The roles of phylogeny and body size in avian heat stress physiology, and how they interact to set the upper limits to heat dissipation capacity, are largely unexplored. Determining thermal end points and maximum capacity for evaporative heat dissipation in species from diverse ecological guilds and evolutionary clades is vital for understanding species-specific vulnerability to future climatic scenarios. I measured evaporative water loss (EWL), resting metabolic rate (RMR) and body temperature (Tb) in three arid-zone passerines and three columbids of varying mass, namely the scaly-feathered weaver (Sporopipes squamifrons, ~11 g, SFW), sociable weaver (Philetairus socius, ~26 g, SW), white-browed sparrow weaver (Plocepasser mahali, ~40 g, WBW), Namaqua dove (Oena capensis, ~37 g, ND), laughing dove (Spilopelia senegalensis, ~89 g, LD) and Cape turtle dove (Streptopelia capicola, ~148 g, CTD) at maximum air temperatures (Ta) of 48–60°C. I found that evaporative water loss increased approximately linearly in all six species above a Ta of ~ 40 °C, which resulted in SFW, SW, WBW, ND, LD and CTD dissipating a maximum of 140, 220, 190, 498, 218 and 231 % of metabolic heat loads at the highest Tas respectively. All six species used facultative hyperthermia at high Tas and were able to regulate Tb up to and just beyond Tb = 45 °C. At the highest Tas experienced, passerines exhibited uncontrolled increases in Tb above 45 °C, resulting in 57, 100 and 100 % of SFW, SW and WBW respectively, reaching thermal limits at Ta = 48, 52 and 54 °C. Very few doves exhibited uncontrolled hyperthermia or reached thermal limits at their highest respective test Tas (Ta = 56, 68 and 60 °C in CTD, LD and ND respectively), suggesting that these birds could potentially survive higher Tas, and that lethal Tb was marginally higher than my conservative estimations. A conventional analysis found significant differences between doves and passerines in the slopes of EWL as well as the magnitude of the change in RMR, EWL and Tb between Ta = 35 and 48 °C. However, once phylogeny was controlled for, these differences were shown to be a result of phylogenetic inertia. Both a conventional analysis and a phylogenetic independent contrast (PIC) found a significant effect of body mass on slope of EWL, change in EWL (PIC only) and change in Tb between Ta = 35 and 48 °C. From the results of this study, I argue that by utilizing high ratios of cutaneous EWL to respiratory EWL, doves generate much less metabolic heat at high Tas than passerines. I suggest that larger passerines are better able to tolerate heat than smaller passerines, whereas the opposite is the case in doves. The lack of data from small doves obscured this finding in the conventional and PIC analyses. Further studies on the upper limits to the avian capacity for evaporative cooling and heat tolerance are critical for larger-scale mechanistic modeling of vulnerability to extreme heat events under current and future climate scenarios. / Dissertation (MSc)--University of Pretoria, 2014. / DST/NRF Centre of Excellence at the Percy FitzPatrick Institute (University of Cape Town) / University of New Mexico / Zoology and Entomology / MSc / Unrestricted

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Human activities and unpredictable changes to environmental habitats impose a suite of stress and challenges to animal homeostatic function. Stress responses are often controlled by the release of glucocorticoids to mobilize energy, primarily corticosterone in small mammals, to help regulate homeostatic function such as heterothermy and changes to body condition, the latter of which serves as a proxy for energy reserves. Adaptive heterothermy is influenced by daily and seasonal patterns, heat produced from daily activity, and has been shown to increase in small mammals during times of environmental stress to conserve energy budgets. Body condition also changes in response to environmental perturbations, mobilization of energy by corticosterone, seasonal changes, and activity. My study aimed to disentangle the effects of environment and activity on homeostatic responses by pharmacologically manipulating corticosterone in kangaroo rats. Kangaroo rats are ecosystem engineers, heterothermic, and their activity periods are functions of their thermoregulatory patterns as well as environmental conditions thus making the species a great candidate for this form of study. I conducted two in-situ field experiments to assess for the effects of stress responses on thermoregulation, activity, and body condition. In my first experiment, I used body temperature (Tb) as a proxy for activity time and examined how pharmacologically increased corticosterone influenced kangaroo rat heterothermy responses to the moon phases and ambient temperatures. I also examined their fat, lean mass, and water content at the end of the study. Moon phase was a significant predictor of activity period as animals typically waited longer during the waxing moon phase to become active and cooled down below activity earlier in the night during the waning moon phase. As nights shortened, activity decreased despite environmental conditions becoming warmer. Corticosterone also significantly decreased total activity time and thus steadily increased heterothermy across the length of my experiment. These results indicate activity, not environment, are stronger drivers of heterothermy patterns. Total fat content (energy content) at the end of the study was not affected by corticosterone. The lack of change in fat content was presumably because kangaroo rat body condition was measured once at the end of the study and likely not a reflection of changed body condition over time. In my second experiment, I examined body condition across a longer period by measuring lean mass, fat content, and total body water across a 2-month period in the summer by pharmacologically increasing corticosterone to test the relationship between stress and body condition and to dissociate the two from environmental factors. Body condition indices generally increased across the summer, but corticosterone implantation did not significantly affect body any of the indices. The loss of heterothermic control and decrease in activity time across the summer suggests that animals are likely conserving energy budgets leading to preservation of condition. Further, banner-tailed kangaroo rats generally breed in the spring where body condition falls due to stress induced by increased competition and then increases across the summer as late summer monsoons promote the growth of primary resources utilized by animals thereby preserving body condition. This study suggests that body condition is driven more by life-history traits, activity time, and environment rather than stress responses.

Bumblebees are experiencing declines and range contractions globally that are, in some cases, independent of anthropogenic pesticide- or land-use change, leaving rising global temperatures as the primary driver of such loses. With ambient temperature (Ta) and thermal limitations being a crucial component in these observed declines, I sought to determine the physiological limitations that high Ta imposes on both individuals and colonies of a temperate bumblebee species, Bombus impatiens. Through Chapter 2, I first established the upper thermal tolerance (CTmax) of the species, testing both adults and larvae to determine which of these colony castes are most thermally sensitive to heat. Collective thermoregulation at the colony-level is then important to ensure that the most heat sensitive individuals are protected from changes in optimal nest temperature (Tn). I thus identified the energetic costs associated with colonial thermoregulation and whether large colonies could successfully achieve thermal homeostasis under various Ta. Chronic bouts of heat stress are also of concern as colonies invest time and energy into thermoregulation, especially given that heatwave events are becoming more frequent. In Chapter 3, I examined whether there exists a trade-off between thermoregulation and foraging effort for colonies under chronic heat stress and how various measures of colony success are impacted. Finally, foraging requires individuals to employ flight for the procurement of resources. In Chapter 4, I investigated if the temperate adaptation of an insulative pile layer would hinder flight performance under high Ta by assessing the metabolic rates of adult castes during flight. I found that larvae were more thermally sensitive compared to bumblebee adults, which emphasizes the importance of colonial thermoregulation – a task successful at low Ta. Under heat stress, however, Tn could not be maintained despite elevated energetic investments (Chapter 2). These findings suggest that Ta which exceeds optimal Tn may pose significant challenges to colonies; not only energetically but also to the health of thermally sensitive larvae within. A trade-off between thermoregulation and foraging effort did not emerge for colonies experiencing chronic exposure to high Ta. Instead, only high incidences of thermoregulation were observed which failed to prevent increases in Tn. Furthermore, a greater number of individuals were found to abandon the colony at high Ta, and fewer offspring were produced (Chapter 3). Here, findings suggest that chronic high Ta may pose the greatest risk to the production of thermally sensitive offspring by way of reduced worker population and failed thermoregulation. Finally, the metabolic output during flight at high Ta was not found to be affected by an insulative layer of pile (Chapter 4), indicating that either pile may play a role in limiting other measures of flight performance at high Ta, or that alternate physiological mechanisms may be responsible instead. Together the findings from this thesis broaden the understanding of how a temperate species of bumblebee responds physiologically to high Ta both at the individual and colonial level, providing further evidence on thermal limitations in a changing climate.

Investigating ecological questions at the scale of individual organisms is necessary to understand and predict the biological consequences of environmental conditions. For small organisms this can be challenging because we need tools with the appropriate accuracy and resolution to record and quantify their ecological interactions. Unfortunately, many of our existing tools are only appropriate for medium to large organisms or those that are wide ranging, inhibiting our ability to investigate the ecology of small organisms at fine scales. In Chapter 1, I tested a novel workflow for recording animal movements at very fine spatial and temporal scales. The workflow combined direct observation and the mapping of locations onto high-resolution uncrewed aerial vehicle (UAV) imagery loaded on hand-held digital devices. Observers identified landscape features they recognized in the imagery and estimated positions relative to those features. I found this approach was approximately twice as accurate as consumer-grade GPS devices with a mean and median error of 0.75 m and 0.30 m, respectively. I also found that performance varied across landscape features, with accuracy highest in areas that had more visual landmarks for observers to use as reference points. In addition to sub-meter accuracy, this method was cost-effective and practical, requiring no bulky equipment and allowing observers to easily record locations away from their own location. While this workflow could be used to record locations in a variety of situations, it will be most cost-effective when also using high-resolution environmental data from a UAV. In Chapter 2, I used the workflow described in Chapter 1 to investigate blunt-nosed leopard lizard (Gambelia sila) thermoregulation at fine-scales. Recent research has suggested that the spatial distribution of temperatures is important to consider for ectotherm thermoregulation, but this work has been limited to simple artificial environments. My goal was to investigate this idea in a complex natural system for the first time. I tracked lizard movement and body temperatures at a desert site from May to July 2021. I used machine learning to combine high-resolution environmental data from a UAV with microclimate temperature data to create a model of the spatial distribution of environmental temperatures over time. I found that including information about the spatial distribution of temperatures improved the models of lizard thermoregulatory accuracy and movement rate. Because these response variables are important aspects of ectotherm energetics, this suggests that the spatial distribution of temperatures may be an important, but often overlooked, component of habitat quality. Going forward, identifying better methods to quantify the spatial distribution of temperatures would provide insights into the specific responses of ectotherms to different spatial distributions. In this work I used recent technological advances in UAVs to investigate ecological questions at the scale of a small organism. The methods developed here provide insights into the importance of the spatial distribution of temperatures for a small ectotherm. Further efforts to develop, test and utilize tools for fine-scale ecological research will advance our ability to understand species’ interactions with current conditions and predict their responses to future changes.

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Skin temperature plays an important role in human thermoregulation together with core temperature. Skin temperature varies to a large extent across the body and this is especially pronounced in cold environments. The variations of skin temperature are also involved in the generation of regional thermal perceptions that can lead to behavioural adjustments. Whilst the temporal and inter-individual variations of skin temperature have been well studied using contact sensors, the knowledge of spatial variations has received less attention in the literature. Infrared thermography is a specific imaging technique particularly valuable for the exploration of the topography or pattern of skin temperature across the body. Most research using this technique has only been case studies or experiments focused in one specific body region. However, extensive regional skin temperature data over the whole-body can be proven useful for different types of applications including the sport clothing industry in combination with other body-mapping data. The primary aim of this thesis was to develop an original and standardised method using infrared thermography enabling whole-body skin temperature data to be compared for the assessment of spatial, temporal and inter-individual variations. A specific methodology for infrared data collection and data processing was successfully developed in order to combine data from a variety of participants varying in anthropometrical characteristics. The main outcomes were the production of several skin temperature body maps, either absolute maps to show the magnitude of the temporal or inter-individual effects, and normalised maps (relative to mean skin temperature) allowing for topographical comparisons between protocol stages, populations or interventions. The second aim of the thesis was to extend the understanding of the skin temperature patterns and how these could relate with thermal perceptions. The body-mapping method gave the opportunity to investigate a large amount of conditions, where various internal or external determinants of skin temperature were be involved. This was mainly done in cool to cold environments (5°C to 20°C) where skin temperature is not uniform but is associated with local and overall comfort. Studies were firstly performed in semi-nude conditions (Chapter 3, 4, 5) and then in clothed conditions (Chapter 6 and 7). The semi-nude studies were designed to explore the potential sexdifferences in regional skin temperature responses whilst running (Chapter 3) with a special interest in the role of skinfold thickness, this was further extended with a group of males at rest having a large variety of fat content and thickness (Chapter 4). The influence of exercise type and air temperature on skin temperature patterns was studied with a rowing exercise (Chapter 5). Studies were then performed in clothed conditions (Chapter 5, 6). The influence of real-life conditions on skin temperature patterns and associated perceptual responses was observed during a hiking scenario (Chapter 6). Following these descriptive studies, manipulation of skin temperature patterns was performed using clothing in order to determine the presence of any relevant effect on thermal comfort (Chapter 7). Our results demonstrated that the skin temperature pattern over the whole-body is relatively universal with several features being consistently found regardless of the conditions or the populations. The upper body is usually warmer than the lower body and the body creases (orbital, elbow regions etc.) are also warmer than surrounding regions. A Y-shape of colder temperatures has been highlighted over the anterior torso as well as a T- or Y-shape of warmer temperature over the posterior torso. There are yet some specificities that can be displayed due to active muscles during exercise such as the warmer skin overlying the trapezius and biceps muscles in rowing (Chapter 5), the influence of the backpack construction with up to 3°C warmer skin temperature in the lower back (Chapter 6) or the importance of additional clothing insulation minimizing the anterior Y-shape of colder skin temperatures (Chapter 7). Beyond the thermal patterns, absolute skin temperature differences have been observed between sexes with females displaying 2°C colder skin during semi-nude running (Chapter 3) and 1°C colder skin during clothed walking (Chapter 6)compared to males. The skin temperature difference can also be as large as 6°C colder skin for an obese male compared to a very lean male (40% vs 7% body fat). Despite these differences, there were almost no significant differences in overall and regional thermal sensations and comfort between sexes or between males with varying body fat. Our results focused on body fat revealed that overall fat content and sum of skinfolds was inversely associated with the mean skin temperature response during various protocols (Chapter 4, 6, 7). Local skinfold thickness explained the inter-individual variability of local skin temperature for resting (Chapter 4) and exercising males (Chapter 7) in most body regions. In terms of intra-individual variations, the distribution of skinfold thickness across the anterior torso explained the distribution of skin temperature in this segment solely in conditions with strong regional contrasts (Chapter 3, 4 and 7). When the whole-body skin temperature pattern is considered, our body-mapping approach failed to show relationships between skin temperature distribution across the body and regional skinfold thickness distribution neither at rest nor during exercise. The relative contribution of other internal determinants such as local heat production,local blood flow distribution and local anthropometry should be further investigated to fully elucidate the spatial skin temperature variations depending on the climate, clothing and the body thermal state. Lastly, there was a trend towards improved thermal comfort during rest and exercise in the cold through a manipulation of skin temperature patterns targeting the naturally cold body regions with high insulation, therefore obtaining a more homogeneous skin temperature distribution across the body (Chapter 7). The present work will benefit the sport goods industry. The descriptive results of skin temperature variations will be useful in order to validate multi-segmental model of human thermoregulation. Further work can include pattern predictions for exercise types and conditions not covered by the present thesis. The skin temperature maps will mainly feed the general body-mapping approach for clothing design taking into account several other body mapping data such as sweat mapping and the combination of cold, warm and wetness sensitivity mappings. Lastly, the present results have highlighted the interest for targeted solutions and also the need for more evolutive systems in the field of cold weather apparel.

Exercise or activity in high ambient temperatures offers a particular challenge to the thermoregulatory system. It is likely that mechanisms such as sweat evaporation alone are not sufficient for maintaining body temperature within a safe limit (~36.5-38.5˚C) and below 40˚C, which may result in impaired physiological function and performance. Exogenous cooling may be of benefit prior to, during and after events that place increased thermal strain due to increased metabolic heat production and elevated environmental temperatures upon the thermoregulatory system. Conversely, in situations where it is not possible to maintain body temperature via either continued physical activity or elevated ambient temperatures, exogenous heating may be required in order to allow optimal physiological performance. Few studies have directly aligned cooling devices with data detailing effective target regions for cooling to allow a pre-cooling garment to be of minimal weight but maximal cooling efficiency. Conversely, no study has considered the effect of muscle temperature maintenance during rest periods on subsequent power-based activities. The aim of this thesis was to determine ways in which body temperature manipulation is capable of improving exercise performance in both power and endurance-based events. It was hypothesised that the manipulation of body temperature will result in subsequent changes in body temperature that would improve performance. Specifically, the use of pre-cooling would result in a reduction of body temperature and improve endurance exercise performance. Conversely, maintaining Tm following warm up completion would have a beneficial effect on sprint and power related performance. Study one set out to determine differences in regional body heat loss in 12 individual anatomical zones using a water perfused suit. Data obtained from this initial study allowed for the specific targeting of regions that were identified as having high rates of heat loss in subsequent studies that focused on pre-cooling and performance. The anatomical regions identified as having high potential affinity for heat exchange with the surrounding environment and cooling devices were the hands, forearms, upper and lower back and torso. Subsequent studies demonstrated that cooling of these areas was capable of lowering thermal sensation and improving thermal comfort prior to and during exercise in moderate environmental conditions (24˚C, 50% RH). In these moderate conditions, there was no statistically significant improvement in treadmill based self-paced 5000m running performance. However, in hot conditions (35˚C 50% RH), the use of a cooling vest and sleeves did yield a significant improvement in cycling time trial performance, which equated to 4.8%. This leads to the suggestion that there may be a threshold ambient temperature, above which pre-cooling becomes an important tool in maximizing performance potential. A parallel area of investigation, on the other side of the temperature spectrum, was the effect of muscle temperature manipulation on power-based exercise performance. The relationship between increased muscle temperature and power output is well established, however little is known about the effect of enforced rest or recovery between two bouts of exercise. Therefore, two studies were conducted to establish what affect a delay between warm up completion and exercise has on muscle temperature and subsequent sprint cycling performance. It was shown that with 30-minutes of rest between exercise bouts wearing tracksuit trousers, muscle temperature declined significantly (~1-1.5˚C). This decline was attenuated with the use of external passive electrical heating during the recovery compared to recovery completed in tracksuit trousers alone. The attenuated decline in muscle temperature following the use of the heated trousers resulted in an improvement in sprint cycling performance (~9%), with the use of insulated trousers having no effect on any variables measured, all relative to wearing tracksuit trousers in the rest period. In a follow-up study, the effect of implementing the heated trousers during the warm up and in addition to the rest period had on muscle temperature increase and sprint performance. A secondary area of investigation in this study was to determine the linearity of muscle temperature decline following warm up cessation. This study demonstrated that there was no additional benefit of combining passive heating with an active warm up on either muscle temperature elevations or subsequent sprint performance compared to the active warm up alone. It was shown that when the no heating was used at any stage, muscle temperature declined exponentially. However, when the heated trousers were used during recovery and/or during warm up, muscle temperature levelled off at a higher value towards the end of the recovery period. This study was also able to show significant improvements in absolute, relative and mean power output following the use of the heated trousers in the warm up and recovery, or the recovery alone. This thesis has identified ways in which body temperature may be manipulated in order to benefit both sprint and endurance exercise performance, using both pre-cooling and active heating. A novel concept for minimizing muscle temperature decline during periods of inactivity between different rounds of competition was shown to maximize sprint performance yielding significant improvements in peak and mean power outputs.

This thesis investigated whether highly concentrated sodium solutions ([HS] 126-164 mmol.L-1 NaCl) could provide viable strategies before during and after exercise in the heat to improve cardiovascular and thermoregulatory functioning and exercise performance. To do this it also examined the gustatory responses to HS drinks before, during and after exercise. All studies compared HS with a low sodium control ([LS] 10-27 mmol.L-1). Chapter 4 found that during 3 h recovery from dehydration, ingestion of 120 % body mass losses of HS restored fluid balance to a greater extent (121 vs. 84 %) than LS. Chapter 7 was the first to investigate the effects of ingesting HS during exercise in the heat and in an untrained population. HS attenuated the decline in stroke volume [SVDrift] and increase in heart rate [HRDrift], but did not affect rectal temperature [TRec], cardiac output, or oxygen uptake during the second of two consecutive 45 min bouts at 55% . In Chapters 8 and 9 untrained participants ingested either HS or LS during 30-45 min pre-exercise rest. HS reduced HRDrift and SVDrift but did not affect TRec during 45-60 min exercise at 10% of the difference between and gas exchange threshold [∆]. HS also increased both time to exhaustion and exercise toleranceduring subsequent exercise bouts at 60-70% ∆. Chapters 5 and 6 found that taste perceptions act as physiological regulators, in this case, one reflecting the priority to restore hyperosmolality over hypovolemia. Exercise-induced dehydration increased the palatability of water, and decreased the palatability HS, when measured before, immediately after and during 3 h recovery. The changes were highly correlated with physiological indicators of fluid balance. The ingestion of highly concentrated sodium solutions can be both an efficient and acceptable means to improve hydration, reduce cardiovascular stress, and improve exercise performance in the heat. Whilst highly effective, caution should apply since the unpleasant taste evoked by these solutions persists for at least three hours post exercise.