Application of the integrated threat theory to conservation law enforcement.

Interactions between law enforcement agents in conservation (e.g., rangers) and illegal resource users (e.g., illegal hunters) can be violent and sometimes fatal, which negatively affects conservation efforts and people's well-being. Models from social psychology, such as integrated threat theory (ITT) (intergroup interactions shape intergroup emotions, prejudices and perceived threats leading to hostile attitudes or behaviors between groups), are useful in addressing such interactions. Conservation approaches relying mainly on law enforcement have never been investigated using this framework. Using a structured questionnaire, we collected data from 282 rangers in protected and unprotected areas (n = 50) in northern Iran. We applied Bayesian structural equation modeling in an assessment of rangers' affective attitudes (i.e., emotions or feelings that shape attitudes toward a person or object) toward illegal hunters in an ITT framework. Rangers' positive perceptions of illegal hunters were negatively associated with intergroup anxiety (emotional response to fear) and negative stereotypes about a hunter's personality, which mediated the relationship between negative contact and affective attitudes. This suggests that negative contact, such as verbal abuse, may lead rangers to perceive illegal hunters as arrogant or cruel, which likely forms a basis for perceived threats. Rangers' positive contact with illegal hunters, such as playing or working together, likely lowered their perceived realistic threats (i.e., fear of property damage). Perceived realistic threats of rangers were positively associated with negative contacts (e.g., physical harm). The associations we identified suggest that relationships based on positive interactions between rangers and illegal hunters can reduce fear and prejudice. Thus, we suggest that rangers and hunters be provided with safe spaces to have positive interactions, which may help lower tension and develop cooperative conservation mechanisms.

Aplicación de la teoría integrada de la amenaza a la implementación de las leyes de conservación Resumen Las interacciones entre los agentes de la ley de la conservación (p. ej.: guardabosques) y los usuarios ilegales de recursos (p. ej.: cazadores ilegales) pueden ser violentas y a veces fatales, lo que afecta negativamente los esfuerzos de conservación y el bienestar de las personas. Los modelos de la psicología social, como la teoría integrada de la amenaza (TIA) (una amenaza percibida que deriva en prejuicios entre los grupos), tienen un uso potencial para tratar estas interacciones. Nunca se ha usado este marco para investigar las estrategias de conservación que dependen principalmente de la implementación de la ley. Usamos un cuestionario estructurado para recolectar datos de 282 guardabosques en áreas protegidas y no protegidas (n = 50) en el norte de Irán. Aplicamos el modelo de ecuación estructural bayesiano a la evaluación de las actitudes afectivas que tienen los guardabosques (es decir, emociones o sentimientos que forjan la actitud hacia una persona o un objeto) hacia los cazadores ilegales en un marco de TIA. La percepción negativa que tienen los guardabosques de los cazadores ilegales estuvo asociada negativamente con ansiedad intergrupal (la respuesta emocional al miedo) y estereotipos negativos de la personalidad de los cazadores, las cuales mediaron la relación entre el contacto negativo y las actitudes afectivas. Esto sugiere que el contacto negativo, como el abuso verbal, puede causar que los guardabosques perciban a los cazadores ilegales como arrogantes o crueles, lo que probablemente forma una base para las amenazas percibidas. El contacto positivo entre los guardabosques y los cazadores ilegales, como jugar o trabajar juntos, probablemente disminuyó la percepción de las amenazas realistas (es decir, miedo al daño material). La percepción que tienen los guardabosques de las amenazas realistas estuvieron asociadas positivamente con los contactos negativos (p. ej.: daño físico). Las asociaciones que identificamos sugieren que las relaciones basadas en las interacciones positivas entre los guardabosques y los cazadores ilegales pueden reducir el miedo y los prejuicios. Por lo tanto, sugerimos que se les proporcionen espacios seguros a los guardabosques y a los cazadores ilegales para que puedan tener interacciones positivas, lo que podría ayudar a reducir tensiones y a desarrollar mecanismos cooperativos de conservación.

Has the Australian Endemic Grey Falcon the Most Extreme Dietary Specialization among all Falco Species?

A clear understanding of a species' diet is crucial in understanding its spatio-temporal dynamics, and is, therefore, pertinent to conservation considerations. The diet of the Grey Falcon (Falco hypoleucos), a rare and threatened predator endemic to the Australian arid and semi-arid zone, is subject to diverging assertions; therefore, we studied its diet through direct observation of food ingestion during more than 17 years of fieldwork across the species' distribution. We found that Grey Falcons of all ages fed almost exclusively on a single type of food, namely, birds, and non-avian food items never constituted a substantial portion of any individual's diet. The extraordinary circumstances that were associated with the ingestion of non-avian food suggest strongly that, across its vast distribution, throughout the year, and throughout its life, the Grey Falcon feeds almost exclusively on birds. Further, we compared the diets of all Falco species and found that the dietary specialization is most extreme in the Grey Falcon, more so than even in the Taita (F. fasciinucha) and Peregrine Falcons (F. peregrinus). Based on aspects of the species' environment and relative prey availability, we offer an evolutionary explanation of the apparently unique dietary specialization of the arid-adapted Grey Falcon.

Low activity levels are an adaptation to desert-living in the Grey Falcon, an endotherm that specializes in pursuing highly mobile prey.

Endothermic animals that live permanently in hot deserts must avoid harmful hyperthermia when their body temperature increases from heat gained through external and internal sources. This is true particularly for endotherms that are exclusively diurnal. We investigated the Grey Falcon (Falco hypoleucos), a predatory Australian endemic restricted to the hot arid/semi-arid zone. To understand how this species' entire population persists exclusively and permanently in this extreme environment we examined its activity levels and compared these with equivalent variables from the Peregrine Falcon (F. peregrinus), a cosmopolitan species that inhabits similar environments without being restricted to them. Further, we compared, across a selected group of Falco species, specific plumage characteristics (measured on museum specimens) that we anticipated would enhance the Grey Falcons' ability to cope with high heat loads. We found no morphological or physiological characteristics that would allow them to cope with heat better than other birds, but the chicks seem to have unusually high thermal tolerances. Grey Falcons do, however, possess a suite of unusual behavioural adaptations that, as we propose, enable them to cope with climatic extremes in arid environments. Specifically, throughout their lives Grey Falcons keep activity levels and thus physical exertion low. This behaviour contrasts strikingly with that of the Peregrine Falcon, which also actively hunts birds in flight. Keeping activity levels low is expected to minimize endogenous heat production and thus ease the Grey Falcon's thermoregulation during periods of high heat load. These birds may rely on low levels of relative humidity for efficient evaporative cooling, and this may explain their absolute restriction to hot arid/semi-arid zones.

Delayed juvenile behavioral development and prolonged dependence are adaptations to desert life in the grey falcon.

Rapid learning in the young of most endothermic animals can be expected to be favored by natural selection because early independence reduces the period of vulnerability. Cases of comparatively slow juvenile development continue, therefore, to attract scientific attention. In most species of birds, including raptors, the young depend on their parents for some time after fledging for the provisioning of food and for protection while they learn to become nutritionally and otherwise independent. Among raptors, post-fledging dependence periods that exceed 6 months are exclusive to the largest species and these have reproductive cycles that exceed 12 months. By contrast, young of the medium-sized grey falcon Falco hypoleucos have been reported in close company with their parents up to 12 months after fledging, that is, at a time when the adults are expected to breed again. We investigated the occurrence and characteristics of prolonged adult-juvenile association relative to other falcons and similar-sized raptors. We found that the behavioral development of grey falcon young is extremely delayed, and that they even depend nutritionally on their parents for up to 12 months after fledging. We suggest that these 2 distinctive features are, ultimately, adaptations of the grey falcon to its extreme environment, Australia's arid and semi-arid zone, one of the hottest environments in the world.

A nomadic avian predator displays flexibility in prey choice during episodic outbreaks of rodents in arid Australia.

In environments driven by unpredictable resource pulses, populations of many consumer species experience dramatic fluctuations in abundance and spatial extent. Predator-prey relationships in these acyclic systems are poorly understood in particular with respect to the level of prey specialisation shown by nomadic predators. To understand the dynamics of such a system I examined the response to rodent outbreaks by the letter-winged kite (Elanus scriptus) in the Simpson Desert, Australia; a region that experiences major pulses in primary productivity, driven by unpredictable rainfall events. The kite feeds on small mammals and is the only night-hunting species in the Accipitridae. Letter-winged kites irrupted in the area on only three occasions during 20 years of sampling (1999-2019) and remained for a maximum of 20 months. Each period of kite occupation occurred only during the increase and/or peak phase of rodent population cycles (which occurred three times during the study). During each period kite diet was dominated by small (10-50 g body mass) quadrupedal rodents (Pseudomys australis, P. hermannsburgensis, Mus musculus). Abundance of these species varied across the three outbreaks and kites typically captured them in proportion to availability. The large body mass (134 g) long-haired rat (Rattus villosissimus) was abundant during one outbreak but was infrequently consumed. The bipedal spinifex hopping-mouse (Notomys alexis) was within the kites' favoured prey size range (35 g) but was consistently avoided. The flexibility in prey selection by letter-winged kites appears to be an important adaptation for survival and reproduction by species exploiting acyclic rodent outbreaks.

Dynamics of bird assemblages in response to temporally and spatially variable resources in arid Australia.

Bird assemblages in arid Australia are often characterized as being highly variable through time in response to boom and bust dynamics, although the importance of habitat in structuring assemblages at a local-scale is also recognized. We use a novel approach to investigate the importance of rainfall variability in structuring bird assemblages in a resource-limited environment. Monthly bird surveys were conducted at ten plots for 8 years at a botanical and zoological park in central Australia, including five irrigated plots within a fenced area and five natural plots outside. Irrigation-used to promote growth, flowering, and fruiting of plants-created an artificial resource-enhanced environment against which the response of birds to natural fluctuations in season and rainfall were compared. Species richness was generally maintained at a higher level in resource-enhanced plots during dry times but was higher in natural plots when rainfall was high, mainly due to increases in granivores and insectivores. Honeyeaters were consistently more abundant at irrigated plots. Rainfall was important in structuring bird assemblages at all plots; however, assemblages were more stable in irrigated plots and did not respond as dramatically to a period of very high rainfall. The comparative smoothing of fluctuations in the composition and abundance of birds in irrigated areas highlights the importance of primary productivity, normally tied to rainfall, in driving temporal change in arid-zone bird communities. There was also evidence that different plots in differing habitats supported distinct bird assemblages and that this spatial distinctiveness persisted irrespective of rainfall and determined, to some extent, the response to rainfall. Our study is one of few long-term studies of arid bird assemblages and highlights the importance of both long-term cycles of productivity driven by rain and season as well as site differences in the dynamics of arid-zone bird communities. These insights are particularly valuable as climate change further exacerbates rainfall variability worldwide and initiatives to conserve avifauna in increasingly extreme environments may be required.

Are physiological and behavioural responses to stressors displayed concordantly by wild urban rodents?

Understanding wild animal responses to stressors underpins effective wildlife management. In order for responses to stressors to be correctly interpreted, it is critical that measurements are taken on wild animals using minimally invasive techniques. Studies investigating wild animal responses to stressors often measure either a single physiological or behavioural variable, but whether such responses are comparable and concordant remains uncertain. We investigated this question in a pilot study that measured responses of wild-caught urban brown and black rats (Rattus norvegicus, Rattus rattus) to fur-based olfactory cues from a predator, the domestic cat (Felis catus); a novel herbivore, the koala (Phascolarctos cinereus); and a familiar herbivore and competitor, the common brushtail possum (Trichosurus vulpecula). Physiological responses, measured by assaying faecal glucocorticoid metabolites, were compared to behavioural responses observed via video recordings. We found that physiological and behavioural responses to stressors were expressed concordantly. There was no sizeable physiological response observed, and the behavioural response when considered across the night was negligible. However, the behavioural response to the predator and competitor cues changed across the observation period, with activity increasing with increasing hours of exposure. Our results indicate that responses of wild rodents to cues are nuanced, with stress responses modulated by behaviour changes that vary over time according to the severity of the perceived threat as animals gather further information. If the physiological response alone had been assessed, this moderated response may not have been evident, and in terms of wildlife management, vital information would have been lost.

Fear and stressing in predator-prey ecology: considering the twin stressors of predators and people on mammals.

Predators induce stress in prey and can have beneficial effects in ecosystems, but can also have negative effects on biodiversity if they are overabundant or have been introduced. The growth of human populations is, at the same time, causing degradation of natural habitats and increasing interaction rates of humans with wildlife, such that conservation management routinely considers the effects of human disturbance as tantamount to or surpassing those of predators. The need to simultaneously manage both of these threats is particularly acute in urban areas that are, increasingly, being recognized as global hotspots of wildlife activity. Pressures from altered predator-prey interactions and human activity may each initiate fear responses in prey species above those that are triggered by natural stressors in ecosystems. If fear responses are experienced by prey at elevated levels, on top of responses to multiple environmental stressors, chronic stress impacts may occur. Despite common knowledge of the negative effects of stress, however, it is rare that stress management is considered in conservation, except in intensive ex situ situations such as in captive breeding facilities or zoos. We propose that mitigation of stress impacts on wildlife is crucial for preserving biodiversity, especially as the value of habitats within urban areas increases. As such, we highlight the need for future studies to consider fear and stress in predator-prey ecology to preserve both biodiversity and ecosystem functioning, especially in areas where human disturbance occurs. We suggest, in particular, that non-invasive in situ investigations of endocrinology and ethology be partnered in conservation planning with surveys of habitat resources to incorporate and reduce the effects of fear and stress on wildlife.

Habitat as a mediator of mesopredator-driven mammal extinction.

A prevailing view in dryland systems is that mammals are constrained by the scarcity of fertile soils and primary productivity. An alternative view is that predation is a primary driver of mammal assemblages, especially in Australia, where 2 introduced mesopredators-feral cat (Felis catus) and red fox (Vulpes vulpes)-are responsible for severe declines of dryland mammals. We evaluated productivity and predation as drivers of native mammal assemblage structure in dryland Australia. We used new data from 90 sites to examine the divers of extant mammal species richness and reconstructed historic mammal assemblages to determine proportional loss of mammal species across broad habitat types (landform and vegetation communities). Predation was supported as a major driver of extant mammal richness, but its effect was strongly mediated by habitat. Areas that were rugged or had dense grass cover supported more mammal species than the more productive and topographically simple areas. Twelve species in the critical weight range (CWR) (35-5500 g) that is most vulnerable to mesopredator predation were extirpated from the continent's central region, and the severity of loss of species correlated negatively with ruggedness and positively with productivity. Based on previous studies, we expect that habitat mediates predation from red foxes and feral cats because it affects these species' densities and foraging efficiency. Large areas of rugged terrain provided vital refuge for Australian dryland mammals, and we predict such areas will support the persistence of CWR species in the face of ongoing mammal declines elsewhere in Australia.

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.

Alignment between values of dryland pastoralists and conservation needs for small mammals.

Policies for conservation outside protected areas, such as those designed to address the decline in Australian mammals, will not result in net improvements unless they address barriers to proenvironmental behavior. We used a mixed-methods approach to explore potential value-action gaps (disconnects between values and subsequent action) for small mammal conservation behaviors among pastoralists in dryland Australia. Using semistructured surveys and open-ended interviews (n = 43), we explored values toward small mammals; uptake of a range of current and intended actions that may provide benefit to small mammals; and potential perceived barriers to their uptake. Pastoralists assigned great conservation value to small mammals; over 80% (n = 36) agreed to strongly agreed that small mammals on their property were important. These values did not translate into stated willingness to engage in voluntary cessation of wild-dog control (r2 = 0.187, p = 0.142, n = 43). However, assigning great conservation value to small mammals was strongly related to stated voluntary willingness to engage in the proenvironmental behavior most likely to result in benefits to small mammals: cat and fox control (r2 = 0.558, p = 0.000, n = 43). There was no significant difference between stated voluntarily and incentivized willingness to engage in cat and fox control (p = 0.862, n = 43). The high levels of willingness to engage in voluntary cat and fox control highlight a potential entry point for addressing Australia's mammal declines because the engagement of pastoralists in conservation programs targeting cat and fox control is unlikely to be prevented by attitudinal constraints. Qualitative data suggest there is likely a subpopulation of pastoralists who value small mammals but do not wish to engage in formal conservation programs due to relational barriers with potential implementers. A long-term commitment to engagement with pastoralists by implementers will thus be necessary for conservation success. On-property cat and fox control programs that build and leverage trust, shared goals, collaboration, and shared learning experiences between stakeholders and that explicitly recognize the complexity of small mammal dynamics and the property-level ecological knowledge of pastoralists are more likely to gain traction.

Photographic identification of individuals of a free-ranging, small terrestrial vertebrate.

Recognition of individuals within an animal population is central to a range of estimates about population structure and dynamics. However, traditional methods of distinguishing individuals, by some form of physical marking, often rely on capture and handling which may affect aspects of normal behavior. Photographic identification has been used as a less-invasive alternative, but limitations in both manual and computer-automated recognition of individuals are particularly problematic for smaller taxa (<500 g). In this study, we explored the use of photographic identification for individuals of a free-ranging, small terrestrial reptile using (a) independent observers, and (b) automated matching with the Interactive Individual Identification System (I(3)S Pattern) computer algorithm. We tested the technique on individuals of an Australian skink in the Egernia group, Slater's skink Liopholis slateri, whose natural history and varied scale markings make it a potentially suitable candidate for photo-identification. From 'photographic captures' of skink head profiles, we designed a multi-choice key based on alternate character states and tested the abilities of observers - with or without experience in wildlife survey - to identify individuals using categorized test photos. We also used the I(3)S Pattern algorithm to match the same set of test photos against a database of 30 individuals. Experienced observers identified a significantly higher proportion of photos correctly (74%) than those with no experience (63%) while the I(3)S software correctly matched 67% as the first ranked match and 83% of images in the top five ranks. This study is one of the first to investigate photo identification with a free-ranging small vertebrate. The method demonstrated here has the potential to be applied to the developing field of camera-traps for wildlife survey and thus a wide range of survey and monitoring applications.

Prey availability affects daily torpor by free-ranging Australian owlet-nightjars (Aegotheles cristatus).

Food availability, ambient temperatures (T(a)), and prevailing weather conditions have long been presumed to influence torpor use. To a large extent, this is based on measurements in the laboratory of animals placed on restricted diets and kept at low T (a). Information on the determinants of torpor employment in the field is limited. We assessed winter torpor by insectivorous, free-ranging Australian owlet-nightjars (Aegotheles cristatus; 22 birds, 834 bird-days over six winters). Birds in three habitats were investigated to test whether torpor use is affected by annual T(a), rainfall, and arthropod abundance. Owlet-nightjars entered daily torpor regularly at all sites. Torpor frequency, depth and bout duration were greatest during two periods with lower arthropod abundance, providing rare evidence of the link between food availability and torpor patterns of wild birds. Temporal organization of torpor was similar among sites, and nocturnal torpor was more frequent than previously reported. Our findings quantitatively demonstrate that reduced food resources affect torpor usage independently from T(a), and support the view that food availability is a primary ecological determinant of torpor use in the wild.

Roost type influences torpor use by Australian owlet-nightjars.

Australian owlet-nightjars (Aegotheles cristatus; ∼50 g) are one of only a few avian species that roost in cavities year-round and regularly enter torpor. Cavity roosts act as thermal buffers, and roost type likely affects energy expenditure of small birds. We used radiotelemetry to locate diurnal winter roost sites of owlet-nightjars in central Australia and to measure body (T (b)) and skin (T (skin)) temperature. We also recorded ambient temperature inside (T (IN)) and outside roosts. Individual owlet-nightjars used one to seven different roosts (tracking time 3-10 weeks), selecting either rock crevices (four birds) or tree hollows (four birds), or switching between the two roost types (seven birds). Rock crevices (T (IN) +9°C to +33°C) were warmer and thermally more stable than tree hollows (T (IN) -4.0°C to +37°C). Torpor, often expressed by a reduction of T (skin)/T (b) by >10°C for 3-4 h at dawn, was influenced by roost selection; torpor use in tree hollows was almost twice that in rock crevices. Despite the potential energy savings accrued from roosting in well-insulated cavities, owlet-nightjars roosted in tree hollows more often (65% bird days, n = 398) than in rock crevices (35% bird days, n = 211). Lower costs of arousal from torpor via passive rewarming and basking and decreased risk of predation are two possible explanations for the preference to roost in tree hollows. We provide the first evidence for the influence of cavity roost selection on torpor use in a free-ranging bird and show that roost selection and thermal biology are strongly interrelated in determining energy expenditure.

Torpor and activity in a free-ranging tropical bat: implications for the distribution and conservation of mammals?

Bats are most diverse in the tropics, but there are no quantitative data on torpor use for energy conservation by any tropical bat in the wild. We examined the thermal biology, activity patterns and torpor use of two tree-roosting long-eared bats (Nyctophilus geoffroyi, 7.8 g) in tropical northern Australia in winter using temperature telemetry. Bats commenced activity about 20 min after sunset, ended activity about 2.5 h before sunrise and entered torpor everyday in the early morning even when minimum ambient temperatures (T (a)) were as high as 23°C. On average, bats remained torpid for almost 5 h, mean minimum skin temperature (T (skin)) measured was 22.8 ± 0.1°C and daily T (skin) minima were correlated with T(a). Our study shows that even in the tropics, torpor is frequently employed by bats, suggesting that worldwide most bat species are heterothermic and use torpor for energy conservation. We propose that the ability of employing torpor and the resulting highly plastic energy requirements may partially explain why these small insectivorous bats can inhabit almost the entire Australian continent despite vastly different climatic and likely trophic conditions. Reduced energy requirements also may permit survival in degraded or modified habitats, reduce the need for foraging and reduce exposure to predators. Thus, the ability to employ torpor may be one important reason for why most Australian bats and other heterothermic mammals have not gone extinct whereas many obligatory homeothermic mammals that cannot employ torpor and have high energy and foraging requirements have suffered high rates of extinctions.

Vertebrate diet decreases winter torpor use in a desert marsupial.

One of the energetic benefits of daily torpor over prolonged hibernation is that it enables animals to regularly forage and, therefore, replenish food reserves between bouts of torpor. However, little is known about the diet of predators undergoing torpor or whether differences in prey composition among individuals influence torpor characteristics. Here, we test the hypothesis that prey composition affects winter torpor use and patterns of a population of carnivorous marsupial, the brush-tailed mulgara (Dasycercus blythi), in the Great Sandy Desert, Australia. Mulgaras in the study population captured a wide range of prey including vertebrates (mammals, reptiles, birds), seven insect orders, spiders and centipedes. The proportion of vertebrates in the diet was negatively correlated with both frequency of torpor use and maximum bout duration. This variation in torpor use with diet can be explained by the higher energetic content of vertebrates as well as their larger size. Even assuming uniform intake of prey biomass among individuals, those that subsisted on an invertebrate-dominated diet during winter apparently suffered energetic shortages as a result of the scarcity of invertebrate taxa with high energy content (such as insect larvae). Our study is the first to demonstrate a link between diet composition and daily torpor use in a free-ranging mammal.

Surviving cave bats: auditory and behavioural defences in the Australian noctuid moth, Speiredonia spectans.

The Australian noctuid moth, Speiredonia spectans shares its subterranean day roosts (caves and abandoned mines) with insectivorous bats, some of which prey upon it. The capacity of this moth to survive is assumed to arise from its ability to listen for the bats' echolocation calls and take evasive action; however, the auditory characteristics of this moth or any tropically distributed Australian moth have never been examined. We investigated the ears of S. spectans and determined that they are among the most sensitive ever described for a noctuid moth. Using playbacks of cave-recorded bats, we determined that S. spectans is able to detect most of the calls of two co-habiting bats, Rhinolophus megaphyllus and Miniopterus australis, whose echolocation calls are dominated by frequencies ranging from 60 to 79 kHz. Video-recorded observations of this roost site show that S. spectans adjusts its flight activity to avoid bats but this defence may delay the normal emergence of the moths and leave some 'pinned down' in the roosts for the entire night. At a different day roost, we observed the auditory responses of one moth to the exceptionally high echolocation frequencies (150-160 kHz) of the bat Hipposideros ater and determined that S. spectans is unable to detect most of its calls. We suggest that this auditory constraint, in addition to the greater flight manoeuvrability of H. ater, renders S. spectans vulnerable to predation by this bat to the point of excluding the moth from day roosts where the bat occurs.

Thermal biology, torpor, and activity in free-living mulgaras in arid zone Australia during the winter reproductive season.

Little is known about the energy conservation strategies of free-ranging marsupials living in resource-poor Australian deserts. We studied activity patterns and torpor of free-living mulgaras (Dasycercus blythi) in arid central Australia during the winter of 2006. Mulgaras are small (approximately 75 g), nocturnal, insectivorous marsupials, with a patchy distribution in hummock grasslands. Mulgaras (six males, three females) were implanted intraperitoneally with temperature-sensitive transmitters and monitored for 6-55 d. Temperature profiles for different microhabitats and the thermal properties of soil and a number of burrows were also measured. Air temperature ranged from -3 degrees C at night to 30 degrees C during the day. Although burrows buffered temperature extremes, the thermal diffusivity of the sandy soil was high, and many burrows were shallow. Hence, soil and burrow temperatures averaged about 15 degrees C. The activity of mulgaras was often restricted to a few hours after sunset, before they retired into their burrows. Mulgaras employed torpor frequently, often entering torpor during the night and arousing around midday, with arousals occurring later on cooler days. Shallow burrows allowed cooling below mean T(soil). Consequently, body temperatures as low as 10.8 degrees C were observed. The longest torpor bout was 20.8 h. Torpor patterns changed seasonally and differed between males and females. From June to August, females entered torpor almost daily despite mating and gestation, but from the end of the gestation period onward, they remained normothermic. In contrast, males showed only shallow and short torpor during the mating season, but from mid-July, a transition to more frequent and deeper torpor resembling that of females was observed. Apparently, in both sexes, the reproductive effort entails energetic costs, but torpor, as an energy-saving mechanism, and reproduction are not exclusive in mulgaras. In a resource-poor environment during the least productive part of the year, frequent torpor seems to provide the means to compensate for the increased energetic costs associated with reproduction.

Basking and torpor in a rock-dwelling desert marsupial: survival strategies in a resource-poor environment.

Australian deserts are characterized by unpredictability, low primary productivity, and high temperature fluctuations. Despite these adverse conditions the diversity of small insectivorous marsupials of the family Dasyuridae is surprisingly high. We quantified the thermal biology of the dasyurid Pseudantechinus madonnellensis (body mass approximately 30 g) in the wild to gain some understanding of whether the success of dasyurids in the arid zone may be related to some extent to their use of energy conservation strategies. In winter, most free-ranging Pseudantechinus frequently (58.3% of 131 animal days) entered daily torpor after midnight (mean 0157 hours) in rock crevices when outside ambient temperatures (T (a)) were low. Most animals remained torpid until the next morning when they moved while still torpid from rock crevices to sun-exposed basking sites. We visually observed basking during rewarming from torpor (mean commencement at 0943 hours) at body temperatures (T (b)) as low as 19.3 degrees C when radiant heat was high and T (a) was rising. Basking continued for the rest of the day. Torpor use was not strongly correlated with T (a), but the temporal organization of daily torpor and activity were apparently linked to the thermal characteristics of basking sites. Our study suggests that by frequently employing daily torpor and basking and by appropriately coordinating their thermal biology with that of specific locations in their environment, Pseudantechinus can reduce daily energy expenditure and thus can live and reproduce in a challenging environment. It is likely that the success of other small dasyurids and perhaps many other small mammals living in deserts is linked to employment of torpor and basking for energy conservation.