Thermal biology,torpor and behaviour in sugar gliders: a laboratory-field comparison

Most studies on animal physiology and behaviour are conducted in captivity without verification that data are representative of free-ranging animals. We provide the first quantitative comparison of daily torpor, thermal biology and activity patterns, conducted on two groups of sugar gliders (Petaurus breviceps, Marsupialia) exposed to similar thermal conditions, one in captivity and the other in the field. Our study shows that activity in captive gliders in an outdoor aviary is restricted to the night and largely unaffected by weather, whereas free-ranging gliders omit foraging on cold/wet nights and may also forage in the afternoon. Torpor occurrence in gliders was significantly lower in captivity (8.4% after food deprivation; 1.1% for all observations) than in the field (25.9%), mean torpor bout duration was shorter in captivity (6.9 h) than in the field (13.1 h), and mean body temperatures during torpor were higher in captivity (25.3°C) than in the field (19.6°C). Moreover, normothermic body temperature as a function of air temperature differed between captive and free-ranging gliders, with a >3°C difference at low air temperatures. Our comparison shows that activity patterns, thermal physiology, use of torpor and patterns of torpor may differ substantially between the laboratory and field, and provides further evidence that functional and behavioural data on captive individuals may not necessarily be representative of those living in the wild.     

Some like it cold: summer torpor by freetail bats in the Australian arid zone

Bats are among the most successful groups of Australian arid-zone mammals and, therefore, must cope with pronounced seasonal fluctuations in ambient temperature (T _a), food availability and unpredictable weather patterns. As knowledge about the energy conserving strategies in desert bats is scant, we used temperature-telemetry to quantify the thermal physiology of tree-roosting inland freetail bats (Mormopterus species 3, 8.5 g, n = 8) at Sturt National Park over two summers (2010–2012), when T _a was high and insects were relatively abundant. Torpor use and activity were affected by T _a. Bats remained normothermic on the warmest days; they employed one “morning” torpor bout on most days and typically exhibited two torpor bouts on the coolest days. Overall, animals employed torpor on 67.9 % of bat-days and torpor bout duration ranged from 0.5 to 39.3 h. At any given T _a, torpor bouts were longer in Mormopterus than in bats from temperate and subtropical habitats. Furthermore, unlike bats from other climatic regions that used only partial passive rewarming, Mormopterus aroused from torpor using either almost entirely passive (68.9 % of all arousals) or active rewarming (31.1 %). We provide the first quantitative data on torpor in a free-ranging arid-zone molossid during summer. They demonstrate that this desert bat uses torpor extensively in summer and often rewarms passively from torpor to maximise energy and water conservation.     

Induction of reproductive diapause in Bruchidius atrolineatus during the dry season in a Sahelian zone

Bruchidius atrolineatus (Pic) (Bruchidae: Coleoptera) is a bruchid beetle infesting Vigna unguiculata (Walp). During the dry season, adults are in reproductive diapause. Studies of adult populations in a traditional store containing contaminated pods of V. unguiculata show that the reproductive diapause is induced during the first larval instars. The temperature decrease from November to January probably is the main climatic factor inducing this diapause. The percentage of diapausing adults depends on the duration of the larval development at low temperatures. Experiments in climatic boxes confirm these results. At 12 h 25 °T-12 h 20° C 11–13 LD, a high proportion of the adults emerging from the V. unguiculata seeds are in reproductive diapause and the duration of their growth is variable. L1 L2 L3 larvae probably are sensitive stages to the thermal variations and the processes allowing emergence of diapausing adults probably are induced during these larval instars.

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Résumé Bruchidius atrolineatus (Pic) est un Coléoptère Bruchidae se développant en zone sahélienne aux dépens d'une légumineuse Vigna unguiculata. Pendant la saison sèche, les adultes sont en diapause reproductrice. Un suivi des populations d'adultes a été réalisé dans un grenier traditionnel en terre séchée contenant des gousses récoltées dans la nature et portant des ufs de B. atrolineatus. Il a permis de constater que la diapause reproductrice était induite durant les premières phases du développement larvaire. La décroissance des températures de Novembre à Janvier avec des minima de 15 ° C et des maxima de 25 °C est probablement le principal facteur permettant l'induction de cette diapause. L'introduction dans le grenier de gousses portant des ufs venant d'être émis à différentes périodes de la saison sèche montre que le pourcentage d'adultes diapausants dépend de la durée du développement larvaire aux températures basses. Des expériences réalisées dans des étuves programmées confirment ces résultats. A 12 h 25 °T/12 h 20 °C 11–13 LD, un taux élevé d'adultes émergeant des graines est en diapause reproductrice et la durée de leur développement post-embryonnaire est très variable. En conditions plus chaudes 12 h 40 °C/12 h 25 °C 11–13 LD, le taux d'adultes diapausants est plus réduit et la durée du développement plus courte. L'analyse de la croissance larvaire montre que les premiers stades larvaires sont probablement des stades sensibles aux variations thermiques. Les modifications physiologiques permettant l'émergence d'adultes diapausants ou sexuellement actifs seraient induites durant ces stades.

     

Thermal biology, torpor use and activity patterns of a small diurnal marsupial from a tropical desert: sexual differences

Many small desert dasyurids employ torpor almost daily during winter, because cold nights and low food availability impose high energetic costs. However, in Western Australia the arid zone extends into tropical, coastal regions, where winter temperature conditions are far less severe. We studied the thermal biology and activity patterns of free-ranging kaluta (~27 g), a dasyurid restricted to these tropical spinifex deserts, during the Austral winter (June–July) and in addition quantified activity patterns in captivity. Unlike most dasyurids, wild and captive kalutas were almost exclusively diurnal and retreated into underground burrows during the night. Despite being active during the warmer part of the day, kalutas entered torpor daily. However, torpor patterns differed remarkably between males and females. While females spent most of the night torpid at body temperatures (T _b) as low as 21°C, close to soil temperature, males entered multiple short and shallow bouts (T _b > 25°C) during the night. Males also maintained higher T _bs during the early morning when active, occupied larger home ranges and covered greater distances while foraging than females. Hence, males appear to expend more energy than the similar-sized females both while foraging and during the rest phase. We propose that physiological as well as behavioural preparations for the September mating season that culminate in a complete male die-off might already impose energetic costs on males during winter.     

Sex differences in the use of daily torpor and foraging time by big brown bats (Eptesicus fuscus) during the reproductive season

Temperate zone bats can use daily torpor as a means of saving energy. Some argue, however, that torpor is costly for both males and females and that individuals should only use it during times of poor foraging conditions. Others hypothesize that the costs are greater for females and that males should enter torpor more regularly. We tested these alternative hypotheses by using temperature-sensitive radiotransmitters to monitor use of torpor and foraging by free-ranging big brown bats ( Eptesicus fuscus ). During the pregnancy period, males used torpor at night more and foraged less often than did females. Males also went into deep torpor more often and remained in torpor longer than did females. When they foraged, males and females were away from the roosts for equal periods of time. During the lactation period, males and females rarely failed to forage and foraging times were again no different between the sexes, although males may roost at night away from the maternity colonies. Males again used torpor and deep torpor more often and for longer than females did. These results support the hypothesis that the fitness costs of using torpor are lower for males than for reproductive females and that males regularly use torpor as an energy-saving mechanism. Females enter torpor only when foraging conditions are poor, presumably because torpor prolongs gestation and slows neonatal growth thereby leaving less time for females and their young to prepare for hibernation.     

Energy expenditure increases during the active season in the small,free-living hibernator Muscardinus avellanarius

Little is known about strategies employed by small mammals to reduce energy expenditure during the summer. To understand whether ambient conditions impact euthermic energy demands in a small free-living hibernator, we measured metabolic rate of hazel dormice (Muscardinus avellanarius) in the field. Furthermore, we aimed to reveal which variables influence torpor use. Our results show that hazel dormice altered euthermic energy expenditure during summer but not as expected as a response to environmental conditions. Euthermic resting metabolic rate was lowest directly after emergence from hibernation and increased by about 95% until the end of August. A considerable part of this increase was presumably caused by the changing influence of gender and rain on energy demands during different months, variation in food quality and quantity, and reversible size changes of organs that had been atrophied during hibernation. Torpor use in hazel dormice occurred more frequently when it was colder, earlier during the day, and in lighter individuals. Torpor was used routinely in males and non-reproductive females. We show that torpor is used more frequently than previously suggested by studies that only used visual proof of torpor use by surveying nest boxes.     

Differential movement and activity patterns of sexes in a biparental beetle during the reproductive season

1. Biparental care is stabilised if parents perform different tasks during care. Specialised parental roles may require different time and energy budgets that in turn are expected to influence the activity and space use of sexes. 2. Here we investigate movement patterns of the biparental Lethrus apterus beetle using a grid of pitfall traps in their natural habitat. 3. Sexes of the burrow building L. apterus perform different roles during caregiving, as females collect most of the leaves, which serve as food for the offspring while paired males stay mostly in the burrow. We hypothesised that sex differences in mate search and parental activities are reflected in movement patterns. 4. We found that females frequently travelled short distances, whereas males were detected less often but when detected, they travelled significantly longer distances than females. 5. Our results are consistent with the notion that efficient parental food provisioning requires more localised movement and activity patterns. Furthermore, the long distance movements of some males may indicate active mate searching behaviour.     

Chronological changes in terrestrial insect assemblages in the arid zone of Australia

Arid and semiarid environments are characterized by highly unpredictable and 'pulsed' availability of essential biological resources. The 'boom and bust' response of many vertebrates is commonly invoked for invertebrates and especially insects. This perception of the Australian arid zone is exacerbated by the lack of long-term surveys of insects identified at high levels of taxonomic resolution. From an 18 mo continuous survey of insects in central Australia I determine the phenology of many insect taxa, and clarify which climatic variables most influenced the activity of these taxa. Total abundance and taxon richness were higher in the warmer months and lower in the cooler months. Minimum temperature, rainfall during the survey month, and rainfall during the previous month had significant effects on phenology, demonstrating that there is pronounced and predictable activity of many species in the absence of rain, although rainfall has a marked effect on the activity of some species. Other species were more active or only active in the coolest months. These findings have implications for the most productive time for surveys in the Australian arid zone, the availability of insects as prey or pollinators, and for the potential effects of climate change.     

Spatial indicators of fire risk in the arid and semi-arid zone of Australia

Fire plays a role in determining the shape of the earth's ecosystems, impacts socio-economic issues, and influences our climate. In arid and semi-arid Australia (70% of the continent), individual fires frequently exceed 1 million ha, and have collectively burnt up to 9% of the total area in a single year, associated with antecedent periods of above average rainfall which boost the fuel load. People affected by these fires – Federal and State governments, pastoralists, Aboriginal communities, larger towns, conservation park managers and tourist operators – all have different outlooks and priorities about these phenomena. Little objective information about the fire regime and its drivers has been available for this vast area with its very low population density. A predictive understanding of the spatial and temporal pattern of risk of large uncontrollable fires is needed to promote pro-active management.We present a conceptual framework which serves both to summarise existing knowledge and to reduce the complexity for a quantitative statistical analysis. This conceptual framework contains four main groups of independent variables; biomass, curing, ignition source, and fire weather. For these groups of variables we identified direct data sources or spatial surrogates. To quantify different aspects of the fire regime, interpretation of NOAA-AVHRR satellite imagery was employed, which identifies both fire hotspots (FHS) and fire affected area (FAA). For temporal variables, we present a surface displaying relationships for different combinations of lag/phase. This highlights different patterns for each region, and the most appropriate timeframes to use in modelling.Results of exploratory regression analysis in arid and semi-arid Australia show that the strongest influence is exerted by biomass or fuel load. As this is highly dependent on antecedent rainfall, we can anticipate a strong effect of climate change on the fire regime. The strongest combinations of relationships may be used as spatial indicators in the development of long-lead fire risk models for these areas. This can help improve the timing of pro-active strategies to manage fire, and in the allocation of sparse funds and resources. Our analysis has highlighted regional patterns of fire across different land tenures. Heightened awareness of these patterns may encourage a more cooperative and coordinated approach to fire management amongst stakeholders.     

Life history patterns in lizards of the arid and semiarid zone of Australia

Summary Studies on the life histories and population dynamics of lizards in the semiarid/arid zone of Australia are reviewed to identify the influence of size (female mean snout-vent length), phylogeny (family effects) and ecological parameters on the evolution of life history traits of these species. Species producing more than one clutch per year are larger than single-clutched ones. In an ANCOVA, significant effects of size and phylogeny on clutch size and on age at sexual maturity were found. Microhabitat (arboreal, terrestrial, and subterranean life style) also had an effect on clutch size, but only mediated through a significant interaction with size. However, results of the ANCOVAs depend on the families and ecological parameters included in the analyses. Therefore, caution is necessary in interpreting or generalizing the results; in any case, size and phylogeny explain only a small percentage of the observed variation. Nevertheless, a direct comparison of a set of syntopic/paratopic desert lizards supports and extends the main conclusions of the ANCOVA. A significant but small phylogenetic effect was found, and arboreal microhabitat was associated with greater age at sexual maturity. Activity (diurnal versus nocturnal) influenced yearly mortalities and clutch frequencies. For both, microhabitat and activity, predation levels and size-dependent mortality were the likely selective factors causing these correlations. The demographic environment explains the paucity of duurnal lizard species with fixed clutch sizes in the semiarid/arid zone of Australia. Possible causes for the evolution of fixed clutch sizes are discussed.     

Water sources accessed by arid zone riparian trees in highly saline environments,Australia

The flow regimes of arid zone rivers are often highly variable, and shallow groundwater in the alluvial aquifers can be very saline, thus constraining the availability and quality of the major water sources available to riparian trees—soil water, shallow groundwater and stream water. We have identified water sources and strategies used by riparian trees in more highly saline and arid conditions than previously studied for riparian trees of arid zone rivers. Our research focused on the riparian species Eucalyptus coolabah, one of the major riparian trees of ephemeral arid zone rivers in Australia. The water sources available to this riparian tree were examined using δ¹⁸O isotope data from xylem, soil water, groundwater and surface water. Additionally, soil chloride and matric potential data were used to infer zones of water availability for root uptake. Despite the saline conditions, the trees used a mixture of soil water and groundwater sources, but they did not use surface water directly. The study identified three strategies used to cope with typically high groundwater and soil water salinities. Firstly, the trees preferentially grow in zones of most frequent flushing by infiltrating streamflow, such as the bank-tops of channels. Secondly, the trees limit water use by having low transpiration rates. Thirdly, the trees are able to extract water at very low osmotic potentials, with water uptake continuing at chloride concentrations of at least 20,000–30,000 mg L⁻¹.     

Thermal and hydric implications of diurnal activity by a small tropical frog during the dry season

Amphibians are typically intolerant of high temperatures and dehydrating conditions, and small species are particularly susceptible to desiccation. The rockhole frog, Litoria meiriana (Hylidae), is diurnal and is often observed on rocks in the sun near streams in tropical Australia. These hot, desiccating conditions are avoided by most frog species. We measured the microclimate in the areas used by frogs and the activity, body temperatures and hydric state of free‐ranging individuals of this small frog. We also used plaster models to further explore the dynamic nature of hydric state by combining estimates of water loss and water uptake with behavioural observations of activity and microhabitat selection. Both direct measures and estimates of dynamic hydric state indicated that free‐ranging frogs generally maintained a hydric state above 95% of full hydration, but occasionally, particularly during the afternoon, frogs allowed their hydric state to fall as low as 85%. Body temperatures of frogs remained below the critical thermal maximum (CT_max) even when the frogs were in the sun, because this species has no cutaneous resistance to evaporative water loss and so they cool by evaporation. However, during the hotter part of the day, on dry sunny substrates, the hydric state of the frogs could fall to near lethal hydration states (approximately 70% of full hydration) within a short period (approximately 20 min). Thus, the threat of desiccation appears to be more limiting than the threat of overheating. These diurnal frogs rely on frequent bouts of rehydration to support their ability to venture onto hot, dry rocks during the day.     

Thermoregulation in free-ranging Nycteris thebaica (Nycteridae) during winter: No evidence of torpor

Bats are among the most heterothermic mammals, with nearly all species investigated under free-ranging conditions to date exhibiting some degree of daily torpor and/or hibernation. We investigated thermoregulation during late winter by seven Nycteris thebaica in a warm, semi-arid habitat in northern South Africa, using temperature-sensitive transmitters to measure skin temperature (T_skin). Unexpectedly, we found no evidence for any expression of daily torpor or hibernation based on a total of 86 days of data from 7 bats (one male and six females), despite air temperatures as low as ～10 °C. Instead, daytime T_skin was distributed unimodally with most values in the 33–35 °C range, and a minimum T_skin of 28.4 °C at a roost temperature of 24.6 °C. There are several possible reasons why N. thebaica may avoid torpor, including predation in roosts, and the long nightly foraging periods of this species compared to many others.     

Torpor during the reproductive season in a free-ranging subtropical bat, Nyctophilus bifax

During times of energetic stress many small mammals reduce their body temperature and metabolic rate, a state known as torpor. Whereas torpor is effective in energy conservation it also entails costs, such as reduced foetal development in pregnant females. Because it is currently not known how subtropical bats deal with energetic challenges during the reproductive season, the thermal biology of free-ranging non-reproductive male and pregnant female Nyctophilus bifax was examined during spring. Males entered torpor much more frequently than pregnant females. However, night time activity periods were similar in both sexes. My results show that even in the subtropics torpor is used regularly during the reproductive period in spring by non-reproductive male N. bifax to conserve energy, but is used rarely by pregnant females likely to prevent slowed foetal development.     

Seasonal emergence, host-seeking activity, age composition and reproductive biology of the mosquito Aedes punctor

Abstract. 1. Seasonal patterns of adult emergence, host-seeking and reproductive ageing in a population of the woodland mosquito Aedes punctor (Kirby) are described from field studies in northern Britain in 1984 and 1985.
2. The pattern of emergence showed little variation between years but duration did vary. Males started emerging before females. Sex ratios at emergence showed a marked female bias. A late, second period of emergence apparently represented a second generation in this normally univoltine species.
3. Host-seeking females were first caught at human bait within a few days of the start of emergence and peak numbers at bait occurred 3–4 weeks later. High numbers lasted for c. 2 weeks. The duration of host-seeking activity was related to weather. The second emergence in 1985 resulted in a second peak of activity in early September and probably extended the flight season.
4. As the season progressed the proportion of females that were parous increased. However, this proportion never reached 1.0, indicating variation in individual reproductive success, which differed between years.
5. The reproductive biology of individuals within the population was investigated. No autogeny was detected, suggesting that the population was entirely anautogenous.     

Biogeography and speciation of a direct developing frog from the coastal arid zone of Western Australia

Within the southwestern Australian biodiversity hotspot, the Shark Bay region displays high levels of plant and animal endemism, particularly in the herpetofauna. The region has been subjected to dramatic climatic fluctuations and has been geologically active from the Late Miocene to the present. The myobatrachid frog Arenophryne rotunda, a Shark Bay endemic, provides an ideal opportunity to examine the relative effects of fluctuating climates and geological activity on the biota of Shark Bay. A comprehensive phylogeographic analysis of A. rotunda, based on data comprising 1154 bp of the mitochondrial gene ND2, is presented. My results demonstrate a major genetic break that divides this species at the northern edge of the Victorian Plateau into northern and southern species lineages, dating to the Late Miocene, with a further division of the southern species lineage across the Murchison Gorge dating to the Plio-Pleistocene border. Both of these periods are related to prominent geological activity and climatic shifts in the Shark Bay region. Interpretation of phylogeographic results point to the prominent role of fluctuating Pleistocene climates and associated coastal landscape evolution in the generation of phylogeographic structure within the distinct A. rotunda species lineages. Similar processes have been invoked to explain the diversity of other Shark Bay biota.     

Adaptive polymorphism inIpomoea pes-tigridis (Convolvulaceae), a common rainy season weed of the Indian arid zone

Ipomoea pes-tigridis is an autogamous and genetically polymorphic weed. In the vicinity of Jodhpur at least 9 forms can be distinguished which differ by leaf shape, presence or absence of latex canals, starch and anthocyanin in shoots, and particularly by size, weight, colour, indumentum, dormancy, and germination requirements of seeds. The different forms prefer those micro-habitats to which they are best adapted.