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.

Prey availability and intraguild competition regulate the spatiotemporal dynamics of a modified large carnivore guild.

Effective conservation management requires an understanding of the spatiotemporal dynamics driving large carnivore density and resource partitioning. In African ecosystems, reduced prey populations and the loss of competing guild members, most notably lion (Panthera leo), are expected to increase the levels of competition between remaining carnivores. Consequently, intraguild relationships can be altered, potentially increasing the risk of further population decline. Kasungu National Park (KNP), Malawi, is an example of a conservation area that has experienced large-scale reductions in both carnivore and prey populations, leaving a resident large carnivore guild consisting of only leopard (Panthera pardus) and spotted hyena (Crocuta crocuta). Here, we quantify the spatiotemporal dynamics of these two species and their degree of association, using a combination of co-detection modeling, time-to-event analyses, and temporal activity patterns from camera trap data. The detection of leopard and spotted hyena was significantly associated with the detection of preferred prey and competing carnivores, increasing the likelihood of species interaction. Temporal analyses revealed sex-specific differences in temporal activity, with female leopard activity patterns significantly different to those of spotted hyena and male conspecifics. Heightened risk of interaction with interspecific competitors and male conspecifics may have resulted in female leopards adopting temporal avoidance strategies to facilitate coexistence. Female leopard behavioral adaptations increased overall activity levels and diurnal activity rates, with potential consequences for overall fitness and exposure to sources of mortality. As both species are currently found at low densities in KNP, increased risk of competitive interactions, which infer a reduction in fitness, could have significant implications for large carnivore demographics. The protection of remaining prey populations is necessary to mitigate interspecific competition and avoid further alterations to the large carnivore guild.

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.

Beneficial Land Management for Hedgehogs (Erinaceus europaeus) in the United Kingdom.

Hedgehogs (Erinaceus europaeus) are traditionally thought of as being a rural dwelling species, associated with rural and agricultural landscapes across Europe. However, recent studies have highlighted that hedgehogs are more likely to be found in urban than rural habitats in the United Kingdom. Here, we review the status of rural hedgehog populations across the UK and evaluate the potential benefits of agri-environment schemes for hedgehog persistence, while highlighting a lack of empirical evidence that agri-environment options will benefit hedgehog populations. Our synthesis has implications for future conservation strategies for hedgehogs and insectivorous mammals living in agricultural landscapes, and calls for more empirical studies on agri-environment options and their potential benefits to hedgehogs.

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.

Vehicular tracks and the influence of land use and habitat protection in the British uplands.

Uplands cover around 25% of the Earth's land surface and comprise highly sensitive soils and habitats of global ecological importance that together provision a suite of valuable ecosystem services. Access to upland areas for land management and recreation is increasingly dependent on the use of motorised vehicles. However, despite the widely acknowledged detrimental consequences of vehicle track development and use, this activity is not currently quantified or monitored. In this study, surfaced and unsurfaced vehicular tracks, footpaths and land cover in an approximately 5% sample of mainland British uplands (1910 km2) were mapped using aerial imagery dating between 2007 and 2016. An information theory approach was used to identify models that best predicted the presence and extent of surfaced tracks as a function of land cover, protected status, extent of blanket peat, proximity to human population and altitude. A total of 2104 km of track were recorded, with 27% defined as being surfaced. This equates to a mean (±SE) track density of 1.10 ± 0.15 km km-2 in current use by vehicles across upland Britain. Areas of managed heather and grass-dominated habitat were the primary predictors of the presence and extent of surfaced tracks and we found no evidence that areas of conservation interest with protective designation were less susceptible to track construction. These data indicate that wide-ranging vehicular track networks exist in ecologically sensitive environments across the British uplands, with implications for the integrity of sensitive habitats and soils. Land use and management activities that are not facilitating ecological improvement or conservation appear to be driving the presence of surfaced tracks in 'protected' areas. The findings suggest that: a) urgent review and revision of upland track legislation is required in Great Britain; and b) wider assessment of upland tracks should be undertaken to inform global conservation strategies for these environments.

Human disturbance has contrasting effects on niche partitioning within carnivore communities.

Among species, coexistence is driven partly by the partitioning of available resources. The mechanisms of coexistence and competition among species have been a central topic within community ecology, with particular focus on mammalian carnivore community research. However, despite growing concern regarding the impact of humans on the behaviour of species, very little is known about the effect of humans on species interactions. The aim of this review is to establish a comprehensive framework for the impacts of human disturbance on three dimensions (spatial, temporal and trophic) of niche partitioning within carnivore communities and subsequent effects on both intraguild competition and community structure. We conducted a systematic review of the literature on carnivore niche partitioning (246 studies) and extracted 46 reported effects of human disturbance. We found evidence that human disturbance impacts resource partitioning, either positively or negatively, in all three niche dimensions. The repercussions of such variations are highly heterogeneous and differ according to both the type of human disturbance and how the landscape and/or availability of resources are affected. We propose a theoretical framework of the three main outcomes for the impacts of human disturbance on intraguild competition and carnivore community structure: (i) human disturbance impedes niche partitioning, increasing intraguild competition and reducing the richness and diversity of the community; (ii) human disturbance unbalances niche partitioning and intraguild competition, affecting community stability; and (iii) human disturbance facilitates niche partitioning, decreasing intraguild competition and enriching the community. We call for better integration of the impact of humans on carnivore communities in future research on interspecific competition.

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.

A citizen science based survey method for estimating the density of urban carnivores.

Globally there are many examples of synanthropic carnivores exploiting growth in urbanisation. As carnivores can come into conflict with humans and are potential vectors of zoonotic disease, assessing densities in suburban areas and identifying factors that influence them are necessary to aid management and mitigation. However, fragmented, privately owned land restricts the use of conventional carnivore surveying techniques in these areas, requiring development of novel methods. We present a method that combines questionnaire distribution to residents with field surveys and GIS, to determine relative density of two urban carnivores in England, Great Britain. We determined the density of: red fox (Vulpes vulpes) social groups in 14, approximately 1km2 suburban areas in 8 different towns and cities; and Eurasian badger (Meles meles) social groups in three suburban areas of one city. Average relative fox group density (FGD) was 3.72 km-2, which was double the estimates for cities with resident foxes in the 1980's. Density was comparable to an alternative estimate derived from trapping and GPS-tracking, indicating the validity of the method. However, FGD did not correlate with a national dataset based on fox sightings, indicating unreliability of the national data to determine actual densities or to extrapolate a national population estimate. Using species-specific clustering units that reflect social organisation, the method was additionally applied to suburban badgers to derive relative badger group density (BGD) for one city (Brighton, 2.41 km-2). We demonstrate that citizen science approaches can effectively obtain data to assess suburban carnivore density, however publicly derived national data sets need to be locally validated before extrapolations can be undertaken. The method we present for assessing densities of foxes and badgers in British towns and cities is also adaptable to other urban carnivores elsewhere. However this transferability is contingent on species traits meeting particular criteria, and on resident responsiveness.

Do power lines and protected areas present a catch-22 situation for Cape vultures (Gyps coprotheres)?

Cape vulture Gyps coprotheres populations have declined across their range due to multiple anthropogenic threats. Their susceptibility to fatal collisions with the expanding power line network and the prevalence of carcasses contaminated with illegal poisons and other threats outside protected areas are thought to be the primary drivers of declines in southern Africa. We used GPS-GSM units to track the movements and delineate the home ranges of five adult (mean ±SD minimum convex polygon area = 121,655±90,845 km(2)) and four immature (mean ±SD minimum convex polygon area = 492,300±259,427 km(2)) Cape vultures to investigate the influence of power lines and their use of protected areas. The vultures travelled more than 1,000 km from the capture site and collectively entered five different countries in southern Africa. Their movement patterns and core foraging ranges were closely associated with the spatial distribution of transmission power lines and we present evidence that the construction of power lines has allowed the species to extend its range to areas previously devoid of suitable perches. The distribution of locations of known Cape vulture mortalities caused by interactions with power lines corresponded to the core ranges of the tracked vultures. Although some of the vultures regularly roosted at breeding colonies located inside protected areas the majority of foraging activity took place on unprotected farmland. Their ability to travel vast distances very quickly and the high proportion of time they spend in the vicinity of power lines and outside protected areas make Cape vultures especially vulnerable to negative interactions with the expanding power line network and the full range of threats across the region. Co-ordinated cross-border conservation strategies beyond the protected area network will therefore be necessary to ensure the future survival of threatened vultures in Africa.

Identifying indicators of illegal behaviour: carnivore killing in human-managed landscapes.

Managing natural resources often depends on influencing people's behaviour, however effectively targeting interventions to discourage environmentally harmful behaviours is challenging because those involved may be unwilling to identify themselves. Non-sensitive indicators of sensitive behaviours are therefore needed. Previous studies have investigated people's attitudes, assuming attitudes reflect behaviour. There has also been interest in using people's estimates of the proportion of their peers involved in sensitive behaviours to identify those involved, since people tend to assume that others behave like themselves. However, there has been little attempt to test the potential of such indicators. We use the randomized response technique (RRT), designed for investigating sensitive behaviours, to estimate the proportion of farmers in north-eastern South Africa killing carnivores, and use a modified logistic regression model to explore relationships between our best estimates of true behaviour (from RRT) and our proposed non-sensitive indicators (including farmers' attitudes, and estimates of peer-behaviour). Farmers' attitudes towards carnivores, question sensitivity and estimates of peers' behaviour, predict the likelihood of farmers killing carnivores. Attitude and estimates of peer-behaviour are useful indicators of involvement in illicit behaviours and may be used to identify groups of people to engage in interventions aimed at changing behaviour.