A test of ecological and sexual selection hypotheses for geographical variation in coloration and morphology of golden-crowned kinglets (Regulus satrapa)

Aim To date, studies on geographical variation have extensively investigated Bergmann’s rule, yet Gloger’s rule remains infrequently tested, and climatic predictors of variation in carotenoid coloration have not yet been studied. In addition, hypotheses based on sexual selection, which predict that sexual dimorphism should vary with population density and climatic conditions, have received little attention. Our goals were to characterize geographical variation in the coloration and morphology of golden‐crowned kinglets, Regulus satrapa (Passeriformes, Regulidae), and to investigate possible ecological and sexual selection correlates of this variation. Location The entire species range of golden‐crowned kinglets, comprising North and Central America. Methods We collected data from 511 museum specimens, dating from 1847 to 2006, encompassing all five subspecies of golden‐crowned kinglets. We used reflectance spectrometry to quantify crown and mantle coloration, and measured wing‐chord and tarsus length to approximate body size. We obtained geographical and climatic data from online databases, and population density estimates from the literature. Results There were significant subspecific and gender differences in crown coloration and morphology: male kinglets were generally larger and more colourful. Our data revealed mixed support for Bergmann’s rule: tarsus length decreased with increasing latitude, while patterns of variation in wing‐chord and tarsus length showed conflicting results with temperature. Mantle coloration exhibited an opposite trend to that predicted by Gloger’s rule: upper parts became lighter with increasing relative humidity. Crown coloration was negatively correlated with actual evapotranspiration, suggesting that levels of primary productivity are not directly linked to carotenoid abundance. Sexual dimorphism and dichromatism generally increased with greater population density, lower latitudes and elevations, and warmer temperatures, supporting a previously observed pattern of variation in sexual dimorphism. Main conclusions Geographical variation in golden‐crowned kinglets yielded mixed support for Bergmann’s rule and contradicted Gloger’s rule, suggesting that other mechanisms may be operating. Allen’s rule is likely to be a stronger factor influencing tarsus length. Differences in the degree of sexual dimorphism and dichromatism in varying climatic conditions suggest that the intensity of sexual selection differs between habitats. Further studies on geographical variation in sexual dimorphism in various taxa may reveal a previously unrecognized ecogeographical rule.     

Adaptation of Pelage Color and Pigment Variations in Israeli Subterranean Blind Mole Rats,Spalax Ehrenbergi

Background

Concealing coloration in rodents is well established. However, only a few studies examined how soil color, pelage color, hair-melanin content, and genetics (i.e., the causal chain) synergize to configure it. This study investigates the causal chain of dorsal coloration in Israeli subterranean blind mole rats, Spalax ehrenbergi.

Methods

We examined pelage coloration of 128 adult animals from 11 populations belonging to four species of Spalax ehrenbergi superspecies (Spalax galili, Spalax golani, Spalax carmeli, and Spalax judaei) and the corresponding coloration of soil samples from the collection sites using a digital colorimeter. Additionally, we quantified hair-melanin contents of 67 animals using HPLC and sequenced the MC1R gene in 68 individuals from all four mole rat species.

Results

Due to high variability of soil colors, the correlation between soil and pelage color coordinates was weak and significant only between soil hue and pelage lightness. Multiple stepwise forward regression revealed that soil lightness was significantly associated with all pelage color variables. Pelage color lightness among the four species increased with the higher southward aridity in accordance to Gloger''s rule (darker in humid habitats and lighter in arid habitats). Darker and lighter pelage colors are associated with darker basalt and terra rossa, and lighter rendzina soils, respectively. Despite soil lightness varying significantly, pelage lightness and eumelanin converged among populations living in similar soil types. Partial sequencing of the MC1R gene identified three allelic variants, two of which were predominant in northern species (S. galili and S. golani), and the third was exclusive to southern species (S. carmeli and S. judaei), which might have caused the differences found in pheomelanin/eumelanin ratio.

Conclusion/Significance

Darker dorsal pelage in darker basalt and terra rossa soils in the north and lighter pelage in rendzina and loess soils in the south reflect the combined results of crypsis and thermoregulatory function following Gloger''s rule.     

Pelage color variation in pocket gophers (Rodentia: Geomyidae) in relation to sex,age and differences in habitat

Pelage coloration is a phenotypic characteristic in mammals that could be associated with an individual's survival and fitness; thus coloration gains adaptive importance. In geomyids, pelage coloration shows a relationship with the color of soils on which they live, mainly freshly dug soil of their burrows. This characteristic could be due to a camouflage adaptation to avoid predators. Pocket gophers disperse aboveground to establish a territory before they reach reproductive condition, and males disperse longer distances than females. The aim of this study was to evaluate pelage color variation in pocket gophers (Thomomys anitae) in relation to habitat differences, sex, and age, and determine its association to the color of the soil on which they live. Brightness of T. anitae's dorsal pelage coloration and that of soil samples from five different habitats in the Baja California peninsula, Mexico were measured to test four hypotheses: (1) Subadults show a wide coloration range, but extreme colors are lost in adulthood due to natural selection. (2) Males are more vulnerable to depredation than females; therefore, males’ coloration is more homogenous as a protective camouflage. (3) In open habitats pocket gophers are more exposed to being detected by predators, therefore their pelage coloration pattern is less variable than that of individuals from habitats with more vegetation cover. (4) Pelage coloration better matches soil coloration in moist conditions similar to that of freshly dug soil of their burrows. The results confirmed our predictions; however, selection does not impose an equal pressure on pelage coloration on the five habitats evaluated; other factors such as population density and predator presence need to be assessed. The strongest effects are found in the most open habitat, and there is less strong support for predictions in habitats where predator assemblage is diminished.     

SHORT COMMUNICATION: Little evidence for Cope’s rule from Bayesian phylogenetic analysis of extant mammals

According to Cope’s rule, lineages tend to evolve towards larger body size, possibly because of selective advantages of being large. The status of Cope’s ‘rule’ remains controversial as it is supported in some but not all large‐scale fossil studies. Here, we test for Cope’s rule by Bayesian analyses of average body masses of 3253 extant mammal species on a dated phylogenetic tree. The data favour a model that does not assume Cope’s rule. When Cope’s rule is assumed, the best estimate of its strength is an average ancestor‐descendant increase in body size of only 0.4%, which sharply contrasts with the 9% bias estimated from fossil mammals. Thus, we find no evidence for Cope’s rule from extant mammals, in agreement with earlier analyses of existing species, which also did not find support for Cope’s rule.     

Photoperiodic and hormonal influences on fur density and regrowth in two hamster species

Temperate and boreal mammals undergo seasonal changes in pelage that facilitate thermoregulation in winter and summer. We investigated photoperiodic influences on pelage characteristics of male Siberian and Syrian hamsters. Fur density (mg fur/cm2 skin) was measured by weighing the shavings of fur patches removed from the dorsal and ventral surfaces of hamsters maintained in long days (LDs) or transferred to short days (SDs). Patches were reshaved 3 wk later to assess fur regrowth (mg regrown fur/cm2 skin). Fur density was greater in SD than in LD Siberian hamsters after 11 wk of differential phototreatment. The onset of increased fur density in SDs was accompanied by a transient increase in fur regrowth (11-14 wk on the dorsal surface and 7-10 and 11-14 wk on the ventral surface), suggestive of a seasonal molting process. Fur density, body mass, and pelage color of Siberian hamsters returned to values characteristic of LD males after a similar duration of prolonged (>27 wk) SD treatment and appear to be regulated by a similar or common interval-timing mechanism. In Syrian hamsters, dorsal fur density, fur regrowth, and hair lengths were greater in SD than in LD males. Castration increased and testosterone (T) treatment decreased dorsal and ventral fur regrowth in LD and SD hamsters, but the effects of T manipulations on fur density were limited to a decrease in dorsal fur density after T treatment. Decreased circulating T in SDs likely contributes to the seasonal molt of male hamsters by increasing the rate of fur growth during the transition to the winter pelage.     

Bergmann′s rule in mammals: a cross‐species interspecific pattern

Although Bergmann’s rule – stating that among closely related species, the bigger ones will inhabit the colder climates/higher latitudes – was formulated for inter‐specific comparisons, most analyses that tested this pattern in mammals were on an intra‐specific level. To date, no large‐scale taxonomy‐driven cross‐species evaluation of the pattern predicted by Bergmann exists. Here we show, in a dataset comprising 3561 mammal species from 26 orders, that while there is no significant correlation between latitude and body mass using conventional methods, this correlation is highly significant when the phylogenetic structure of the dataset is accounted for, thus supporting Bergmann’s claim that the rule only applies to closely related species. Analyses of different subsets indicate that the Bergmann’s rule is evident across a variety of latitude ranges. In many taxonomic subsets, when analysed alone, there is no significant correlation between body mass and latitude. In combination with both the significant relationship in the overall dataset and with results of intra‐specific analyses from the literature, this suggests that Bergmann’s rule describes a fundamental principle within mammals, but that its expression has been modified by a variety of factors during mammalian diversification yet to be resolved.     

On the validity of Bergmann's rule

Aim We reviewed the occurrence of Bergmann's rule in birds (ninety‐four species) and mammals (149 species), using only studies where statistical significance of the results was tested. We also tested whether studies using different characters as surrogates of body size have a different tendency to conform to Bergmann's rule, whether body size and nest type (in birds) have an influence on the tendency to conform to the rule, and whether sedentary birds conform to the rule more than migratory birds. Location Worldwide. Methods We reviewed published data on geographic and temporal variation in body size, using only studies where the statistical significance of the results was tested. We asked how many species conform to the rule out of all species studied in each order and family. Results Over 72% of the birds and 65% of the mammal species follow Bergmann's rule. An overall tendency to follow the rule occurs also within orders and families. Studies using body mass in mammals show the greatest tendency to adhere to Bergmann's rule (linear measurements and dental measurements show a weaker tendency); while in birds, studies using body mass and other surrogates (linear measurements and egg size) show a similar tendency. Birds of different body mass categories exhibit a similar tendency to follow Bergmann's rule, while in mammals the lower body size categories (4–50 and 50–500 g) show a significantly lower tendency to conform to the rule. Sedentary birds tend to conform to Bergmann's rule more than migratory species. Nest type does not affect the tendency to conform to Bergmann's rule. Main conclusions Bergmann's rule is a valid ecological generalization for birds and mammals.     

Colors of primate pelage and skin: objective assessment of conspicuousness

We present a quantitative means of assessing the conspicuousness of animal coats or other objects in terms of the color vision of each possible observer. We measured reflectance spectra from the fur and skin of many primate species in order to provide an objective survey of the possibilities of pelage coloration found in extant primates. We show that the orange coloration displayed by many platyrrhine and some strepsirhine primates, while being conspicuous to humans, would be cryptic amongst foliage to all males and many females of their own species. In relation to this finding, we briefly review what is known of the color vision of birds that prey on primates, and assess how conspicuous the orange pelage would be to these predators.     

A review of Gloger's rule,an ecogeographical rule of colour: definitions,interpretations and evidence

Gloger's rule is an ecogeographical rule that links animal colouration with climatic variation. This rule is named after C.W.L. Gloger who was one of the first to summarise the associations between climatic variation and animal colouration, noting in particular that birds and mammals seemed more pigmented in tropical regions. The term ‘Gloger's rule' was coined by B. Rensch in 1929 and included different patterns of variation from those described by Gloger. Rensch defined the rule in two ways: a simple version stating that endothermic animals are predicted to be darker in warmer and humid areas due to the increased deposition of melanin pigments; and a complex version that includes the differential effects of humidity and temperature on both main types of melanin pigments – eu‐ and phaeo‐melanin. The blackish eu‐melanins are predicted to increase with humidity, and decrease only at extreme low temperatures, while the brown‐yellowish phaeomelanins prevail in dry and warm regions and decrease rapidly with lower temperatures. A survey of the literature indicates that there is considerable variation/confusion in the way Gloger's rule is understood (based on 271 studies that define the rule). Whereas the complex version is hardly mentioned, only a quarter of the definitions are consistent with the simple version of Gloger's rule (darker where warm and wet), and most definitions mention only the effects of humidity (darker where wet). A smaller subset of studies define the rule based on other correlated climatic and environmental variables such as vegetation, latitude, altitude, solar radiation, etc., and a few even contradict the original definition (darker where cold). Based on the literature survey, I synthesised the qualitative (N = 124 studies) and quantitative (meta‐analytically, N = 38 studies, 241 effects) evidence testing the simple version of Gloger's rule (I found no tests of the complex version). Both lines of evidence supported the predicted effects of humidity (and closely linked variables) on colour variation, but not the effects of temperature. Moreover, humidity effects are not restricted to birds and mammals, as the data indicate that these effects also apply to insects. This suggests that the simple version of Gloger's rule as originally defined may not be valid, and possibly that the rule should be re‐formulated in terms of humidity effects only. I suggest, however, that more data are needed before such a reformulation, due to potential publication biases. In conclusion, I recommend that authors cite Rensch when referring to Gloger's rule and that they make clear which version they are referring to. Future research should concentrate on rigorously testing the validity and generality of both versions of Gloger's rule and establishing the mechanism(s) responsible for the patterns it describes. Since humidity seems to be the core climatic variable behind Gloger's rule, I suggest that the two most plausible mechanisms are camouflage and protection against parasites/pathogens, the latter possibly through pleiotropic effects on the immune system. Understanding the processes that lead to climatic effects on animal colouration may provide insights into past and future patterns of adaptation to climatic change.     

哺乳动物毛被传热性能及其影响因素

毛被能够加强或减弱动物向周围环境的热量散失,毛被的形态结构和颜色是传热性能的决定因素,其传热过程往往是传导、对流和辐射3个过程的耦合。以往研究发现环境因子中,风可增加机体向环境中的散热速率,且散失量与风速正相关,且动物通过调节在风场中的姿态来适应不同风向。动物体与环境间的温差是影响散热速率的另一因素,不同环境中的动物通过改变毛被结构来适应温差变化。毛被含水率上升会引起导热和蒸发冷却作用加强,动物通过行为或毛被结构变化来调节毛被含水率。毛色决定毛被吸收和反射热辐射的能力。毛被传热性能直接把动物的生理特点与环境因子关联起来,这对揭示动物的适应、进化都具有重要意义。同时提出,毛被结构和传热性能的研究还有助于仿生学意义的挖掘。因此,今后应重点在毛被结构和物理性能、研究技术与方法以及毛被生物学和仿生学意义等方面开展研究。     

The adaptive significance of coloration in lagomorphs

Lagomorph pelage coloration was matched to habitat type, geographical region, altitude and behaviour to explore the adaptive significance of coloration patterns in this little-studied order of mammals. Analyses were conducted with and without taking phylogeny into account. The former analyses were based on a weighted, phylogenetic supertree for all extant species of lagomorphs that we constructed using morphological and molecular data from 146 papers in the literature. Although our analyses represent an initial, somewhat crude investigation, several clear trends are evident. First, overall body coloration across lagomorphs tends to match the background as shown for pale and red coloration and perhaps seasonal pelage change. The case for countershading being a method of concealment is far less strong. Second, ear tips appear to have a communicative role since they are conspicuous in many different habitats. Third, hypotheses for tail tips having a communicative role, for extremities being dark for physiological reasons, and for Gloger's rule received only partial support.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79, 309–328.     

Morphological Variation in Wild Marmosets (Callithrix penicillata and C. geoffroyi) and Their Hybrids

Evolutionary theory and observation predict wider phenotypic variation in hybrids than parental species. Emergent phenotypic novelty in hybrids may in turn drive new adaptations or speciation by breaking parental phenotypic constraints. Primate hybridization is often documented through genetic evidence, but knowledge about the primate hybrid phenotype remains limited due to a small number of available studies on hybrid primate morphology. Here, we examine pelage and morphometric variation in two Brazilian marmoset species (Callithrix penicillata and C. geoffroyi) and their hybrids. Hybrids were sampled in an anthropogenic hybrid zone in the municipality of Viçosa, Minas Gerais state, Brazil. We analyzed hybrid facial and body pelage color variation, and compared 13 morphometric measures between hybrids and parental species. Five different hybrid facial morphotypes were observed, varying from intermediate to parental-like. Hybrid facial morphotypes were biased towards C. penicillata, suggesting that the pelage of this species may be dominant to that of C. geoffroyi in this context, and indicating that mate preference, and therefore gene flow/introgression, may be biased towards C. penicillata within the hybrid zone. Hybrid morphometric features were on average intermediate to parental species traits, but transgressive hybrids were also observed, suggesting that morphometric variation for the studied traits is consistent with Rieseberg’s complementary allele model. Finally, we observed a decoupling of facial patterning and size/shape in hybrids, relative to parent phenotypes, suggesting that an important factor driving phenotypic novelty within the Viçosa marmoset hybrid zone might be the loosening of evolutionary constraints on phenotypic trait integration.     

Gloger's rule in plants: The species and ecosystem levels

Gloger''s rule posits that darker birds are found more often in humid environments than in arid ones, especially in the tropics. Accordingly, desert-inhabiting animals tend to be light-colored. This rule is also true for certain mammalian groups, including humans. Gloger''s rule is manifested at 2 levels: (1) at the species level (different populations of the same species have different pigmentation at different latitudes), and (2) at the species assembly level (different taxa at a certain geography have different pigmentation than other taxa found at different habitats or latitudes). Concerning plants, Gloger''s rule was first proposed to operate in many plant species growing in sand dunes, sandy shores and in deserts, because of being white, whitish, or silver colored, based on white trichomes, because of sand grains and clay particles glued to sticky glandular trichomes, or because of light-colored waxes. Recently, Gloger''s rule was shown to also be true at the intraspecific level in relation to protection of anthers from UV irradiation. While Gloger''s rule is true in certain plant taxa and ecologies, there are others where “anti-Gloger” coloration patterns exist. In some of these the selective agents are known and in others they are not. I present both Gloger and “anti-Gloger” cases and argue that this largely neglected aspect of plant biology deserves much more research attention.     

Contrasting coloration in terrestrial mammals

Here I survey, collate and synthesize contrasting coloration in 5000 species of terrestrial mammals focusing on black and white pelage. After briefly reviewing alternative functional hypotheses for coloration in mammals, I examine nine colour patterns and combinations on different areas of the body and for each mammalian taxon to try to identify the most likely evolutionary drivers of contrasting coloration. Aposematism and perhaps conspecific signalling are the most consistent explanations for black and white pelage in mammals; background matching may explain white pelage. Evidence for contrasting coloration is being involved in crypsis through pattern blending, disruptive coloration or serving other functions, such as signalling dominance, lures, reducing eye glare or in temperature regulation has barely moved beyond anecdotal stages of investigation. Sexual dichromatism is limited in this taxon and its basis is unclear. Astonishingly, the functional significance of pelage coloration in most large charismatic black and white mammals that were new to science 150 years ago still remains a mystery.     

Variation of coat color in house mice throughout Asia

Coat color variation due to melanin pigment synthesis in house mice Mus musculus in Asia is described and found to be consistent with Gloger's rule, which states that individuals of endothermic animals are darker in humid habitats than those in drier habitats. Three properties of coat color (hue, value and chroma) were measured, and a lightness variable was derived from a principal components analysis using 428 skin specimens representing three subspecies from 85 localities. Dorsal coat color ranged from yellow through brown to black, whereas ventral coat color ranged from white to black. Dorsal coat color varied less than the ventral color. In our samples, the variation in coat color in natural populations was far less than that observed in the laboratory. We found a significant correlation between the lightness variable of dorsal coat color and precipitation. Dark coat color was observed in more humid and closed habitats (darker background color), and pale coat color in drier, more open habitats (lighter background color). This result might imply the role of concealment as a selective force affecting dorsal coat color that was observed in house mice. We also discussed other selective forces that could affect the coat color variation in house mice, such as resistance to bacterial degradation and thermoregulation. In addition, the color spectra of the dorsal pelage among the three subspecies were different, the major distinction being the environmental background color of the habitats in which they are distributed.     

Aposematic signaling and seasonal variation in dorsal pelage in a venomous mammal

In mammals, colouration patterns are often related to concealment, intraspecific communication, including aposematic signals, and physiological adaptations. Slow lorises (Nycticebus spp.) are arboreal primates native to Southeast Asia that display stark colour contrast, are highly territorial, regularly enter torpor, and are notably one of only seven mammal taxa that possess venom. All slow loris species display a contrasting stripe that runs cranial‐caudally along the median sagittal plane of the dorsum. We examine whether these dorsal markings facilitate background matching, seasonal adaptations, and intraspecific signaling. We analyzed 195 images of the dorsal region of 60 Javan slow loris individuals (Nycticebus javanicus) from Java, Indonesia. We extracted greyscale RGB values from dorsal pelage using ImageJ software and calculated contrast ratios between dorsal stripe and adjacent pelage in eight regions. We assessed through generalized linear mixed models if the contrast ratio varied with sex, age, and seasonality. We also examined whether higher contrast was related to more aggressive behavior or increased terrestrial movement. We found that the dorsal stripe of N. javanicus changed seasonally, being longer and more contrasting in the wet season, during which time lorises significantly increased their ground use. Stripes were most contrasting in younger individuals of dispersal age that were also the most aggressive during capture. The dorsal stripe became less contrasting as a loris aged. A longer stripe when ground use is more frequent can be related to disruptive colouration. A darker anterior region by younger lorises with less fighting experience may allow them to appear larger and fiercer. We provide evidence that the dorsum of a cryptic species can have multimodal signals related to concealment, intraspecific communication, and physiological adaptations.     

Transformation of bird and mammal faunas in steppe ecosystems under the impact of plowing: The example of Saratov steppes

The impact of plowing on the bird and mammal faunas of steppe landscapes is considered using the example of steppes in the Saratov Region. It is shown that the faunas of ground-nesting birds and small mammals in crop-rotation fields represent impoverished variants of the same faunas in natural steppe land-scapes with their characteristic diversity of habitats. As a rule, dominance in the fields belongs to bird and mammal species that, prior to extensive plowing of the steppes, inhabited intrazonal landscape elements. Only a few species characteristic of zonal steppe habitats have adapted to living in crop-rotation fields.     

Responses of Primates to Landscape Change in Amazonian Land‐bridge islands—a Multi‐scale Analysis

Large hydroelectric dams are one of the current drivers of habitat loss across Amazonian forests. We investigated how the primate community at a hydroelectric dam in Brazilian Amazonia responded to changes in the landscape and local habitat structure of land‐bridge islands after 21 yr of post‐isolation history. The Balbina Dam, constructed in 1986, inundated 3129 km² of primary forests and created more than 3500 variable‐sized islands. We conducted primate and habitat structure surveys on 20 islands from 5 to 1815 ha, and extracted forest patch and landscape metrics for each island. The number of primate species per island varied between 0 and 7 species. Primate composition varied substantially according to both island area and forest cover remaining within the landscape, whereas island area alone was the most significant predictor of richness. Locally, tree density and vertical stratification were the most significant explanatory variables of primate composition and richness. A model containing area effects had the most explanatory power regarding site occupancy for most species. Individually, each species responded differently, with howler and brown capuchin monkeys showing greater tolerance to cope with habitat changes. Body size was also an important predictor of primate occupancy. We recommend protecting large fragments and enhancing the suitability of surrounding habitats to ensure primate conservation in most Neotropical fragmented landscapes. Given the flat topography of hydroelectric reservoirs, which mainly favors the formation of small islands, and the escalating hydropower development plans in Amazonia, our findings provide evidence for pervasive detrimental impacts of dams on primate communities.     

Support for the elevational Rapoport's rule among seed plants in Nepal depends on biogeographical affinities and boundary effects

As one of the most important hypotheses on biogeographical distribution, Rapoport's rule has attracted attention around the world. However, it is unclear whether the applicability of the elevational Rapoport's Rule differs between organisms from different biogeographical regions. We used Stevens’ method, which uses species diversity and the averaged range sizes of all species within each (100 m) elevational band to explore diversity‐elevation, range‐elevation, and diversity‐range relationships. We compared support for the elevational Rapoport's rule between tropical and temperate species of seed plants in Nepal. Neither tropical nor temperate species supported the predictions of the elevational Rapoport's rule along the elevation gradient of 100–6,000 m a.s.l. for any of the studied relationships. However, along the smaller 1,000–5,000 m a.s.l. gradient (4,300 m a.s.l. for range‐elevation relationships) which is thought to be less influenced by boundary effects, we observed consistent support for the rule by tropical species, although temperate species did not show consistent support. The degree of support for the elevational Rapoport's rule may not only be influenced by hard boundary effects, but also by the biogeographical affinities of the focal taxa. With ongoing global warming and increasing variability of temperature in high‐elevation regions, tropical taxa may shift upward into higher elevations and expand their elevational ranges, causing the loss of temperate taxa diversity. Relevant studies on the elevational Rapoport's rule with regard to biogeographical affinities may be a promising avenue to further our understanding of this rule.     

Heterothermy in large mammals: inevitable or implemented?

Advances in biologging techniques over the past 20 years have allowed for the remote and continuous measurement of body temperatures in free‐living mammals. While there is an abundance of literature on heterothermy in small mammals, fewer studies have investigated the daily variability of body core temperature in larger mammals. Here we review measures of heterothermy and the factors that influence heterothermy in large mammals in their natural habitats, focussing on large mammalian herbivores. The mean 24 h body core temperatures for 17 species of large mammalian herbivores (>10 kg) decreased by ～1.3°C for each 10‐fold increase in body mass, a relationship that remained significant following phylogenetic correction. The degree of heterothermy, as measured by the 24 h amplitude of body core temperature rhythm, was independent of body mass and appeared to be driven primarily by energy and water limitations. When faced with the competing demands of osmoregulation, energy acquisition and water or energy use for thermoregulation, large mammalian herbivores appear to relax the precision of thermoregulation thereby conserving body water and energy. Such relaxation may entail a cost in that an animal moves closer to its thermal limits for performance. Maintaining homeostasis requires trade‐offs between regulated systems, and homeothermy apparently is not accorded the highest priority; large mammals are able to maintain optimal homeothermy only if they are well nourished, hydrated, and not compromised energetically. We propose that the amplitude of the 24 h rhythm of body core temperature provides a useful index of any compromise experienced by a free‐living large mammal and may predict the performance and fitness of an animal.