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1.
Ben Smit  Andrew E. McKechnie 《Ibis》2015,157(4):774-786
Endothermic animals resident in hot, arid terrestrial environments are likely to face a trade‐off between their ability to obtain water and elevated thermoregulatory water requirements. We assessed whether daily water flux (DWF) is higher on hot days, reflecting increases in evaporative cooling demands, in an arid‐zone bird that obtains its water through food intake. We obtained measurements of DWF (partitioned into water influx and efflux rates) in 71 White‐browed Sparrow‐Weavers Plocepasser mahali at a desert site and a semi‐desert site, during summer in the Kalahari Desert of southern Africa. We found no evidence that DWF varied with maximum daily air temperature (Tair, range = 27.6–39.2 °C). Instead, DWF was lower during dry periods than in the wet season at the semi‐desert site. Furthermore, birds showed deficits in water balance (water influx/water efflux) during the dry periods at both sites. Our data show that DWF is low in a non‐drinking bird that obtains its water through food, and that demands for evaporative water loss on very hot days (maximum Tair of 40–44 °C) may exceed water intake rates during hot and dry periods. Species that do not have opportunities to drink will experience strong trade‐offs between thermoregulation, hydration state and activity levels as temperatures increase.  相似文献   

2.
Sexual size dimorphism (SSD) is a common phenomenon in animals. In many species females are substantially larger than males. Because body size plays a central role in modulating the body temperature (T b) of ectotherms, intersexual differences in body size may lead to important intersexual differences in thermoregulation. In addition, because SSD is realized by differences in growth rate and because growth rate is strongly temperature dependent in ectotherms, a conflict between male reproductive behaviour and thermoregulation may affect the expression of SSD. In this study, we investigated the thermal implications of SSD in a reptile exhibiting spectacular female-biased SSD: the northern map turtle (Graptemys geographica). Over three seasons, we collected >150,000 measurements of T b in free-ranging adult and juvenile northern map turtles using surgically implanted miniature temperature loggers. Northern map turtles exhibited seasonal patterns of thermoregulation typical of reptiles in northern latitudes, but we found that large adult females experienced a lower daily maximum T b and a narrower daily range of T b than adult males and small juvenile females. In addition, despite more time spent basking, large adult females were not able to thermoregulate as accurately as small turtles. Our findings strongly suggest that body size limits the ability to thermoregulate accurately in large females. By comparing thermoregulatory patterns between adult males and juvenile females of similar body size, we found no evidence that male reproductive behaviours are an impediment to thermoregulation. We also quantified the thermal significance of basking behaviour. We found, contrary to previous findings, that aerial basking allows northern map turtles to raise their T b substantially above water temperature, indicating that basking behaviour likely plays an important role in thermoregulation.  相似文献   

3.
Differences in predator‐inspection behaviour between gravid and non‐gravid female as well as between male and female three‐spined sticklebacks Gasterosteus aculeatus were investigated. Gravid females confronted with a live rainbow trout Oncorhynchus mykiss showed bolder inspection behaviour than non‐gravid ones. The behaviour of gravid females was comparable with that of males, maybe because both face a high risk of predation. The results indicate that antipredator behaviour in female G. aculeatus is not fixed but adjusted to their reproductive state.  相似文献   

4.
The objective of this study was to estimate reproductive and population parameters of the spiny dogfish Squalus acanthias for the south‐western Atlantic Ocean. In total, 2714 specimens (1616 males and 1098 females) were collected from surveys carried out using research vessels. Males ranged from 225 to 861 mm total length (LT) and females from 235 to 925 mm LT. The size at maturity of females (651 mm) was significantly greater than that of males (565 mm). The maximum proportion of mature individuals (Pmax) of the gestation ogive was <1, which indicates that a proportion of mature females was not in gestation. This inactivity may be explained by the occurrence of resting periods between cycles or by the asynchrony of the reproductive cycle. The estimated Pmax for the maternity ogive suggested that about one third of mature females were in the maternity stage (i.e. with embryos >156 mm). The temporal and spatial co‐occurrence of non‐gravid adult females at different stages of ovarian development, as well as gravid females at all embryonic development stages would indicate that the female reproductive cycle in the south‐western Atlantic Ocean is asynchronous. The results indicate that S. acanthias is susceptible to fishing pressure on account of its length at maturity, extended reproductive cycles and low fecundity.  相似文献   

5.
It has been proposed that there is a thermal cost of the mane to male lions, potentially leading to increased body surface temperatures (Ts), increased sperm abnormalities, and to lower food intake during hot summer months. To test whether a mane imposes thermal costs on males, we measured core body temperature (Tb) continuously for approximately 1 year in 18 free‐living lions. There was no difference in the 24‐hr maximum Tb of males (n = 12) and females (n = 6), and males had a 24‐hr mean Tb that was 0.2 ± 0.1°C lower than females after correcting for seasonal effects. Although feeding on a particular day increased 24‐hr mean and 24‐hr maximum Tb, this phenomenon was true of both male and female lions, and females had higher 24‐hr mean and 24‐hr maximum Tb than males, on both days when lions did not feed, and on days when lions did feed. Twenty‐four‐hour Tb was not influenced by mane length or color, and 24‐hr mean Tb was negatively correlated with mane length. These data contradict the suggestion that there exists a thermal cost to male lions in possessing a long dark mane, but do not preclude the possibility that males compensate for a mane with increased heat loss. The increased insulation caused by a mane does not necessarily have to impair heat loss by males, which in hot environments is primarily through respiratory evaporative cooling, nor does in necessarily lead to increased heat gain, as lions are nocturnal and seek shade during the day. The mane may even act as a heat shield by increasing insulation. However, dominant male lions frequent water points more than twice as often as females, raising the possibility that male lions are increasing water uptake to facilitate increased evaporative cooling. The question of whether male lions with manes compensate for a thermal cost to the mane remains unresolved, but male lions with access to water do not have higher Tb than females or males with smaller manes.  相似文献   

6.
Life‐history theory predicts that access to limited resources leads to trade‐offs between competing body functions. Women, who face higher costs of reproduction when compared to men, should be especially vulnerable to these trade‐offs. We propose the ‘cognitive costs of reproduction hypothesis’, which states that energy trade‐offs imposed by reproduction may lead to a decline in maternal cognitive function during gestation. In particular, we hypothesize that the decline in cognitive function frequently observed during pregnancy is associated with the allocation of resources between the competing energetic requirements of the mother's brain and the developing foetus. Several distinctive anatomical and physiological features including a high metabolic rate of the brain, large infant size, specific anatomical features of the placenta and trophoblast, and the lack of maternal control over glucose flow through the placenta make the occurrence of these trade‐offs likely. Herein, we review several lines of evidence for trade‐offs between gestation and cognition that are related to: (i) energy metabolism during reproduction; (ii) energy metabolism of the human brain; (iii) links between energy metabolism and cognitive function; and (iv) links between gestation and cognitive function. We also review evidence for the important roles of cortisol, corticotropin‐releasing hormone and sex hormones in mediating the effects of gestation on cognition, and we discuss possible neurophysiological mechanisms underlying the observed effects. The evidence supports the view that energy trade‐offs between foetal growth and maternal endocrine and brain function lead to changes in maternal cognition, and that this phenomenon is mediated by neuroendocrine mechanisms involving the hypothalamic–pituitary–adrenal axis, brainstem nucleus locus coeruleus and hippocampus.  相似文献   

7.
Previous leaf‐scale studies of carbon assimilation describe short‐term resource‐use efficiency (RUE) trade‐offs where high use efficiency of one resource requires low RUE of another. However, varying resource availabilities may cause long‐term RUE trade‐offs to differ from the short‐term patterns. This may have important implications for understanding canopy‐scale resource use and allocation. We used continuous gas exchange measurements collected at five levels within a Norway spruce, Picea abies (L.) karst., canopy over 3 years to assess seasonal differences in the interactions between shoot‐scale resource availability (light, water and nitrogen), net photosynthesis (An) and the use efficiencies of light (LUE), water (WUE) and nitrogen (NUE) for carbon assimilation. The continuous data set was used to develop and evaluate multiple regression models for predicting monthly shoot‐scale An. These models showed that shoot‐scale An was strongly dependent on light availability and was generally well described with simple one‐ or two‐parameter models. WUE peaked in spring, NUE in summer and LUE in autumn. However, the relative importance of LUE for carbon assimilation increased with canopy depth at all times. Our results suggest that accounting for seasonal and within‐canopy trade‐offs may be important for RUE‐based modelling of canopy carbon uptake.  相似文献   

8.
Natural populations respond to selection pressures like increasing local temperatures in many ways, including plasticity and adaptation. To predict the response of ectotherms like lizards to local temperature increase, it is essential to estimate phenotypic variation in and determine the heritability of temperature‐related traits like average field body temperature (Tb) and preferred temperature (Tp). We measured Tp of Uta stansburiana in a laboratory thermal gradient and assessed the contribution of sex, reproductive status and throat color genotype to phenotypic variation in Tb of adult lizards. Females had higher Tp than males. However, they temporarily preferred lower temperature when gravid than when nongravid. Using a nested half‐sib design for genetic crosses in the laboratory, we estimated relative contributions of additive genetic variation and maternal effects to Tp of hatchlings. Our results show that maternal effects, but not additive genetic variation, influence Tp of hatchlings in U. stansburiana. Maternal Tp and the presence or absence of blue throat color alleles significantly influenced Tp of hatchlings. We discuss ecological and evolutionary consequences of these maternal effects in the context of rapid climate change and natural selection that we measure on progeny survival to maturity as a function of maternal Tp.  相似文献   

9.
Deania calcea (n = 420) were collected from the catch of deep‐water trawlers in the southern and eastern scalefish and shark fishery in southern Australia during the years 2008–2011. The total length (LT) range varied between sexes, females being larger (n = 264; 280–1530 mm) than males (n = 156; 310–921 mm). The reproductive cycle in this population is non‐continuous and asynchronous. The estimated LT at which 50% of males are mature is 807 mm and is 914 mm for females. Populations of D. calcea in higher latitudes appear to mature at a larger size than conspecifics in lower latitudes, in both the northern and southern hemispheres. Litters ranged from three to 10 embryos with a 1:1 sex ratio, but litter size does not increase with maternal LT. Deania calcea shows geographical variability in its biological parameters and gathering information on life‐history traits of populations is vital to understand the trade‐offs made by this species in response to environmental conditions and to predict intraspecific spatial differences. Such information is a basis for specific spatial management to protect populations from excessive fishing.  相似文献   

10.
Endotherms allocate large amounts of energy and water to the regulation of a precise body temperature (Tb), but can potentially reduce thermoregulatory costs by allowing Tb to deviate from normothermic levels. Many data on heterothermy at low air temperatures (Ta) exist for caprimulgids, whereas data on thermoregulation at high Ta are largely absent, despite members of this taxon frequently roosting and nesting in sites exposed to high operative temperatures. We investigated thermoregulation in free‐ranging rufous‐cheeked nightjars Caprimulgus rufigena and freckled nightjars Caprimulgus tristigma in the southern African arid zone. Individuals of both species showed labile Tb fluctuating around a single modal Tb (Tb‐mod). Average Tb‐mod was 39.7°C for rufous‐cheeked nightjars and 39.0°C for freckled nightjars. In both species, diurnal Tb increased with increasing Ta. At Ta ≥ 38°C, rufous‐cheeked nightjar mean Tb increased to 42°C, equivalent to 2.3°C above Tb‐mod. Under similar conditions, freckled nightjar Tb was on average only 1.1°C above Tb‐mod, with a mean Tb of 40.0°C. Freckled nightjars are one of the most heterothermic caprimulgids investigated to date, but our data suggest that during hot conditions this species maintains Tb within a narrow range above Tb‐mod, possibly reflecting an evolutionary tradeoff between decreased thermal sensitivity to lower Tb but increased sensitivity to high Tb. These findings reveal how general thermoregulatory patterns at similar Ta can vary even among closely related species.  相似文献   

11.
Life‐history traits such as fecundity and offspring size are shaped by investment trade‐offs faced by mothers and mediated by environmental conditions. We use a 21‐year time series for three populations of wild sockeye salmon (Oncorhynchus nerka) to test predictions for such trade‐offs and responses to conditions faced by females during migration, and offspring during incubation. In years when their 1100 km upstream migration was challenged by high water discharges, females that reached spawning streams had invested less in gonads by producing smaller but not fewer eggs. These smaller eggs produced lighter juveniles, and this effect was further amplified in years when the incubation water was warm. This latter result suggests that there should be selection for larger eggs to compensate in populations that consistently experience warm incubation temperatures. A comparison among 16 populations, with matching migration and rearing environments but different incubation environments (i.e., separate spawning streams), confirmed this prediction; smaller females produced larger eggs for their size in warmer creeks. Taken together, these results reveal how maternal phenotype and environmental conditions can shape patterns of reproductive investment and consequently juvenile fitness‐related traits within and among populations.  相似文献   

12.
The evolution of floral display is thought to be constrained by trade‐offs between the size and number of flowers; however, empirical evidence for the trade‐off is inconsistent. We examined evidence for trade‐offs and hierarchical allocation of resources within and between two populations each of the monocarpic perennials, Cardiocrinum cordatum and C. giganteum. Within all populations, flower size–number trade‐offs were evident after accounting for variation in plant size. In addition, variation in flower size explained much variation in flower‐level allocation to attraction, and female and male function, a pattern consistent with hierarchical allocation. However, between population differences in flower size (C. cordatum) and number (C. giganteum) were not consistent with size–number trade‐offs or hierarchical allocation. The population‐level difference in C. cordatum likely reflects the combined influence of a time lag between initiation and maturation of flowers, and higher light levels in one population. Thus, our study highlights one mechanism that may account for the apparent independence of flower size and number in many studies. A prediction of sex allocation theory was also supported. In C. giganteum: plants from one population invested more mass in pistils and less in stamens than did plants from the other population. Detection of floral trade‐offs in Cardiocrinum may be facilitated by monocarpic reproduction, production of a single inflorescence and ease of measuring plant size.  相似文献   

13.
Precise measures of phenology are critical to understanding how animals organize their annual cycles and how individuals and populations respond to climate-induced changes in physical and ecological stressors. We show that patterns of core body temperature (T b) can be used to precisely determine the timing of key seasonal events including hibernation, mating and parturition, and immergence and emergence from the hibernacula in free-living arctic ground squirrels (Urocitellus parryii). Using temperature loggers that recorded T b every 20 min for up to 18 months, we monitored core T b from three females that subsequently gave birth in captivity and from 66 female and 57 male ground squirrels free-living in the northern foothills of the Brooks Range Alaska. In addition, dates of emergence from hibernation were visually confirmed for four free-living male squirrels. Average T b in captive females decreased by 0.5–1.0°C during gestation and abruptly increased by 1–1.5°C on the day of parturition. In free-living females, similar shifts in T b were observed in 78% (n = 9) of yearlings and 94% (n = 31) of adults; females without the shift are assumed not to have given birth. Three of four ground squirrels for which dates of emergence from hibernation were visually confirmed did not exhibit obvious diurnal rhythms in T b until they first emerged onto the surface when T b patterns became diurnal. In free-living males undergoing reproductive maturation, this pre-emergence euthermic interval averaged 20.4 days (n = 56). T b-loggers represent a cost-effective and logistically feasible method to precisely investigate the phenology of reproduction and hibernation in ground squirrels.  相似文献   

14.
Despite being ectotherms, reptiles have an ability to thermoregulate which is enhanced by adopting a variety of behavioural mechanisms. Different behavioural postures, the use of retreat sites and selection of microhabitats enable reptiles to maintain their core body temperatures (Tb) above that of ambient temperatures (Ta) in winter or below Ta maximum in summer. This study describes the daily activity patterns of leopard tortoises (Stigmochelys pardalis) in relation to Tb and Ta, and the extent to which leopard tortoises can manipulate their Tb in response to seasonal changes in Ta. Ten and nine leopard tortoises were radio-tracked in 2002 and 2003, respectively and cloacal Tb and behaviours observed. Core Tbs were measured using Thermocron iButtons surgically implanted into the body cavities of 4 and 5 adult telemetered tortoises for summer and winter 2003, respectively. There were seasonal differences in the extent to which certain behaviours were practiced and the time of day that these occurred. Leopard tortoises generally had unimodal activity patterns in winter and bimodal ones in summer. In winter tortoises were active at lower Tbs, and at lower Ta, than in summer. Tortoises maintained their core Tb well below Ta minimum profiles in summer and well above these in winter. Core Tb closely followed the increase in Ta minimum profiles in the mornings, however tortoises exhibited an extended thermal lag when Ta minimum profiles cooled overnight. By using different behavioural mechanisms in summer and winter, leopard tortoises maintained their core Tb at different levels compared with Ta minimum and maximum profiles. Consequently although they are ectotherms, they maintained their core Tb independent of Ta.  相似文献   

15.
The jerboa (Jaculus orientalis) has been described in the past as a hibernator, but no reliable data exist on the daily and seasonal rhythmicity of body temperature (T b). In this study, T b patterns were determined in different groups of jerboas (isolated males and females, castrated males and grouped animals) maintained in captivity during autumn and winter, and submitted to natural variations of light and ambient temperature (T a). T b and T a variations were recorded with surgically implanted iButton temperature loggers at 30-min intervals for two consecutive years. About half (6/13) of isolated female jerboas hibernated with a T b < 33°C, with hibernation bouts interspersed with short periods of normothermy from November to February. Hibernation bout durations were longer (4–5 days) than those of normothermia phases (1–4 days). During hibernation, the minimum T b was low (T bmin ~10.7°C). In contrast, one of the 12 isolated males showed short hibernation bouts of ca. 2 days late in the hibernation season, February–March. The males had T bmin values of 15.1°C. In contrast to predictions, no castrated males hibernated. When jerboas were grouped, females and males exhibited concomitant torpor bouts. In males, the longest bouts were observed during the late hibernation season. These data suggest complex regulation of hibernation in jerboas.  相似文献   

16.
Energy allocation theory predicts that a lactating female should alter the energetic demands of its organ systems in a manner that maximizes nutrient allocation to reproduction while reducing nutrient use for tasks that are not vital to immediate survival. We posit that organ‐specific plasticity in the function of mitochondria plays a key role in mediating these energetic trade‐offs. The goal of this project was to evaluate mitochondrial changes that occur in response to lactation in two of the most energetically demanding organs in the body of a rodent, the liver and skeletal muscle. This work was conducted in wild‐derived house mice (Mus musculus) kept in seminatural enclosures that allow the mice to maintain a natural social structure and move within a home range size typical of wild mice. Tissues were collected from females at peak lactation and from age‐matched nonreproductive females. Mitochondrial respiration, oxidative damage, antioxidant, PGC‐1α, and uncoupling protein levels were compared between lactating and nonreproductive females. Our findings suggest that both liver and skeletal muscle downregulate specific antioxidant proteins during lactation. The liver, but not skeletal muscle, of lactating females displayed higher oxidative damage than nonreproductive females. The liver mass of lactating females increased, but the liver displayed no change in mitochondrial respiratory control ratio. Skeletal muscle mass and mitochondrial respiratory control ratio were not different between groups. However, the respiratory function of skeletal muscle did vary among lactating females as a function of stage of concurrent pregnancy, litter size, and mass of the mammary glands. The observed changes are predicted to increase the efficiency of skeletal muscle mitochondria, reducing the substrate demands of skeletal muscle during lactation. Differences between our results and prior studies highlight the role that an animals’ social and physical environment could play in how it adapts to the energetic demands of reproduction.  相似文献   

17.
Adaptation of one set of traits is often accompanied by attenuation of traits important in other selective environments, leading to fitness trade‐offs. The mechanisms that either promote or prevent the emergence of trade‐offs remain largely unknown, and are difficult to discern in most systems. Here, we investigate the basis of trade‐offs that emerged during experimental evolution of Methylobacterium extorquens AM1 to distinct growth substrates. After 1500 generations of adaptation to a multi‐carbon substrate, succinate (S), many lineages had lost the ability to use one‐carbon compounds such as methanol (M), generating a mixture of M+ and M? evolved phenotypes. We show that trade‐offs in M? strains consistently arise via antagonistic pleiotropy through recurrent selection for loss‐of‐function mutations to ftfL (formate‐tetrahydrofolate ligase), which improved growth on S while simultaneously eliminating growth on M. But if loss of FtfL was beneficial, why were M trade‐offs not found in all populations? We discovered that eliminating FtfL was not universally beneficial on S, as it was neutral or even deleterious in certain evolved lineages that remained M+. This suggests that sign epistasis with earlier arising mutations prevented the emergence of mutations that drove trade‐offs through antagonistic pleiotropy, limiting the evolution of metabolic specialists in some populations.  相似文献   

18.
The seasonal availability of food resources is an important factor shaping the life‐history strategies of organisms. During times of nutritional restriction, physiological trade‐offs can induce periods of immune suppression, thereby increasing susceptibility to infectious disease. Our goal was to provide a conceptual framework describing how the endemic level bovine brucellosis (Brucella abortus) may be maintained in Yellowstone bison based on the seasonality of food resources and the life‐history strategies of the host and pathogen. Our analysis was based on active B. abortus infection (measured via bacterial culture), nutritional indicators (measured as metabolites and hormones in plasma), and carcass measurements of 402 slaughtered bison. Data from Yellowstone bison were used to investigate (1) whether seasonal changes in diet quality affect nutritional condition and coincide with the reproductive needs of female bison; (2) whether active B. abortus infection and infection intensities vary with host nutrition and nutritional condition; and (3) the evidence for seasonal changes in immune responses, which may offer protection against B. abortus, in relation to nutritional condition. Female bison experienced a decline in nutritional condition during winter as reproductive demands of late gestation increased while forage quality and availability declined. Active B. abortus infection was negatively associated with bison age and nutritional condition, with the intensity of infection negatively associated with indicators of nutrition (e.g., dietary protein and energy) and body weight. Data suggest that protective cell‐mediated immune responses may be reduced during the B. abortus transmission period, which coincides with nutritional insufficiencies and elevated reproductive demands during spring. Our results illustrate how seasonal food restriction can drive physiological trade‐offs that suppress immune function and create infection and transmission opportunities for pathogens.  相似文献   

19.
Correlated responses to bi‐directional selection on thorax length, examined on several life‐history traits and chromosome inversion polymorphisms, have revealed apparent novel trade‐offs in Drosophila ananassae. We provide evidence of trade‐offs between hatching time and pupal period, pupal period and egg‐pupa development time, and pupal period and larval development time (LDT). Body size shows positive correlations with ovariole number, LDT and DT (egg–fly). We provide evidence of sexual dimorphism for trade‐offs between longevity and body size and starvation and longevity in females only. Trade‐offs between wing/thorax (W/T) ratio and longevity, W/T ratio and starvation, and DT (egg‐ fly) and longevity are evident in males only. Sexual dimorphism is also evident for inversion polymorphism with body size and longevity. A longevity assay suggests that low line females outlived high line females whereas high line males outlived low line males. The mean longevity in males is negatively correlated with the 2L‐ST and 3R‐ST arrangement frequencies whereas the 3L‐ST arrangement frequency is positively correlated with the mean longevity in males but opposite arrangements are found in females. Absolute starvation resistance is negatively correlated with 2L‐ST and 3R‐ST chromosome arrangements and results in a trade‐off between longevity and absolute starvation resistance in females. Analyses of fecundity, hatchability, and viabilities based on age intervals in both G10 and G13 suggest that the early reproduction is favoured in D. ananassae. The productivity percentage is highest in the high line and there is no effect of late reproduction on it. Overall, we provide some unravelled trade‐offs and striking sex differences, which may help in understanding the life‐history evolution of the species. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 90 , 669–685.  相似文献   

20.
J. C. Lee 《Oecologia》1980,44(2):171-176
Summary In a habitat judged to be energetically costly for thermoregulation, mean body temperatures (MBT's) ofAnolis sagrei are significantly higher than those ofA. distichus. As indexed by the slope of the regression of body temperatures (T b ) on substrate temperature (T s ),A. sagrei is more dependent upon environmental temperatures thanA. distichus.In a habitat judged to be less costly for thermoregulation and where interspecific competition for perch sites may be less, MBT's ofA. sagrei are significantly higher, proportionally more lizards occupy sunny perches, and the slope of the regression of T b on T s is significantly less, than for conspecifics in the costly habitat.As indexed by length-specific fat body weights, well-nourished lizards in the costly habitat have T b 's which are independent of environmental temperature; T b 's of poorly-nourished lizards are highly dependent upon environmental temperature. This relationship does not hold for lizards in the low-cost habitat.These results corroborate the hypothesis that energetic costs are important in controlling the extent to which lizards thermoregulate. In high-cost habitats lizards thermoregulate less precisely than in low-cost habitats. Lizards that exploit the habitat as if it were highly productive thermoregulate more precisely than lizards that exploit the environment as if it were of low productivity.  相似文献   

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