Torpor in dark times: patterns of heterothermy are associated with the lunar cycle in a nocturnal bird

Many studies have shown that endotherms become more heterothermic when the costs of thermoregulation are high and/or when limited energy availability constrains thermoregulatory capacity. However, the roles of many ecological variables, including constraints on foraging opportunities and/or success, remain largely unknown. To test the prediction that thermoregulatory patterns should be related to foraging opportunities in a heterothermic endotherm, we examined the relationship between the lunar cycle and heterothermy in Freckled Nightjars (Caprimulgus tristigma), which are visually orienting, nocturnal insectivores that are dependent on ambient light to forage. This model system provides an opportunity to assess whether variation in foraging opportunities influences the expression of heterothermy. The nightjars were active and foraged for insects when moonlight was available but became inactive and heterothermic in the absence of moonlight. Lunar illumination was a much stronger predictor of the magnitude of heterothermic responses than was air temperature (T(a)). Our data suggest that heterothermy was strongly related to variation in foraging opportunities associated with the lunar cycle, even though food abundance appeared to remain relatively high throughout the study period. Patterns of thermoregulation in this population of Freckled Nightjars provide novel insights into the environmental and ecological determinants of heterothermy, with the lunar cycle, and not T(a), being the strongest predictor of torpor use.     

Adaptive thermoregulation in endotherms may alter responses to climate change

Climate change is one of the major issues facing natural populations and thus a focus of recent research has been to predict the responses of organisms to these changes. Models are becoming more complex and now commonly include physiological traits of the organisms of interest. However, endothermic species have received less attention than have ectotherms in these mechanistic models. Further, it is not clear whether responses of endotherms to climate change are modified by variation in thermoregulatory characteristics associated with phenotypic plasticity and/or adaptation to past selective pressures. Here, we review the empirical data on thermal adaptation and acclimatization in endotherms and discuss how those factors may be important in models of responses to climate change. We begin with a discussion of why thermoregulation and thermal sensitivity at high body temperatures should be co-adapted. Importantly, we show that there is, in fact, considerable variation in the ability of endotherms to tolerate high body temperatures and/or high environmental temperatures, but a better understanding of this variation will likely be critical for predicting responses to future climatic scenarios. Next, we discuss why variation in thermoregulatory characteristics should be considered when modeling the effects of climate change on heterothermic endotherms. Finally, we review some biophysical and biochemical factors that will limit adaptation and acclimation in endotherms. We consider both long-term, directional climate change and short-term (but increasingly common) anomalies in climate such as extreme heat waves and we suggest areas of important future research relating to both our basic understanding of endothermic thermoregulation and the responses of endotherms to climate change.     

Heterothermy in two mole-rat species subjected to interacting thermoregulatory challenges

Maintaining a high and constant body temperature (T(b) ) is often viewed as a fundamental benefit of endothermy, but variation in T(b) is likely the norm rather than an exception among endotherms. Thus, attempts to elucidate which factors cause T(b) of endotherms to deviate away from the T(b) that maximizes performance are becoming more common. One approach relies on an adaptive framework of thermoregulation, used for a long time to predict variation in T(b) of ectotherms, as a starting point to make predictions about the factors that should lead to thermoregulatory variation in endotherms. Here we test the predictions that when confronted with thermoregulatory challenges endotherms should (1) become more heterothermic, (2) lower their T(b) setpoint, and/or (3) increase behavioral thermoregulation (e.g., activity levels or social thermoregulation). We exposed two species of relatively homeothermic mole-rats to two such challenges: (a) ambient temperatures (T(a)) well below the thermoneutral zone and (b) increased heat loss caused by the removal of dorsal fur. In general, our results support the adaptive framework of endothermic thermoregulation with each species conforming to some of the predictions. For example, Mashona mole-rats (Fukomys darlingi) increased heterothermy as T(a) decreased, highveld mole-rats (Cryptomys hottentotus pretoriae) displayed lower T(b) 's after shaving, and both species increased behavioral thermoregulation as T(a) decreased. This suggests that there is some merit in extending the adaptive framework to endotherms. However, none of the three predictions we tested was supported under all experimental conditions, reiterating that attempts to determine universal factors causing variation in T(b) of endotherms may prove challenging.     

A cluster of diagnostic Hsp68 amino acid sites that are identified in Drosophila from the melanogaster species group are concentrated around beta-sheet residues involved with substrate binding.

The coding region of the hsp68 gene has been amplified, cloned, and sequenced from 10 Drosophila species, 5 from the melanogaster subgroup and 5 from the montium subgroup. When the predicted amino acid sequences are compared with available Hsp70 sequences, patterns of conservation suggest that the C-terminal region should be subdivided according to predominant secondary structure. Conservation levels between Hsp68 and Hsp70 proteins were high in the N-terminal ATPase and adjacent beta-sheet domains, medium in the alpha-helix domain, and low in the C-terminal mobile domain (78%, 72%, 41%, and 21% identity, respectively). A number of amino acid sites were found to be "diagnostic" for Hsp68 (28 of approximately 635 residues). A few of these occur in the ATPase domain (385 residues) but most (75%) are concentrated in the beta-sheet and alpha-helix domains (34% of the protein) with none in the short mobile domain. Five of the diagnostic sites in the beta-sheet domain are clustered around, but not coincident with, functional sites known to be involved in substrate binding. Nearly all of the Hsp70 family length variation occurs in the mobile domain. Within montium subgroup species, 2 nearly identical hsp68 PCR products that differed in length are either different alleles or products of an ancestral hsp68 duplication.     

The latitudinal cline in the In(3R)Payne inversion polymorphism has shifted in the last 20 years in Australian Drosophila melanogaster populations

Clinal variation has been described in a number of inversions in Drosophila but these clines are often characterized by cytological techniques using small sample sizes, and associations with specific genes are rarely considered. Here we have developed a molecular assay for In(3R)Payne in Drosophila melanogaster from eastern Australia populations. It shows in repeated samples that the inversion cline is very tightly associated with latitude and is almost fixed in tropical populations while relatively rare in temperate populations. This steep cline has shifted in position in the last 20 years. The heat shock gene, hsr-omega, located centrally inside the inversion sequence, shows a different clinal pattern to In(3R)Payne. These results suggest strong ongoing selection on In(3R)Payne over the last 100 years since the colonization of Australia that is partly independent of hsr-omega.     

Biopedturbation by an island ecosystem engineer: Burrowing volumes and litter deposition by sooty shearwaters (Puffinus griseus)

Abstract

Seabirds can influence entire island ecosystems through the effects of their burrowing and of their underground deposition of vegetation on biotic and abiotic island processes. This study quantifies the extent of these effects at three sooty shearwater breeding islands in southern New Zealand, with the aim of assessing the importance of this species as an ecosystem engineer. Mean burrow volumes ranged between 158.2 and 528.1 m³ ha^–1. Between 18 and 34% of the ground surface was undermined by burrow space on the three islands. This extent of burrowing is comparable to that of fossorial mammals, widely recognised as ecosystem engineers. Mean vegetation inputs (dry weight), transported underground by birds and incorporated into nests, varied between 33 and 96 g m^‐2 The implications of the biopedturbation caused by sooty shearwater burrowing to the extent measured in this study may be profound for some ecosystem processes, and certainly warrants further research.     

Data quality problems undermine analyses of endotherm upper critical temperatures

In an analysis of avian and mammalian thermal tolerances recently published in this journal, Khaliq et al. ( 2015 ) reported that endotherm thermal niches are phylogenetically conserved in tropical, but not temperate, regions. However, closer examination of the data upon which this analysis was based reveals that many of the upper critical temperature (UCT) data are not valid. Approximately 55% and 42% of avian and mammalian UCT data, respectively, originated from studies in which animals were not exposed to air temperatures high enough to elicit an increase in metabolic rate above minimum levels; the cited UCT values are merely the highest air temperatures at which measurements took place. An additional 18% and 25% of avian and mammalian UCT data, respectively, represent values based on just one individual per species and/or measurements at too few air temperatures above the thermoneutral zone (TNZ) to reliably estimate the UCT.     

Phenotypic flexibility in basal metabolic rate and the changing view of avian physiological diversity: a review

Comparative analyses of avian energetics often involve the implicit assumption that basal metabolic rate (BMR) is a fixed, taxon-specific trait. However, in most species that have been investigated, BMR exhibits phenotypic flexibility and can be reversibly adjusted over short time scales. Many non-migrants adjust BMR seasonally, with the winter BMR usually higher than the summer BMR. The data that are currently available do not, however, support the idea that the magnitude and direction of these adjustments varies consistently with body mass. Long-distance migrants often exhibit large intra-annual changes in BMR, reflecting the physiological adjustments associated with different stages of their migratory cycles. Phenotypic flexibility in BMR also represents an important component of short-term thermal acclimation under laboratory conditions, with captive birds increasing BMR when acclimated to low air temperatures and vice versa. The emerging view of avian BMR is of a highly flexible physiological trait that is continually adjusted in response to environmental factors such as temperature. The within-individual variation observed in avian BMR demands a critical re-examination of approaches used for comparisons across taxa. Several key questions concerning the shapes and other properties of avian BMR reaction norms urgently need to be addressed, and hypotheses concerning metabolic adaptation should explicitly account for phenotypic flexibility.