download Water Vole Conservation Handbook (): NHBS - Rob Strachan, Tom Moorhouse and Merryl Gelling, Wild Cru. CONSERVATION HANDBOOK As with many mammals, a direct sighting of the water vole is Water vole burrow entrances are typically wider than high. Epub 09/12/ doi: doi/bestthing.info Strachan R, Moorhouse T, Gelling M. Water Vole Conservation Handbook, Third Edition. third edition.
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Scotland or in Scottish waters. 10 Restoration is increasingly accepted as an essential part of conservation. Local reintroductions of water voles carried out over. Mar 24, Water voles were housed in laboratory cages containing between one .. Watervole Conservation Handbook: Wildlife Conservation Research. The ePub format uses eBook readers, which have several "ease of reading" features already built in. Water voles had been extinct in the Monnow Catchment since the s. for American mink control has been conservation of the water vole (Arvicola amphibius; Strachan R. Water vole conservation handbook.
Thus, while there is a long-term decline in dormouse populations in the UK precipitating a national monitoring programme, habitat restoration efforts and population reintroductions Bright et al.
Our objectives were to 1 provide phylogeographic coverage of common dormouse populations in the UK relative to its continental European range; 2 assess the geographical and temporal patterns of genetic variation within the UK, and between the UK and continental European populations, and; 3 assess competing post-glacial expansion hypotheses EMH vs.
LMH by estimating the timing of dispersal in this species. Materials and Methods Sampling Collection Non-invasive genetic sampling of hair was conducted during summer nest-box surveys in and This sample size is commensurate with current standards for biogeographical genetic studies with the goal of estimating long-term divergence of lineages see Gillespie, ; Mouton et al. Study sites were chosen to represent the current natural range of the common dormouse in the UK.
Three of our study sites were from reintroduced populations within the UK Table 1 as part of the dormouse reintroduction programme, to enable us to quantify genetic differentiation, if any, between these and natural populations.
We analyzed UK data we produced along with previously published sequences available on GenBank from continental Europe Mouton et al. Geographic location of sampling sites.
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Geographical distribution of the common dormouse, Muscardinus avellinarius, and samples collected around the UK from 25 sites samples and haplotype distribution for concatenated sequences in Figure 2B. Key indicates the genetic grouping of populations and three reintroduced populations.
DNA degradation was an issue affecting amplification using these primers. DNA sequencing was then performed using BigDye v3. The robustness of the trees was assessed by bootstrap replications Felsenstein, Bandelt et al. The net distance between groups and average distances within groups were calculated using MEGA 6.
This also allowed us to estimate the time of divergence for other genetic clades found in continental Europe Central Northern Europe, Turkey, Balkans and Western Europe, Belgium. The UK clade was given a normal distribution truncated at lower and upper limits of 7.
The analyses were repeated without the prior on the UK clade to test the effect of the priors on the posterior distributions. Multimodal distributions are considered to correspond to a condition of demographic stability or multiple colonization, whereas recent sudden population expansions would be observed by unimodal patterns Slatkin and Hudson, Results Phylogenetic Analysis and Genetic Diversity A total of nucleotide positions of the target Cytb were resolved in all individuals from around the UK.
We found a total of five haplotypes, four of which are unique to known European haplotypes all sequences have been deposited in GenBank; Table 1 , Figure 2A , and all positions were parsimony informative. Haplotype diversity we found was similar to that reported in continental Europe UK 0. Haplotype map minimum spanning network. Numbers along the branches correspond to the number of mutational steps observed between haplotypes, no number is one mutational step, size of the circles correspond to the number of populations for each haplotype.
A Number for UK haplotypes given in the circles correspond to Table 1. B D-loop haplotypes are colored as found in key. In total, UK dormice show a 0.
The Cumbria sample the most northerly extant population in England forms a cluster with this central haplotype group. Haplotype 39 Table 1 was found in 11 populations and is the most frequent group sampled in this study that also form a grouping with CNE clade Figure 2A.
Maximum-likelihood ML topology for concatenated sequences Cytb, D-loop, and bfibr. The numbers on the branches indicate the bootstrap values. A bp fragment of D-loop was sequenced for individuals Table 1. Haplotype diversity 0. The bfibr bp nuclear marker was sequenced for all individuals around the UK, Lithuania and France no other sequences are available on GenBank for common dormice. This gene was found to be monomorphic across all populations in the UK, however there was divergence between the UK and continental European populations.
As such bfibr and D-loop sequences were only further analyzed as concatenated sequences along with Cytb. Concatenated sequence analysis bp showed a total of 7 unique haplotypes, and had higher resolution to reveal regional genetic variation than that of Cytb alone Figure 2B.
For Cytb, only one mutation is observed between sequences as seen in both the ML tree and haplotype network Figures 2 , 3 , whilst for D-loop there are four mutational steps present Figure 2B. Samples sequenced from Suffolk in East England show a close genetic grouping with each other, especially in the D-loop haplotype network Figure 2B. Two populations sampled in Suffolk were part of a national re-introduction programme and clustered separately from natural populations in Suffolk by a single mutation.
Based on our Cytb analysis, these reintroduced populations group with the South East and North England clade Figure 2A , likely due to the genetic source of these populations from Southern England. Based on our D-loop analysis, one reintroduced Suffolk population Bradfield falls into the CE clade and the second reintroduced population groups with another Suffolk population Bonny; Table 1 , Figure 2B.
The last population is geographically close less than 1 km with available connective habitat to the population in which it forms a haplotype connection with see Figure 2B , but to assess whether there is evidence of gene flow or whether this is consistent with ancestral polymorphism, additional data and analysis would be required.
Test runs with alternative prior distributions did not influence posterior estimates of this parameter. Estimates also place a Balkan and Turkey clade divergence at Mismatch distribution analysis Figure 5 showed a unimodal distribution for the UK and CNE indicating a recent, rapid population expansion, which is consistent with an expansion at the end of the LGM when a land bridge facilitated dispersal of dormice to the UK.
Maximum clade credibility chronogram of the Cytb dataset. Numbers below branches are Bayesian posterior probabilities BPPs. Blue highlights period when UK was isolated from continental Europe by rising sea levels.
The bottom bar indicates the approximate timings of the two colonization hypotheses early migration hypothesis EMH and late migration hypothesis LMH.
Discussion Genetic Structure between UK and Continental Europe Here, we present the first evidence of genetic divergence between UK common dormouse populations and those in continental Europe. Our sequence analysis of Cytb, in the context of published data from continental Europe Mouton et al. Further, it is shown that these UK lineages are most closely related to populations located in CNE Figure 2 and the within-population variation detected here is similar to that identified within regional subgroups in continental Europe by Mouton et al.
The nuclear gene bfibr, although monomorphic in the UK, shows genetic subdivision from continental Europe with a similar mutational difference to that of Cytb and D-loop studied here. Comparisons of genetic diversity for mitochondrial genes observed in the UK are comparable to that found in the genetic clades of continental Europe which may be a result of recent genetic divergence in the order of magnitude in thousands of years.
Because the entire mitochondrial genome is inherited as a unit, sequencing more than one mitochondrial gene, as we have done, whilst offering more resolution on the history of the maternal lineage, still represents a single lineage and thus has some inherent limitations.
For this reason the addition of a nuclear gene in our analysis helps to confirm the identification of divergence between continental Europe and the UK.
While there is evidence for reciprocal monophyly between continental European and UK dormice, further study of adaptive genetic variation in UK dormice is perhaps needed to inform management of the UK common dormouse as distinct. Genetic Structure within the UK We describe the genetic structure of the common dormouse within the UK for the first time.
Here, we present evidence of regional genetic clustering of populations around the UK based on mtDNA variation clustering Figure 3. This regional variation is possibly explained by gross geographical features; the UK has several major rivers and uplands in the North, West, and South of the country which may be important geographical boundaries leading to the further genetic clustering seen in this study.
Although, the nuclear gene bfibr was monomorphic in our samples, the allelic sequence variant we report is unique to the UK. Mitochondrial DNA evolves at a much faster rate 5—10 fold and as such mtDNA is more suitable for resolving contemporary events and defining evolutionary significant units Wan et al.
While the extent of regional genetic variation we describe is modest, it suggests the possibility of local adaptation and genetic differentiation which warrants further study at a finer geographical scale.
The phylogeographic structure we describe contrasts with that of previous small mammal studies. This is possibly explained by a two-stage colonization where the initial colonists were largely replaced by the second wave, leaving peripheral populations in Northern and Western areas of the UK.
A pattern of single colonization was observed in dormice.
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This is possibly explained by the relatively low dispersal capability of the common dormouse. Haplotype network analysis shows a clustering between the Isle of Wight and the reintroduced Wych population in North England. Based on the geographical distance between the Isle of Wight and Wych over km , the relatively close genetic relatedness of these populations is surprising.
The Isle of Wight island population is relatively isolated off the south coast of England. However, the Wych population was reintroduced in , with founding individuals originating from the Isle of White pers. Only few studies have investigated the impacts of conservation activities on wildlife health and welfare 8 — Difficulties or the inability to cope with environmental pressures can lead to stress and hence potential negative impacts on animal health and well-being as well as decreased resilience 14 — Moberg 17 proposed that determining to which extent an animal is impacted from stress due to changes in its biological functions, thereby entering a pre-pathological state, is the only defensible measurement of well-being in animals 17 , Accordingly, the definition of potential stressors and the further development of methods to measure and assess stress responses are crucial for the evaluation of wildlife welfare 19 — More recently the allostasis concept was extended within the reactive scope model, which integrates the importance of species developmental strategies and their potential long-lasting impact in priming and programming later life stress responses Beyond the mere definition of stress, which due to the complexity and multi-dimensionality of the phenomenon may be hard to frame, the main physiological systems for coping with stressors are relatively well-studied.
There are two major mediators orchestrating the stress response in vertebrates: i catecholamine's controlled by the sympathetic nervous system SNS and ii glucocorticoid stress hormones [GCs; corticosterone in amphibians, reptiles and birds, cortisol in most fish and mammals— 27 ] modulated by the Hypothalamic-Pituitary-Adrenal axis HPA-axis. The HPA axis response is slower within minutes and acts on various physiological pathways to adjust essential bio-regulatory mechanisms in response to stressors, such as extreme weather conditions, predator exposure, or shortages of food 15 , This is primarily achieved by up-regulating key body functions, including cardiac-, respiratory- and brain-activity as well as energy mobilization at the expense of other processes such as growth, reproduction, immunity or the balance between oxygen radicals and the antioxidant system 29 — However, nature, duration and magnitude of stressful events are likely to be fundamental in determining the biological benefits or costs of exposure to stress 29 , There is growing evidence suggesting that the long-term and repeated exposure to moderately challenging stressors is associated with positive, rather than negative, organismal outcomes, improving survival and delaying the onset of reproductive senescence 34 , It is therefore key to assess and quantify how and to which extent differing stressors such as those provoked during wildlife and conservation management activities i.
Measuring Stress Stress responses vary vastly among species as well as within individuals of the same species 28 , 38 — 40 , are modulated by season, time of day 41 and can be triggered by a great variety of stressors Moreover, stress responses involve several physiological processes in parallel and are therefore difficult to measure and to assess, particularly with the small sample sizes typical in field studies of wildlife species Currently physiological stress responses in wildlife are assessed with a variety of techniques 20 including measuring GCs in various tissues 44 — 46 , changes in blood chemistry and hematology 47 and behavioral alterations, such as exploratory or avoidance behaviors Measuring GCs has generally been adopted as a standard procedure to estimate individual stress levels.
However an elevation of GCs does not necessarily always indicate a state of stress or discomfort, as baseline and stress GCs levels can fluctuate hugely among an individual's life history stages 49 , Therefore, the use of GCs as a single metric to gain a comprehensive understanding of individual stress conditions is limited While there has been an over-reliance upon GCs, other pathways of the stress response, such as endocrine-immune interactions as proxies for stress and animal welfare, are surprisingly understudied.
In order to better understand the causalities and complex mechanisms within the stress response and its implications for animal welfare, it is imperative to integrate different approaches to better assess and interpret the phenomenon of stress 43 , Immune Markers as a Potential Proxy for Stress and Animal Welfare Several studies provide solid evidence for the strong and reciprocal interaction between immune processes and stress 52 — It is now widely accepted that the immune system and the neuroendocrine system form an integrated and evolutionary highly conserved element of physiology across phyla 55 , Therefore, direct and indirect stress-induced effects on quantitative and functional immune parameters can serve as additional markers to assess stress and wildlife welfare.
The best established and most commonly used immune parameter applied across all five vertebrate taxa is the stress-related change in immune cell distribution i. Higher stress levels are associated with an increase of neutrophil granulocytes heterophils in bird and reptile species and a decreae of lymphocytes in the bloodstream and hence an increase in neutrophil to lymphocyte ratio [N:L; 57 ] [for review see 47 , 58 ].
Another interesting immunological marker for stress is neopterin, a pteridine derivate synthetized by monocytes and macrophages upon inflammatory cytokine stimulation.
Another promising immune marker appears to be Immunoglobulin A IgA and in particular its secretory form SIgA the major antibody of mucosal immune defense in mamals and birds.
The review by Staley et al. In contrast, situations with good or enhanced welfare, lead to increased SIgA levels suggesting that this marker can be a suitable immunological proxy for animal welfare.
An example emphasizing the biological significance of this innate immune reaction is chronic granulomatous disease CGD , an inherited immunodeficiency in humans, where PMNLs are not able to generate ROS upon stimulation. CGD and an insufficient oxidative burst response in general, are characterized by recurrent bacterial and fungal infections and a set of inflammatory complications with not uncommonly, lethal outcome 67 , In wildlife the initial stress-induced oxidative burst of PMNLs acts as immediate protection against invading pathogens in the case of injury by a predator 69 , However, the capability of PMNLs to produce further ROS after the initial stress induced burst is curtailed to protect the organism from over-activation of PMNLs while reducing free radical damage of surrounding tissues 71 , On the other hand, if stress conditions persist, this innate immune response is diminished to depleted with detrimental impacts for the health, welfare and survival of the individual 70 , 73 — McLaren et al.
PMNLs have over different receptors which are sensitive to varied stress signals in the organism, including plasma endocrine factors, changes in blood biochemistry and red cell hemodynamics, changes of cytokine levels and mediators released by the HPA axis and the SNS This synchronous sensitivity to several stress mediators and an array of stress- related physiological changes emphasizes PMNLs as excellent indicators in evaluating stress levels The technique relies on the observation that PMNLs of stressed individuals have a reduced capacity to produce ROS in response to a secondary chemical external stimulus Thus, low LCC levels in an individual indicate a decreased innate immune response and increased stress levels.
Despite the sensitivity of PMNLs to an array of constituent mediators of the stress response, the physiological relevance of the method is promising for the following reasons: i PMNLs remain in their natural environment, i. The method provides several additional technical advantages: i a relatively small amount of blood i.
There are several aspects which require further experimental testing to establish the diagnostic efficacy of the methodology. It should be noted that studies investigating the relationship of LCC to more commonly used proxies for stress e. An additional explanation may be the synchronous sensitivity of PMNLs to several stress related changes During infection and disease a multitude of immunological factors are altered.
Gonadal steroids e.
Future studies will need to assess stress hormone, and gonadal steroid effects on the LCC response in order to better elucidate functional endocrine-immune interactions that could be linked with animal welfare. We also need further studies to elucidate the down-stream mechanisms triggering PMNL activation and relevant time windows in which these pathways do operate.
However, there are no clear physiological profiles of ensured welfare within a species or even between individuals. Hence future studies should aim for a systematic, multivariate approach including several parameters of physiological and behavioral nature to gain more insight toward the validity of potential tools such as LCC to assess stress and welfare 89 — Capture and handling of wildlife species often involve anesthesia of individuals with varying protocols which are constantly adapted for animal safety and welfare reasons 93 , Anesthetic agents have the potential to decrease PMNL oxidative burst capacity in humans.
This decrease has been shown for opioids morphine , thiopental, propofol, midazolam, volatile anesthetics i. However, despite some studies in humans [e. Studies Inferring LCC as a Valid Proxy to Assess Stress in the Context of Welfare A review on phagocyte photon emission in response to stress and disease noted that the capability of PMNLs to emit ROS reflects the pathophysiological state of the host and that the magnitudes of stress as well as the presence of pathogens and disease processes can be estimated A later study in Atlantic salmon Salmo salar revealed that fish subjected to a 2 h period of confinement stress had a reduction in oxygen free radical production in isolated PMNLs and therefore a lower oxidative burst capacity and a debilitated innate immune response In Table 1 , we review a sample of studies inferring LCC as a valid proxy to assess stress and animal welfare.
The study showed that transported individuals ca. These data indicate that transport is likely to be a compounding stressor beyond the capture event A study on bank voles Clethrionomys glareolus and wood mice Apodemus sylvaticus indicates that handling per se is likely to alter LCC responses.
Handled animals only for 20 s showed remarkable reduction in LCC in comparison to non-handled animals In non-anesthetized European Roe deer Capreolus capreolus LCC levels were negatively impacted by the time of human presence at the capture site prior to the actual handling procedure, suggesting that human presence at the trapping site prior to handling should be minimized The LCC technique was used to investigate the stress response caused by capture and subsequent abdominal surgery of free-ranging brown bears Ursus arctos and to evaluate whether variation in LCC co-varied with other proxies of metabolic and physiological stress, such as heart rate, N:L—ratio, blood glucose and circulating cortisol concentrations Their main result revealed that LCC values following capture were lower in solitary bears when compared to females with cubs and lower in bears in poorer body condition when compared to those in good body condition.
LCC levels did not seem to be influenced by the actual surgical procedure under anesthesia A recent study comparing blood glucocorticoid levels, hematology, LCC, scrotal, and perineal temperature, scrotal lesion, and a pain score in two groups of male calves Bos Taurus , a ring castration and a sham castration control group, suggests LCC as an innovative tool for stress and pain assessment Overview of studies inferring LCC as a valid proxy to assess stress and welfare in animals.
Within a reintroduction program for conservation purposes Moorhouse et al. The authors found a larger decrease in LCC levels between week 1 and 2 for individuals that were radio-collared while this was not the case in non-collared individuals, suggesting that radio-collaring could be an additional stressor, at least in this species. In this experiment one group of individuals were housed in outdoor enclosures and the other group in indoor laboratory cages. LCC values of both groups decreased constantly over the 6-week study period, but interestingly, animals housed indoors and individually in laboratory cages showed lower LCC values despite the fact that they usually do not live in large groups and are territorial in the wild This result partially contrasts results from , who examined short-term social stress by means of body weight change and LCC to test the effects of group size in captive-bred water voles destined for release within a reintroduction program.
LCC scores were negatively correlated with group size, suggesting that individuals held in larger groups experienced higher relative levels of stress and therefore showed a greater decline in LCC As such there is consensus that species dispersed into Northern regions and the UK from 23 kya onwards as glacial ice caps retreated and tundra steppes were no longer frozen.
Effects of cage area were accounted for by including them as a separate variable in the analysis see Data analysis, below. Accordingly, the definition of potential stressors and the further development of methods to measure and assess stress responses are crucial for the evaluation of wildlife welfare 19 — The sampling programme was carefully arranged to ensure that equal sex ratios were sampled on any given day and that the distribution of sampling effort was equally distributed between cages of different sizes containing different numbers to exclude the possibility of temporal bias.
While there has been an over-reliance upon GCs, other pathways of the stress response, such as endocrine-immune interactions as proxies for stress and animal welfare, are surprisingly understudied.