Expanding ranges of wild and feral deer in Great Britain

1. It is often stated that wild deer in Great Britain are increasing in range and number. This study presents the first quantified estimate of national range expansion for all six species. From national surveys of deer presence in 10 km squares between 1972 and 2002, the red deer range is estimated to have expanded at a compound rate of 0.3% per year, fallow deer at 1.8%, Chinese water deer at 2.0%, roe deer at 2.3%, Japanese sika at 5.3% and Reeves’ muntjac at 8.2%. 2. Each species is expected to expand its range further for the foreseeable future. The most widespread species, roe deer, is predicted to be present within 79% of all 10 km squares in mainland Britain within 10 years. 3. Deer range expansion brings a combination of benefits and costs, and some degree of management is considered necessary to limit both environmental damage and deer suffering.     

Low genotyping error rates in non-invasively collected samples from roe deer of the Bavarian Forest National Park

Genetic wildlife monitoring is increasingly carried out on the basis of non-invasively collected samples, whereby the most commonly used DNA sources are skin appendages (hairs, feathers) and faeces. In order to guide decisions regarding future adequate ways to monitor the roe deer (Capreolus capreolus) population of the Bavarian Forest National Park in Germany, we tested these two different types of DNA source materials to compare their suitability for genetic monitoring. We determined the haplotypes (d-loop) of 19 roe deer and genotyped each individual (tissue, hairs, faeces) across 12 microsatellite loci. The amount of missing and erroneous microsatellite alleles obtained from hair and faeces samples, respectively, was estimated based on comparisons with the corresponding tissue sample control. We observed no missing alleles in hair samples, but in fecal samples PCR failed in 30 out of 228 instances (19 individuals x 12 loci), corresponding to a frequency of missing alleles of 13.2% across all loci and individuals. In genotypes generated from hairs erroneous alleles were detected in 2 out of 228 instances (0.9%), while genotypes retrieved from fecal samples displayed erroneous alleles in 6 out of 198 remaining instances (3%). We conclude that both hair and fecal samples are generally well suited for genetic roe deer monitoring, but that fecal sample based analyses require a larger sample size to account for higher PCR failure rates.     

The relative densities of red (Cervus elaphus) and roe (Capreolus capreolus) deer and their relationship in Scottish plantation forests

Population densities of red and roe deer were estimated in 20 conifer plantations throughout central and northern Scotland. Ten forests were selected on the basis of culling records that were considered likely to cover the range of possible densities. Ten further forests were selected at random to investigate the relationship between red and roe deer densities. Density assessment was by standing-crop dung-group counts, on 1.5 x 100 m transects. Dung decay rates were estimated in four areas and used to convert the values to deer densities. Red deer densities were estimated to range from 0.3-35 km⁻²; roe deer densities ranged from 0.5-25 km⁻². Preferences for different structural habitats within individual forests (i.e. blocks with different age classes of tree) were inferred from dung densities. Red deer preferred unplanted ground, open-thicket, and prethicket; roe deer likewise preferred prethicket (highly) and open-thicket. Both species avoided closed-canopy habitats. Red and roe deer densities were negatively correlated and the relationship was significant when the data were log transformed ( P < 0.05).     

Malignant catarrhal fever in free-ranging cervids associated with OvHV-2 and CpHV-2 DNA

Pathologic lesions were summarized in 18 free-ranging cervids (15 moose [Alces alces], two roe deer [Capreolus capreolus], and one red deer [Cervus elaphus]) diagnosed with malignant catarrhal fever (MCF) after examination at the National Veterinary Institute, Oslo 1982-2005. Eye lesions (conjunctivitis, corneal opacity, fibrin clots in the anterior eye chamber) were the most frequent gross finding. Erosive-ulcerative mucosal lesions in the nose and mouth were also commonly found. Histopathology revealed a nonpurulent vasculitis and perivasculitis in the central nervous system (CNS) typical of MCF in 16 of the cases. The diagnosis in the remaining two animals was based upon histologic eye lesions consistent with MCF (CNS not available for examination). Polymerase chain reaction was run on samples from 15 individuals for evidence of MCF-virus DNA, and ovine herpesvirus-2 (OvHV-2) DNA was detected in five moose, one roe deer, and one red deer, and caprine herpesvirus-2 (CpHV-2) DNA was detected in two moose and one roe deer. Sera from 1,000 free-ranging cervids were tested for specific antibodies to MCF-associated viruses (MCFV) by competitive inhibition enzyme-linked immunosorbent assay. The seroprevalences were: red deer 5%, reindeer (Rangifer tarandus) 4%, roe deer 2%, and moose 0.4% (n = 250 for all four species). The results indicate that sheep and goat MCFV may cause serious disease in wild moose, roe deer, and red deer. The seropositive cervids most likely represent individuals infected with either OvHV-2 or CpHV-2, but may also reflect infections with other related MCFV.     

Tick burden on European roe deer (<Emphasis Type="Italic">Capreolus capreolus</Emphasis>)

In our study we assessed the tick burden on roe deer (Capreolus capreolus L.) in relation to age, physical condition, sex, deer density and season. The main objective was to find predictive parameters for tick burden. In September 2007, May, July, and September 2008, and in May and July 2009 we collected ticks on 142 culled roe deer from nine forest departments in Southern Hesse, Germany. To correlate tick burden and deer density we estimated deer density using line transect sampling that accounts for different detectability in March 2008 and 2009, respectively. We collected more than 8,600 ticks from roe deer heads and necks, 92.6% of which were Ixodes spp., 7.4% Dermacentor spp. Among Ixodes, 3.3% were larvae, 50.5% nymphs, 34.8% females and 11.4% males, with significant seasonal deviation. Total tick infestation was high, with considerable individual variation (from 0 to 270 ticks/deer). Adult tick burden was positively correlated with roe deer body indices (body mass, age, hind foot length). Significantly more nymphs were found on deer from forest departments with high roe deer density indices, indicating a positive correlation with deer abundance. Overall, tick burden was highly variable. Seasonality and large scale spatial characteristics appeared to be the most important factors affecting tick burden on roe deer.     

Variation in cohort mandible size as an index of roe deer (Capredus capreolus) densities and population trends

This paper examines the effects of increasing population density on the skeletal development of roe deer as indexed by variation in the size of the mandible. We tested for density-dependence in total jaw length and minimum diastema height among cohorts in a single, intensively studied population in France. A strong relationship of average adult jaw length of both males and females with deer density in the year of birth of a given cohort was observed, and presumably reflects inadequate nutrition of juveniles at high population density. No such relationship was observed for the minimum diastema height which remained constant over the range of densities studied. This relationship was exploited to investigate population trends of roe deer in Britain. In nine populations, where equivalent data were available for between 16 and 28 separate cohorts, a consistent pattern of variation in cohort jaw size was observed, with a decrease in average total length, but not diastema height, towards the present. This trend is discussed in relation to changes in habitat structure in the coniferous forests of Britain. The utility of cohort jaw length as a management tool to index population trends of roe deer is highlighted.     

Temporal shifts of DNA‐microsatellite allele profiles in roe deer (Capreolus capreolus L.) within three decades

DNA‐microsatellite polymorphism (four loci) was studied in 56 male roe deer (Capreolus capreolus) from a 900‐ha hunting territory in the Vosges du Nord Mountains (France), culled over 34 years (1956–1990). Changed allele frequencies at two loci within this period, and increased allelic diversity, were traced to a phase of reduced population density and subsequent immigration. Decadic population samples collected within 900‐ha were distinguished by higher genetic variability measures than were certain geographical samples across Central Europe (4–900 km). On average, the decadic cohorts were distinguished by a gene diversity index of G_ST = 0.0286, and a genetic distance of D = 0.0938, which reflect 54% (G_ST) and 69% (D) of the respective geographic (350 km) differentiation indices of roe deer in Central Europe. The importance of demography and population ecology effects for microevolution in a large mammal is demonstrated, as is the risk of artefact by composing population samples of deer over several years. Population genetic screening should cover various demes of roe deer from the same general region, and be based on many unlinked polymorphic loci, to minimize the distorting effects of genetic dynamics at the small spatial scale.     

Unexpected population genetic structure of European roe deer in Poland: an invasion of the mtDNA genome from Siberian roe deer

Introgressive hybridization is a widespread evolutionary phenomenon which may lead to increased allelic variation at selective neutral loci and to transfer of fitness‐related traits to introgressed lineages. We inferred the population genetic structure of the European roe deer (Capreolus capreolus) in Poland from mitochondrial (CR and cyt b) and sex‐linked markers (ZFX, SRY, DBY4 and DBY8). Analyses of CR mtDNA sequences from 452 individuals indicated widespread introgression of Siberian roe deer (C. pygargus) mtDNA in the European roe deer genome, 2000 km from the current distribution range of C. pygargus. Introgressed individuals constituted 16.6% of the deer studied. Nearly 75% of them possessed haplotypes belonging to the group which arose 23 kyr ago and have not been detected within the natural range of Siberian roe deer, indicating that majority of present introgression has ancient origin. Unlike the mtDNA results, sex‐specific markers did not show signs of introgression. Species distribution modelling analyses suggested that C. pygargus could have extended its range as far west as Central Europe after last glacial maximum. The main hybridization event was probably associated with range expansion of the most abundant European roe deer lineage from western refugia and took place in Central Europe after the Younger Dryas (10.8–10.0 ka BP). Initially, introgressed mtDNA variants could have spread out on the wave of expansion through the mechanism of gene surfing, reaching high frequencies in European roe deer populations and leading to observed asymmetrical gene flow. Human‐mediated introductions of C. pygargus had minimal effect on the extent of mtDNA introgression.     

Antibodies to granulocytic Ehrlichia in moose, red deer, and roe deer in Norway

Serum samples from 104 moose (Alces alces), 124 red deer (Cervus elaphus) and 114 roe deer (Capreolus capreolus), collected from different counties in southern Norway from 1994 to 2000, were analysed by an indirect immunofluorescent antibody staining method for antibodies to Ehrlichia equi. The overall seroprevalences for granulocytic Ehrlichia spp. in moose, red deer, and roe deer from Ixodes ricinus infested counties were 43%, 55%, and 96%, respectively. Antibody prevalence was significantly higher in roe deer than in moose and red deer (P < 0.001). Mean antibody titers (log10 +/- SD) to E. equi in sera from moose, red deer, and roe deer were 1:1,497 (3.17 +/- 0.646), 1:234 (2.37 +/- 0.424) and 1:676 (2.83 +/- 0.404), respectively. The present work indicates that all these wild ruminant species are exposed to granulocytic Ehrlichia in Norway.     

Quantitative analysis of genetic parameters in populations of European (Capreolus capreolus L.) and Siberian (Capreolus pygargus Pall.) roe deer with RAPD markers

A technique for obtaining unbiased estimates of genetic parameters (allelic frequencies of RAPD loci, heterozygosity (H), Wright's F statistic, and Nei's genetic distances) in populations of the European (Capreolus capreolus L.) and Siberian (Capreolus pygargus Pall.) roe deer is presented. The technique employs jackknifing and multiple comparative analysis based on a modified Holmes's procedure for Bonferroni's test. It was demonstrated that samples from local groups of roe deer in the Trans-Ural region did not differ significantly in allelic frequencies (0.8, 0.81, and 0.78; P > 0.447) or Nei's genetic distances (0.0056, 0.0273, and 0.0218; P = 0.26), but they could be differentiated based on Wright's F statistic (0.0346, 0.0519, and 0.0450; P = 10(-9)). The parameters of intrapopulation heterozygosity (from 0.18 to 0.042) formed a gradient from the east to the west. Calibration estimates of molecular evolution rate in the family Cervidae obtained based on published data and Jukes-Cantor genetic distances estimated in this study demonstrated that the Siberian roe deer has split into two subspecies, C. pygargus pygargus Pall. and C. pygargus tianschanicus Satunin in the interval between 229 and 462.3 thousand years ago. The species formation of the Siberian and European roe deer was dated between 1.375 and 2.75 Myr ago. Based on the results obtained we recommend the approaches used in the study for analysis of population genetic structure and phylogenetic relationships between populations, subspecies, species, and higher taxa.     

Wolf predation on moose and roe deer: chase distances and outcome of encounters

We examined chase distances of gray wolves Canis lupus Linnaeus, 1758 hunting moose Alces alces and roe deer Capreolus capreolus, and recorded details of encounters between wolves and prey on the Scandinavian Peninsula, 1997–2003. In total, 252 wolf attacks on moose and 64 attacks on roe deer were registered during 4200 km of snow tracking in 28 wolf territories. Average chase distances were 76 m for moose and 237 m for roe deer, a difference likely due to variation in body size and vigilance between prey species. A model including prey species, outcome of the attack, and snow depth explained 15–19% of the variation found in chase distances, with shorter chase distances associated with greater snow depth and with successful attacks on moose but not on roe deer. Wolf hunting success did not differ between prey species (moose 43%, roe deer 47%) but in 11% of the wolf attacks on moose at least one moose was injured but not killed, whereas no injured roe deer survived. Compared with most North American wolf studies chase distances were shorter, hunting success was greater, and fewer moose made a stand when attacked by wolves in our study. Differences in wolf encounters with moose and roe deer likely result from different anti-predator behaviour and predator-prey history between prey species.     

In vitro digestibility of seasonal forages used by wapiti and roe deer in northeastern China

Seasonal changes in in vitro dry matter digestibility (IVDMD) of five forage classes (browses, sedges, forbs. ferns and the bark of Ulmus spp.) used by wapiti and roe deer were determined in relation to chemical composition, using the two-stage technique of in vitro trial with rumen inocula from one wapiti and one roe deer captured in winter. Wapiti and roe deer showed remarkable fluctuation in IVDMD among seasons and forage classes. IVDMD decreased from 43.8-64.3% in spring to 29.4-40.5% in winter for wapiti, and from 52.4-77.4% in spring to 25.9-36.5% in winter for roe deer. There was significant variation in IVDMD among species of browses for wapiti and roe deer. IVDMD ranged from 28.5% to 58.7% in twigs and from 39.8% to 58.2% in leaves for wapiti, and from 24.1% to 67.7% in twigs and from 40.0% to 71.7% in leaves for roe deer. Roe deer had higher IVDMD (40.0-71.7%) in both leaves and twigs of browses than wapiti (35.9-58.7%) during snow-free seasons, but IVDMD of twigs was lower for roc deer (25.9-36.5%) than wapiti (29.4-38.4%) in winter. Significant multiple correlation was found between NDF and ADF, and IVDMD in sedges, forbs and ferns for both wapiti and roe deer, whereas only roe deer showed the significant multiple correlation for leaves of browses.     

Intolerance amongst deer species at feeding: roe deer are uneasy banqueters

The function of inter-specific interactions in large herbivore assemblies is under debate. We have studied inter-specific interactions between roe and fallow deer on pastures, to assess whether competition between these Cervidae occurs through behavioural interference. Roe were displaced by fallow deer (i.e. the former moved away from the latter at a distance of >50 m) in 83% of cases (N = 127). Ninety-four percentage (N = 83) of displacement events occurred while roe deer were feeding; in 50% of these cases (N = 78), roe stopped grazing and left the feeding ground. Even when fallow deer did not show any sign of direct aggression to roe, these moved away from fallow in 72% of cases (N = 127). Vigilance rate was significantly greater in roe than in fallow deer, irrespectively of the presence of the other species. When roe and fallow deer grazed within 50 m from each other, vigilance rate increased significantly in roe, but not in fallow deer. Roe deer, in a group, were significantly more tolerant of the presence of fallow deer (even in group), than when solitary. Fallow deer seemed to be able to exclude roe deer from feeding sites through behavioural intolerance.     

Large Impact of Eurasian Lynx Predation on Roe Deer Population Dynamics

The effects of predation on ungulate populations depend on several factors. One of the most important factors is the proportion of predation that is additive or compensatory respectively to other mortality in the prey, i.e., the relative effect of top-down and bottom-up processes. We estimated Eurasian lynx (Lynx lynx) kill rate on roe deer (Capreolus capreolus) using radio-collared lynx. Kill rate was strongly affected by lynx social status. For males it was 4.85 ± 1.30 S.E. roe deer per 30 days, for females with kittens 6.23 ± 0.83 S.E. and for solitary females 2.71 ± 0.47 S.E. We found very weak support for effects of prey density (both for Type I (linear) and Type II (non-linear) functional responses) and of season (winter, summer) on lynx kill rate. Additionally, we analysed the growth rate in a roe deer population from 1985 to 2005 in an area, which lynx naturally re-colonized in 1996. The annual roe deer growth rate was lower after lynx re-colonized the study area, but it was also negatively influenced by roe deer density. Before lynx colonized the area roe deer growth rate was λ = 1.079 (± 0.061 S.E.), while after lynx re-colonization it was λ = 0.94 (± 0.051 S.E.). Thus, the growth rate in the roe deer population decreased by Δλ = 0.14 (± 0.080 S.E.) after lynx re-colonized the study area, which corresponded to the estimated lynx predation rate on roe deer (0.11 ± 0.042 S.E.), suggesting that lynx predation was mainly additive to other mortality in roe deer. To conclude, this study suggests that lynx predation together with density dependent factors both influence the roe deer population dynamics. Thus, both top-down and bottom-up processes operated at the same time in this predator-prey system.     

Annual changes in roe deer (Capreolus capreolus L.) diet in the Bohemian Forest, Czech Republic/Germany

The composition of roe deer diet in the Bohemian Forest was analysed with the aim to assess its role in forest habitat altered by bark beetle outbreaks and wind calamities. The annual diet of roe deer was studied at both, Czech and German, sides of the Bohemian Forest using microscopic analyses of faeces. On average, the largest part of the roe deer diet consisted of forbs (32%), followed by three other components—grasses (17%), coniferous trees (13%) and broadleaved trees (11%). Overall the results show that the composition of roe deer diet in the Bohemian Forest is that of a typical concentrate selector.     

Microsatellite investigation of roe deer (Capreolus capreolus) in Scandinavia reveals genetic differentiation of a Baltic Sea Island population

After a decline in the early 19th century, a remnant population of the Scandinavian roe deer (Capreolus capreolus) was protected in southern Sweden around 1840. The roe deer quickly recovered and recolonized most of the Scandinavian Peninsula. In this study, I analyze microsatellite variation in 14 roe deer populations in Scandinavia to (1) investigate if all Swedish roe deer trace their ancestry to southern Sweden, (2) define other areas where roe deer may have remained during the population decline, and (3) examine if the large geographic distances during the expansion phase have influenced the degree and sorting of genetic variation. The results show that the mainland roe deer most likely trace their ancestry to southern Sweden, but interestingly, the two neighboring populations Övedskloster and Wittskövle in the south are significantly different from each other (p<0.01). Furthermore, the population in Wittskövle and roe deer from Köpingsvik at the Baltic Sea Island Öland are significantly different from all other populations (p<0.1). Thus, potentially, there are more than one founder population that contributed to the current Swedish population. The particular differentiation of roe deer from Öland might be an effect of random population genetic processes during the population expansion phase, potentially in combination with introductions of roe deer from other areas than the Swedish mainland. The results are discussed in relation to management of roe deer in Sweden.     

黑龙江省胜山林场冬季驼鹿、马鹿和狍的种间关系

1987—1989年的两个冬季在黑龙江省黑河市胜山林场,对驼鹿、马鹿和狍的食性、采食高度和栖息地进行了研究。驼鹿的主要食物是柳(22.4％)、榛(16.0％)、红松(14.3％)和桦(13.8％);马鹿为杨(36.0％)、桦(22.7％)和红松(17.3％);狍为桦(26.5％)、柳(19.0％)、杨(12.7％)。三种动物采食高度的频次为正态分布。驼鹿的采食高度(标准差)为137.4(±39.0)厘米;马鹿为136.2(±37.0)厘米;狍为84.2(±29.2)厘米。驼鹿和马鹿对红松林的利用程度最高,而狍主要利用沼泽植物和红松林。驼鹿与狍在食物维和栖息地维上的生态位重叠最大,驼鹿与马鹿在采食高度维上的重叠最大。三种动物在食物、采食高度和栖息地三维生态位上的重叠大小的顺序为:驼鹿和马鹿>驼鹿和狍>马鹿和狍。     

Genetic diversity and genetic structure of the Siberian roe deer (<Emphasis Type="Italic">Capreolus pygargus</Emphasis>) populations from Asia

Background

The roe deer, Capreolus sp., is one of the most widespread meso-mammals of Palearctic distribution, and includes two species, the European roe deer, C. capreolus inhabiting mainly Europe, and the Siberian roe deer, C. pygargus, distributed throughout continental Asia. Although there are a number of genetic studies concerning European roe deer, the Siberian roe deer has been studied less, and none of these studies use microsatellite markers. Natural processes have led to genetic structuring in wild populations. To understand how these factors have affected genetic structure and connectivity of Siberian roe deer, we investigated variability at 12 microsatellite loci for Siberian roe deer from ten localities in Asia.

Results

Moderate levels of genetic diversity (H _E = 0.522 to 0.628) were found in all populations except in Jeju Island, South Korea, where the diversity was lowest?(H _E?= 0.386). Western populations showed relatively low genetic diversity and higher degrees of genetic differentiation compared with eastern populations (mean Ar = 3.54 (east), 2.81 (west), mean F _ST = 0.122). Bayesian-based clustering analysis revealed the existence of three genetically distinct groups (clusters) for Siberian roe deer, which comprise of the Southeastern group (Mainland Korea, Russian Far East, Trans-Baikal region and Northern part of Mongolia), Northwestern group (Western Siberia and Ural in Russia) and Jeju Island population. Genetic analyses including AMOVA (F _RT = 0.200), Barrier and PCA also supported genetic differentiation among regions separated primarily by major mountain ridges, suggesting that mountains played a role in the genetic differentiation of Siberian roe deer. On the other hand, genetic evidence also suggests an ongoing migration that may facilitate genetic admixture at the border areas between two groups.

Conclusions

Our results reveal an apparent pattern of genetic differentiation among populations inhabiting Asia, showing moderate levels of genetic diversity with an east-west gradient. The results suggest at least three distinct management units of roe deer in continental Asia, although genetic admixture is evident in some border areas. The insights obtained from this study shed light on management of Siberian roe deer in Asia and may be applied in conservation of local populations of Siberian roe deer.

     

Age estimation of the roe deer (Capreolus capreolus) mandibles from the Mesolithic site of Star Carr, Yorkshire, based on radiographs of mandibular tooth development

A scheme developed by Brown & Chapman, (1991a), which allocates scores to stages of tooth development from radiographs, has been applied to 12 of the 28 roe deer ( Capreolus capreolus ) mandibular rami recovered from the early Mesolithic site of Star Carr, N. Yorkshire ( c. 7500 BC). The mandibles of 35 known-age roe deer kids from Britain and Denmark were also radiographed. The developmental stages of all premolars and molars were analysed and assigned a score using the scheme. After radiographing, four of the Star Carr rami could be paired with a fair degree of confidence, reducing the sample size from 12 to 10 roe deer. Comparisons of scores between the archaeological and modem populations revealed that seven of the Star Carr mandibular rami are aged approximately 10 to 11 months. The youngest is nine months, and the oldest 12 months. These results have significant implications for determining the seasonal presence of humans at the site. Contrary to the received view of a purely late spring/summer occupation of the site, it is now apparent that Star Carr was also visited during the late winter and early spring.     

Landscape connectivity influences gene flow in a roe deer population inhabiting a fragmented landscape: an individual-based approach

Changes in agricultural practices and forest fragmentation can have a dramatic effect on landscape connectivity and the dispersal of animals, potentially reducing gene flow within populations. In this study, we assessed the influence of woodland connectivity on gene flow in a traditionally forest-dwelling species--the European roe deer--in a fragmented landscape. From a sample of 648 roe deer spatially referenced within a study area of 55 x 40 km, interindividual genetic distances were calculated from genotypes at 12 polymorphic microsatellite loci. We calculated two geographical distances between each pair of individuals: the Euclidean distance (straight line) and the 'least cost distance' (the trajectory that maximizes the use of wooded corridors). We tested the correlation between genetic pairwise distances and the two types of geographical pairwise distance using Mantel tests. The correlation was better using the least cost distance, which takes into account the distribution of wooded patches, especially for females (the correlation was stronger but not significant for males). These results suggest that in a fragmented woodland area roe deer dispersal is strongly linked to wooded structures and hence that gene flow within the roe deer population is influenced by the connectivity of the landscape.