Selection of food patches by sympatric herbivores in response to concealment and distance from a refuge

Small herbivores face risks of predation while foraging and are often forced to trade off food quality for safety. Life history, behaviour, and habitat of predator and prey can influence these trade‐offs. We compared how two sympatric rabbits (pygmy rabbit, Brachylagus idahoensis; mountain cottontail, Sylvilagus nuttallii) that differ in size, use of burrows, and habitat specialization in the sagebrush‐steppe of western North America respond to amount and orientation of concealment cover and proximity to burrow refuges when selecting food patches. We predicted that both rabbit species would prefer food patches that offered greater concealment and food patches that were closer to burrow refuges. However, because pygmy rabbits are small, obligate burrowers that are restricted to sagebrush habitats, we predicted that they would show stronger preferences for greater cover, orientation of concealment, and patches closer to burrow refuges. We offered two food patches to individuals of each species during three experiments that either varied in the amount of concealment cover, orientation of concealment cover, or distance from a burrow refuge. Both species preferred food patches that offered greater concealment, but pygmy rabbits generally preferred terrestrial and mountain cottontails preferred aerial concealment. Only pygmy rabbits preferred food patches closer to their burrow refuge. Different responses to concealment and proximity to burrow refuges by the two species likely reflect differences in perceived predation risks. Because terrestrial predators are able to dig for prey in burrows, animals like pygmy rabbits that rely on burrow refuges might select food patches based more on terrestrial concealment. In contrast, larger habitat generalists that do not rely on burrow refuges, like mountain cottontails, might trade off terrestrial concealment for visibility to detect approaching terrestrial predators. This study suggests that body size and evolutionary adaptations for using habitat, even in closely related species, might influence anti‐predator behaviors in prey species.     

Spatio-Temporal Factors Shaping Diurnal Space Use by Pygmy Rabbits

Abstract: Factors influencing patterns of space use by pygmy rabbits (Brachylagus idahoensis) are poorly understood. We studied diurnal space use by adult pygmy rabbits during multiple breeding and nonbreeding seasons at 3 sites in the Lemhi Valley, Idaho, USA, during 2004–2005. Pygmy rabbits used larger areas than predicted by allometric models and documented by some previous investigations. Sex and season strongly influenced space use by rabbits. Males used larger home ranges and core areas, more burrow systems, and more widely dispersed burrow systems than did female rabbits. We also documented significant differences among study sites in many movement parameters, which suggested that local resource distribution also might influence how pygmy rabbits use space. Our results indicated that pygmy rabbits use large areas and exhibit seasonal, sex, and site-specific variation in patterns of movement and space use. Therefore, larger areas of habitat may be needed to conserve pygmy rabbits to accommodate seasonal, regional, and potentially annual variation in resource availability and to maintain linkages among populations.     

Sagebrush Characteristics Influencing Foraging Patterns of Pygmy Rabbits

The pygmy rabbit (Brachylagus idahoensis) is endemic to the sagebrush steppe landscapes in the western United States. Pygmy rabbits have adapted to this region by depending on big sagebrush (Artemisia tridentata) as a source of nutrition and concealment from predators. Being a central place forager, and a dietary specialist makes pygmy rabbits an ideal subject to study foraging tradeoffs. Our objectives were to determine if pygmy rabbit foraging patterns are influenced by the nutrient content of individual sagebrush, or the size and location of individual sagebrush. We first developed a near infrared spectroscopy assay that can be conducted in the field, with whole plant foliage, to determine individual sagebrush crude protein (CP) and acid detergent fiber (ADF) contents. We then measured the size, location, protein, and fiber contents of >1,500 foraged and non-foraged sagebrush plants surrounding pygmy rabbit burrows in 2 field sites in northern Utah, USA. We found significant site differences in sagebrush CP levels and significant season differences in sagebrush ADF levels; sagebrush closer to central burrows were higher in protein and lower in fiber. Pygmy rabbits preferred to forage on taller sagebrush plants that were closer to the central burrow but only marginally higher in CP (0.2%) and lower in ADF (1%). Sagebrush plants with a >50% chance of being foraged, were <5.6 m from the burrow, >0.67 m tall, >10.4% CP, and <34.95% ADF. The selection of closer, larger, and more nutrient-dense sagebrush may be influenced by pygmy rabbits' need for concealment from predators or a way to minimize foraging effort. © 2020 The Wildlife Society.     

Effects of sagebrush treatments on multi-scale resource selection by pygmy rabbits

The effects of widespread sagebrush removal treatments on pygmy rabbits (Brachylagus idahoensis) are not well understood. Due to reliance on sagebrush, pygmy rabbits are among the species for which these treatments may be detrimental. Our objectives were to evaluate the effects of experimental sagebrush treatment on 8 radio-collared pygmy rabbits between and within home range habitat selection using Monte Carlo simulation from null models. Pygmy rabbits were not extirpated from plots containing habitat treatments, and we found no evidence that treatments affected home range placement. The mean treatment distance of observed home range centers did not differ from repeated trials of random points. However, we found evidence of within home range selection against treatments from 2 of 8 rabbits located close to the treatments. The mean treatment distance of all observed locations for these 2 rabbits was greater than expected based on a null model. We also used snow tracking to show that pygmy rabbits entered treatments in 4 out of 21 trials, which was less often than expected by chance (G² = 8.662, P = 0.003). Conservatively, sagebrush removal treatments should not be conducted on active or recently active pygmy rabbit burrows. Elsewhere near known pygmy rabbit sites, treated patches should be small and connected by untreated corridors to prevent potentially limiting movement of rabbits among the untreated habitat. © 2011 The Wildlife Society.     

Survival of Juvenile Pygmy Rabbits

ABSTRACT Until recently, natal behavior of pygmy rabbits (Brachylagus idahoensis) was largely unknown, and no information on survival of free-ranging juveniles was available. We evaluated survival of radiotagged juvenile pygmy rabbits at 2 sites in east-central Idaho, USA, during 2004 and 2005. We captured juveniles (n 58) shortly after they emerged from natal burrows. Mortality rates were high and variable, ranging from 27% for females during 2004 to 63% for males during 2005. Approximately 69% of mortalities were attributed to predation. We evaluated variables influencing juvenile survival through 18 weeks old using known-fate models in Program MARK. We expected survival to decline around the age of natal dispersal and to be lower for young born later in the season. We evaluated 14 candidate models that included sex, year, study area, and relative date of birth within each year. Model selection results did not indicate strong support for any single combination of variables, and 8 competing models all included effects of relative date of birth, year, and study area. These results revealed substantial variability in survival of juveniles across multiple factors, and we documented similar patterns for adult pygmy rabbits. Such high variability in survival over relatively small spatial and temporal scales might contribute to marked fluctuations in populations of pygmy rabbits and, hence, managers interested in monitoring this species might consider monitoring multiple populations across broader geographic areas to assess regional trends in numbers.     

Density of rabbits (

The density of rabbits (Oryctolagus cuniculus) in the modified tussock grasslands of the Mackenzie Basin, South Island, New Zealand, in August-September 1991 was determined within 26 I-ha quadrats spread over 1000 ha. The area was poisoned with 1080- carrot baits and dead and live rabbits counted. The overall kill rate was 93%. Wide variability in rabbit densities amongst the quadrats was correlated with burrow density, but vegetation was not a significant predictor of rabbit numbers. High density quadrats were not all spatially clumped together. Variation amongst quadrats of 0-43% of rabbits dying underground shows that searching burrows as well as the surface will provide the most accurate rabbit densities. Poisoning efficacy was greater than or equal to 90% in 77% of the quadrats, but two low- density quadrats recorded kill rates <70%. Nine per cent of carcasses had been partly eaten by predators, suggesting a potential for the predators to be killed during rabbit control operations via secondary poisoning. The average density of 19 rabbits per ha confirms the classification of this area as a moderate to high rabbit-prone zone, with the rabbit population imposing grazing pressure equivalent to at least 1-2 sheep per ha at the beginning of spring. Sustainable agriculture cannot be attained in these semi-arid regions of New Zealand without cheap, widespread and effective rabbit control.     

European rabbits as ecosystem engineers: warrens increase lizard density and diversity

Mammals that build extensive open burrow systems are often classified as ecosystem engineers, since they have the potential to modulate the availability of resources for themselves and other organisms. Lizards may benefit from the heterogeneity created by these structures, especially if coupled with an increased offer of sites for refuge and thermoregulation. However, information about these engineering effects by burrowing animals is scarce. We investigated the influence of European rabbit burrows on several parameters of a Mediterranean lizard community (abundance, density, diversity and body condition) in three different habitats (open pastures, holm oak and scrub patches). We found that lizards were positively associated with burrows, and that burrows determined lizard presence at otherwise unfavourable habitats. Moreover, community parameters such as density and species richness were higher in sites with burrows. Burrows influenced lizard species in different ways, and were also relevant for other Mediterranean vertebrates, as revealed by questionnaires to experts. We also explored the possible resources provided by burrows for lizards. Warrens offer relatively abundant prey and appropriate retreat sites for refuge and thermoregulation. Warrens may have further implications within the ecosystem, acting as stepping stones, allowing lizards to reach otherwise inaccessible habitat patches. This study shows that European rabbit warrens have a positive influence on lizard density and diversity, and confirms the role of rabbits as ecosystem engineers. This reinforces the need for appropriate conservation measures for rabbits, especially given their threatened status in the Iberian Peninsula. Furthermore, our study highlights that taking into account the influence of engineering activities increases our awareness of species interactions, and may translate into more adequate conservation measures for the preservation of biodiversity.     

Model of burrow selection predicts pattern of burrow switching by Leach's Storm‐Petrels

Patterns of nest site selection exhibited at the scale of a population should result from initial preferences of individuals occupying nest sites as well as preferences exhibited by individuals moving between nest sites. We tested whether nest‐site preferences measured at the population scale were predictive of patterns of burrow switching by Leach's Storm‐Petrels (Oceanodroma leucorhoa), a long‐lived seabird that nests in underground burrows. Breeding pairs generally choose from the pool of available existing burrows rather than constructing new burrows, and a portion of the burrows in a colony remains unused in any breeding season. We quantified burrow preference at a colony on Kent Island, New Brunswick, over four breeding seasons. We used a classification and regression tree analysis to build a predictive model of nest‐site selection. Preferentially occupied burrows were drier, longer, had larger nest chambers, and were in areas of higher burrow density. To measure preferences during burrow switching, we tracked individuals that switched burrows, comparing characteristics of the burrows in which these birds were originally found to those they inhabited at the end of the study period. Characteristics preferred by switching individuals were a subset of those observed at the scale of the population; individuals moved to burrows that were drier, longer, and had larger nest chambers. Our results show how preferences of individuals that move between nest sites contribute to nest site preferences exhibited at the population scales commonly tested.     

Diet of the killifish Rivulus marmoratus collected from land crab burrows,with further ecological notes

Synopsis One-hundred eleven specimens of the cyprinodont fish Rivulus marmoratus were trapped from burrows of the great land crab Cardisoma guanhumi in east central Florida and the entire alimentary tract removed for analysis of food items. Three categories of burrows (inactive, active and sealed) were sampled to examine possible relationships between burrow type and presence of R. marmoratus. Forty percent of the guts examined contained no food items, while the remaining 60% contained items of both terrestrial and aquatic origin. Capture rates among burrow categories did not vary significantly, and most collections consisted of 1 or 2 individuals per burrow. Mean standard length of fish within a burrow declined when more than 3 specimens were captured from a burrow. Supplemental observations on reproductive status and burrow water temperature are noted.     

Cape ground squirrels as ecosystem engineers: modifying habitat for plants,small mammals and beetles in Namib Desert grasslands

Burrowing and foraging of semi‐fossorial rodents can affect species distribution and composition. Ground squirrels dig large burrow systems for refuge from predators and temperature extremes. Burrowing and foraging around burrows by squirrels may affect habitat and resource distributions for other organisms. We examined the impact of Cape ground squirrels (Xerus inauris) on vegetation, small mammals and beetles during winter and summer in grasslands on the edge of the Namib Desert. At each burrow system and paired control site without burrows, we estimated plant cover and height using quadrats (N = 8 paired sites), small mammal abundance and species richness using mark‐recapture techniques (N = 8 paired sites) and beetle abundance and species richness using pitfall traps (N = 6 paired sites, winter only). Squirrel burrowing and foraging activities resulted in lower plant cover and height, higher small mammal abundance and lower beetle abundance and species richness. Squirrels also reduced more plant cover in winter compared to summer, but had no effect on small mammal species richness. Furthermore, plant cover and height were higher in summer, whereas small mammal abundance and species richness were higher in winter. Our results suggest that Cape ground squirrels are important ecosystem engineers that influence plant and animal communities in the Namib Desert grasslands.     

Pests controlling pests: does predator control lead to greater European rabbit abundance in Australasia?

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Environmental heterogeneity and habitat use of the red viscacha rat in drylands of west-central Argentina

Environmental heterogeneity shapes the traits of natural populations, including space use and burrowing behavior. We studied the red viscacha rat (Tympanoctomys barrerae), a semifossorial rodent with patchy distribution in drylands of west-central Argentina, to provide ecological data on habitat and burrow features of this specialist species. We collected data on 165 burrows and 55 individuals from 2017–2018 and observed correlations between different vegetation types, vegetation diversity, and physical attributes (e.g., soil hardness). A high number of individual red viscacha rats was negatively associated with herbs. The number of burrow entrances was positively associated with harder soils and negatively associated with vegetation cover. Our results suggest that the species' patchy distribution is strongly associated with environmental heterogeneity of these arid environments, and consequently provide an essential step towards conserving their populations across their geographic range.     

Location of long‐term communal burrows of a threatened arid‐zone lizard in relation to soil and vegetation

The great desert skink (Liopholis kintorei) of the Egerniinae subfamily (Reptilia: Scincidae) is a communal burrowing lizard that inhabits arid spinifex grasslands in central Australia. Great desert skink activity is centred in and around the burrows which are inhabited for many years. However, it is not known whether skinks select burrow sites with specific attributes or how continuing occupancy of burrows is influenced by the surrounding habitat; especially post‐fire, when plant cover is reduced. Here, we test whether great desert skink burrows in areas burnt 2 years previously and in longer unburnt areas are associated with particular habitat attributes, and whether there are differences between occupied and recently abandoned burrow sites. Vegetation composition, cover and soil surface characteristics at 56 established great desert skink burrows, including occupied and recently unoccupied burrows, were compared with 56 random nearby non‐burrow control sites. Burrow sites had higher plant cover compared with the surrounding landscape in both recently burnt and longer unburnt areas and were more likely to be associated with the presence of shrubs. Soil stability and infiltration were also higher at burrow sites. However, we found no evidence that burrows with lower cover were more likely to be abandoned. Our results suggest that great desert skinks may actively select high cover areas for burrow construction, although differences between burrow and control sites may also partly reflect local changes to plant cover and composition and soil properties resulting from burrow construction and long‐term habitation of a site. Further research should determine if burrows with shrubs or higher plant cover provide greater protection from predators, more structural stability for burrow construction, increased prey abundance or other benefits. We recommend that maintenance of areas with relatively higher plant cover be prioritized when managing great desert skink habitat.     

Obligate crayfish burrow use and core habitat requirements of crawfish frogs

Crawfish frogs (Lithobates areolatus) have experienced declines across large portions of their former range. These declines are out of proportion to syntopic wetland-breeding amphibian species, suggesting losses are resulting from unfavorable aspects of non-breeding upland habitat. Crawfish frogs get their common name from their affinity for crayfish burrows, although the strength of this relationship has never been formally assessed. We used radiotelemetry to address 4 questions related to upland burrow dwelling in crawfish frogs: 1) what burrow types are used and how do they function to affect crawfish frog survivorship; 2) what are the physical characteristics and habitat associations of crawfish frog burrows; 3) what are the home range sizes of crawfish frogs when burrow dwelling; and 4) where are crawfish frog burrows situated with respect to breeding wetlands? We tracked crawfish frogs to 34 burrows, discovered another 7 occupied burrows, and therefore report on 41 burrows. Crawfish frogs exclusively occupied crayfish burrows as primary burrows, which they inhabited for an average of 10.5 months of the year. With one exception, crawfish frogs also used crayfish burrows as secondary burrows—temporary retreats occupied while exhibiting breeding migrations or ranging forays. Burrows were exclusively located in grassland habitats, although crawfish frogs migrated through narrow woodlands and across gravel roads to reach distant grassland primary burrow sites. Home range estimates while inhabiting burrows were 0.05 m² (the area of the burrow entrance plus the associated feeding platform) or 0.01 m³ (the estimated volume of their burrow). Crawfish frog burrows were located at distances up to 1,020 m from their breeding wetlands. To protect crawfish frog populations, we recommend a buffer (core habitat plus terrestrial buffer) of at least 1.2 km around each breeding wetland. Within this buffer, at least 3 critical habitat elements must be present: 1) extensive grasslands maintained by prescribed burning and/or logging, 2) an adequate number of upland crayfish burrows, and 3) no soil disturbance of the sort that would destroy crayfish burrow integrity. © 2012 The Wildlife Society.     

Pattern of Dispersion of Young Wild Rabbits,Oryctolagus cuniculus L., in Relation to Burrows

The development of dispersion in relation to burrows of young rabbits, Oryctolagus cuniculus L., was studied in a sand dune habitat between May and September 1984–1985. Generally, young rabbits did not show a close association with their original burrow. From the first week of life on the surface they used different burrows as well as the original one. No significant age-related changes in the mean distance from different kinds of burrows were observed. The mean distance from the nearest burrow remained always under 3 m, but this distance may have been due largely to the high density of burrows. The apparent freedom of movements of young rabbits around different burrows may be related to the social system of the adults in a sand dune habitat.     

The depth of Sooty Shearwater Ardenna grisea burrows varies with habitat and increases with competition for space

The Sooty Shearwater Ardenna grisea, an abundant but declining petrel, is one of many seabird species that construct breeding burrows, presumably because these confer protection from predators and the elements. Little is known about the causes of variation in Sooty Shearwater burrow architecture, which can differ markedly both within and between breeding sites. We hypothesize that burrow architecture varies in response to habitat type and competition for space. To address these hypotheses, we recorded Sooty Shearwater burrow dimensions on Kidney Island, the largest Sooty Shearwater colony in the Falkland Islands, South Atlantic, and modelled these as functions of burrow density (a proxy for competition) and habitat indices. Our models suggest that Sooty Shearwaters burrow further underground in response to competition for breeding space, and that soil underlying dense tussac grass Poa flabellata is more easily excavated than other substrates, indicating how vegetation restoration could aid the conservation of this species.     

Ecological Predictors of Range Areas and Use of Burrow Systems in the Diurnal Rodent, Octodon degus

Variation in animal space use patterns may be linked to numerous ecological factors affecting survival and reproduction. We examined the relationship between ecology and above‐ and below‐ground components of space use by Octodon degus, a semi‐fossorial rodent in Chile. We monitored the daytime minimum convex polygon and adaptive kernel range areas of 26 individuals and determined the number of burrow systems used by degus during night‐time radiotelemetry and trapping of burrow systems on two study grids at Rinconada de Maipú, a semi‐arid Matorral in central Chile. We quantified food biomass, soil hardness, distance to overhead vegetative cover, and density of burrow openings at putative nest burrows. Degus living on the grid with more shrub cover had larger range areas than degus living on the grid with less cover. The range areas of degus decreased with increasing distance from overhead vegetative cover. There was a weak (but statistically significant) negative relationship between the number of burrow systems used by degus and the distance to vegetative cover and density of burrow openings at burrow systems. Male and female degus had similar range areas. Our results suggest that overhead cover decreases the risk of predation to male and female degus. Degus probably balance the benefits of numerous burrow openings (reduced predation risk) with time and energy requirements of burrow construction and maintenance. Models of space use that consider the effect of multiple ecological variables should measure different dimensions of space use.     

When to Run and When to Hide: The Influence of Concealment,Visibility, and Proximity to Refugia on Perceptions of Risk

An animal's ability to avoid predation likely depends on its ability to detect approaching predators, conceal itself, and seek refuge or protection from predators. Habitat, especially vegetation structure, can influence all of these factors concurrently. Binary categorical assessments of habitat as ‘open’ or ‘closed’, however, confound at least two functions of habitat structure that could influence the perceived risk of predation: concealment, which functions to hide an individual, and visibility, which enhances detection of a potential predator. Both can influence predation risk independently and s imultaneously. In this study, we decoupled these functional properties of vegetation and studied the effects of concealment, visibility, and proximity to a refuge on the distance at which pygmy rabbits (Brachylagus idahoensis) fled from an approaching threat (flight initiation distance; FID). Concealment by vegetation decreased perceptions of risk; however, pygmy rabbits exhibited elevated risk at high levels of visibility, regardless of the amount of concealment. Proximity to burrow entrances also influenced perceptions of risk, such that risk was significantly lower when rabbits were on or near burrow systems. Disentangling the functional properties of habitat can provide a more comprehensive understanding of the factors that influence perceived risk and escape behaviors of prey and provide insight into how habitat structure mechanistically relates to predation risk.     

The use of cover and burrows by a population of rabbits (Mammalia: Oryctolagus cuniculus) in eastern Scotland

Adult rabbits were trapped in the spring on an area of grass and broom in the east of Scotland. They were ear-tagged and a sample of males and females were fitted with radio-collars. These were followed for two months to see whether they used burrows or ground cover for shelter during the day. Rabbits found on the surface were disturbed to see whether they bolted to burrows or stayed on the surface. The population was then re trapped to measure the population density, which was estimated to be 12.6 adults/ha in March, allowing for an apparent trapping bias against older females. One hundred and twelve burrow entrances/ha were counted from random quadrats. Males were more likely to be found on the surface than females, and rabbits that were more often on the surface were less likely to bolt to burrows when disturbed. Even when rabbits did bolt to burrows they frequently left them shortly afterwards and returned to the cover of the broom. Males that were most often found on the surface were lighter in weight than those found in burrows. This was not the case for females.     

Is shadier better? The effect of agroforestry management on small mammal diversity

Agroforestry systems play fundamental roles for wildlife conservation, but are prone to disturbances from management practices aiming at increasing local productivity. This work investigates the small mammal assemblages present in cacao agroforests, which differ in shade tree density. We tested the prediction that higher tree density increases shade level, analyzed how some environmental variables important for small mammals (vegetation complexity, tree basal area, and invertebrate biomass) vary across the shade level gradient, and how the assemblages respond to these variations. We also tested the effect of the environmental variables on the abundance of the three most common species: Rhipidomys mastacalis, Hylaeamys seuanezi, and Marmosa murina. We captured 651 individuals belonging to 18 species. A positive relationship was observed between an abundance of non-forest specialists and tree basal area, while species diversity within this group showed positive association with vegetation complexity. Assemblage structure (described by a matrix of species abundance per site) was not affected by our environmental variables, but R. mastacalis was more abundant in sites with lower vegetation complexity. Higher shade levels in cacao agroforest tended to occur in sites with greater tree basal area, which was not a good predictor of small mammal diversity. This suggests that environmental management to reduce shade with the purpose of increasing cacao productivity is not necessarily negative for small mammal conservation. Species diversity was favored by structurally complex systems, a possible response to greater niche diversity.