Spatio‐temporal gradients in food supply help explain the short‐term colonisation dynamics of the critically endangered central rock‐rat (Zyzomys pedunculatus)

Currently, the impact of introduced predators on small mammal population decline is a focal research direction in the Australian desert literature. In all likelihood though, single‐factor explanation of population dynamics is inadequate, leaving gaps in our knowledge of the multitude of potential influences on small mammal abundance and occupancy patterns in time and space. Here, we investigated floristic gradients across four potential refuge sites of the central rock‐rat, Zyzomys pedunculatus, a granivore rodent (50–120 g) that is endemic to central Australia and is categorised as critically endangered. The study took place in Tjoritja/West MacDonnell National Park in the MacDonnell Ranges bioregion. Floristic sampling was allocated across the four sites, the locations of which were predetermined by an established monitoring and management programme for the central rock‐rat. Our aim was to examine the relationship between environmental gradients and floristic composition across the four sites, and thereby test the extent to which the patterns of food type and food availability can inform central rock‐rat spatio‐temporal dynamics. We found high site‐scale floristic patterning that related foremost to elevation and then to antecedent rainfall and time‐since‐fire and fire‐severity effects. To interpret these results, we applied the principles of refuge theory and we described a gradient from core refuge habitat to intermittent and then marginal habitat within the current central rock‐rat stronghold area. Overall, our results implied a strong floristic basis to central rock‐rat site occurrence, and they thus compel us to take explicit account of spatial (elevation) and temporal (rainfall–productivity and fire‐disturbance) influences on the food axis of potential refuge sites of this critically endangered species.     

The role of refuges in the persistence of Australian dryland mammals

Irruptive population dynamics are characteristic of a wide range of fauna in the world's arid (dryland) regions. Recent evidence indicates that regional persistence of irruptive species, particularly small mammals, during the extensive dry periods of unpredictable length that occur between resource pulses in drylands occurs as a result of the presence of refuge habitats or refuge patches into which populations contract during dry (bust) periods. These small dry‐period populations act as a source of animals when recolonisation of the surrounding habitat occurs during and after subsequent resource pulses (booms). The refuges used by irruptive dryland fauna differ in temporal and spatial scale from the refugia to which species contract in response to changing climate. Refuges of dryland fauna operate over timescales of months and years, whereas refugia operate on timescales of millennia over which evolutionary divergence may occur. Protection and management of refuge patches and refuge habitats should be a priority for the conservation of dryland‐dwelling fauna. This urgency is driven by recognition that disturbance to refuges can lead to the extinction of local populations and, if disturbance is widespread, entire species. Despite the apparent significance of dryland refuges for conservation management, these sites remain poorly understood ecologically. Here, we synthesise available information on the refuges of dryland‐dwelling fauna, using Australian mammals as a case study to provide focus, and document a research agenda for increasing this knowledge base. We develop a typology of refuges that recognises two main types of refuge: fixed and shifting. We outline a suite of models of fixed refuges on the basis of stability in occupancy between and within successive bust phases of population cycles. To illustrate the breadth of refuge types we provide case studies of refuge use in three species of dryland mammal: plains mouse (Pseudomys australis), central rock‐rat (Zyzomys pedunculatus), and spinifex hopping‐mouse (Notomys alexis). We suggest that future research should focus on understanding the species‐specific nature of refuge use and the spatial ecology of refuges with a focus on connectivity and potential metapopulation dynamics. Assessing refuge quality and understanding the threats to high‐quality refuge patches and habitat should also be a priority. To facilitate this understanding we develop a three‐step methodology for determining species‐specific refuge location and habitat attributes. This review is necessarily focussed on dryland mammals in continental Australia where most refuge‐based research has been undertaken. The applicability of the refuge concept and the importance of refuges for dryland fauna conservation elsewhere in the world should be investigated. We predict that refuge‐using mammals will be widespread particularly among dryland areas with unpredictable rainfall patterns.     

Resource pulses affect prey selection and reduce dietary diversity of dingoes in arid Australia

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Influence of fire history on small mammal distributions: insights from a 100‐year post‐fire chronosequence

Aim Fire affects the structure and dynamics of ecosystems world‐wide, over long time periods (decades and centuries) and at large spatial scales (landscapes and regions). A pressing challenge for ecologists is to develop models that explain and predict faunal responses to fire at broad temporal and spatial scales. We used a 105‐year post‐fire chronosequence to investigate small mammal responses to fire across an extensive area of ‘tree mallee’ (i.e. vegetation characterized by small multi‐stemmed eucalypts). Location The Murray Mallee region (104,000 km²) of semi‐arid Australia. Methods First, we surveyed small mammals at 260 sites and explored the fire responses of four species using nonlinear regression models. Second, we assessed the predictive accuracy of models using cross‐validation and by testing with independent data. Third, we examined our results in relation to an influential model of animal succession, the habitat accommodation model. Results Two of four study species showed a clear response to fire history. The distribution of the Mallee Ningaui Ningaui yvonneae, a carnivorous marsupial, was strongly associated with mature vegetation characterized by its cover of hummock grass. The occurrence of breeding females was predicted to increase up to 40–105 years post‐fire, highlighting the extensive time periods over which small mammal populations may be affected by fire. Evaluation of models for N. yvonneae demonstrated that accurate predictions of species occurrence can be made from fire history and vegetation data, across large geographical areas. The introduced House Mouse Mus domesticus was the only species positively associated with recently burnt vegetation. Main conclusions Understanding the impact of fire over long time periods will benefit ecological and conservation management. In this example, tracts of long‐unburnt mallee vegetation were identified as important habitat for a fire‐sensitive native mammal. Small mammal responses to fire can be predicted accurately at broad spatial scales; however, a conceptual model of post‐fire change in community structure developed in temperate Australia is not, on its own, sufficient for small mammals in semi‐arid systems.     

Threatened mammals become more predatory after small‐scale prescribed fires in a high‐rainfall rocky savanna

Understanding mechanisms underlying fire regime effects on savanna fauna is difficult because of a wide range of possible trophic interactions and feedbacks. Yet, understanding mechanisms underlying fauna dynamics is crucial for conservation management of threatened species. Small savanna mammals in northern Australia are currently undergoing widespread declines and regional extinctions partly attributable to fire regimes. This study investigates mammal trophic and ecosystem responses to fire in order to identify possible mechanisms underlying these declines. Mammal trophic responses to fire were investigated by surveying mammal abundance, mammal diet, vegetation structure and non‐mammal fauna dynamics in savannas six times at eight sites over a period of 3 years. Known site‐specific fire history was used to test for trophic responses to post‐fire interval and fire frequency. Mammal and non‐mammal fauna showed only minor responses of post‐fire interval and no effect of fire frequency. Lack of fauna responses differed from large post‐fire vegetation responses. Dietary analysis showed that two mammal species, Dasyurus hallucatus and Isoodon auratus, increased their intake of large prey groups in recently burnt, compared to longer unburnt vegetation. This suggests a fire‐related change in trophic interactions among predators and their prey, after removal of ground‐layer vegetation. No evidence was found for other changes in food resource uptake by mammals after fire. These data provide support for a fire‐related top‐down ecosystem response among savanna mammals, rather than a bottom‐up resource limitation response. Future studies need to investigate fire responses among other predators, including introduced cats and dingoes, to determine their roles in fire‐related mammal declines in savannas of northern Australia.     

Testing top‐down and bottom‐up effects on arid zone beetle assemblages following mammal reintroduction

Species extinctions and declines are occurring globally and commonly have cascading effects on ecosystems. In Australia, mammal extinctions have been extensive, particularly in arid areas, where precipitation drives ecosystems. Many ecologically extinct mammals feed on soil‐dwelling insects. However, how this top‐down pressure affected their prey and how this contrasts with the bottom‐up impacts of fluctuating precipitation remains unclear. We constructed a long‐term exclusion experiment in a multi‐species mammal reintroduction zone in semi‐arid Australia to test how top‐down (reintroduced mammals) and bottom‐up (precipitation) factors affect root‐feeding chafer beetles (Coleoptera: Melolonthinae). We used emergence traps in ten replicate 20 × 20 m plots of control, exclusion and procedural control treatments to trap chafers biannually from 2009 to 2015. Annual precipitation during this period varied from 173 to 481 mm. Mammal exclusion did not affect chafers, indicating that top‐down regulation was not important. Instead, chafer abundance, species density and biomass increased with precipitation. Chafer body size and assemblage composition were best predicted by sampling year, suggesting that random drift determined species abundances. Increased resource availability therefore favoured all species similarly. We thus found no evidence that mammal predation alters chafer populations and conclude that they may be driven primarily by bottom‐up processes. Further research should determine if the cascading effects of species loss are less important for herbivores generally than for higher level trophic groups and the role of ecosystem stability in mediating these patterns.     

Not all predators are equal: a continent‐scale analysis of the effects of predator control on Australian mammals

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Arid Recovery – A comparison of reptile and small mammal populations inside and outside a large rabbit,cat and fox‐proof exclosure in arid South Australia

Australian arid zone mammal species within the Critical Weight Range (CWR) of 35 g–5.5 kg have suffered disproportionately in the global epidemic of contemporary faunal extinctions. CWR extinctions have been attributed largely to the effects of introduced or invasive mammals; however, the impact of these threatening processes on smaller mammals and reptiles is less clear. The change in small mammal and reptile assemblages after the removal of rabbits, cats and foxes was studied over a 6‐year period in a landscape‐scale exclosure in the Australian arid zone. Rodents, particularly Notomys alexis and Pseudomys bolami, increased to 15 times higher inside the feral‐proof Arid Recovery Reserve compared with outside sites, where rabbits, cats and foxes were still present. Predation by cats was thought to exert the greatest influence on rodent numbers owing to the maintenance of the disparity in rodent responses through dry years and the differences in dietary preferences between rabbits and P. bolami. The presence of introduced Mus domesticus or medium‐sized re‐introduced mammal species did not significantly affect resident small mammal or reptile abundance. Abundance of most dasyurids and small lizards did not change significantly after the removal of feral animals although reductions in gecko populations inside the reserve may be attributable to second order trophic interactions or subtle changes in vegetation structure and cover. This study suggests that populations of rodent species in northern South Australia below the CWR may also be significantly affected by introduced cats, foxes and/or rabbits and that a taxa specific model of Australian mammal decline may be more accurate than one based on body weight.     

Communities at the extreme: Aquatic food webs in desert landscapes

Studying food webs across contrasting abiotic conditions is an important tool in understanding how environmental variability impacts community structure and ecosystem dynamics. The study of extreme environments provides insight into community‐wide level responses to environmental pressures with relevance to the future management of aquatic ecosystems. In the western Lake Eyre Basin of arid Australia, there are two characteristic and contrasting aquatic habitats: springs and rivers. Permanent isolated Great Artesian Basin springs represent hydrologically persistent environments in an arid desert landscape. In contrast, hydrologically variable river waterholes are ephemeral in space and time. We comprehensively sampled aquatic assemblages in contrasting ecosystem types to assess patterns in community composition and to quantify food web attributes with stable isotopes. Springs and rivers were found to have markedly different invertebrate communities, with rivers dominated by more dispersive species and springs associated with species that show high local endemism. Qualitative assessment of basal resources shows autochthonous carbon appears to be a key basal resource in both types of habitat, although the particular sources differed between habitats. Food‐web variables such as trophic length, trophic breadth, and community isotopic niche size were relatively similar in the two habitat types. The basis for the similarity in food‐web structure despite differences in community composition appears to be broader isotopic niches for predatory invertebrates and fish in springs as compared with rivers. In contrast to published theory, our findings suggest that the food webs of the hydrologically variable river sites may show less dietary generalization and more compact food‐web modules than in springs.     

Leftover prey remains: a new taphonomic mode from the Late Miocene Cerro Azul Formation of Central Argentina

The Cerro Azul Formation (La Pampa Province, Argentina) comprises a rich vertebrate fauna of small mammals dominated by notoungulates and rodents. The fauna pertains to the Late Miocene specifically to the Huayquerian Stage/Age. Taphonomic analysis of micro‐mammals from Estancia Ré locality evidenced that the faunal assemblage was accumulated by the activity of a predator. This assemblage was compared with others from the Cerro Azul Formation in Telén and Caleufú localities (La Pampa Province), previously interpreted as products of predator activities. These microfossil accumulations differ from assemblages attributed to pellets and faeces produced by modern predators (nocturnal and diurnal bird raptors and carnivore mammals). However, due to their anatomical representation, degree of bone breakage and presence of tooth marks, they are interpreted as accumulations of uneaten prey remains discarded by the predator. The predator involved could not be determined with certainty, although the presence of tooth marks in some skeletal remains and the presence of coprolites in Telén and Caleufú suggest that it could be a carnivore mammal. Similarities in the accumulation mechanism, patterns of preservation and sedimentary contexts in the three assemblages support the recognition of a new taphonomic mode, termed ‘leftover prey remains’.     

Ecological correlates of small mammal community structure in the semi-arid Karoo, South Africa

The understanding of the determinants of small mammal community structure in arid and semiarid ecosystems is of importance, both in the light of the role that small mammals play, and the impact of livestock grazing on the flora of these systems. In a study aimed at identifying these determinants, small mammal assemblages and environmental features were quantified at six localities (a gradient of floristic structure, with constant annual rainfall) across the southern Karoo, South Africa. Stepwise variable regression indicated that small mammal diversity was correlated with plant and rock cover, as well as plant cover and horizontal foliage diversity at intermediate heights (40–60 cm). Initially, small mammal diversity increased with increasing plant cover, but decreased at cover levels greater than 30%. This relationship is similar to that found in other desert systems, although the peak in diversity found here is at higher levels of plant cover than found previously. I suggest that this higher peak may be owing to the lack of reliance on granivory by these animals, which are relatively omnivorous. This model may therefore explain the conflicting reports on the impact of livestock grazing on desert small mammals, with small mammal diversity decreasing with grazing below the peak, and increasing with grazing above the peak.     

Intraindividual variability in behavior shapes fitness landscapes

Although intraindividual variability (IIV) in behavior is fundamental to ecological dynamics, the factors that contribute to the expression of IIV are poorly understood. Using an individual‐based model, this study examined the effects of stochasticity on the evolution of IIV represented by the residual variability of behavior. The model describes a population of prey with nonoverlapping generations, in which prey take refuge upon encountering a predator. The strategy of a prey is characterized by the mean and IIV (i.e., standard deviation) of hiding duration. Prey with no IIV will spend the same duration hiding in a refuge at each predator encounter, while prey with IIV will have variable hiding durations among encounters. For the sources of stochasticity, within‐generation stochasticity (represented by random predator encounters) and between‐generation stochasticity (represented by random resource availability) were considered. Analysis of the model indicates that individuals with high levels of IIV are maintained in a population in the presence of between‐generation stochasticity even though the optimal strategy in each generation is a strategy with no IIV, regardless of the presence or absence of within‐generation stochasticity. This contradictory pattern emerges because the mean behavioral trait and IIV do not independently influence fitness (e.g., the sign of the selection gradient with respect to IIV depends on the mean trait). Consequently, even when evolution eventually leads toward a strategy with no IIV (i.e., the optimal strategy), greater IIV may be transiently selected. Between‐generation stochasticity consistently imposes such transient selection and maintain high levels of IIV in a population.     

Apex predator suppression is linked to restructuring of ecosystems via multiple ecological pathways

Removal of apex predators can drive ecological regime shifts owing to compensatory positive and negative population level responses by organisms at lower trophic levels. Despite evidence that apex predators can influence ecosystems though multiple ecological pathways, most studies investigating apex predators’ effects on ecosystems have considered just one pathway in isolation. Here, we provide evidence that lethal control of an apex predator, the dingo Canis dingo, drives shifts in the structure of Australia's tropical‐savannah ecosystems. We compared mammal assemblages and understorey structure at seven paired‐sites. Each site comprised an area where people poisoned dingoes and an area without dingo control. The effects of dingo control on mammals scaled with body size. Where dingoes were poisoned, we found greater activity of herbivorous macropods and feral cats, a mesopredator, but sparser understorey vegetation and lower abundances of native rodents. Our study suggests that ecological cascades arising from apex predators’ suppressive effects on herbivores and mesopredators occur simultaneously. Concordant effects of dingo removal across tropical‐savannah, forest and desert biomes suggest that dingoes once exerted ubiquitous top–down effects across Australia and provides support for calls that top–down forcing should be considered a fundamental process governing ecosystem structure.     

Comparative population dynamics of large and small mammals in the Northern Hemisphere: deterministic and stochastic forces

Deterministic feedbacks within populations interact with extrinsic, stochastic processes to generate complex patterns of animal abundance over time and space. Animals inherently differ in their responses to fluctuating environments due to differences in body sizes and life history traits. However, controversy remains about the relative importance of deterministic and stochastic forces in shaping population dynamics of large and small mammals. We hypothesized that effects of environmental stochasticity and density dependence are stronger in small mammal populations relative to their effects in large mammal populations and thus differentiate the patterns of population dynamics between them. We conducted an extensive, comparative analysis of population dynamics in large and small mammals to test our hypothesis, using seven population parameters to describe general dynamic patterns for 23 (14 species) time series of observations of abundance of large mammals and 38 (21 species) time series for small mammals. We used state‐space models to estimate the strength of direct and delayed density dependence as well as the strength of environmental stochasticity. We further used phylogenetic comparative analysis to detect differences in population dynamic patterns and individual population parameters, respectively, between large and small mammals. General population dynamic patterns differed between large and small mammals. However, the strength of direct and delayed density dependence was comparable between large and small mammals. Moreover, the variances of population growth rates and environmental stochasticity were greater in small mammals than in large mammals. Therefore, differences in population response to stochastic forces and strength of environmental stochasticity are the primary factor that differentiates population dynamic patterns between large and small mammal species.     

Is the loss of Australian digging mammals contributing to a deterioration in ecosystem function?

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Eco‐evolutionary feedback promotes Red Queen dynamics and selects for sex in predator populations

Although numerous hypotheses exist to explain the overwhelming presence of sexual reproduction across the tree of life, we still cannot explain its prevalence when considering all inherent costs involved. The Red Queen hypothesis states that sex is maintained because it can create novel genotypes with a selective advantage. This occurs when the interactions between species induce frequent environmental change. Here, we investigate whether coevolution and eco‐evolutionary feedback dynamics in a predator‐prey system allows for indirect selection and maintenance of sexual reproduction in the predator. Combining models and chemostat experiments of a rotifer‐algae system we show a continuous feedback between population and trait change along with recurrent shifts from selection by predation and competition for a limited resource. We found that a high propensity for sex was indirectly selected and was maintained in rotifer populations within environments containing these eco‐evolutionary dynamics; whereas within environments under constant conditions, predators evolved rapidly to lower levels of sex. Thus, our results indicate that the influence of eco‐evolutionary feedback dynamics on the overall evolutionary change has been underestimated.     

Are dingoes a trophic regulator in arid Australia? A comparison of mammal communities on either side of the dingo fence

The direct and indirect interactions that large mammalian carnivores have with other species can have far‐reaching effects on ecosystems. In recent years there has been growing interest in the role that Australia's largest terrestrial predator, the dingo, may have in structuring ecosystems. In this study we investigate the effect of dingo exclusion on mammal communities, by comparing mammal assemblages where dingoes were present and absent. The study was replicated at three locations spanning 300 km in the Strzelecki Desert. We hypothesized that larger species of mammal subject to direct interactions with dingoes should increase in abundance in the absence of dingoes while smaller species subject to predation by mesopredators should decrease in abundance because of increased mesopredator impact. There were stark differences in mammal assemblages on either side of the dingo fence and the effect of dingoes appeared to scale with body size. Kangaroos and red foxes were more abundant in the absence of dingoes while Rabbits and the Dusky Hopping‐mouse Notomys fuscus were less abundant where dingoes were absent, suggesting that they may benefit from lower red fox numbers in the presence of dingoes. Feral cats and dunnarts (Sminthopsis spp.) did not respond to dingo exclusion. Our study provides evidence that dingoes do structure mammal communities in arid Australia; however, dingo exclusion is also associated with a suite of land use factors, including sheep grazing and kangaroo harvesting that may also be expected to influence kangaroo and red fox populations. Maintaining or restoring populations of dingoes may be useful strategies to mitigate the impacts of mesopredators and overgrazing by herbivores.     

Can protective attributes of artificial refuges offset predation risk in lizards?

Artificial refuges are often used to supplement habitat in areas where natural shelters have been degraded or removed. Although artificial refuges are intended to support particular species, they may be equally attractive and accessible to others, including predators. We explored the influence of snake predation risk and shelter attributes on the overnight use of different artificial refuges (timber, tiles, and iron) using the predator‐prey relationship between Boulenger's skink, Morethia boulengeri and the curl snake, Suta suta. We collected adult M. boulengeri from two bioregions in south‐eastern Australia: the Riverina, where the two species co‐occur, and the South Western Slopes, where S. suta does not occur. Two adult S. suta were collected for use as chemical donors. We conducted four experiments on overnight refuge choice to determine: (i) predator‐scent avoidance, (ii) artificial refuge preferences, (iii) a trade‐off between a preferred refuge and predator‐avoidance, and (iv) the effect of gap height on refuge preference. We found that skinks avoided predator‐scented refuges in favour of identical, but unscented refuges. Skinks preferred timber refuges, and most skinks maintained this preference when predator‐scent was added. However, when gap height was manipulated, skinks shifted to the refuge with the smallest gap. Skinks displayed complex regional variation in behaviour; skinks from both bioregions avoided predator‐scent, but in the trade‐off experiment, skinks from the South Western Slopes were less likely to avoid predator‐scented timber refuges than those from the Riverina. Our findings suggest that protective refuge attributes, such as small gap height, can offset the risk implied by predator‐scent within a refuge. This study highlights the need to consider predator‐prey interactions when designing and using artificial refuges for habitat restoration or biological monitoring.     

Primary productivity can affect mammalian body size frequency distributions

Frequency distributions of mammal body sizes in large‐scale assemblages have often been found to show a positive skew. In an attempt to explain this pattern, a model has been put forward which incorporates energetic constraints on fitness and thereby predicts optimal body sizes corresponding to the mode of the distribution. A key assumption of the model is that energy is unlimited. However, if energy is limited, the input of energy into a herbivorous mammal community should influence the shape of the frequency distribution. Thus, I propose that increases in primary productivity will decrease the variation of body size and increase the mean body size in a distribution. So, in low‐productivity environments we should see a predominance of small‐sized species, but with a great variation of body sizes due to limitations of resources (energy). I tested this hypothesis using the herbivorous mammal fauna (rodents, bats and marsupials) in seven biomes of Australia. Because herbivorous marsupials generally are fairly large‐bodied while rodents and bats are small‐sized and because marsupials also have a different mode of reproduction from placental mammals, the hypothesis was also tested on placental mammals and marsupials separately. There was no clear mode for the entire assemblage in any biome, but as primary productivity increased, the variation of body masses decreased and the mean body mass of the distribution increased. Body mass distributions of both placental mammals and marsupials displayed clear modes. Placental mammals also showed an increase in mean body mass. The variation in body mass of marsupials was highest for the intermediately productive biomes. Primary productivity does seem to have some effect on mammalian body mass in this case, but the results here need to be complemented with studies of other assemblages before any general conclusions can be drawn. It is also important to distinguish which taxa are affected in a heterogeneous assemblage like the Australian herbivorous mammal fauna.     

Can recreational hunting contribute to pest mammal control on public land in Australia?

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