and Zachos and Hartl (2011) ...
Article
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Eurasian deer are characterized by the extraordinary diversity of their vocal repertoires. Male sexual calls range from roars with relatively low fundamental frequency (hereafter fo) in red deer Cervus elaphus, to moans with extremely high fo in sika deer Cervus nippon, and almost infrasonic groans with exceptionally low fo in fallow deer Dama dama. Moreover, while both red and fallow males are capable of lowering their formant frequencies during their calls, sika males appear to lack this ability. Female contact calls are also characterized by relatively less pronounced, yet strong interspecific differences. The aim of this study is to examine the anatomical bases of these inter-specific and inter-sexual differences by identifying if the acoustic variation is reflected in corresponding anatomical variation. To do this, we investigated the vocal anatomy of male and female specimens of each of these three species. Across species and sexes, we find that the observed acoustic variability is indeed related to expected corresponding anatomical differences, based on the source-filter theory of vocal production. At the source level, low fo is associated with larger vocal folds, whereas high fo is associated with smaller vocal folds: sika deer have the smallest vocal folds and male fallow deer the largest. Red and sika deer vocal folds do not appear to be sexually dimorphic, while fallow deer exhibit strong sexual dimorphism (after correcting for body size differences). At the filter level, the variability in formants is related to the configuration of the vocal tract: in fallow and red deer, both sexes have evolved a permanently descended larynx (with a resting position of the larynx much lower in males than in females). Both sexes also have the potential for momentary, call-synchronous vocal tract elongation, again more pronounced in males than in females. In contrast, the resting position of the larynx is high in both sexes of sika deer and the potential for further active vocal tract elongation is virtually absent in both sexes. Anatomical evidence suggests an evolutionary reversal in larynx position within sika deer, that is, a secondary larynx ascent. Together, our observations confirm that the observed diversity of vocal behaviour in polygynous deer is supported by strong anatomical differences, highlighting the importance of anatomical specializations in shaping mammalian vocal repertoires. Sexual selection is discussed as a potential evolutionary driver of the observed vocal diversity and sexual dimorphisms.
... L'augmentation de l'aire de répartition a également été observée pour les espèces de plaine. Le chevreuil et le cerf comptent aujourd'hui parmi les mammifères les plus largement distribués en Europe (Linnell & Zachos, 2010 ; Figure 1D&E) grâce à leurs capacités à occuper une large gamme d'habitats tels que les forêts tempérées ou les milieux du pourtour méditerranéen (Lovari, 2018), et dans le cas du cerf à de nombreuses réintroductions (Zachos and Hartl, 2011). Au cours de la fin du XIX e et au début du XX e siècle, une forte baisse du nombre de chevreuils en Europe et de sa distribution ont été observées en raison de la chasse, et dans certains pays, l'espèce était au bord de l'extinction -comme par exemple en Angleterre (Hewison and Staines, 2008) ou en Suède (Liberg et al., 2010). ...
Thesis
Les effets des changements globaux sur les habitats naturels sont de plus en plus perceptibles, et comprendre comment les animaux y répondent est nécessaire pour une meilleure gestion de leurs populations. C’est en effet à travers leur impact sur l’environnement, et essentiellement sur les habitats, que les activités humaines ont souvent le plus grand effet sur les écosystèmes, à travers le changement climatique, la fragmentation, la destruction de l'habitat, les changements dans l'utilisation des terres ou la surexploitation des ressources. Les ongulés constituent un exemple marquant de progression numérique et spatiale d’une guilde d’espèces dans des écosystèmes impactés par l’Homme. Cet essor démographique est à l’origine d’un nombre croissant d’interactions entre Homme et faune et place la gestion de ces espèces au cœur des préoccupations des politiques publiques. Dans ce contexte, j’ai étudié cinq espèces de grands ongulés sauvages : le chamois, le mouflon, le bouquetin, le chevreuil et le cerf, dans le cadre du projet Mov-It (Ungulates MOVing across heterogeneous landscapes: identifying behavioural processes linking global change to spatially-explicIT demographic performance and management), soutenu par l’Agence Nationale de la Recherche (ANR). Dans un premier temps, je mets en évidence les liens entre variations intraspécifiques de la taille du domaine vital saisonnier des ongulés, le paysage (i.e. les ressources, le risque et l’hétérogénéité) et les traits d’histoire de vie de ces espèces. Je me suis ensuite intéressée plus particulièrement à l’influence des structures linéaires anthropiques et naturelles du paysage sur l'utilisation individuelle de l'espace. Je montre ainsi que les grands herbivores utilisent des structures linéaires du paysage pour délimiter leur domaine vital mensuel, mais que l'importance relative de ces structures linéaires dans la délimitation du domaine vital mensuel diminuait à mesure que leur densité augmentait dans le paysage local. Je mets également en évidence le caractère risqué des structures anthropiques pour les ongulés, en particulier l'effet de l'intensité de l’utilisation humaine de ces structures sur le nombre de traversées par les mouflons. Enfin, l’importance de la prise en compte du paysage du risque et des ressources sur l’organisation sociale est démontré. En effet, la formation de dyades (i.e. paires d’individus) est plus probable dans les milieux ouverts riche en ressources et lorsque le risque, incluant prédation et dérangement, est le plus fort (i.e. le jour). L’ensemble des résultats présentés dans ce travail de thèse a permis d’améliorer notre compréhension des effets de la structure du paysage et de la socialité sur la sélection d’habitat et le mouvement chez différentes espèces d’ongulés.
... Red deer (Cervus elaphus L.) is currently one of the most widespread European ungulate species [1]. Nowadays, the New Zealand is the kingdom of deer farms [2], but they have also been developed in Europe during the last twenty years. ...
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The aim was to compare the blastocyst stages of red deer embryos in respect of in vitro fertilization (IVF) efficiency, morphology, apoptotic and proliferative abilities, and antioxidative potential according to the reproductive status of hinds. We used three experimental groups, including the ovaries collected post mortem on the 4th and 13th days of the estrous cycle and during pregnancy (n = 18). After oocyte maturation, frozen-thawed epididymal semen was used for IVF. Blastocyst quality, apoptotic potential by determining the mRNA expression of BAX, BCL-2, OCT4, SOX2, and placenta-specific 8 gene (PLAC8), and antioxidative potential of blastocysts were evaluated by determining the mRNA expression of CuSOD, MnSOD, and GPX as well as the enzymatic activity of superoxide dismutase and reduced glutathione. The highest development rate of expanded blastocyst, mRNA expression of BCL-2, OCT4, SOX2, and PLAC8 and mRNA expression and enzymatic activity of the antioxidative factors increased (p < 0.05) in blastocysts developed from the oocytes collected on the 4th day, compared to those developed from the oocytes collected on the 13th day of the cycle and during pregnancy. Our study indicates that the 4th day of the estrous cycle is the most effective period for oocyte collection for IVF and embryo development in hinds, considering quality parameters and antioxidative potential of the blastocysts.
Article
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A molecular-genetic analysis of a control region and cytochrome b gene of the mtDNA of red deer (Cervus elaphus) from East-Central Europe (Pannonian Basin), South-Eastern Europe (South-Eastern part of the Balkan Peninsula) and in East European Plain (Northern Pontic region) has been carried out. The manifested genetic diversity of red deer, based on an analysis of the sequences of the cytochrome b gene indicates the presence of five deeply divergent mitochondrial lineages: lineages (A), (B) and (C), known from previous research, and two newly found unique mitochondrial mtDNA lineages (D) and (E). The mitochondrial lineages in the investigated red deer shown by D-loop were consistent with those identified by cytochrome b analysis (lineage (E) not included in the data set). The generalized analysis of the geographic pattern of biodiversity of C. elaphus on population level in Europe reveals that it is much more complex than previously assumed. Therefore, additional information on red deer population history, biology, ecology and recent genetic status must be collected in more biogeographical regions in Europe to clarify in details the conservation genetics of European red deer, as one of the urgent measures to protect its population biodiversity on the continent.
Technical Report
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Ce rapport, portant sur 125 espèces au total, reprend l'évaluation réalisée par espèce dans le cadre de la Liste rouge des mammifères de métropole de 2017 (Comité français de l'Union internationale pour la conservation de la nature, Muséum national d'Histoire naturelle, Société française pour l'étude et la protection des mammifères & Office national de la chasse et de la faune sauvage).
Chapter
This chapter is best viewed as an attempt at going a little bit deeper into various aspects of the species problem while at the same time serving the overall aim of this book, namely, to provide biologists with a distillation, as it were, of the species debate(s). Therefore, none of the subchapters claim to present an exhaustive discussion of its topic or species concept. Rather, they are best viewed as short commentary sections. As in many other sections of this book, the reader will see that often seemingly different views actually have a lot in common. I am not denying that there are important differences between various species concepts, but sometimes they are exaggerated, and the theoretical solution to the species problem (or at least an important part of it) in the form of a hierarchy of species concepts is proof that there is enough common ground for reconciliation. Cracraft (2000, p. 7) was right when he remarked on the similarity of species concepts saying “Similarities: all else is rhetoric” and “A student of species concepts must be able to sort through the rhetoric, unless, of course, the goal is to use it for one’s own gain”. In this book I argue that all species concepts are based on biological realities. Thus, none of them can simply be wrong. Some may be more general or more consistent with research results in different disciplines, and one or several may be superior to others (as indeed, I think, is the case with the Evolutionary Species Concept or General Lineage/Unified Species Concept), but all are real in the sense that they capture biological phenomena. This should be kept in mind because as Stamos (2003, p. 355) put it: “And indeed there is ‘something natural and something beautiful’ in each and every species concept, which taken together in their diversity reveal a conceptual world as rich and as breathtaking, in its own way, as anything to be found in the biological world. And if symbiosis in the biological world is truly a source of evolutionary innovation, there is no reason why it cannot also be so in the conceptual world of theories. All the more reason, then, to value rather than slash and burn the diversity of solutions to the species problem, for out of that diversity endosymbiotic innovations may be born”. This is not to say, and Stamos also adds this, that there are not better and less good solutions, but it is worth remembering when species debates become heated.
Chapter
The introductory quotation by Aristotle is not only a nice encapsulation of the idea of a great chain of being or , but it also shows that, from the birth of Western science and philosophy, it has always been obvious that nature has fuzzy boundaries and that demarcation of similar entities is a difficult task. In Sect. 5. 2 a potential theoretical solution to the species problem has been introduced: a hierarchical view that distinguishes between a true ontological or primary species concept and secondary operational species criteria. The ontological species concept (telling us what species taxa really are) is based on conceiving of species as population-level lineages or segments of such lineages as defined by either the Evolutionary Species Concept or the General Lineage/Unified Species Concept which are very similar. However, these concepts are all non-operational, they just provide the framework within which species can and should be identified. Identification and delimitation of species taxa is what taxonomists are doing, and it is important that they are doing it within a consistent theoretical framework. The practice of species delimitation, however, is not guided by the lineage framework beyond the condition that species must be lineages. Since the Tree of Life (and even more so the Web of Life in the prokaryotic world) is made up of lineages within lineages, the question whether there is an objective level of the species crucially depends on whether there is some (population) level within this fractal pattern that stands out. If that is not the case, then the species category, at least a single universal species category, is a myth: “What if there are theoretically important differences between the various segments of population lineages that we are identifying as species? Perhaps there are crucial differences between vertebrates, invertebrates, fungi and bacteria such that they should not all be regarded as forming the same kinds of species. What if, on our best theoretical understanding, there really do seem to be different kinds of species?” (Richards 2013, p. 60). If species taxa cannot be unified under the same kind of species category, then we would have to accept species pluralism. Yet, even if we accept, for the time being, the hierarchical solution of the species problem as far as the theoretical side of the coin is concerned, species delimitation is a fuzzy business. I will argue that completely objective, non-arbitrary species delimitation is impossible and that the approach coming closest to this unattainable ideal is not feasible and biologically irrelevant, which leaves us with the insight that taxonomy, even at its best, is only going to be an approximation of the real natural pattern and, importantly, that species delimitation resulting in biologically meaningful entities can only be done in hindsight. In other words, it is theoretically impossible to arrive at a fully consistent, completely non-arbitrary and biologically meaningful classification of life. The main reason for this is that taxonomy is essentially binary—species or no (sub-/super-)species, one or two (sub-/super-)species, etc.—while the evolutionary process is continuous and will create inherently fuzzy boundaries. The practical side of the species problem, the actual delimitation of species, is what biologists are dealing with every day—either directly as taxonomists or indirectly as “users” of taxonomy when they count species, analyse and compare conspecific and interspecific populations or manage species. It is therefore hardly surprising that species delimitation has received increasing attention, boosted undoubtedly by the methodological revolution in both genomics and bioinformatics. A Scopus search revealed that the number of published papers that had “species delimitation” in their title, abstract or keywords increased from 93 between 2001 and 2005 through 321 between 2006 and 2010 to 885 between 2011 and 2015 (see Fig. 1 in Camargo and Sites, 2013, for a similar finding). The available analytical approaches to species delimitation are diverse and comprise tree-based and non-tree-based methods. A more recent development is the implementation of coalescent theory, but there are also old ideas, such as the one regarding species as fields for recombination which originally goes back to Carson (1957) but has been revived by first Doyle (1995) and then more recently by Flot et al. (2010), which delimit pools of alleles that recombine and co-occur in individuals. An overview and discussion of existing methods can be found in the following publications (and references therein): Sites and Marshall (2003, 2004), Knowles and Carstens (2007), Wiens (2007), Petit and Excoffier (2009), Birky et al. (2010), Ence and Carstens (2011), Fujita et al. (2012), Hey and Pinho (2012), Birky (2013), Camargo and Sites (2013), Carstens et al. (2013), Zhang et al. (2013). There is even evidence that, across eukaryotic groups, a very simple correlation seems to exist between sexual compatibility and so-called compensatory base changes (CBC) in the internal transcribed spacer 2 (ITS2) transcript secondary structure such that the presence of even a single CBC seems to coincide very reliably (>90 %) with sexual incompatibility (Müller et al. 2007; Coleman 2009 and references therein). However, no matter if simple methods are based on genetic distances or morphological gaps or gene flow across hybrid zones or more complex methods such as tree-based coalescent models or cohesion tests, what all these approaches have in common is that they aim at consistently identifying lineages. What they do not do is objectively mark the level which is considered the species level: they make grouping (more) objective, but not ranking! This is why Sites and Marshall (2004, p. 220) conclude “virtually all [methods of species delimitation] will require researchers to make qualitative judgments. For example, there is no objective criterion for how much morphological divergence is enough to delimit a species, what threshold frequency of intermediates is needed to delimit species by genotypic clusters […], what proportion of unlinked loci are needed to delimit coalescent species […], or what frequency cutoff most appropriately indicates that no significant gene flow is occurring between populations”. There simply is no silver bullet for species delimitation. The available species delimitation methods become ever more sophisticated, but they only help us identify lineages; none of them can answer the question of where the line of the species lineage should be drawn. They will always entail a decision on some kind of cut-off criterion. But an (arbitrary!) cut-off criterion that can be measured or tested for objectively is not the same as an objective delimitation criterion! The sooner we recognize that some arbitrariness is naturally inherent in species delimitation, the earlier we can focus on biological phenomena, not names. The grey area during lineage divergence is a field of exciting evolutionary and ecological phenomena and for research that should not be spoiled by endless and ultimately futile debates about how to make species delimitation absolutely objective. I will not discuss concrete species delimitation algorithms here but rather focus on the more general aspects and difficulties surrounding delimitation in a world of fuzzy boundaries.
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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.
Article
The browsing habits of sika deer (Cervus nippon) in Japan have caused serious ecological problems. Appropriate management of sika deer populations requires understanding the different genetic structures of local populations. In the present study, we used 10 microsatellite polymorphisms to explore the genetic structures of sika deer populations (162 individuals) living in the Kanto region. The expected heterozygosity of the Tanzawa mountain range population (Group I) was lower than that of the populations in the Kanto mountain areas (Group II). Our results suggest that moderate gene flow has occurred between the sika deer populations in the Kanto mountain areas (Group II), but not to or from the Tanzawa mountain range population (Group I). Also, genetic structure analysis showed that the Tanzawa population was separated from the other populations. This is probably attributable to a genetic bottleneck that developed in the Tanzawa sika deer population in the 1950s. However, we found that the Tanzawa population has since recovered from the bottleneck situation and now exhibits good genetic diversity. Our results show that it is essential to periodically evaluate the genetic structures of deer populations to develop conservation strategies appropriate to the specific structures of individual populations at any given time.
Preprint
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Systematic relationships of cervids have been controversial for decades. Despite new input from molecular systematics, consensus could only be partially reached. The initial, gross (sub)classification based on morphology and comparative anatomy was mostly supported by molecular data. The rich fossil record of cervids has never been extensively tested in phylogenetic frameworks concerning potential systematic relationships of fossil cervids to extant cervids. The aim of this work was to investigate the systematic relationships of extant and fossil cervids using molecular and morphological characters and make implications about their evolutionary history based on the phylogenetic reconstructions. To achieve these objectives, molecular data were compiled consisting of five nuclear markers and the complete mitochondrial genome of 50 extant and one fossil cervid species. Several analyses using different data partitions, taxon sampling, partitioning schemes, and optimality criteria were undertaken. In addition, the most extensive morphological character matrix for such a broad cervid taxon sampling was compiled including 168 cranial and dental characters of 41 extant and 29 fossil cervid species. The morphological and molecular data were analysed in a combined approach and other comprehensive phylogenetic reconstructions. The results showed that most of the Miocene cervids were more closely related to each other than to any other cervids. They were often positioned between the outgroup and all other cervids or as the sister taxon to Muntiacini. Two Miocene cervids were frequently placed within Muntiacini. Plio- and Pleistocene cervids could often be affiliated to Cervini, Odocoileini or Capreolini. The phylogenetic analyses of this work provide new insights into the evolutionary history of cervids. Several fossil cervids could be successfully related to living representatives, confirming previously assumed affiliations based on comparative morphology and introducing new hypotheses. New systematic relationships were observed, some uncertainties persisted and resolving systematics within certain taxa remained challenging.
Article
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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.
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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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Chapter
There are about 30 species concepts in the literature. The exact number depends on what one considers a full or acknowledged concept (sometimes a comment or definition in the literature may not be called a concept and/or may not have made it into the canon of “official” species concepts), and it also depends on where one draws the line between similar or nearly identical concepts. The list below of 31 species concepts is therefore neither exhaustive nor the only way of listing species concepts. I mainly follow Mayden (1997, 1999), the list in Wilkins (2009b) which is based on Wilkins (2006a) and Wilkins’ website http:// scienceblogs. com/ evolvingthoughts / 2006/ 10/ 01/ a-list-of-26-species-concepts/ (Wilkins 2006b) but have included the General Lineage Species Concept and the Unified Species Concept by de Queiroz (1998, 1999, 2005b, 2007), added Stamos’ (2003) Biosimilarity Species Concept, Baum and Shaw’s (1995) Genealogical Species Concept and finally the Differential Fitness Species Concepts which was published more recently (Hausdorf 2011). I also sometimes use slightly different names than those given in other lists.
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Molecular forensic methods are being increasingly used to help enforce wildlife conservation laws. Using multilocus genotyping, illegal translocation of an animal can be demonstrated by excluding all potential source populations as an individual's population of origin. Here, we illustrate how this approach can be applied to a large continuous population by defining the population genetic structure and excluding suspect animals from each identified cluster. We aimed to test the hypothesis that recreational hunters had illegally introduced a group of red deer into a hunting area in Luxembourg. Reference samples were collected over a large area in order to test the possibility that the suspect individuals might be recent immigrants. Due to isolation-by-distance relationships in the data set, inferring the number of genetic clusters using Bayesian methods was not straightforward. Biologically meaningful clusters were only obtained by simultaneously analysing spatial and genetic information using the program baps 4.1. We inferred the presence of three genetic clusters in the study region. Using partial Mantel tests, we detected barriers to gene flow other than distance, probably created by a combination of urban areas, motorways and a river valley used for viticulture. The four focal animals could be excluded with a high certainty from the three genetic subpopulations and it was therefore likely that they had been released illegally.
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The dwarfing of large mammals on islands occurred repeatedly in the Pleistocene. Elephants, deer, hippopotami and other species became dwarfed on islands in Indonesia, the Mediterranean, the east Pacific and elsewhere. In most cases, the full-sized ancestral form can be recognized among the adjacent mainland fauna, but evolutionary rates cannot be estimated because the entry of the ancestor onto the island, and appearance of the dwarf form, are poorly dated. Here I give the first example in which the island dwarf is well dated, the full-sized ancestor is found in demonstrably older deposits on the island, and a good estimate can be made for the duration of the isolation leading to dwarfing. In the Last Interglacial, red deer on Jersey, Channel Islands, became reduced to one sixth of their body weight in less than six thousand years.
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Intra-specific Y-chromosomal sequence variation is useful for analysing the male contribution to a species’ spatial genetic structure. In red deer (Cervus elaphus) this is especially relevant, because geographic dispersal and game translocations occur mainly through the males. However, Y-chromosomal markers for wild organisms are scarce and frequently non-polymorphic within species. We assessed the intra-specific variation of two Y-chromosomal introns in red deer, one in the DBY (or DDX3Y) gene and the other in the UBE1Y gene. The introns were amplified using previously published exonic primers and directly sequenced in individuals of five red deer subspecies from across Eurasia. However, no nucleotide polymorphism was observed, which rebuts the usefulness of these introns for studies of red deer phylogeography and on illegal transport of red deer within this region. Male-based phylogeographic studies should thus be focused on other Y-chromosomal markers for this species.
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Screening of 43 protein loci was used to evaluate the level of genetic variability in 42 red deer Cervus elaphus Linnaeus, 1758 from Mesola Wood, the only native population presently occurring in peninsular Italy. The survey revealed polymorphism at four loci. In one of them evidence was provided for a significant deficiency of heterozygotes. Average expected heterozygosity and the proportion of polymorphic loci were He = 0.025 and P = 0.093, respectively. Although these values are low, they are consistent with those reported for other European red deer populations. Repeated bottlenecks and slow recoveries along with prolonged selective removal of stags were indicated as affecting genetic variation as a consequence of random drift. A genetic factor may have influenced female fertility and antler conformation. Comparisons with free-ranging deer from the Alps, and with an enclosed population yielded estimates of absolute (mean Nei's 1972 D = 0.004, SD = 0.002; mean modified Rogers' D = 0.063, SD = 0.013) and relative (Wright's FST = 0.094) genetic distance typical for red deer populations. Given the biological value of Mesola red deer, genetic results are discussed in relation to both population history and conservation. A strategy for urgent management interventions is also proposed.
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Morphological characters in red deer ( Cervus elaphus), which serve as criteria for selective hunting, were examined in relation to electrophoretic variation in three populations from the Vosges in eastern France. From the polymorphic loci examined, certain alleles at Idh-2, Me-1 and Acp-1 showed significant associations with a special development of body and antler characters selected for by hunters. Idh-2125 was associated with larger hind foot length in females and a higher number of antler points in males. Me-190 and Acp-1100 were associated with small spikes. The populations studied differed from one another in the duration and intensity of selective hunting and the increase or decrease in the respective allele frequencies could be explained by selection for large body size, a high number of antler points and against small spikes in yearlings, rather than by genetic drift. Among other morphological characters examined, the length of the main beam was significantly associated with the allele Acp-2100. In contrast, no associations could be detected between overall heterozygosity and the development or the degree of asymmetry (in paired structures) of any of the morphological traits in question. Although no obvious differences in the overall values of polymorphism or heterozygosity were found between the populations, selective hunting leads towards a change in allele frequencies and eventually to the loss of one or the other rare allele.
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The Mesola red deer Cervus elaphus in the Po delta are the only native red deer population on the Italian mainland and have been the focus of conservationists and wildlife biologists for some time. In our study, we present a genetic analysis of 25 Mesola red deer on the basis of 20 polymorphic microsatellite loci, aiming at estimating the population's genetic diversity and at providing information for a future genetic screening. In addition, we carried out a population viability analysis (PVA) with demographic and life-history data available from a long-term population survey, simulating different management scenarios. Genetic diversity was very low compared to the rest of Europe (observed and expected heterozygosity 0.50 and 0.61, respectively), and an overall excess of homozygosity was indicative of inbreeding. Calculations of the probability of identity and genotype mismatch frequencies suggested that between five and seven highly informative loci were sufficient to resolve individuals with reasonable certainty. The PVA yielded a generally poor outlook, but at the same time, it showed that management measures already taken significantly increased population viability. A sensitivity analysis revealed that inbreeding depression and possible catastrophes had a huge impact on the population's prospects. However, the establishment of two subpopulations and successful attempts at reducing the consequences of catastrophic events were able to significantly mitigate the harmful effects of both inbreeding and environmental stochasticity. These results, in particular the splitting of the population, may be of general interest to conservationists dealing with unique threatened populations.
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The Endangered Corsican red deer Cervus elaphus corsicanus was extirpated from Corsica in the early 1970s, at which time the Sardinian population fell to <250 individuals. The Sardinian authorities agreed to protect this subspecies and to secure its reintroduction in Corsica, a natural choice, considering ethological and historical descriptions. Since the beginning of 1985, when the first deer destined for captive breeding and eventual reintroduction arrived in Corsica, the population increased from 13 Sardinian founders to 106 captive animals under constant monitoring in three enclosures (Quenza, Casabianda and Ania di Fium'Orbu). The sites of Quenza, Chisà and Santo Pietro di Venaco were selected by the Regional Nature Park of Corsica for the reintroduction into the wild that began in 1998. Currently the size of the whole Corsican population is c. 250 individuals. These deer are still closely monitored and studied, both in enclosures and in the wild, to secure the long-term conservation of this subspecies. The Corsican and Sardinian populations together now total slightly >1,000, and the subspecies could therefore be downgraded to Near Threatened on the IUCN Red List.
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ABSTRACT • The European roe deer Capreolus capreolus is a typical faunal element of the Holocene. It was already present in Europe at least 600 000 years ago and it has been known from both glacial and interglacial phases since then. With nearly 3000 fossil and subfossil records, it is one of the most frequent mammals in the Late Quaternary. • During the Middle and Late Weichselian Pleniglacial, the distribution of the roe deer was not restricted to the Mediterranean peninsulas but repeatedly reached regions of central Europe. In contrast to that, roe deer records from the Last Glacial Maximum (LGM, 21.0–14.5 ka 14C BP) are largely confined to the Mediterranean peninsulas – with the exception of south-western France and the surroundings of the Carpathians where several records attest to its occurrence during the LGM. • During the Greenland Interstadial 1 (12.5–10.8 ka 14C BP), the species' distribution extended further north and the roe deer appeared north of the Alps and reached regions of central Germany. This seems to be correlated with the abrupt change to more favourable environmental conditions during this period. It is very likely that the roe deer disappeared north of the Alps during the Younger Dryas cooling (10.8–10.0 ka14C BP). The northern regions of the central European lowlands were recolonized by roe deer during the late Preboreal 9.7–9.5 ka 14C BP for the first time since the Weichselian Glacial. • The combined pattern of genetic data and fossil records of European roe deer suggests several regions in the Iberian peninsula, southern France, Italy and the Balkans as well as in the Carpathians and/or eastern Europe as glacial refugia. It further suggests that C. capreolus might have recolonized most parts of central-northern Europe out of one or more eastern European (not Balkan) and/or Carpathian refugia. This recolonization wave might have blocked immigration from the traditional Mediterranean areas.
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The red deer (Cervus elaphus) population in Denmark became almost extinct in recent historical times due to over-hunting. The species has subsequently recovered within remote areas, but non-Danish individuals have been introduced at several localities. To assess genetic structure, past demographic history, and the possibility of a still existing original stock, we analysed 349 specimens from 11 geographically separate areas and from three enclosed areas, genotyping 11 microsatellite loci. Moreover, an 826-bp fragment of the control region of the mitochondrial DNA was sequenced for 116 recent specimens and seven museum specimens. There was a significant difference in mean expected heterozygosity (HE) between the three enclosed areas and the 11 unenclosed areas. Significant departures from Hardy–Weinberg equilibrium were observed in the three enclosed areas and in nine of the unenclosed areas. The overall degree of genetic differentiation among all 14 areas was significant (FST = 0.09, P < 0.01), primarily because the mean pairwise FST for the three enclosed areas was significantly higher than that for the 11 unenclosed areas. A Bayesian clustering procedure detected three genetically distinct populations and indicated reduced gene flow between the enclosed and unenclosed areas. The individuals in the unenclosed areas show genotypic mixture, presumably as a result of gene flow among them. Markov Chain Monte Carlo simulations, based on the genealogical history of the microsatellite alleles, suggest a drastic decline in the effective population size of the enclosed areas some 188–474 years ago. Mitochondrial DNA analysis of the recent specimens showed seven haplotypes. Individuals from the enclosed Jægersborg Dyrehave contain haplotypes that occur all over Denmark and also are found in Western Europe. A close relationship between Scandinavian and Western European red deer is most likely. Only individuals from the unenclosed Lindenborg Estate and the enclosed Tofte Skov did not group with any other Danish individuals. As six of seven museum specimens had haplotypes also found in modern Danish samples, the current population of red deer in Denmark is genetically close to the original Danish red deer. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95, 688–701.
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Hunting can affect animal populations not only by increasing mortality but also by introducing selection components associated with particular features of individuals. In addition to the most widespread hunting system in Spain for Iberian red deer stags (Cervus elaphus hispanicus) called montería, there are also selective monterías aimed at culling poor-trophy males in order to improve the average quality of the trophies for commercial hunt. This way of removing poor-trophy males contrasts with the most common procedure of shooting individual males by selective stalking that is used in other areas of Europe. Also, due to the hunting procedure by which most deer are shot while running chased by dogs, it is doubtful whether hunters are actually producing a selective impact on deer populations. In this paper, we compare data of males shot in commercial montería and in selective montería in Southern Spain. We found that males in selective montería were smaller in body size and in antler size than in commercial hunts, even correcting by age, although the selective effect was stronger at some ages. We discuss the implications of this practice for sustainable use and conservation.
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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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For several centuries, game management has involved translocations of non-native individuals of many species to reinforce local native populations. However, there are few quantitative studies of potentially negative effects on population viability as expected when taxa with different local adaptations hybridise. The European red deer has been subject to particularly many translocations. Around 1900, a total of 17 red deer of Hungarian (Cervus elaphus hippelaphus) and German (C. e. germanicus) origin were introduced onto the island of Otterøya in Norway where few native red deer (C. e. atlanticus) remained (n~13). To assess interbreeding, the present stock on Otterøya and the indigenous Norwegian and Hungarian populations were characterised in 14 microsatellite loci and in the control region of mtDNA. An intermediate level of genetic variation in the Otterøya population and the presence of population specific alleles from both the indigenous Norwegian and the Hungarian population demonstrate that the introduced red deer interbred with the native. Even distributions of one indigenous and one non-indigenous mtDNA haplotype in the Otterøya population and two point estimates of admixture indicate similar genetic contributions from the two parental populations into the hybrid stock. Low numbers of migrants identified with Bayesian assignment tests demonstrate low recent gene flow from Otterøya into the Norwegian mainland population. The Otterøya hybrid stock has grown vastly in numbers during recent decades, suggesting a high population viability. We observed that the body mass of red deer on Otterøya was similar or greater than in adjacent indigenous Norwegian stocks, indicating that population performance has not been reduced in the hybrid stock and that gene flow probably has not had any negative effects. KeywordsTranslocation-Hybrid stock-Introgression-Admixture-Dispersal
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The red deer (Cervus elaphus) populationof Bavaria in Southern Germany was severelyreduced during the 19th century due toover-hunting. The species has since recoveredwithin designated areas. Subsequent habitatfragmentation presumably has changed thegenetic structure of Bavarian red deer.In order to assess the genetic diversity, weanalysed samples obtained from nine differentBavarian and two adjacent (Thueringen andCzech-Republic) red deer populations,genotyping 19 microsatellite loci. Our analysesrevealed moderate and significant differencesin diversity. Referring to assignment tests,the genetic differentiation of Bavarian reddeer was sufficient to assign an individual'sorigin to the correct population at an averageof 91.6%. The correlation of genetic andgeographic distance matrices revealed noevidence for isolation by distance. Thecoalescent model analysis suggests that thegenetic structure used to be characterized by adrift–gene flow equilibrium and is nowinfluenced by drift and disruption of the geneflow. Only three of the examined populationsshowed a probability of less than 10% that twogenes within these populations share a commonancestor (F IS-value). Twopopulations had high F IS values,indicating the influence of drift.Additionally, the intrapopulation indicesrevealed a low variability in thesepopulations. The estimated effective populationsizes (N e) were generally in thesame range as the actual population sizes. Theinbreeding rates, based on the estimated N e, and the inbreeding coefficientssuggested that the Bavarian red deerpopulations are in a stable state.
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The Corsican red deer, a sub-species of the European red deer endemic to Sardinia and Corsica, was abundant on both islands at the beginning of 1900. It went extinct in Corsica and reached a minimum of 100 individuals in Sardinia by 1970. Numbers have recovered in Sardinia with more than 1,000 rutting males now present; in the 1980s the deer was reintroduced to Corsica, but the Sardinian population remains fragmented. We developed a potential distribution model in Sardinia using Ecological Niche Factor Analysis. To assess the deer's protection status we compared the model with the existing and proposed conservation areas and investigated diVerent conservation scenarios in relation to the expansion of its current range and resilience to future changes in land use and predicted trends of desertiWcation. According to our results over 70% of Sardinia is unsuitable to the deer, nevertheless high suitability areas (Mediterranean forests away from main roads) are available throughout the island, particularly in the south and in the central-eastern part. Existing protected areas do not provide for the conservation of the deer but public owned forests, where hunting is prohibited, extend some level of protection, and the protected areas proposed by the Regional administration, if implemented, will be increasing this protection. Three main areas have emerged as conservation priorities to guarantee adequate conservation potential in the future. Our approach provides valuable data to inform conservation policy, and could be easily replicated in other parts of the Mediterranean.
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Abstract The biometry, demography and genetics of red deer Cervus elaphus of Mesola Wood (NE Italy), are presented and discussed in relation to the conservation of this population. Modest body size, low stature, oversimplified antlers and a low reproductive performance characterise red deer from Mesola Wood. The mitochondrial genome showed a private haplotype, different from other red deer in Italy and central Europe. The uniqueness of this nucleus and its biogeographic importance make a long-term conservation plan particularly urgent. Management measures such as fallow deer reduction, winter feeding and pasture mowing were tested, giving promising results. The physical condition of the animals improved, calf and adult mortality declined, and a few cases of antlers with bez tine or crown were reported in this study after four decades. Riassunto Il Cervo della Mesola: caratteristiche fisiche, dinamica di popolazione e prospettive di conservazione La biometria, la demografia e la genetica del cervo Cervus elaphus del Gran Bosco della Mesola (Italia nord-orientale), vengono presentate e discusse in relazione alla salvaguardia di questa popolazione. Il cervo della Mesola risulta caratterizzato dalle modeste dimensioni corporee, dalla struttura semplificata dei palchi e da un basso rendimento riproduttivo. L&apos;analisi del genoma mitocondriale ha evidenziato un aplotipo privato, diverso da quello degli altri cervi italiani e centroeuropei. L&apos;unicità di questo nucleo e la sua importanza biogeografica rendono particolarmente urgente un piano di conservazione a lungo termine. Sono stati verificati interventi gestionali quali la riduzione numerica dei daini, il foraggiamento invernale e lo sfalcio delle superfici a pascolo, con risultati promettenti. Le condizioni fisiche degli animali sono migliorate, la mortalità tra i piccoli e gli adulti è diminuita, e sono stati registrati alcuni casi di palchi dotati di ago o corona per la prima volta dopo quattro decenni.
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European red deer are known to show a conspicuous phylogeographic pattern with three distinct mtDNA lineages (western, eastern and North-African/Sardinian). The western lineage, believed to be indicative of a southwestern glacial refuge in Iberia and southern France, nowadays covers large areas of the continent including the British Isles, Scandinavia and parts of central Europe, while the eastern lineage is primarily found in southeast-central Europe, the Carpathians and the Balkans. However, large parts of central Europe and the whole northeast of the continent were not covered by previous analyses. To close this gap, we produced mtDNA control region sequences from more than 500 red deer from Denmark, Germany, Poland, Lithuania, Belarus, Ukraine and western Russia and combined our data with sequences available from earlier studies to an overall sample size of almost 1,100. Our results show that the western lineage extends far into the European east and is prominent in all eastern countries except for the Polish Carpathians, Ukraine and Russia where only eastern haplotypes occurred. While the latter may actually reflect the natural northward expansion of the eastern lineage after the last ice age, the present distribution of the western lineage in eastern Europe may in large parts be artificial and a result of translocations and reintroduction of red deer into areas where the species became extinct in historical times.
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There is growing concern about the evolutionary consequences of human harvesting on phenotypic trait quality in wild populations. Undesirable consequences are especially likely with trophy hunting because of its strong bias for specific phenotypic trait values, such as large antlers in cervids and horns in bovids. Selective hunting can cause a decline in a trophy trait over time if it is heritable, thereby reducing the long-term sustainability of the activity itself. How can we build a sustainable trophy hunting tradition without the negative trait-altering effects? We used an individual-based model to explore whether selective compensatory culling of ‘low quality’ individuals at an early life stage can facilitate sustainability, as suggested by information from managed game populations in eastern and central Europe. Our model was rooted in empirical data on red deer, where heritability of sexual ornaments has been confirmed and phenotypic quality can be assessed by antler size in individuals as young as 1 year. Simulations showed that targeted culling of low-quality yearlings could counter the selective effects of trophy hunting on the distribution of the affected trait (e.g. antler or horn size) in prime-aged individuals. Assumptions of trait heritability and young-to-adult correlation were essential for compensation, but the model proved robust to various other assumptions and changes to input parameters. The simulation approach allowed us to verify responses as evolutionary changes in trait values rather than short-term consequences of altered age structure, density and viability selection. We conclude that evolutionarily enlightened management may accommodate trophy hunting. This has far reaching implications as income from trophy hunting is often channelled into local conservation efforts and rural economies. As an essential follow-up, we recommend an analysis of the effects of trophy hunting in conjunction with compensatory culling on the phenotypic and underlying genetic variance of the trophy trait.
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The Norwegian red deer population (Cervus elaphus) was from the mid eighteenth to the early twentieth century drastically reduced in size and distribution but has the last century expanded both demographically and spatially. We have investigated genetic variation, differentiation and admixture in this spatially expanding ungulate population, using 14 microsatellites. The present genetic structure is moderate to strong with an average F ST = 0.08. Low M-ratios indicate loss of genetic variation in all localities and signals of a recent bottleneck was identified in 14 of 15 localities. Genetic distances between the localities indicate two main routes of dispersal during expansion, from the north–west and south–west, respectively. Bayesian assignment tests verify a break of the dataset in two, and demonstrate 99.9% probability for the existence of five sub-populations, which coincide well with five relict populations described by historic records. Computer simulations suggest that the observed genetic differentiation is recent rather than ancient, and that it may be explained by models of fragmentation or of founder events and subsequent merging rather than by models of recent bottlenecks in some particular demes within an ancient genetic structure.
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The largest population of red deer (Cervus elaphus) in Europe is found in Scotland. However, human impacts through hunting and introduction of foreign deer stock have disturbed the population's genetics to an unknown extent. In this study, we analysed mitochondrial control region sequences of 625 individuals to assess signatures of human and natural historical influence on the genetic diversity and population structure of red deer in the Scottish Highlands. Genetic diversity was high with 74 haplotypes found in our study area (115 x 87 km). Phylogenetic analyses revealed that none of the individuals had introgressed mtDNA from foreign species or subspecies of deer and only suggested a very few localized red deer translocations among British localities. A haplotype network and population analyses indicated significant genetic structure (Phi(ST)=0.3452, F(ST)=0.2478), largely concordant with the geographical location of the populations. Mismatch distribution analysis and neutrality tests indicated a significant population expansion for one of the main haplogroups found in the study area, approximately dated c. 8200 or 16 400 years ago when applying a fast or slow mutation rate, respectively. Contrary to general belief, our results strongly suggest that native Scottish red deer mtDNA haplotypes have persisted in the Scottish Highlands and that the population retains a largely natural haplotype diversity and structure in our study area.
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Red deer representing the four different European subspecies Cervus elaphus atlanticus, C. e. elaphus, C. e. germanicus, and C. e. scoticus were examined for allozyme variability at 35 enzyme loci. The proportion of polymorphic loci within populations (P) ranged from 0 to 13.8 per cent and the average heterozygosity (H) from 0 to 3.6 per cent. These estimates are within the range previously observed among mammalian species. Significant allele frequency differences were found both within and between subspecies. The mean genetic distance between subspecies (D = 0.0164) was smaller than the differentiation at similar taxonomic levels among other ungulates, probably because of a shorter time since divergence. Within subspecies the genetic differences between populations were similar to those reported between populations within closely related species in the same geographic region. Cluster analysis based on genetic distances indicated a major genetic dichotomy between the British C. e. scoticus and the Norwegian C. e. atlanticus on one hand and the Swedish C. e. elaphus and the continental C. e. germanicus on the other. Populations of pure C. e. elaphus were not found to differ genetically in any substantial way from Swedish populations of possible heterogeneous subspecific origin. An allele unique to C. e. scoticus was found in a Swedish enclosed population where imports of British deer are known to have taken place. A population established to preserve the genetic characteristics of the C. e. elaphus subspecies appeared to have lost 36 per cent of the electrophoretically measurable heterozygosity.
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In this article we describe the structure of a hybrid zone in Argyll, Scotland, between native red deer (Cervus elaphus) and introduced Japanese sika deer (Cervus nippon), on the basis of a genetic analysis using 11 microsatellite markers and mitochondrial DNA. In contrast to the findings of a previous study of the same population, we conclude that the deer fall into two distinct genetic classes, corresponding to either a sika-like or red-like phenotype. Introgression is rare at any one locus, but where the taxa overlap up to 40% of deer carry apparently introgressed alleles. While most putative hybrids are heterozygous at only one locus, there are rare multiple heterozygotes, reflecting significant linkage disequilibrium within both sika- and red-like populations. The rate of backcrossing into the sika population is estimated as H = 0.002 per generation and into red, H = 0.001 per generation. On the basis of historical evidence that red deer entered Kintyre only recently, a diffusion model evaluated by maximum likelihood shows that sika have increased at approximately 9.2% yr-1 from low frequency and disperse at a rate of approximately 3.7 km yr-1. Introgression into the red-like population is greater in the south, while introgression into sika varies little along the transect. For both sika- and red-like populations, the degree of introgression is 30-40% of that predicted from the rates of current hybridization inferred from linkage disequilibria; however, in neither case is this statistically significant evidence for selection against introgression.