Improving predictions of the location and use of warrens in sensitive rabbit populations

The location and use of warrens is a key factor in the population dynamics of the European rabbit Oryctolagus cuniculus L., which can influence stability and persistence in the long term. Within the species' original distribution range, its numbers have declined sharply in recent decades, which is a serious problem for the conservation of Mediterranean ecosystems. Artificial warrens are commonly used to bolster rabbit populations and the characterization of natural warrens in those areas might improve their efficiency. In this study, we use binomial generalized linear models to identify the factors associated with the location and use of warrens in a low-density area and we evaluate the effect of including the spatial structure of the data in the model. In addition, we generate a map that predicts the most suitable areas for artificial warrens. Contrary to what was expected, habitat variables are only secondary factors, and the location and use of warrens is influenced mainly by spatial factors, such as proximity to nearby warrens. Furthermore, the aggregated spatial pattern of warrens suggests that, at the local scale, for example, the hunting estate, intra-specific interactions might be playing a primary role in these low-density populations. To identify the most suitable sites for artificial warrens and, thereby, improve the efficiency of artificial warren building for conservation purposes, information about spatial structures should be included in models predicting natural warrens of European rabbits.     

Effectiveness of habitat management in the recovery of low-density populations of wild rabbit

Understanding the relationship between spatial patterns of landscape attributes and population presence and abundance is essential for understanding population processes as well as supporting management and conservation strategies. This study evaluates the influence of three factors: environment, habitat management, and season on the presence and abundance of the wild rabbit (Oryctolagus cuniculus), an important prey species for Mediterranean endangered predator species. To address this issue, we estimated wild rabbit presence and abundance by latrine counting in transects located in 45 plots within a 250?×?250 m grid from June 2007 until June 2009 in a 1,200 ha hunting area in southern Portugal. We then analyzed how wild rabbit presence and abundance correlate with the aforementioned factors. Our results showed that the main variable influencing wild rabbit presence and abundance was the distance to the artificial warrens. North and northeast slope directions were negatively related to wild rabbit presence. Conversely, rabbit presence was positively correlated with short distances to ecotone, artificial warrens, and spring. Regarding rabbit abundance, in addition to artificial warrens, soft soils, bushes, and season also had a positive effect. We found that environmental variables, management practices, and season each affect wild rabbit presence and abundance differently at a home range scale in low-density population. Thus, our major recommendations are reducing the distance to artificial warrens and ecotone, ideally to less than 100 m, and promoting habitat quality improvement on slopes with plenty of sun exposure.     

Age and season determine European rabbit habitat use in Mediterranean ecosystems

Knowledge about the factors determining habitat use is especially interesting for herbivores living under seasonal climates as they have to deal with food shortage during the drought season. In this context, different-aged individuals are expected to respond differently to seasonal variations because nutritional requirements and predation risk can vary with age. We investigated adult and juvenile European rabbit (Oryctolagus cuniculus) habitat use in a Mediterranean ecosystem of central Spain, during spring, summer and winter. Relationships between adult and juvenile rabbit pellet abundances and 11 environmental variables related to food availability and refuge density were analysed by means of multiple regression, and evaluated using information theory to identify the set of models best supported by the data. Density of warren entrances was the more constant predictor of habitat use for juvenile rabbits in all the seasons. Herbaceous vegetation volume had a negative influence and was the strongest predictor for adult rabbit habitat use in spring and winter. In summer, green vegetation cover became the strongest positive habitat use predictor. These results suggest that adults prefer to forage in low volume swards ensuring a wide sensory range for the detection of approaching predators. However, the arrival of summer and its associated food depletion forces them to shift toward more open productive areas where green vegetation persists, but at the expense of higher predation risk. Seasonal variation induces minor changes in juvenile habitat use due to their strong dependence on warrens. Thus, our results show that rabbit habitat use is influenced by animal age and seasonal variations in resources.     

Responses of naïve and experienced European rabbits to predator odour

The response of prey species to predator scent has been investigated in many mammalian species; however, there is little information about the responses of European wild rabbits at the population level. Therefore, we conducted a simple experiment to investigate the behavioural response of a rabbit population to native predator cues in the wild. We compared the response to the scent of a predator (red fox) in a wild rabbit population bred in semi-natural conditions and naïve to terrestrial predators with the response of a population in a similar environment where terrestrial predators were present. The response to predators was based on rabbit abundance, inferred from pellet counts and measured by the defecation rate per day (DRD). Our results indicate that rabbits responded to the odour of fox faeces in the treatment warrens, resulting in a lower DRD. The main anti-predator behaviour observed was spatial avoidance (warren abandonment), which seemed to be more accentuated for rabbits who had not previously had contact with foxes in the plot where terrestrial predators were excluded. In both the fenced and the unfenced plot, the differences in the effect of the predator odour between the control and treatment warrens disappeared after cessation of treatment, suggesting a flexible and adaptive behaviour of rabbits to predator cues.

     

Conservation of native fauna in highly invaded systems: managing mammalian predators in New Zealand

Programs to conserve native fauna in invaded ecosystems often aim to reduce the impacts of alien predators. This approach can lead to unexpected outcomes in the native and the remaining invasive components of restored ecosystems. In New Zealand, suppression and eradication of invasive mammalian predators are well‐established conservation strategies, particularly on offshore islands and in mainland ecosanctuaries. Predator control has achieved important conservation gains over increasingly large areas but these can be offset by the ecological release of other uncontrolled pest species. In addition, novel ecosystems created by selective predator control and reintroductions of locally extinct or depleted native species may have unexpected trajectories as they evolve. Effective conservation requires new techniques for controlling entire suites of invasive predators over large areas, routine monitoring of the conservation outcomes of predator control, and better understanding of how modified, and in some cases reconstructed, seminatural ecosystems change when invasive predators are removed.     

When more is less: the negative effect of European rabbit release upon local warren occupancy

European rabbit translocation is an extended practice in Spain, France, and Portugal, for both conservation and hunting purposes. Some of these translocations are carried out with the aim of reinforcing existing rabbit populations. In these cases, some of the new rabbits are released into warrens already occupied by resident conspecifics. This could have a negative impact upon both the released and the resident individuals owing to the “dear enemy” effect and the territoriality of the resident rabbits. In this study, we evaluated the effect of rabbit release into occupied warrens, in small areas populated by low-density resident rabbit populations. We observed negative effects at two different levels: the number of active entrances per recipient warren and the number of active warrens per reinforced plot, in addition to a general lack of increase in rabbit abundance in the area and, therefore, the failure of the reinforcement actions. Our results strongly suggest that the release of European rabbits into warrens occupied by resident rabbits is contraindicated if the objective is to recover rabbit populations in the area.     

Free housing for declining populations: Optimizing the provision of artificial breeding structures

The improvement of habitat quality and reproductive success through the implementation of artificial breeding structures is one of the most widespread in situ conservation strategies applied to the recovery of declining wildlife populations. Several past studies have monitored the use of artificial breeding structures in the wild, but virtually none of them have investigated which demographic and environmental factors actually determine their effectiveness in facilitating reproductive success. Therefore, the aim of the present study was to identify those factors influencing breeding success in artificial structures. With this purpose we surveyed a declining population of a keystone species of Mediterranean ecosystems, namely the European wild rabbit. In Doñana National Park, we sampled during the course of two years the breeding success of wild rabbits from 47 artificial warrens, experimentally provided under different demographic and environmental conditions. In order to determine the relative importance of such factors, we applied an AIC model selection procedure to alternative biological hypothesis groups, postulating that breeding success in artificial structures would primarily depend on population density, food availability and/or predator limitation. We found that the efficiency of artificial warrens in facilitating rabbit reproductive success can vary, with an increase associated with greater pasture availability. Predator limitation by fencing also positively affected rabbit reproduction, but only in combination with high pasture availability. On their own, fencing treatment and population reinforcements had low and null effectiveness facilitating reproduction, respectively, as well as entailing significant economic and biological costs. The best way to improve breeding success in artificial structures, therefore, was through their strategic placement in potential high-quality habitats, i.e. with suitable pasture availability. We suggest that vegetation management can effectively increase carrying capacity and consequently the target species’ reproductive success, especially in fast breeding species like the European wild rabbit. The provision of artificial breeding structures for conservation purposes may significantly benefit from the elaboration of concurrent experimental studies similar to the present research, which can help establish useful guidelines for optimizing future conservation efforts.     

Habitat use and movement patterns by dependent and independent juvenile Grasshopper Sparrows during the post‐fledging period

The post‐fledging period is a critical life stage for young grassland birds. Habitat selection by recently fledged birds may differ from that of adults and may change as juveniles transition from the care and protection of parents to independence. To describe patterns of habitat selection during these important life stages, we studied habitat use by juvenile Grasshopper Sparrows (Ammodramus savannarum) in a Conservation Reserve Program grassland in Maryland. We used radio‐telemetry to track daily movement patterns of two age classes of Grasshopper Sparrows during the post‐fledging period. Sparrows were classified as either dependent (<32‐d‐old) or independent (≥32‐d‐old). We characterized the vegetation at 780 vegetation plots (390 plots where birds were located and 390 paired random plots). Microhabitats where dependent birds were found had significantly more bare ground, litter, and plant species richness than paired random plots. In addition, dependent birds were found in plots with less bare ground, more warm‐season grass cover, more total vegetation cover, and more forb cover than plots used by independent birds. Plots where independent birds were located also had significantly more bare ground than random plots. Dependent birds are less able to escape from predators because their flight feathers are not fully grown so they may benefit from remaining in areas of greater vegetation cover. However, juveniles transitioning from dependence to independence must forage on their own, possibly explaining their increased use of more open areas where foraging may be easier. To properly manage habitat for grassland birds, management strategies must consider the changing needs of birds during different stages of development. Our results highlight the importance of diverse grassland ecosystems for juvenile grassland birds during the transition to independence.     

Restocking a keystone species in a biodiversity hotspot: Recovering the European rabbit on a landscape scale

Rabbit populations in Iberia remain at low densities in several areas in which their endangered predators still coexist, and the recovery of these populations is therefore urgent if the integrity of Iberian Mediterranean ecosystems is to be maintained. The enhancement of wild rabbit populations has been attempted through the use of in situ extensive rabbit captive breeding enclosures (restocking plots), which reduce mortality caused by terrestrial predators and dispersal movements and permit the breeding of young individuals which can then naturally disperse to settle in the surrounding areas. However, their effectiveness, the role of its size, the optimal habitat management that should be promoted around them and the habitat features remains uncertain. Here, we show results from a four year study of an ambitious rabbit restocking plan on a landscape scale. We measured rabbit abundance in a vast area in which thirty-two restocking plots were built to create an initial rabbit population for further dispersion, in addition to an intensive habitat management program. We also compared rabbit abundance between managed and unmanaged UTM cells of 2.5 km × 2.5 km. Our results showed that rabbit abundance was three times higher in managed cells, but four years after restocking, rabbit abundances had only reached the threshold needed to support stable Iberian lynx presence (at least 10 latrines per km^?1) in 9 of the 23 managed cells. Rabbit abundance was strongly affected by habitat treatment and scrub coverage. The increase of shelter was useful in low cover areas but ineligible in places with high scrub cover, where the increase of refuge plus scrub clearing to create pastures improve rabbit abundance more effectively. In the light of our results, restocking plots should be built only in places with suitable habitat, whereas pastures should be created in dense scrublands and refuge in low cover areas.     

Rabbit (Oryctolagus cuniculus) abundance and protected areas in central-southern Spain: why they do not match?

European rabbits (Oryctolagus cuniculus), a keystone species in the Iberian Mediterranean ecosystem, are the staple prey of the Iberian lynx (Lynx pardinus) and the Spanish imperial eagle (Aquila adalberti). These predators require medium to high rabbit densities and a low degree of human disturbance. We compared rabbit abundances in areas of central-southern Spain under three levels of protection and management: protected areas, intensively managed (nonprotected) hunting estates, and other nonprotected areas. We used pellet abundance indices to estimate rabbit density in 118 surveys conducted during the summers of 2002 and 2003. We observed greater rabbit abundance in intensively managed hunting estates compared to protected areas and other nonprotected areas, perhaps because policy makers did not consider rabbit numbers when selecting priority areas. Alternatively, differences in game management practices (e.g., predator control or habitat management) may explain the higher rabbit densities observed in managed hunting estates. Our results suggest that the best feeding conditions for the Iberian lynx and the Spanish imperial eagle occur in intensively managed hunting areas, where such predators are frequently persecuted. The conservation of these endangered predators may require efforts to increase rabbit densities in protected areas.     

Burrowing by translocated boodie (Bettongia lesueur) populations alters soils but has limited effects on vegetation

Digging and burrowing mammals modify soil resources, creating shelter for other animals and influencing vegetation and soil biota. The use of conservation translocations to reinstate the ecosystem functions of digging and burrowing mammals is becoming more common. However, in an increasingly altered world, the roles of translocated populations, and their importance for other species, may be different. Boodies (Bettongia lesueur), a commonly translocated species in Australia, construct extensive warrens, but how their warrens affect soil properties and vegetation communities is unknown. We investigated soil properties, vegetation communities, and novel ecosystem elements (specifically non‐native flora and fauna) on boodie warrens at three translocation sites widely distributed across the species’ former range. We found that soil moisture and most soil nutrients were higher, and soil compaction was lower, on warrens in all sites and habitat types. In contrast, there were few substantial changes to vegetation species richness, cover, composition, or productivity. In one habitat type, the cover of shrubs less than 1 m tall was greater on warrens than control plots. At the two sites where non‐native plants were present, their cover was greater, and they were more commonly found on boodie warrens compared to control plots. Fourteen species of native mammals and reptiles were recorded using the warrens, but, where they occurred, the scat of the non‐native rabbit (Oryctolagus cuniculus) was also more abundant on the warrens. Together, our results suggest that translocated boodie populations may be benefiting both native and non‐native flora and fauna. Translocated boodies, through the construction of their warrens, substantially alter the sites where they are released, but this does not always reflect their historic ecosystem roles.     

Responses of juvenile terrestrial salamanders to introduced (Lithobius forficatus) and native centipedes (Scolopocryptops sexspinosus)

When introduced species invade ecosystems, alterations in community structure can emerge from the competitive and predatory interactions that occur between introduced and native guild members. Because a number of recent studies have shown that large predatory invertebrates can both compete with and prey on small vertebrates and because introductions of non-native species may play a role in amphibian declines, the effects of introduced centipedes Lithobius forficatus and native centipedes Scolopocryptops sexspinosus on juveniles of the red-backed salamander Plethodon cinereus were examined. In laboratory arenas, juvenile salamanders exhibited submissive behaviour in response to the odours of both species of centipede. There were no significant differences in salamander response to the two centipede odour treatments, but compared to controls, juveniles of P. cinereus spent significantly more time in escape and in a flattened submissive posture when presented with native centipede odours. Despite significant size differences between centipedes and juvenile salamanders, no predation of salamanders by either species of centipede occurred in any pairings. Juveniles exhibited more chemosensory behaviour towards native centipedes and towards their odours and exhibited marked reductions in aggressive posturing when centipedes were present. Field and laboratory data suggest that juveniles of P. cinereus and centipedes were negatively associated. In laboratory trials, the native centipede excluded juvenile salamanders from cover objects and we found fewer instances of co-occurrence in the field than expected. These studies are the first to examine the behavioural interactions between juveniles of P. cinereus and invertebrate predators, one introduced and one native, of eastern deciduous forest-floor food webs.     

Short-Term Vegetation and Soil Responses to Mechanical Destruction of Rabbit (Oryctolagus cuniculus L.) Warrens in an Australian Box Woodland

We studied the impact of disturbance by rabbits on plants and soils along a gradient out from the center of ripped rabbit warrens in an Australian semiarid woodland. Five years after the warrens were ripped, the impact of rabbits was still apparent. The cover of bare soil declined, and the cryptogam cover increased with increasing distance from the warren mound. However, litter cover, plant cover, and plant diversity remained unchanged with increasing distance from the mounds. Differences in plant composition were apparent with increasing distance from the mounds, with three species, Schismus barbatus, Salsola kali var. kali, and Chenopodium melanocarpum dominating the mounds, whereas the perennial grass Austrostipa scabra dominated the nonwarren control surfaces. Two species, Crassula sieberana and S. barbatus, dominated the active soil seed bank on ripped warrens. The mounds had the lowest number of species in the soil seed bank, whereas the warren edge microsite had the greatest. Ripped and unripped warrens differed substantially in their complement of species, and ripped warrens contained an order of magnitude fewer active warren entrances compared with unripped warrens. Ripped warrens also had significantly more plant cover than unripped warrens. Taken together, our results reinforce the view that rabbits have a destructive effect on surface soils and vegetation in semiarid woodlands and suggest that restoration of the original woodland vegetation after warren ripping is likely to be a slow and ongoing process.     

European rabbit population trends and associated factors: a review of the situation in the Iberian Peninsula

• 1 European rabbits are considered a keystone species in the Iberian Peninsula. Their populations have sharply declined over the past century, mainly due to habitat loss and the arrival of two viral diseases: myxomatosis in the 1950s and rabbit haemorrhagic disease (RHD) at the end of the 1980s. For the conservation of the Iberian Mediterranean ecosystem, it is important to determine whether rabbit populations are recovering two decades after the RHD outbreak, and to identify the factors associated with population recovery.
• 2 Here, we review the current knowledge on recent rabbit population trends in the Iberian Peninsula and the factors associated with these trends.
• 3 Although most rabbit populations are still declining in the Iberian Peninsula, a few seem to have recovered. In general, positive trends have been recorded in species‐friendly habitats characterized by non‐fragmented landscapes, interspersed patches of Mediterranean scrubland, good pastures and/or crops, soft soils that are suitable for warren construction and a Mediterranean climate with relatively high rainfall. Additionally, rabbits seem to be recovering better in areas where management practices (e.g. low hunting pressure, habitat management and predator control) are applied to increase their numbers.
• 4 From these findings, it is possible to identify five broad objectives for rabbit conservation in the Iberian Peninsula. First, it is clearly necessary to establish a long‐term programme for monitoring rabbit abundance and trends on a large scale. Second, the conservation and restoration of open Mediterranean scrubland should be a priority for stabilizing and maintaining existing healthy rabbit populations. Third, despite the lack of experimental evidence, management activities aimed at increasing the quantity and quality of both refuge and food should continue to be implemented. Fourth, legislation on the timing of the hunting season should be revisited following recommendations made by scientists. Finally, experimental approaches are required to investigate whether the control of generalist predators is a successful strategy to allow rabbit populations to recover.

     

Age and seasonal effects on predator-sensitive foraging in squirrel monkeys (Saimiri sciureus): a field experiment

A field experiment was conducted to examine the effect of perceived predation risk on the use of foraging areas by juvenile and adult primates under different conditions of local food abundance. Wild squirrel monkeys, Saimiri sciureus, were observed in an experiment conducted during the dry and the wet seasons at a site in Eastern Amazonia, Brazil. Animals were presented with feeding platforms that differed in food quantity and exposure to aerial predators through varying vegetative cover. In the dry season, juveniles and adults chose platforms based solely on food quantity. However, in the wet season, juveniles foraged preferentially on high-reward platforms only if cover level also was high (i.e., potentially offered greater concealment from predators). In contrast, adults showed the same pattern of platform use regardless of season. These results indicate that age and local resource availability based on seasonality affect whether primates forage in a predator-sensitive manner. Juveniles may be more sensitive to predation risk when foraging, and individuals may take fewer risks when resource abundance is high in their environment.     

Localised control of an introduced predator: creating problems for the future?

Introduced mammalian predators have had significant impacts on many native prey species. Although control of such predators for conservation management is becoming increasingly commonplace, it is often undertaken at a relatively small scale in relation to the overall predator population. Processes such as immigration mean that it remains difficult to determine the effectiveness of control measures. We investigated the impacts of feral ferret Mustela furo removal on the entire feral ferret population on Rathlin Island, UK. Removal of ferrets prior to breeding led to a substantial increase in the post-dispersal population through the enhanced survival of juveniles. Despite increased numbers, overwinter survival remained high, potentially aided by the reduced territoriality shown by this feral species compared to wild carnivores. The response of this ferret population to control is a further illustration of the complex ecological processes and outcomes arising from the anthropogenic disruption of wildlife populations. It highlights how partial or localised management may prove ineffective, and at worst might exacerbate the problems that management was designed to avert.     

Radio‐transmitters have no impact on survival of pre‐fledged American Woodcocks

American Woodcocks (Scolopax minor) are a high priority species of conservation need across most of their breeding range due to long‐term population declines. Survival of juveniles may be key to understanding these population declines, but there have been few direct estimates of juvenile woodcock survival rates, and no recent assessment of the possible effect of radio‐tagging on juvenile survival. In 2011 and 2012, we radio‐tagged 73 juvenile American Woodcocks in west‐central Minnesota and compared survival rates of radio‐tagged (N = 58) and non‐radio‐tagged (N = 82) juveniles during the period from hatching to fledging. We compared survival rates of juveniles with known fates and used logistic‐exposure models to assess the potential impact of radio‐transmitters on survival. We evaluated variables related to juvenile survival including age, hatch date, maximum and minimum temperature, precipitation, and year to assess the possible effects of radio‐transmitters. The best‐supported model of survival rate of juvenile American Woodcocks included the interaction of age and year and a negative effect of precipitation (β = ?0.76, 85% CI: ?1.08 to ?0.43), but did not include a negative effect of transmitters. Our results suggest that radio‐transmitters did not impact survival of juvenile American Woodcocks and that transmitters are a reliable tool for studying survival of juvenile American Woodcocks, and perhaps other precocial shorebirds.     

Habitat complexity affects how young of the year Atlantic cod Gadus morhua perceive predation threat from older conspecifics

The response of age 0+ year juvenile Atlantic cod Gadus morhua to the presence of age 1+ and age 3+ year conspecifics was measured with and without cover available. Juveniles reacted by aggregating more closely and maintaining distance from older conspecifics in an experimental setting without cover but only to age 3+ year conspecifics when cover was available. The results indicate that prior residence of older juveniles can affect age 0+ year juveniles during benthic settlement and highlights the conservation value of structurally complex nursery habitats.     

Varying demographic impacts of different fisheries on three Mediterranean seabird species

Fisheries have an enormous economic importance, but reconciling their socio‐economic features with the conservation and sustainability of marine ecosystems presents major challenges. Bycatch mortality from fisheries is clearly among the most serious global threats for marine ecosystems, affecting a wide range of top predators. Recent estimates report ca. 200,000 seabirds killed annually by bycatch in European waters. However, there is an urgent need to rigorously estimate actual mortality rates and quantify effects of bycatch on populations. The Mediterranean Sea is one of the most impacted regions. Here, we estimate for the first time both bycatch mortality rates and their population‐level effects on three endemic and vulnerable Mediterranean taxa: Scopoli's shearwater, Mediterranean shag, and Audouin's gull, that die in different types of fishing gears: longlines, gillnets and sport trolling, respectively. We use multi‐event capture–recapture modelling to estimate crucial demographic parameters, including the probabilities of dying in different fishing gears. We then build stochastic demography models to forecast the viability of the populations under different management scenarios. Longline bycatch was particularly severe for adults of Scopoli's shearwaters and Audouin's gulls (ca. 28% and 23% of total mortality, respectively) and also for immature gulls (ca. 90% of mortality). Gillnets had a lower impact, but were still responsible for ca. 9% of juvenile mortality on shags, whereas sport trolling only slightly influenced total mortality in gulls. Bycatch mortality has high population‐level impacts in all three species, with shearwaters having the highest extinction risk under current mortality rates. Different life‐history traits and compensatory demographic mechanisms between the three species are probably influencing the different bycatch impact: for shearwaters, urgent conservation actions are required to ensure the viability of their populations. Results will be very useful for guiding future seabird conservation policies and moving towards an ecosystem‐based approach to sustainable fisheries management.     

Keystone effects of an alien top-predator stem extinctions of native mammals

Alien predators can have catastrophic effects on ecosystems and are thought to be much more harmful to biodiversity than their native counterparts. However, trophic cascade theory and the mesopredator release hypothesis predict that the removal of top predators will result in the reorganization of trophic webs and loss of biodiversity. Using field data collected throughout arid Australia, we provide evidence that removal of an alien top-predator, the dingo, has cascading effects through lower trophic levels. Dingo removal was linked to increased activity of herbivores and an invasive mesopredator, the red fox (Vulpes vulpes), and to the loss of grass cover and native species of small mammals. Using species distribution data, we predict that reintroducing or maintaining dingo populations would produce a net benefit for the conservation of threatened native mammals across greater than 2.42 × 10⁶ km² of Australia. Our study provides evidence that an alien top predator can assume a keystone role and be beneficial for biodiversity conservation, and also that mammalian carnivores more generally can generate strong trophic cascades in terrestrial ecosystems.