Small Tropical Mammals Can Take the Heat: High Upper Limits of Thermoneutrality in a Bornean Treeshrew.

Tropical ectotherms are generally believed to be more vulnerable to global heating than temperate species. Currently, however, we have insufficient knowledge of the thermoregulatory physiology of equatorial tropical mammals, particularly of small diurnal mammals, to enable similar predictions. In this study, we measured the resting metabolic rates (via oxygen consumption) of wild-caught lesser treeshrews (Tupaia minor, order Scandentia) over a range of ambient temperatures. We predicted that, similar to other treeshrews, T. minor would exhibit more flexibility in body temperature regulation and a wider thermoneutral zone compared with other small mammals because these thermoregulatory traits provide both energy and water savings at high ambient temperatures. Basal metabolic rate was on average 1.03±0.10 mL O2 h-1 g-1, which is within the range predicted for a 65-g mammal. We calculated the lower critical temperature of the thermoneutral zone at 31.0°C (95% confidence interval: 29.3°-32.7°C), but using metabolic rates alone, we could not determine the upper critical temperature at ambient temperatures as high as 36°C. The thermoregulatory characteristics of lesser treeshrews provide a means of saving energy and water at temperatures well in excess of their current environmental temperatures. Our research highlights the knowledge gaps in our understanding of the energetics of mammals living in high-temperature environments, specifically in the equatorial tropics, and questions the purported lack of variance in the upper critical temperatures of the thermoneutral zone in mammals, emphasizing the importance of further research in the tropics.

Obligatory Nocturnalism in Triassic Archaic Mammals: Preservation of Sperm Quality?

Archaic mammals became exclusively nocturnal by the Late Triassic, and today, the majority of extant mammals remain nocturnal. Although there is ample morphological and physiological evidence supporting Late Mesozoic nocturnalism, a succinct hypothesis for why mammals became nocturnal remains elusive. Here, I propose a hypothesis that, with the onset of body size miniaturization in the Triassic and the concomitant evolution of fur and increased mass-specific metabolic rate and hence body temperature, small mammals became obligatorily nocturnal in order to avoid poor sperm quality, hyperthermia, and high rates of evaporative water loss and to maximize foraging time. The hypothesis hinges heavily on the assumption that, with the absence of externalized testes, the maximum optimum temperature of about 36°C for spermatogenesis was subject to strong stabilizing selection that placed a ceiling on increases in metabolic rate and body temperature. Heat-dissipating capacity during the daytime during the Triassic, Jurassic, and Cretaceous was thereby compromised. The release from the constraint of the optimum temperature of spermatogenesis occurred in placental mammals only with the advent of the externalization of testes in the scrotum in Boreotheria in the Cenozoic or, with the recent claim that the scrotum is plesiomorphic in mammals, as early as the Jurassic with the origin of the marsupials.

Staying hot to fight the heat-high body temperatures accompany a diurnal endothermic lifestyle in the tropics.

Much of our knowledge of the thermoregulation of endotherms has been obtained from species inhabiting cold and temperate climates, our knowledge of the thermoregulatory physiology of tropical endotherms is scarce. We studied the thermoregulatory physiology of a small, tropical mammal, the large treeshrew (Tupaia tana, Order Scandentia) by recording the body temperatures of free-ranging individuals, and by measuring the resting metabolic rates of wild individuals held temporarily in captivity. The amplitude of daily body temperature (~ 4 °C) was higher in treeshrews than in many homeothermic eutherian mammals; a consequence of high active-phase body temperatures (~ 40 °C), and relatively low rest-phase body temperatures (~ 36 °C). We hypothesized that high body temperatures enable T. tana to maintain a suitable gradient between ambient and body temperature to allow for passive heat dissipation, important in high-humidity environments where opportunities for evaporative cooling are rare. Whether this thermoregulatory phenotype is unique to Scandentians, or whether other warm-climate diurnal small mammals share similar thermoregulatory characteristics, is currently unknown.

Searching for the Haplorrhine Heterotherm: Field and Laboratory Data of Free-Ranging Tarsiers.

The observation of heterothermy in a single suborder (Strepsirrhini) only within the primates is puzzling. Given that the placental-mammal ancestor was likely a heterotherm, we explored the potential for heterothermy in a primate closely related to the Strepsirrhini. Based upon phylogeny, body size and habitat stability since the Late Eocene, we selected western tarsiers (Cephalopachus bancanus) from the island of Borneo. Being the sister clade to Strepsirrhini and basal in Haplorrhini (monkeys and apes), we hypothesized that C. bancanus might have retained the heterothermic capacity observed in several small strepsirrhines. We measured resting metabolic rate, subcutaneous temperature, evaporative water loss and the percentage of heat dissipated through evaporation, at ambient temperatures between 22 and 35°C in fresh-caught wild animals (126.1 ± 2.4 g). We also measured core body temperatures in free-ranging animals. The thermoneutral zone was 25-30°C and the basal metabolic rate was 3.52 ± 0.06 W.kg-1 (0.65 ± 0.01 ml O2.g-1.h-1). There was no evidence of adaptive heterothermy in either the laboratory data or the free-ranging data. Instead, animals appeared to be cold sensitive (Tb ~ 31°C) at the lowest temperatures. We discuss possible reasons for the apparent lack of heterothermy in tarsiers, and identify putative heterotherms within Platyrrhini. We also document our concern for the vulnerability of C. bancanus to future temperature increases associated with global warming.

A phenology of the evolution of endothermy in birds and mammals.

Recent palaeontological data and novel physiological hypotheses now allow a timescaled reconstruction of the evolution of endothermy in birds and mammals. A three-phase iterative model describing how endothermy evolved from Permian ectothermic ancestors is presented. In Phase One I propose that the elevation of endothermy - increased metabolism and body temperature (Tb ) - complemented large-body-size homeothermy during the Permian and Triassic in response to the fitness benefits of enhanced embryo development (parental care) and the activity demands of conquering dry land. I propose that Phase Two commenced in the Late Triassic and Jurassic and was marked by extreme body-size miniaturization, the evolution of enhanced body insulation (fur and feathers), increased brain size, thermoregulatory control, and increased ecomorphological diversity. I suggest that Phase Three occurred during the Cretaceous and Cenozoic and involved endothermic pulses associated with the evolution of muscle-powered flapping flight in birds, terrestrial cursoriality in mammals, and climate adaptation in response to Late Cenozoic cooling in both birds and mammals. Although the triphasic model argues for an iterative evolution of endothermy in pulses throughout the Mesozoic and Cenozoic, it is also argued that endothermy was potentially abandoned at any time that a bird or mammal did not rely upon its thermal benefits for parental care or breeding success. The abandonment would have taken the form of either hibernation or daily torpor as observed in extant endotherms. Thus torpor and hibernation are argued to be as ancient as the origins of endothermy itself, a plesiomorphic characteristic observed today in many small birds and mammals.

Mammal survival at the Cretaceous-Palaeogene boundary: metabolic homeostasis in prolonged tropical hibernation in tenrecs.

Free-ranging common tenrecs, Tenrec ecaudatus, from sub-tropical Madagascar, displayed long-term (nine months) hibernation which lacked any evidence of periodic interbout arousals (IBAs). IBAs are the dominant feature of the mammalian hibernation phenotype and are thought to periodically restore long-term ischaemia damage and/or metabolic imbalances (depletions and accumulations). However, the lack of IBAs in tenrecs suggests no such pathology at hibernation Tbs > 22°C. The long period of tropical hibernation that we report might explain how the ancestral placental mammal survived the global devastation that drove the dinosaurs and many other vertebrates to extinction at the Cretaceous-Palaeogene boundary following a meteorite impact. The genetics and biochemistry of IBAs are of immense interest to biomedical researchers and space exploration scientists, in the latter case, those envisioning a hibernating state in astronauts for deep space travel. Unravelling the physiological thresholds and temperature dependence of IBAs will provide new impetus to these research quests.

The energetics of a Malagasy rodent, Macrotarsomys ingens (Nesomyinae): a test of island and zoogeographical effects on metabolism.

This study provides first insights into the energetics of the Nesomyinae, a subfamily of rodents endemic to Madagascar. The ancestral nesomyine colonized Madagascar from Africa ca. 30-15 mya at the onset of Oligocene global cooling. We tested the hypothesis that, contrary to what might be expected from Island Biogeography theory, post-colonization character displacement of thermoregulatory traits was constrained by phylogenetic inertia through climate adaptation. The study was conducted in the Parc National d'Ankarafantsika, Madagascar. We measured the basal metabolic rate (BMR) and body temperature (T b) patterns of naturally warm-acclimated, freshly captured adult long-tailed big-footed mice Macrotarsomys ingens (67.4 g). The mean ± SD BMR of M. ingens was 0.298 ± 0.032 Watts (n = 12), 31.7 % lower than that predicted by a phylogenetically independent allometric equation. Body mass was correlated with BMR. The lower critical limit of thermoneutrality (T lc) was 30.7°C. The mean ± SD T b = 36.1 ± 0.8°C (n = 12) compared well with the mean T b values for myomorph rodents from the Afrotropical zone, but was lower than those of the Neotropical and Palearctic zones. M. ingens became pathologically hypothermic when exposed to ambient temperatures lower than 18°C. The soil temperature at depths of 250 mm and deeper did not decrease below 22°C throughout the austral winter. The thermoregulatory data for M. ingens did not differ from those that characterize mainland Afrotropical rodents. However, BMR and T b were lower than those of Holarctic rodents. Thus, contrary to expectations of Island Biogeography theory that rapid character displacement often occurs in morphological and behavioural traits when mammals colonize islands, M. ingens displayed climate-related physiological traits indicative of phylogenetic inertia. Presumably the tropical conditions that prevailed on Madagascar at the time of colonisation differed very little from those of the African mainland, and hence there was no strong driving force for change. Unlike small tenrecs and lemurs that radiated on Madagascar prior to the Oligocene, traits associated with an insular existence, such as daily torpor and hibernation, were not evident in M. ingens.

Effects of reproductive status and high ambient temperatures on the body temperature of a free-ranging basoendotherm.

Tenrecs (Order Afrosoricida) exhibit some of the lowest body temperatures (T b) of any eutherian mammal. They also have a high level of variability in both active and resting T bs and, at least in cool temperatures in captivity, frequently employ both short- and long-term torpor. The use of heterothermy by captive animals is, however, generally reduced during gestation and lactation. We present data long-term T b recordings collected from free-ranging S. setosus over the course of two reproductive seasons. In general, reproductive females had slightly higher (~32 °C) and less variable T b, whereas non-reproductive females and males showed both a higher propensity for torpor as well as lower (~30.5 °C) and more variable rest-phase T bs. Torpor expression defined using traditional means (using a threshold or cut-off T b) was much lower than predicted based on the high degree of heterothermy in captive tenrecs. However, torpor defined in this manner is likely to be underestimated in habitats where ambient temperature is close to T b. Our results caution against inferring metabolic states from T b alone and lend support to the recent call to define torpor in free-ranging animals based on mechanistic and not descriptive variables. In addition, lower variability in T b observed during gestation and lactation confirms that homeothermy is essential for reproduction in this species and probably for basoendothermic mammals in general. The relatively low costs of maintaining homeothermy in a sub-tropical environment might help shed light on how homeothermy could have evolved incrementally from an ancestral heterothermic condition.

Increased homeothermy during reproduction in a basal placental mammal.

Homeothermic endothermy, the maintenance of a high and stable body temperature (Tb) using heat produced by elevated metabolism, is energetically expensive. There is increasing evidence that the earliest endotherms were heterotherms that, rather than maintaining strict homeothermy, allowed Tb to fluctuate with large variations between active and rest-phase Tb. The high level of homeothermy observed in modern mammals is therefore likely to have evolved from an ancestral heterothermic state. One of the hypotheses for the evolution of endothermy is that homeothermy allows for greater energetic output during reproduction (parental care model). We tested this hypothesis by measuring metabolic rates over a range of ambient temperatures in both reproductive and non-reproductive greater hedgehog tenrecs (Setifer setosus), a physiologically primitive mammal from Madagascar. Tenrecs have some of the lowest metabolic rates and highest levels of Tb variability of any mammal and are therefore good models of the ancestral eutherian state. During pregnancy and lactation, there was an increase in metabolism and Tb below the thermoneutral zone, accompanied by a decrease in Tb variability. The lower critical limit of the thermoneutral zone was estimated at ~25°C. However, whereas increases in resting metabolism were substantial below 20°C (up to 150% higher during reproduction), daytime rest-phase ambient temperatures at the study site rarely reached equivalent low levels. Thus, S. setosus provide an example for how relatively low-cost increases in homeothermy could have led to substantial increases in fitness by allowing for the faster production of young. The mechanisms necessary for increases in thermogenesis during reproduction would have further benefited the development of homeothermy in mammals.

Are tropical small mammals physiologically vulnerable to Arrhenius effects and climate change?

There is some urgency in the necessity to incorporate physiological data into mechanistic, trait-based, demographic climate change models. Physiological responses at the individual level provide the mechanistic link between environmental changes and individual performances and hence population dynamics. Here we consider the causal relationship between ambient temperature (Ta) and metabolic rate (MR), namely, the Arrhenius effect, which is directly affected by global warming through increases in average global air temperatures and the increase in the frequency and intensity of extreme climate events. We measured and collated data for several small, free-ranging tropical arboreal mammals and evaluated their vulnerability to Arrhenius effects and putative heat stress associated with climate change. Skin temperatures (Tskin) were obtained from free-ranging tarsiers (Tarsius syrichta) on Bohol Island, Philippines. Core body temperature (Tb) was obtained from the greater hedgehog tenrec (Setifer setosus) and the gray brown mouse lemur (Microcebus ravelobensis) from Ankarafantsika, Madagascar. Tskin for another mouse lemur, Microcebus griseorufus, was obtained from the literature. All four species showed evidence of hyperthermia during the daytime rest phase in the form of either Tskin or Tb that was higher than the normothermic Tb during the nighttime active phase. Potentially, tropical arboreal mammals with the lowest MRs and Tb, such as tarsiers, are the most vulnerable to sustained heat stress because their Tb is already close to Ta. Climate change may involve increases in MRs due to Arrhenius effects, especially during the rest phase or during torpor and hibernation. The most likely outcome of increased Arrhenius effects with climate change will be an increase in energy expenditure at the expense of other critical functions such as reproduction or growth and will thus affect fitness. However, we propose that these hypothetical Arrhenius costs can be, and in some species probably are, offset by the use of hyperthermic daily torpor, that is, hypometabolism at high Ta.

The evolution of micro-cursoriality in mammals.

In this study we report on the evolution of micro-cursoriality, a unique case of cursoriality in mammals smaller than 1 kg. We obtained new running speed and limb morphology data for two species of elephant-shrews (Elephantulus spp., Macroscelidae) from Namaqualand, South Africa, which we compared with published data for other mammals. Elephantulus maximum running speeds were higher than those of most mammals smaller than 1 kg. Elephantulus also possess exceptionally high metatarsal:femur ratios (1.07) that are typically associated with fast unguligrade cursors. Cursoriality evolved in the Artiodactyla, Perissodactyla and Carnivora coincident with global cooling and the replacement of forests with open landscapes in the Oligocene and Miocene. The majority of mammal species, though, remained non-cursorial, plantigrade and small (<1 kg). The extraordinary running speed and digitigrady of elephant-shrews was established in the Early Eocene in the earliest macroscelid Prodiacodon, but was probably inherited from Paleocene, Holarctic stem macroscelids. Micro-cursoriality in macroscelids evolved from the plesiomorphic plantigrade foot of the possum-like ancestral mammal earlier than in other mammalian crown groups. Micro-cursoriality evolved first in forests, presumably in response to selection for rapid running speeds facilitated by local knowledge, in order to avoid predators. During the Miocene, micro-cursoriality was pre-adaptive to open, arid habitats, and became more derived in the newly evolved Elephantulus and Macroscelides elephant-shrews with trail running.

The evolution of mammalian body temperature: the Cenozoic supraendothermic pulses.

In this study, I investigated the source(s) of variation in the body temperatures of mammals. I also attempted to reconstruct ancestral normothermic rest-phase body temperature states using a maximum parsimony approach. Body temperature at the familial level is not correlated with body mass. For small mammals, except the Macroscelidae, previously identified correlates, such as climate adaptation and zoogeography explained some, but not all, T(b) apomorphies. At the species level in large cursorial mammals, there was a significant correlation between body temperature and the ratio between metatarsal length and femur length, the proxy for stride length and cursoriality. With the exception of two primate families, all supraendothermic (T(b) > 37.9°C) mammals are cursorial, including Artiodactyla, Lagomorpha, some large Rodentia, and Carnivora. The ruminant supraendothermic cursorial pulse is putatively associated with global cooling and vegetation changes following the Paleocene-Eocene Thermal Maximum. Reconstructed ancestral body temperatures were highly unrealistic deep within the mammalian phylogeny because of the lack of fossil T(b) data that effectively creates ghost lineages. However, it is anticipated that the method of estimating body temperature from the abundance of ¹³C-¹8O bonds in the carbonate component of tooth bioapatite in both extant and extinct animals may be a very promising tool for estimating the T(b) of extinct mammals. Fossil T(b) data are essential for discerning derived T(b) reversals from ancestral states, and verifying the dates of supraendothermic pulses.

The allometry of parrot BMR: seasonal data for the Greater Vasa Parrot, Coracopsis vasa, from Madagascar.

In this study we examined the allometry of basal metabolic rate (BMR) of 31 parrot species. Unlike previous reports, we show that parrots per se do not display BMRs that are any different to other captive-raised birds of their body size. An ordinary least squares regression fitted the data best and body mass explained 95% of the variation in BMR. There was no phylogenetic signal in the BMR data. We also provide new data for the Greater Vasa Parrot (Coracopsis vasa) of Madagascar. We tested the hypotheses that C. vasa may, because of its insular existence, display conservative energetic traits (low BMR, use of adaptive heterothermy) similar to those observed in several Malagasy mammals. However, this was not the case. C. vasa had a higher BMR than other parrots, especially during summer, when BMR was up-regulated by 50.5% and was 95.7% higher than predicted from an ordinary least squares (OLS) allometry of parrots (BMR = 0.042M (b) (0.649) , BMR in Watts, M (b) in grammes). Compared with BMR data for 94 captive-raised bird species, the winter and summer BMRs were, respectively, 45.5 and 117.8% higher than predicted by a phylogenetic generalised least squares (PGLS) allometry (BMR = 0.030M (b) (0.687) , BMR in Watts, M (b) in grammes). The summer up-regulation of BMR is the highest recorded for a bird of any size to date. We suggest that the costs of a high summer BMR may be met by the unusual cooperative breeding system of C. vasa in which groups of males feed the female and share paternity. The potential breeding benefits of a high summer BMR are unknown.

The allometry of rodent intestines.

This study examined the allometry of the small intestine, caecum, colon and large intestine of rodents (n = 51) using a phylogenetically informed approach. Strong phylogenetic signal was detected in the data for the caecum, colon and large intestine, but not for the small intestine. Most of the phylogenetic signal could be attributed to clade effects associated with herbivorous versus omnivorous rodents. The herbivorous rodents have longer caecums, colons and large intestines, but their small intestines, with the exception of the desert otomyine rodents, are no different to those of omnivorous rodents. Desert otomyine rodents have significantly shorter small intestines than all other rodents, reflecting a possible habitat effect and providing a partial explanation for the low basal metabolic rates of small desert mammals. However, the desert otomyines do not have shorter colons or large intestines, challenging claims for adaptation to water retention in arid environments. Data for the Arvicolidae revealed significantly larger caecums and colons, and hence longer large intestines, with no compensatory reduction in the length of the small intestine, which may explain how the smallest mammalian herbivores manage to meet the demands of a very high mass-specific metabolic rate. This study provides phylogenetically corrected allometries suitable for future prediction testing.

Modification and miniaturization of Thermochron iButtons for surgical implantation into small animals.

Thermochron iButtons are being used increasingly by animal physiologists to measure long-term patterns of body temperature in reptiles, birds and mammals. Typically, iButtons are surgically implanted into the intraperitoneal cavity where they measure and store body temperature together with the date and time from an onboard real-time clock. In 16-bit resolution, the DS1922L iButton can store a total of 4,096 data points over pre-determined sampling intervals. iButtons have proved invaluable in measuring patterns of torpor and hibernation in animals larger than 70 g. Weighing around 3.5 g after potting with wax, iButtons are too heavy and large to implant into animals smaller than 70 g because their weight exceeds 5% of the animal's total body weight. This paper describes how the stainless steel canister housing the DS 1922L iButton battery and circuit board can be removed to reduce the weight of the components to 1.49 g after waxing (ready for implantation) without compromising the function or battery life of the iButton. The modified iButton can be implanted into animals as small as 20-30 g. Calibration data revealed an offset of ca. 1 degrees C on average, confirming that iButtons must be calibrated prior to implantation.

Age at first reproduction and growth rate are independent of basal metabolic rate in mammals.

This study tested an emergent prediction from the Metabolic Theory of Ecology (MTE) that the age at first reproduction (alpha) of a mammal is proportional to the inverse of its mass-corrected basal metabolic rate: alpha proportional (B / M)-1 The hypothesis was tested with multiple regression models of conventional species data and phylogenetically independent contrasts of 121 mammal species. Since age at first reproduction is directly influenced by an individual's growth rate, the hypothesis that growth rate is proportional to BMR was also tested. Although the overall multiple regression model was significant, age at first reproduction was not partially correlated with either body mass, growth rate or BMR. Similarly, growth rate was not correlated with BMR. Thus at least for mammals in general, there is no evidence to support the fundamental premise of the MTE that individual metabolism governs the rate at which energy is converted to growth and reproduction at the species level. The exponents of the BMR allometry calculated using phylogenetic generalized least squares regression models were significantly lower than the three-quarter value predicted by the MTE.

Torpor and hibernation in a basal placental mammal, the Lesser Hedgehog Tenrec Echinops telfairi.

The patterns of heterothermy were measured in Lesser Hedgehog Tenrecs, Echinops telfairi, under semi-natural conditions in an outdoor enclosure during the austral mid-winter in southwestern Madagascar. The animals were implanted with miniaturized body temperature (Tb) loggers (iButtons) that measured body temperature every 42 min for 2 months (May and June). The tenrecs entered daily torpor on all 60 consecutive days of measurement, that is, on 100% of animal days, with body temperature closely tracking ambient temperature (Ta) during the ambient heating phase. The mean minimum daily Tb of the tenrecs was 18.44 +/- 0.50 degrees C (n = 174, N = 3), and never exceeded 25 degrees C whereas, apart from a few hibernation bouts in one animal, the mean maximum daily Tb was 30.73 +/- 0.15 degrees C (n = 167, N = 3). Thus during winter, tenrecs display the lowest normothermic Tb of all placental mammals. E. telfairi showed afternoon and early evening arousals, but entered torpor before midnight and remained in torpor for 12-18 h each day. One animal hibernated on two occasions for periods of 2-4 days. We consider E. telfairi to be a protoendotherm, and discuss the relevance and potential of these data for testing models on the evolution of endothermy.

Phenotypic flexibility in the basal metabolic rate of laughing doves: responses to short-term thermal acclimation.

Many birds exhibit considerable phenotypic flexibility in maintenance energy requirements, and up- or downregulate basal metabolic rate (BMR) over time scales of days to weeks during thermal acclimation. However, the extent to which individual birds can reverse the direction of BMR adjustments over short time scales remains unknown. In this study, we examined metabolic responses to short-term thermal acclimation in laughing doves Streptopelia senegalensis. In 30 wild-caught doves (mean body mass=92.6 g) divided into three experimental groups of 10 birds each, initial BMR averaged 0.760+/-0.036 W. Thereafter, each group was acclimated to one of three acclimation air temperatures (T(acc)=10, 22 or 35 degrees C) for 21 days, during which time the doves were housed in individual cages. Following the first acclimation period (acclimation I), BMR (W) was significantly lower and was negatively and linearly related to T(acc) [BMR=0.714-0.005T(acc)]. Acclimation I BMR varied from 0.546+/-0.039 W in doves acclimated to T(acc)=35 degrees C to 0.665+/-0.058 W at T(acc)=10 degrees C. A second acclimation period of a further 21 days (acclimation II) revealed that the direction of BMR adjustments could be reversed within individuals, with acclimation II BMR again negatively and linearly related to T(acc). The slope of the relationship between BMR and T(acc) following acclimation II was not significantly different to that following acclimation I. BMR exhibited consistent inter-individual variation, with a low but significant repeatability of 0.113. The within-individual BMR variation of up to 26% that we observed in laughing doves reveals that BMR is a highly flexible trait in this species, and reiterates the need to take phenotypic plasticity into account in comparative analyses of avian energetic parameters.

Lack of torpor in free-ranging southern lesser galagos, Galago moholi: ecological and physiological considerations.

Studies investigating heterothermy under natural conditions are particularly scarce for tropical species. However, heterothermy patterns in tropical and subtropical environments often differ markedly from those observed in arctic and temperate species. The investigation of heterothermy in strepsirhine primates has focussed largely on the Malagasy cheirogaleids. In addition, a physiological verification of torpor occurrence in mainland strepsirhines is important with regard to arguments pertaining to the colonization of Madagascar by strepsirhine primates. We measured body temperatures of 11 free-ranging Galago moholi, between February 2002 and September 2003, for 3 consecutive months for each animal. No incidents of heterothermy were recorded throughout the study period. We considered how physiological and ecological aspects of G. moholi biology might have obviated the use of torpor. It was suggested that the breeding pattern observed in G. moholi prevented torpor use whilst increasing fecundity, and that the ecological costs of torpor far outweighed the energetic costs. This study highlights the need for more studies on free-ranging animals to elucidate the physiological, ecological and phylogenetic constraints and determinants of torpor use. Furthermore, if convincing arguments are to be made regarding the possible role of heterothermy in species dispersal, more data from free-ranging animals are needed.

The power of fitness in mammals: perceptions from the African slipstream.

Evolutionary physiology is the emerging physiological discipline. Unlike environmental physiology or ecophysiology, whose definitions have long been made quite clear, evolutionary physiology has a broader scope of objectives, and its definition lacks a concise treatise. This paper presents the argument that the lack of a common definition of evolutionary physiology is retarding the unification of the mechanistic and amechanistic physiological sciences, a multidisciplinary obligation crucial for a holistic understanding of a physiological basis of fitness. The divide between mechanistic "how" questions, devoted primarily to homeostasis, and evolutionary "why" questions, concerned with understanding phenotypic and genotypic physiological variation, remains broad and is currently not conducive to synergy in the physiological disciplines. Unification may be facilitated, however, by embracing a common currency of measurement and analysis. A likely candidate is the cascade of energy from the environment to offspring and the evolution of physiological form and function, including homeostasis, associated with power management. This currency approach seeks to identify an energetic basis of fitness, namely, whether or how the evolution of life-history traits is influenced by energetic constraints and/or trade-offs.