Part B: Revisions to the COMFA outdoor thermal comfort model for application to subjects performing physical activity

This study assessed the performance of the COMFA outdoor thermal comfort model on subjects performing moderate to vigorous physical activity. Field tests were conducted on 27 subjects performing 30 min of steady-state activity (walking, running, and cycling) in an outdoor environment. The predicted COMFA budgets were compared to the actual thermal sensation (ATS) votes provided by participants during each 5-min interval. The results revealed a normal distribution in the subjects’ ATS votes, with 82% of votes received in categories 0 (neutral) to +2 (warm). The ATS votes were significantly dependent upon sex, air temperature, short and long-wave radiation, wind speed, and metabolic activity rate. There was a significant positive correlation between the ATS and predicted budgets (Spearman’s rho = 0.574, P < 0.01). However, the predicted budgets did not display a normal distribution, and the model produced erroneous estimates of the heat and moisture exchange between the human body and the ambient environment in 6% of the cases.     

Thermal comfort modelling of body temperature and psychological variations of a human exercising in an outdoor environment

Human thermal comfort assessments pertaining to exercise while in outdoor environments can improve urban and recreational planning. The current study applied a simple four-segment skin temperature approach to the COMFA (COMfort FormulA) outdoor energy balance model. Comparative results of measured mean skin temperature ( [`(T)]\nolimits<sub>Msk</sub> \mathop{{\bar{T}}}\nolimits_{{Msk}} ) with predicted [`(T)]\nolimits_sk \mathop{{\bar{T}}}\nolimits_{{sk}} indicate that the model accurately predicted [`(T)]\nolimits<sub>sk</sub> \mathop{{\bar{T}}}\nolimits_{{sk}} , showing significantly strong agreement (r = 0.859, P < 0.01) during outdoor exercise (cycling and running). The combined 5-min mean variation of the [`(T)]\nolimits_sk \mathop{{\bar{T}}}\nolimits_{{sk}} RMSE was 1.5°C, with separate cycling and running giving RMSE of 1.4°C and 1.6°C, respectively, and no significant difference in residuals. Subjects’ actual thermal sensation (ATS) votes displayed significant strong rank correlation with budget scores calculated using both measured and predicted [`(T)]\nolimits<sub>sk</sub> \mathop{{\bar{T}}}\nolimits_{{sk}} (r <sub>s</sub>  = 0.507 and 0.517, respectively, P < 0.01). These results show improved predictive strength of ATS of subjects as compared to the original and updated COMFA models. This psychological improvement, plus [`(T)]\nolimits_sk \mathop{{\bar{T}}}\nolimits_{{sk}} and T _c validations, enables better application to a variety of outdoor spaces. This model can be used in future research studying linkages between thermal discomfort, subsequent decreases in physical activity, and negative health trends.     

Improved predictive ability of climate–human–behaviour interactions with modifications to the COMFA outdoor energy budget model

The purpose of this paper is to implement current and novel research techniques in human energy budget estimations to give more accurate and efficient application of models by a variety of users. Using the COMFA model, the conditioning level of an individual is incorporated into overall energy budget predictions, giving more realistic estimations of the metabolism experienced at various fitness levels. Through the use of VO₂ reserve estimates, errors are found when an elite athlete is modelled as an unconditioned or a conditioned individual, giving budgets underpredicted significantly by −173 and −123 W m⁻², respectively. Such underprediction can result in critical errors regarding heat stress, particularly in highly motivated individuals; thus this revision is critical for athletic individuals. A further improvement in the COMFA model involves improved adaptation of clothing insulation (I _cl), as well clothing non-uniformity, with changing air temperature (T _a) and metabolic activity (M _act). Equivalent T _a values (for I _cl estimation) are calculated in order to lower the I _cl value with increasing M _act at equal T _a. Furthermore, threshold T _a values are calculated to predict the point at which an individual will change from a uniform I _cl to a segmented I _cl (full ensemble to shorts and a T-shirt). Lastly, improved relative velocity (v _r) estimates were found with a refined equation accounting for the degree angle of wind to body movement. Differences between the original and improved v _r equations increased with higher wind and activity speeds, and as the wind to body angle moved away from 90°. Under moderate microclimate conditions, and wind from behind a person, the convective heat loss and skin temperature estimates were 47 W m⁻² and 1.7°C higher when using the improved v _r equation. These model revisions improve the applicability and usability of the COMFA energy budget model for subjects performing physical activity in outdoor environments. Application is possible for other similar energy budget models, and within various urban and rural environments.     

Case study of skin temperature and thermal perception in a hot outdoor environment

Focusing on the understanding and the estimation of the biometeorological conditions during summer in outdoor places, a field study was conducted in July 2010 in Athens, Greece over 6 days at three different sites: Syntagma Square, Ermou Street and Flisvos coast. Thermo-physiological measurements of five subjects were carried out from morning to evening for each site, simultaneously with meteorological measurements and subjective assessments of thermal sensation reported by questionnaires. The thermo-physiological variables measured were skin temperature, heat flux and metabolic heat production, while meteorological measurements included air temperature, relative humidity, wind speed, globe temperature, ground surface temperature and global radiation. The possible relation of skin temperature with the meteorological parameters was examined. Theoretical values of mean skin temperature and mean radiant temperature were estimated applying the MENEX model and were compared with the measured values. Two biometeorological indices, thermal sensation (TS) and heat load (HL)—were calculated in order to compare the predicted thermal sensation with the actual thermal vote. The theoretically estimated values of skin temperature were underestimated in relation to the measured values, while the theoretical model of mean radiant temperature was more sensitive to variations of solar radiation compared to the experimental values. TS index underestimated the thermal sensation of the five subjects when their thermal vote was ‘hot’ or ‘very hot’ and overestimated thermal sensation in the case of ‘neutral’. The HL index predicted with greater accuracy thermal sensation tending to overestimate the thermal sensation of the subjects.     

Assessment of daytime outdoor comfort levels in and outside the urban area of Glasgow,UK

To understand thermal preferences and to define a preliminary outdoor comfort range for the local population of Glasgow, UK, an extensive series of measurements and surveys was carried out during 19 monitoring campaigns from winter through summer 2011 at six different monitoring points in pedestrian areas of downtown Glasgow. For data collection, a Davis Vantage Pro2 weather station equipped with temperature and humidity sensors, cup anemometer with wind vane, silicon pyranometer and globe thermometer was employed. Predictions of the outdoor thermal index PET (physiologically equivalent temperature) correlated closely to the actual thermal votes of respondents. Using concurrent measurements from a second Davis Vantage Pro2 weather station placed in a rural setting approximately 15 km from the urban area, comparisons were drawn with regard to daytime thermal comfort levels and urban–rural temperature differences (?T_u-r) for the various sites. The urban sites exhibited a consistent lower level of thermal discomfort during daytime. No discernible effect of urban form attributes in terms of the sky-view factor were observed on ?T_u-r or on the relative difference of the adjusted predicted percentage of dissatisfied (PPD*).     

Thermal bioclimatic conditions and patterns of behaviour in an urban park in Göteborg,Sweden

People in urban areas frequently use parks for recreation and outdoor activities. Owing to the complexity of the outdoor environment, there have only been a few attempts to understand the effect of the thermal environment on people's use of outdoor spaces. This paper therefore seeks to determine the relationship between the thermal environment, park use and behavioural patterns in an urban area of Sweden. The methods used include structured interviews, unobtrusive observations of the naturally occurring behaviour and simultaneous measurements of thermal comfort variables, i.e., air temperature, air humidity, wind speed and global radiation. The thermal environment is investigated through the mean radiant temperature (T_mrt) and the predicted mean vote (PMV) index. The outcome is compared to the subjective behaviour and thermal sensation of the interviewees. It is found that the thermal environment, access and design are important factors in the use of the park. In order to continue to use the park when the thermal conditions become too cold or too hot for comfort, people improve their comfort conditions by modifying their clothing and by choosing the most supportive thermal opportunities available within the place. The study also shows that psychological aspects such as time of exposure, expectations, experience and perceived control may influence the subjective assessment. Comparison between the thermal sensation of the interviewees and the thermal sensation assessed by the PMV index indicates that steady-state models such as the PMV index may not be appropriate for the assessment of short-term outdoor thermal comfort, mainly because they are unable to analyse transient exposure.     

Evaluating the behaviour of different thermal indices by investigating various outdoor urban environments in the hot dry city of Damascus, Syria

Consideration of urban microclimate and thermal comfort is an absolute neccessity in urban development, and a set of guidelines for every type of climate must be elaborated. However, to develop guidelines, thermal comfort ranges need to be defined. The aim of this study was to evaluate the behaviour of different thermal indices by investigating different thermal environments in Damascus during summer and winter. A second aim was to define the lower and upper limits of the thermal comfort range for some of these indices. The study was based on comprehensive micrometeorological measurements combined with questionnaires. It was found that the thermal conditions of different outdoor environments vary considerably. In general, Old Damascus, with its deep canyons, is more comfortable in summer than modern Damascus where there is a lack of shade. Conversely, residential areas and parks in modern Damascus are more comfortable in winter due to more solar access. The neutral temperatures of both the physiologically equivalent temperature (PET) and the outdoor standard effective temperature (OUT_SET*) were found to be lower in summer than in winter. At 80 % acceptability, the study defined the lower comfort limit in winter to 21.0 °C and the upper limit in summer to 31.3 °C for PET. For OUT_SET*, the corresponding lower and upper limits were 27.6 °C and 31.3 °C respectively. OUT_SET* showed a better correlation with the thermal sensation votes than PET. The study also highlighted the influence of culture and traditions on people’s clothing as well as the influence of air conditioning on physical adaptation.     

The effects of solar radiation on thermal comfort

The aim of this study was to investigate the relationship between simulated solar radiation and thermal comfort. Three studies investigated the effects of (1) the intensity of direct simulated solar radiation, (2) spectral content of simulated solar radiation and (3) glazing type on human thermal sensation responses. Eight male subjects were exposed in each of the three studies. In Study 1, subjects were exposed to four levels of simulated solar radiation: 0, 200, 400 and 600 Wm⁻². In Study 2, subjects were exposed to simulated solar radiation with four different spectral contents, each with a total intensity of 400 Wm⁻² on the subject. In Study 3, subjects were exposed through glass to radiation caused by 1,000 Wm⁻² of simulated solar radiation on the exterior surface of four different glazing types. The environment was otherwise thermally neutral where there was no direct radiation, predicted mean vote (PMV)=0±0.5, [International Standards Organisation (ISO) standard 7730]. Ratings of thermal sensation, comfort, stickiness and preference and measures of mean skin temperature (t_sk) were taken. Increase in the total intensity of simulated solar radiation rather than the specific wavelength of the radiation is the critical factor affecting thermal comfort. Thermal sensation votes showed that there was a sensation scale increase of 1 scale unit for each increase of direct radiation of around 200 Wm⁻². The specific spectral content of the radiation has no direct effect on thermal sensation. The results contribute to models for determining the effects of solar radiation on thermal comfort in vehicles, buildings and outdoors.     

Prediction of air temperature for thermal comfort of people in outdoor environments

Current thermal comfort indices do not take into account the effects of wind and body movement on the thermal resistance and vapor resistance of clothing. This may cause public health problem, e.g. cold-related mortality. Based on the energy balance equation and heat exchanges between a clothed body and the outdoor environment, a mathematical model was developed to determine the air temperature at which an average adult, wearing a specific outdoor clothing and engaging in a given activity, attains thermal comfort under outdoor environment condition. The results indicated low clothing insulation, less physical activity and high wind speed lead to high air temperature prediction for thermal comfort. More accurate air temperature prediction is able to prevent wearers from hypothermia under cold conditions.     

Weather impacts on leisure activities in Halifax,Nova Scotia

The aim of this study was to investigate the impact of daily atmospheric weather conditions on daily leisure activity engagement, with a focus on physically active leisure. The methods capitalize on time diary data that were collected in Halifax, Nova Scotia to calculate objective measures of leisure activity engagement. Daily meteorological data from Environment Canada and daily sunrise and sunset times from the National Research Council of Canada are used to develop objective measures of the natural atmospheric environment. The time diary data were merged with the meteorological data in order to quantify the statistical association between daily weather conditions and the type, participation rate, frequency, and duration of leisure activity engagement. The results indicate that inclement and uncomfortable weather conditions, especially relating to thermal comfort and mechanical comfort, pose barriers to physically active leisure engagement, while promoting sedentary and home-based leisure activities. Overall, daily weather conditions exhibit modest, but significant, effects on leisure activity engagement; the strongest associations being for outdoor active sports and outdoor active leisure time budgets. In conclusion, weather conditions influence the type, participation rate, frequency, and duration of leisure activity engagement, which is an important consideration for health-promotion programming.     

Effects of housing differences upon activity budgets in captive sifakas (Propithecus verreauxi)

Activity budgets of captive sifakas (Propithecus verreauxi coquereli and Propithecus verreauxi verreauxi) were assessed from 500 hours of observational data obtained at the Duke University Primate Center (Durham, NC). Data were examined for behavioral differences according to gender, availability of intergroup contact, subspecies, indoor/outdoor housing, and enclosure size. Results showed few differences between the activity budgets of males and females. Several differences found in conjunction with availability of intergroup contact appeared to relate more to subspecific, than to contact, differences. Sifakas housed outdoors were more active, spending less time resting and more time in locomotion, feeding, and playing than sifakas housed indoors. The findings of this study implicate outdoor housing as a primary factor in stimulating activity in these rare prosimian primates.     

Young But Not Old Adult African Striped Mice Reduce Their Activity in the Dry Season When Food Availability is Low

An individual′s survival and fitness depend on its ability to effectively allocate its time between competing behaviors. Sex, social tactic, season and food availability are important factors influencing activity budgets. However, few field studies have tested their influences. The African striped mouse (Rhabdomys pumilio) lives in highly seasonal habitats in southern Africa, and individuals can adopt different social tactics. We investigated seasonal changes in activity budgets of different tactics and predicted that individuals will reduce their activity in the non‐breeding season to save energy when food availability is low and that young non‐breeding adults (‘philopatrics’) invest mainly in activities related to gaining body mass to increase survival probability. We predicted old adults (‘breeders’), which bred during the previous breeding season, to invest mainly in maintenance of their social status. We conducted 90 focal observations during the non‐breeding season and 73 during the breeding season. Activity budgets of striped mice were season and tactic specific, with philopatrics, but not breeders, reducing activity when food availability was low, possibly to decrease energy expenditure. Philopatrics of both sexes foraged and basked more in the breeding season than during the non‐breeding season. Male philopatrics gained body mass and female philopatrics maintained their body mass in both seasons. Sex‐specific differences occurred during the breeding season, when female breeders foraged more than male breeders, while male breeders chased other individuals more than female breeders. These findings indicate that individuals adopting different social tactics display distinct behaviors to fulfill tactic‐specific energetic needs .     

Outdoor thermal comfort study in a sub-tropical climate: a longitudinal study based in Hong Kong

This paper presents the findings of an outdoor thermal comfort study conducted in Hong Kong using longitudinal experiments—an alternative approach to conventional transverse surveys. In a longitudinal experiment, the thermal sensations of a relatively small number of subjects over different environmental conditions are followed and evaluated. This allows an exploration of the effects of changing climatic conditions on thermal sensation, and thus can provide information that is not possible to acquire through the conventional transverse survey. The paper addresses the effects of changing wind and solar radiation conditions on thermal sensation. It examines the use of predicted mean vote (PMV) in the outdoor context and illustrates the use of an alternative thermal index—physiological equivalent temperature (PET). The paper supports the conventional assumption that thermal neutrality corresponds to thermal comfort. Finally, predictive formulas for estimating outdoor thermal sensation are presented as functions of air temperature, wind speed, solar radiation intensity and absolute humidity. According to the formulas, for a person in light clothing sitting under shade on a typical summer day in Hong Kong where the air temperature is about 28°C and relative humidity about 80%, a wind speed of about 1.6 m/s is needed to achieve neutral thermal sensation.     

Effects of antithymocyte serum on lymph node cells participating in the graft-vs-host reaction.

BALB/c mice were injected with 0.1 ml of antithymocyte serum (ATS) and tested at various times thereafter for graft-versus-host (GVH) reactivity of lymph node cells (LNC) and spleen cells (SC). Splenomegaly produced by LNC or SC injected alone was used as a measure of precursor T cell function and the capacity of such cells to produce synergy when combined with normal thymocytes was used to evaluate amplifier T cell activity. In lymph node, both precursor and amplifier function were strikingly depressed 2 days after ATS administration; by day 11, precursor function showed slight recovery while amplifier activity had recovered to supranormal levels. In spleen, precursor activity recovered two fold between 2 and 11 days after ATS inoculation but no amplifier activity could be detected at day 11. Since donors had not been deliberately stimulated with alloantigens prior to testing of LN and SC GVH acivity, these studies demonstrate (a) that precursors and amplifiers are disinct T cell populations that are committed to express unique functional activities before antigen exposure and (b) that, following depletion with ATS, these two populations recover independently at different rates in separate lymphoid compartments.     

Thermal sensitivity of cold climate lizards and the importance of distributional ranges

One of the fundamental goals in macroecology is to understand the relationship among species’ geographic ranges, ecophysiology, and climate; however, the mechanisms underlying the distributional geographic patterns observed remain unknown for most organisms. In the case of ectotherms this is particularly important because the knowledge of these interactions may provide a robust framework for predicting the potential consequences of climate change in these organisms. Here we studied the relationship of thermal sensitivity and thermal tolerance in Patagonian lizards and their geographic ranges, proposing that species with wider distributions have broader plasticity and thermal tolerance. We predicted that lizard thermal physiology is related to the thermal characteristics of the environment. We also explored the presence of trade-offs of some thermal traits and evaluated the potential effects of a predicted scenario of climate change for these species. We examined sixteen species of Liolaemini lizards from Patagonia representing species with different geographic range sizes. We obtained thermal tolerance data and performance curves for each species in laboratory trials. We found evidence supporting the idea that higher physiological plasticity allows species to achieve broader distribution ranges compared to species with restricted distributions. We also found a trade-off between broad levels of plasticity and higher optimum temperatures of performance. Finally, results from contrasting performance curves against the highest environmental temperatures that lizards may face in a future scenario (year 2080) suggest that the activity of species occurring at high latitudes may be unaffected by predicted climatic changes.     

Predicting urban outdoor thermal comfort by the Universal Thermal Climate Index UTCI—a case study in Southern Brazil

Recognising that modifications to the physical attributes of urban space are able to promote improved thermal outdoor conditions and thus positively influence the use of open spaces, a survey to define optimal thermal comfort ranges for passers-by in pedestrian streets was conducted in Curitiba, Brazil. We applied general additive models to study the impact of temperature, humidity, and wind, as well as long-wave and short-wave radiant heat fluxes as summarised by the recently developed Universal Thermal Climate Index (UTCI) on the choice of clothing insulation by fitting LOESS smoothers to observations from 944 males and 710 females aged from 13 to 91 years. We further analysed votes of thermal sensation compared to predictions of UTCI. The results showed that females chose less insulating clothing in warm conditions compared to males and that observed values of clothing insulation depended on temperature, but also on season and potentially on solar radiation. The overall pattern of clothing choice was well reflected by UTCI, which also provided for good predictions of thermal sensation votes depending on the meteorological conditions. Analysing subgroups indicated that the goodness-of-fit of the UTCI was independent of gender and age, and with only limited influence of season and body composition as assessed by body mass index. This suggests that UTCI can serve as a suitable planning tool for urban thermal comfort in sub-tropical regions.     

Thermal physiology explains the elevational range for a lizard,Eutropis longicaudata,in Taiwan

1The decrease of temperatures along an elevation gradient imposes physiological constraints on reptiles that ultimately determine their distribution ranges. Forest patterns are likely to interact with this process, but very few studies have examined their contribution in determining distribution limits.2We examined the role played by thermal physiology and forest cover in determining the elevational ranges of a lizard, Eutropis longicaudata. We integrated this species’ thermal traits in simulating its maximum activity time under different conditions of forest cover and elevation using a NicheMapR model. In addition, we evaluated the influence of winter temperatures on the range limit by examining the simulated soil temperatures at the occurrence sites.3Laboratory experiments showed that E. longicaudata has a high preferred body temperature and low cold tolerance. The model predicts that maximum activity time decreases with elevation and forest cover. Although unforested areas may provide longer active time in all simulated elevations, mountain areas in Taiwan are heavily forested and are predicted to allow only a very short period of activity above 1000 m elevation.4All sightings were indeed located in areas below 1000 m elevation, in which the predicted average soil temperature is above 10 °C in January in cold years.5Our results show that reptile physiological response does respond strongly to the change of microclimate induced by forest cover and elevation. Overall, this suggests that forest cover is a major determinant of some reptiles’ elevational range.     

Activity budgets and activity rhythms in red ruffed lemurs (Varecia rubra) on the Masoala Peninsula, Madagascar: seasonality and reproductive energetics

The activity budgets and daily activity rhythms of Varecia rubra were examined over an annual cycle according to season and reproductive stage. Given the relatively high reproductive costs and patchy food resources of this species, I predicted that V. rubra would 1) travel less and feed more during seasonal resource scarcity in an attempt to maintain energy balance, and 2) show sex differences in activity budgets due to differing reproductive investment. Contrary to the first prediction, V. rubra does not increase feeding time during seasonal food scarcity; rather, females feed for a consistent amount of time in every season, whereas males feed most during the resource-rich, hot dry season. The results are consistent with other predictions: V. rubra travels less in the resource-scarce cold rainy season, and there are some pronounced sex differences, with females feeding more and resting less than males in every season and in every reproductive stage except gestation. However, there are also some provocative similarities between the sexes when activity budgets are examined by reproductive stage. During gestation, female and male activity budgets do not differ and appear geared toward energy accumulation: both sexes feed and rest extensively and travel least during this stage. During lactation, activity budgets are geared toward high energy expenditure: both sexes travel most and in equal measure, and rest least, although it remains the case that females feed more and rest less than males. These similarities between female and male activity budgets appear related to cooperative infant care. The high energetic costs of reproduction in V. rubra females may require that they allot more time to feeding year round, and that their overall activity budget be more directly responsive to seasonal climate change, seasonal food distribution, and reproductive schedules.     

Thermal sensation and thermophysiological responses to metabolic step-changes

This study investigated the effect on thermal perception and thermophysiological variables of controlled metabolic excursions of various intensities and durations. Twenty-four subjects were alternately seated on a chair or exercised by walking on a treadmill at a temperature predicted to be neutral at sedentary activity. In a second experimental series, subjects alternated between rest and exercise as well as between exercise at different intensities at two temperature levels. Measurements comprised skin and oesophageal temperatures, heart rate and subjective responses. Thermal sensation started to rise or decline immediately (within 1 min) after a change of activity, which means that even moderate activity changes of short duration affect thermal perceptions of humans. After approximately 15–20 min under constant activity, subjective thermal responses approximated the steady-state response. The sensitivity of thermal sensation to changes in core temperature was higher for activity down-steps than for up-steps. A model was proposed that estimates transient thermal sensation after metabolic step-changes. Based on predictions by the model, weighting factors were suggested to estimate a representative average metabolic rate with varying activity levels, e.g. for the prediction of thermal sensation by steady-state comfort models. The activity during the most recent 5 min should be weighted 65%, during the prior 10–5 min 25% and during the prior 20–10 min 10%.     

An ecophysiological perspective on likely giant panda habitat responses to climate change

Threatened and endangered species are more vulnerable to climate change due to small population and specific geographical distribution. Therefore, identifying and incorporating the biological processes underlying a species’ adaptation to its environment are important for determining whether they can persist in situ. Correlative models are widely used to predict species’ distribution changes, but generally fail to capture the buffering capacity of organisms. Giant pandas (Ailuropoda melanoleuca) live in topographically complex mountains and are known to avoid heat stress. Although many studies have found that climate change will lead to severe habitat loss and threaten previous conservation efforts, the mechanisms underlying panda's responses to climate change have not been explored. Here, we present a case study in Daxiangling Mountains, one of the six Mountain Systems that giant panda distributes. We used a mechanistic model, Niche Mapper, to explore what are likely panda habitat response to climate change taking physiological, behavioral and ecological responses into account, through which we map panda's climatic suitable activity area (SAA) for the first time. We combined SAA with bamboo forest distribution to yield highly suitable habitat (HSH) and seasonal suitable habitat (SSH), and their temporal dynamics under climate change were predicted. In general, SAA in the hottest month (July) would reduce 11.7%–52.2% by 2070, which is more moderate than predicted bamboo habitat loss (45.6%–86.9%). Limited by the availability of bamboo and forest, panda's suitable habitat loss increases, and only 15.5%–68.8% of current HSH would remain in 2070. Our method of mechanistic modeling can help to distinguish whether habitat loss is caused by thermal environmental deterioration or food loss under climate change. Furthermore, mechanistic models can produce robust predictions by incorporating ecophysiological feedbacks and minimizing extrapolation into novel environments. We suggest that a mechanistic approach should be incorporated into distribution predictions and conservation planning.