Demographic effects of road mortality on mammalian populations: a systematic review.

In light of rapidly expanding road networks worldwide, there is increasing global awareness of the growing amount of mammalian roadkill. However, the ways in which road mortality affects the population dynamics of different species remains largely unclear. We aimed to categorise the demographic parameters in mammalian populations around the world that are directly or indirectly affected by road mortality, as well as identify the most effective study designs for quantifying population-level consequences of road mortality. We conducted a comprehensive systematic review to synthesise literature published between 2000 and 2021 and out of 11,238 unique studies returned, 83 studies were retained comprising 69 mammalian species and 150 populations. A bias towards research-intensive countries and larger mammals was apparent. Although searches were conducted in five languages, all studies meeting the inclusion criteria were in English. Relatively few studies (13.3%) provided relevant demographic context to roadkill figures, hampering understanding of the impacts on population persistence. We categorised five direct demographic parameters affected by road mortality: sex- and age-biased mortality, the percentage of a population killed on roads per year (values up to 50% were reported), the contribution of roadkill to total mortality rates (up to 80%), and roadkill during inter-patch or long-distance movements. Female-biased mortality may be more prevalent than previously recognised and is likely to be critical to population dynamics. Roadkill was the greatest source of mortality for 28% of studied populations and both additive and compensatory mechanisms to roadkill were found to occur, bringing varied challenges to conservation around roads. In addition, intra-specific population differences in demographic effects of road mortality were common. This highlights that the relative importance of road mortality is likely to be context specific as the road configuration and habitat quality surrounding a population can vary. Road ecology studies that collect data on key life parameters, such as age/stage/sex-specific survival and dispersal success, and that use a combination of methods are critical in understanding long-term impacts. Quantifying the demographic impacts of road mortality is an important yet complex consideration for proactive road management.

What makes a house a home? Nest box use by West European hedgehogs (Erinaceus europaeus) is influenced by nest box placement, resource provisioning and site-based factors.

Artificial refuges provided by householders and/or conservation practitioners potentially represent one mechanism for mitigating declines in the availability of natural nest sites used for resting, breeding and hibernating in urban areas. The effectiveness of such refuges for different species is, however, not always known. In this study, we conducted a questionnaire survey of UK householders to identify factors associated with the use of ground-level nest boxes for West European hedgehogs (Erinaceus europaeus), a species of conservation concern. Overall, the percentage of boxes used at least once varied with season and type of use: summer day nesting (35.5-81.3%), breeding (7.2-28.2%), winter day nesting (20.1-66.5%) and hibernation (21.7-58.6%). The length of time the box had been deployed, the availability of artificial food and front garden to back garden access significantly increased the likelihood that a nest box had been used for all four nesting types, whereas other factors related to placement within the garden (e.g., in a sheltered location, on hardstanding such as paving, distance from the house) and resource provisioning (bedding) affected only some nesting behaviours. The factors most strongly associated with nest box use were the provisioning of food and bedding. These data suggest, therefore, that householders can adopt simple practices to increase the likelihood of their nest box being used. However, one significant limitation evident within these data is that, for welfare reasons, householders do not routinely monitor whether their box has been used. Consequently, future studies need to adopt strategies which enable householders to monitor their boxes continuously. Ultimately, such studies should compare the survival rates and reproductive success of hedgehogs within artificial refuges versus more natural nest sites, and whether these are affected by, for example, the impact of nest box design and placement on predation risk and internal microclimate.

An Estimate of the Scale and Composition of the Hedgehog (Erinaceus europeaus) Rehabilitation Community in Britain and the Channel Islands.

The conservation benefits of wildlife rehabilitation are equivocal, but could be substantial for formerly common species that are declining rapidly but are still commonly admitted to wildlife centres. We used a questionnaire survey to estimate the number of practitioners rehabilitating West European hedgehogs (Erinaceus europaeus) in Britain and the numbers entering hospitals/centres in one benchmark year (2016); practitioners were identified using an internet search and snowball sampling. Overall, 304 rehabilitators were identified: 148 supplied data on their structure, and 174 outlined the number of hedgehogs admitted in 2016. The former comprised 62.6% small (≤50 hedgehogs admitted year−1), 16.7% medium-sized (51−250 yr−1), and 20.7% large (>250 yr−1) hospitals; however, these accounted for 4.8%, 12.4%, and 82.8% of hedgehog admissions, respectively. Small hospitals were less likely to be registered as a charity, have paid staff, have a social media account, to record admissions electronically, or to conduct post-release monitoring. However, they were more likely to operate from their home address and to have been established for ≤5 years. Extrapolations indicate that this rehabilitation community admitted >40,000 hedgehogs in 2016, of which approximately 50% could have been released. These figures suggest that wildlife rehabilitation has potentially been an important factor in the dynamics of hedgehog populations in Britain in the last two decades.

An assessment of a conservation strategy to increase garden connectivity for hedgehogs that requires cooperation between immediate neighbours: A barrier too far?

Urban areas are associated with high levels of habitat fragmentation. For some terrestrial species with limited climbing abilities, property boundaries can pose a significant problem by limiting access to residential gardens. The West European hedgehog (Erinaceus europaeus) has declined markedly in the UK but is commonly found in areas of human habitation, including residential gardens. 'Hedgehog Street' is a public engagement campaign aimed at recruiting volunteers ('Hedgehog Champions') to create access points ('hedgehog highways') across garden boundaries to improve habitat connectivity. In this study, we used a series of questionnaire surveys to explore motivations for and obstacles to the creation of highways. Householders were more likely to have created a highway if they were already aware of the Hedgehog Street campaign, if their garden contained a high number of wildlife-friendly features and if they considered watching wildlife to be important. Hedgehog Champions created, on average, 1.69 highways each with 52.0% creating none; this would equate to an estimated >120,000 across all registered Champions. In comparison, 6.1-29.8% of non-Champions stated that they had made a highway. However, most highways had been created in boundaries that could already be traversed via naturally occurring holes: only 11.4% of garden boundaries could be traversed, and 3.2% of gardens accessed, just via a hedgehog highway. In addition, only 5.0% of gardens were considered totally inaccessible to hedgehogs. The most common reasons cited for not having made a highway were that householders' gardens were already accessible to hedgehogs followed by concerns relating to boundary ownership and / or communicating with neighbours. Future studies need to identify strategies for overcoming these obstacles to maximize citizen engagement, particularly with those householders who are not innately "wildlife-friendly", and to quantify the degree to which networks of highways affect patterns of individual movement and, ultimately, populations.

Patterns of Feeding by Householders Affect Activity of Hedgehogs (Erinaceus europaeus) during the Hibernation Period.

West European hedgehogs (Erinaceus europaeus) are likely to encounter unusual ecological features in urban habitats, such as anthropogenic food sources and artificial refugia. Quantifying how these affect hedgehog behaviour is vital for informing conservation guidelines for householders. We monitored hedgehog presence/absence in gardens in the town of Reading, UK, over the winter of 2017-2018 using a volunteer-based footprint tunnel survey, and collected data on garden characteristics, supplementary feeding (SF) habits, and local environmental conditions. Over a 20-week survey period, hedgehog presence was lowest between January and March. Occupancy analysis indicated that SF significantly affected hedgehog presence/absence before, during, and after hibernation. The number of nesting opportunities available in gardens, average temperatures, and daylength were also supported as important factors at different stages. In particular, our results suggest that SF could act to increase levels of activity during the winter when hedgehogs should be hibernating. Stimulating increased activity at this sensitive time could push hedgehogs into a net energy deficit or, conversely, help some individuals survive which might not otherwise do so. Therefore, further research is necessary to determine whether patterns of feeding by householders have a positive or negative effect on hedgehog populations during the hibernation period.

Over-Winter Survival and Nest Site Selection of the West-European Hedgehog (Erinaceus europaeus) in Arable Dominated Landscapes.

The West-European hedgehog (Erinaceus europaeus) has declined markedly in the UK. The winter hibernation period may make hedgehogs vulnerable to anthropogenic habitat and climate changes. Therefore, we studied two contrasting populations in England to examine patterns of winter nest use, body mass changes and survival during hibernation. No between-site differences were evident in body mass prior to hibernation nor the number of winter nests used, but significant differences in overwinter mass change and survival were observed. Mass change did not, however, affect survival rates; all deaths occurred prior to or after the hibernation period, mainly from predation or vehicle collisions. Hedgehogs consistently nested in proximity to hedgerows, roads and woodlands, but avoided pasture fields; differences between sites were evident for the selection for or avoidance of arable fields, amenity grassland and buildings. Collectively, these data indicate that hibernation was not a period of significant mortality for individuals that had attained sufficient weight (>600 g) pre-hibernation. Conversely, habitat composition did significantly affect the positioning of winter nests, such that different land management practices (historic and current) might potentially influence hibernation success. The limitations of this study and suggestions for future research are discussed.

Impacts and Potential Mitigation of Road Mortality for Hedgehogs in Europe.

Transport infrastructure is a pervasive element in modern landscapes and continues to expand to meet the demands of a growing human population and its associated resource consumption. Road-induced mortality is often thought to be a major contributor to the marked declines of European hedgehog populations. This review synthesizes available evidence on the population-level impacts of road mortality and the threat to population viability for the five hedgehog species in Europe. Local and national studies suggest that road mortality can cause significant depletions in population sizes, predominantly removing adult males. Traffic collisions are a probable cause of fragmentation effects, subsequently undermining ecological processes such as dispersal, as well as the genetic variance and fitness of isolated populations. Further studies are necessary to improve population estimates and explicitly examine the consequences of sex- and age-specific mortality rates. Hedgehogs have been reported to use crossing structures, such as road tunnels, yet evaluations of mitigation measures for population survival probability are largely absent. This highlights the need for robust studies that consider population dynamics and genetics in response to mitigation. In light of ongoing declines of hedgehog populations, it is paramount that applied research is prioritised and integrated into a holistic spatial planning process.

Environmental variation at the onset of independent foraging affects full-grown body mass in the red fox.

The period following the withdrawal of parental care has been highlighted as a key developmental period for juveniles. One reason for this is that juveniles cannot forage as competently as adults, potentially placing them at greater risk from environmentally-induced changes in food availability. However, no study has examined this topic. Using a long-term dataset on red foxes (Vulpes vulpes), we examined (i) dietary changes that occurred in the one-month period following the attainment of nutritional independence, (ii) diet composition in relation to climatic variation, and (iii) the effect of climatic variation on subsequent full-grown mass. Diet at nutritional independence contained increased quantities of easy-to-catch food items (earthworms and insects) when compared with pre-independence. Interannual variation in the volume of rainfall at nutritional independence was positively correlated to the proportion of earthworms in cub diet. Pre-independence cub mass and rainfall immediately following nutritional independence explained a significant proportion of variance in full-grown mass, with environmental variation affecting full-grown mass of the entire cohorts. Thus, weather-mediated availability of easy-to-catch food items at a key developmental stage has lifelong implications for the development of juvenile foxes by affecting full-grown mass, which in turn appears to be an important component of individual reproductive potential.

Reduced occupancy of hedgehogs (Erinaceus europaeus) in rural England and Wales: The influence of habitat and an asymmetric intra-guild predator.

Agricultural landscapes have become increasingly intensively managed resulting in population declines across a broad range of taxa, including insectivores such as the hedgehog (Erinaceus europaeus). Hedgehog declines have also been attributed to an increase in the abundance of badgers (Meles meles), an intra-guild predator. The status of hedgehogs across the rural landscape at large spatial scales is, however, unknown. In this study, we used footprint tracking tunnels to conduct the first national survey of rural hedgehog populations in England and Wales. Single and two-species occupancy modelling was used to quantify hedgehog occupancy in relation to habitat and predator covariates. Hedgehog occupancy was low (22% nationally), and significantly negatively related to badger sett density and positively related to the built environment. Hedgehogs were also absent from 71% of sites that had no badger setts, indicating that large areas of the rural landscape are not occupied by hedgehogs. Our results provide the first field based national survey of hedgehogs, providing a robust baseline for future monitoring. Furthermore, the combined effects of increasing badger abundance and intensive agriculture may have provided a perfect storm for hedgehogs in rural Britain, leading to worryingly low levels of occupancy over large spatial scales.

Changes in the distribution of red foxes (Vulpes vulpes) in urban areas in Great Britain: findings and limitations of a media-driven nationwide survey.

Urbanization is one of the major forms of habitat alteration occurring at the present time. Although this is typically deleterious to biodiversity, some species flourish within these human-modified landscapes, potentially leading to negative and/or positive interactions between people and wildlife. Hence, up-to-date assessment of urban wildlife populations is important for developing appropriate management strategies. Surveying urban wildlife is limited by land partition and private ownership, rendering many common survey techniques difficult. Garnering public involvement is one solution, but this method is constrained by the inherent biases of non-standardised survey effort associated with voluntary participation. We used a television-led media approach to solicit national participation in an online sightings survey to investigate changes in the distribution of urban foxes in Great Britain and to explore relationships between urban features and fox occurrence and sightings density. Our results show that media-based approaches can generate a large national database on the current distribution of a recognisable species. Fox distribution in England and Wales has changed markedly within the last 25 years, with sightings submitted from 91% of urban areas previously predicted to support few or no foxes. Data were highly skewed with 90% of urban areas having <30 fox sightings per 1000 people km(-2). The extent of total urban area was the only variable with a significant impact on both fox occurrence and sightings density in urban areas; longitude and percentage of public green urban space were respectively, significantly positively and negatively associated with sightings density only. Latitude, and distance to nearest neighbouring conurbation had no impact on either occurrence or sightings density. Given the limitations associated with this method, further investigations are needed to determine the association between sightings density and actual fox density, and variability of fox density within and between urban areas in Britain.

Spatio-temporal variation in predation by urban domestic cats (Felis catus) and the acceptability of possible management actions in the UK.

Urban domestic cat (Felis catus) populations can attain exceedingly high densities and are not limited by natural prey availability. This has generated concerns that they may negatively affect prey populations, leading to calls for management. We enlisted cat-owners to record prey returned home to estimate patterns of predation by free-roaming pets in different localities within the town of Reading, UK and questionnaire surveys were used to quantify attitudes to different possible management strategies. Prey return rates were highly variable: only 20% of cats returned ≥4 dead prey annually. Consequently, approximately 65% of owners received no prey in a given season, but this declined to 22% after eight seasons. The estimated mean predation rate was 18.3 prey cat⁻¹ year⁻¹ but this varied markedly both spatially and temporally: per capita predation rates declined with increasing cat density. Comparisons with estimates of the density of six common bird prey species indicated that cats killed numbers equivalent to adult density on c. 39% of occasions. Population modeling studies suggest that such predation rates could significantly reduce the size of local bird populations for common urban species. Conversely, most urban residents did not consider cat predation to be a significant problem. Collar-mounted anti-predation devices were the only management action acceptable to the majority of urban residents (65%), but were less acceptable to cat-owners because of perceived risks to their pets; only 24% of cats were fitted with such devices. Overall, cat predation did appear to be of sufficient magnitude to affect some prey populations, although further investigation of some key aspects of cat predation is warranted. Management of the predation behavior of urban cat populations in the UK is likely to be challenging and achieving this would require considerable engagement with cat owners.

Accumulation of anticoagulant rodenticides in a non-target insectivore, the European hedgehog (Erinaceus europaeus).

Studies on exposure of non-targets to anticoagulant rodenticides have largely focussed on predatory birds and mammals; insectivores have rarely been studied. We investigated the exposure of 120 European hedgehogs (Erinaceus europaeus) from throughout Britain to first- and second-generation anticoagulant rodenticides (FGARs and SGARs) using high performance liquid chromatography coupled with fluorescence detection (HPLC) and liquid-chromatography mass spectrometry (LCMS). The proportion of hedgehogs with liver SGAR concentrations detected by HPLC was 3-13% per compound, 23% overall. LCMS identified much higher prevalence for difenacoum and bromadiolone, mainly because of greater ability to detect low-level contamination. The overall proportion of hedgehogs with LCMS-detected residues was 57.5% (SGARs alone) and 66.7% (FGARs and SGARs combined); 27 (22.5%) hedgehogs contained >1 rodenticide. Exposure of insectivores and predators to anticoagulant rodenticides appears to be similar. The greater sensitivity of LCMS suggests that hitherto exposure of non-targets is likely to have been under-estimated using HPLC techniques.

Effect of British hunting ban on fox numbers.

Pressure to ban the hunting of foxes with hounds in Britain has fuelled debate about its contribution to the control of fox populations. We took advantage of a nationwide one-year ban on fox-hunting during the outbreak of foot-and-mouth disease (FMD) in 2001 to examine this issue and found that the ban had no measurable impact on fox numbers in randomly selected areas. Our results argue against suggestions that fox populations would increase markedly in the event of a permanent ban on hunting.

Modelling the costs of fox predation and preventive measures on sheep farms in Britain.

Economic analysis is a useful tool to aid decisions on what to do about wildlife impacts, such as those of vertebrate predators on livestock farmers. The case-study of lamb predation by foxes in Britain is used to develop a theoretical economic model, with the aim of determining a financially optimal solution to minimise the total costs of livestock predation at the farm-level. Total costs include output losses and expenditure on preventive and control measures, in this case indoor housing and lethal fox control. The model is tested empirically with data from a questionnaire survey of sheep farmers and field data on fox population densities in Britain. Regression analyses are used to determine the relationships between lamb losses and expenditure on indoor housing, fox population density and other non-management characteristics. The effect of fox abundance on the cost of fox control is also assessed. Marginal analysis is used to determine the total cost-minimising solution from the farmer's point-of-view, in terms of how many ewes should be housed indoors and for how long, as well as how many foxes should be killed in addition to any lethal control already carried out. Optimal solutions vary according to farm characteristics, including flock size and the regional location of farms. In all cases, to minimise the costs of predation, as many ewes as possible should be housed. However, it is not worthwhile housing them for more than a day after lambing. Efficient fox predation management does not necessarily mean that lamb losses should be reduced to zero, and additional fox control is not worthwhile on the majority of farms. The analysis provides a framework for future evaluations of wildlife impacts and cost-effective management of these problems.

The current and future management of wild mammals hunted with dogs in England and Wales.

There is increasing concern about the use of lethal methods to control wild mammal populations, especially those methods that may have significant impacts on animal welfare. The continued use of dogs to hunt wild mammals in England and Wales, principally foxes (Vulpes vulpes), red deer (Cervus elaphus), brown hares (Lepus europaeus) and mink (Mustela vison), has become a focus for political debate and has been the subject of a recent UK government inquiry. This paper reports the results of a questionnaire study to quantify the use, effectiveness and acceptability of the different methods currently used to manage these four species, and future changes in management following a possible ban on hunting with dogs. There was no straightforward relationship between culling pressure and perceived pest status of the different species from the questionnaire data. For foxes and brown hares, the proportion of land managers (practitioners) carrying out lethal control was higher than that considering these species to be pests. However, the reverse was the case for mink. The most frequently used and effective control methods, which were most acceptable to practitioners and public alike, were various forms of shooting. The general public perceived hunting with dogs as one of the least acceptable means of control for all four species. Practitioners thought that hunting with dogs for red deer and the use of terriers against foxes were among the least acceptable forms of control, but considered hunting with dogs in other situations and for other species to be relatively acceptable. Most practitioners said a ban on hunting with dogs would make no difference to their management of the four species. A ban on hunting with dogs would have minimal impact on populations of foxes, red deer and mink, but it may be of conservation benefit to hares.