Are humans evolved specialists for running in the heat? Man vs. horse races provide empirical insights.

NEW FINDINGS: What is the central question of this study? Do available comparative data provide empirical evidence that humans are adapted to endurance running at high ambient temperatures? What is the main finding and its importance? Comparing the results of races that pit man against horse, we find that ambient temperature on race day has less deleterious effects on running speed in humans than it does on their quadrupedal adversary. This is evidence that humans are adapted for endurance running at high ambient temperatures. We debate whether this supports the hypothesis that early man was evolutionarily adapted for persistence hunting. ABSTRACT: Many mammals run faster and for longer than humans and have superior cardiovascular physiologies. Yet humans are considered by some scholars to be excellent endurance runners at high ambient temperatures, and in our past to have been persistence hunters capable of running down fleeter quarry over extended periods during the heat of the day. This suggests that human endurance running is less affected by high ambient temperatures than is that of other cursorial ungulates. However, there are no investigations of this hypothesis. We took advantage of longitudinal race results available for three annual events that pit human athletes directly against a hyper-adapted ungulate racer, the thoroughbred horse. Regressing running speed against ambient temperature shows race speed deteriorating with hotter temperatures more slowly in humans than in horses. This is the first direct evidence that human running is less inhibited by high ambient temperatures than that of another endurance species, supporting the argument that we are indeed adapted for high temperature endurance running. Nonetheless, it is far from clear that this capacity is explained by an endurance hunting past because in absolute terms humans are slower than horses and indeed many other ungulate species. While some human populations have persistence hunted (and on occasion still do), the success of this unlikely foraging strategy may be best explained by the application of another adaption - high cognitive capacity. With dedication, experience and discipline, capitalising on their small endurance advantage in high temperatures, humans have a chance of running a more athletic prey to exhaustion.

Inter-stride variability triggers gait transitions in mammals and birds.

Speed-related gait transitions occur in many animals, but it remains unclear what factors trigger gait changes. While the most widely accepted function of gait transitions is that they reduce locomotor costs, there is no obvious metabolic trigger signalling animals when to switch gaits. An alternative approach suggests that gait transitions serve to reduce locomotor instability. While there is evidence supporting this in humans, similar research has not been conducted in other species. This study explores energetics and stride variability during the walk-run transition in mammals and birds. Across nine species, energy savings do not predict the occurrence of a gait transition. Instead, our findings suggest that animals trigger gait transitions to maintain high locomotor rhythmicity and reduce unstable states. Metabolic efficiency is an important benefit of gait transitions, but the reduction in dynamic instability may be the proximate trigger determining when those transitions occur.

Comparative locomotor costs of domestic dogs reveal energetic economy of wolf-like breeds.

The broad diversity in morphology and geographic distribution of the 35 free-ranging members of the family Canidae is only rivaled by that of the domesticated dog, Canis lupus familiaris. Considered to be among nature's most elite endurance athletes, both domestic and wild canids provide a unique opportunity to examine the variability in mammalian aerobic exercise performance and energy expenditure. To determine the potential effects of domestication and selective breeding on locomotor gait and economy in canids, we measured the kinematics and mass-specific metabolism of three large (>20 kg) dog breed groups (northern breeds, retrievers and hounds) of varying morphological and genomic relatedness to their shared progenitor, the gray wolf. By measuring all individuals moving in preferred steady-state gaits along a level transect and on a treadmill, we found distinct biomechanical, kinematic and energetic patterns for each breed group. While all groups exhibited reduced total cost of transport (COT) at faster speeds, the total COT and net COT during trotting and galloping were significantly lower for northern breed dogs (3.0 and 2.1 J kg-1 m-1, respectively) relative to hound (4.2 and 3.4 J kg-1 m-1, respectively) and retriever dogs (3.8 and 3.0 J kg-1 m-1, respectively) of comparable mass. Similarly, northern breeds expended less energy per stride (3.5 J kg-1 stride-1) than hounds or retrievers (5.0 and 4.0 J kg-1 stride-1, respectively). These results suggest that, in addition to their close genetic and morphological ties to gray wolves, northern breed dogs have retained highly cursorial kinematic and physiological traits that promote economical movement across the landscape.

The golden age of bio-logging: how animal-borne sensors are advancing the frontiers of ecology.

Great leaps forward in scientific understanding are often spurred by innovations in technology. The explosion of miniature sensors that are driving the boom in consumer electronics, such as smart phones, gaming platforms, and wearable fitness devices, are now becoming available to ecologists for remotely monitoring the activities of wild animals. While half a century ago researchers were attaching balloons to the backs of seals to measure their movement, today ecologists have access to an arsenal of sensors that can continuously measure most aspects of an animal's state (e.g., location, behavior, caloric expenditure, interactions with other animals) and external environment (e.g., temperature, salinity, depth). This technology is advancing our ability to study animal ecology by allowing researchers to (1) answer questions about the physiology, behavior, and ecology of wild animals in situ that would have previously been limited to tests on model organisms in highly controlled settings, (2) study cryptic or wide-ranging animals that have previously evaded investigation, and (3) develop and test entirely new theories. Here we explore how ecologists are using these tools to answer new questions about the physiological performance, energetics, foraging, migration, habitat selection, and sociality of wild animals, as well as collect data on the environments in which they live.

Dogs as Pets and Pests: Global Patterns of Canine Abundance, Activity, and Health.

Dogs (Canis familiaris) were the first domesticated species and, at an estimated population of 1 billion individuals, are globally ubiquitous today. Describing the tremendous morphometric diversity and evolutionary origins of dogs is a scientific endeavor that predates Darwin, yet our interdisciplinary understanding of the species is just beginning. Here, I present global trends in dog abundance, activity, and health. While the human-dog relationship has for millennia been close, it is also complicated. As pets, companion dogs are often treated as family members and constitute the largest sector of the ever-growing >$200 billion USD global pet care industry. As pests, free-roaming dogs are an emerging threat to native species via both predation and nonconsumptive effects (e.g., disturbance, competition for resources, and hybridization). Furthermore, I briefly discuss mounting evidence of dogs as not only infectious disease reservoirs but also as bridges for the transmission of pathogens between wild animals and humans in zoonotic spillover events, triggering intensive dog population management strategies such as culling. Dog mobility across the urban-wildland interface is an important driver for this and other adverse effects of canines on wildlife populations and is an active topic of disease ecologists and conservation biologists. Other canine scientists, including veterinary clinicians and physiologists, study more mechanistic aspects of dog mobility: the comparative kinetics, kinematics, and energetics of dog locomotor health. I outline the prevalent methodological approaches and breed-specific findings within dog activity and health research, then conclude by recognizing promising technologies that are bridging disciplinary gaps in canine science.

Surviving in steep terrain: a lab-to-field assessment of locomotor costs for wild mountain lions (Puma concolor).

BACKGROUND: Under current scenarios of climate change and habitat loss, many wild animals, especially large predators, are moving into novel energetically challenging environments. Consequently, changes in terrain associated with such moves may heighten energetic costs and effect the decline of populations in new localities. METHODS: To examine locomotor costs of a large carnivorous mammal moving in mountainous habitats, the oxygen consumption of captive pumas (Puma concolor) was measured during treadmill locomotion on level and incline (6.8°) surfaces. These data were used to predict energetic costs of locomotor behaviours of free-ranging pumas equipped with GPS/accelerometer collars in California's Santa Cruz Mountains. RESULTS: Incline walking resulted in a 42.0% ± 7.2 SEM increase in the costs of transport compared to level performance. Pumas negotiated steep terrain by traversing across hillsides (mean hill incline 17.2° ± 0.3 SEM; mean path incline 7.3° ± 0.1 SEM). Pumas also walked more slowly up steeper paths, thereby minimizing the energetic impact of vertical terrains. Estimated daily energy expenditure (DEE) based on GPS-derived speeds of free-ranging pumas was 18.3 MJ day- 1 ± 0.2 SEM. Calculations show that a 20 degree increase in mean steepness of the terrain would increase puma DEE by less than 1% as they only spend a small proportion (10%) of their day travelling. They also avoided elevated costs by utilizing slower speeds and shallower path angles. CONCLUSIONS: While many factors influence survival in novel habitats, we illustrate the importance of behaviours which reduce locomotor costs when traversing new, energetically challenging environments, and demonstrate that these behaviours are utilised by pumas in the wild.

Tourist photographs as a scalable framework for wildlife monitoring in protected areas.

Protected areas are critical to conservation efforts in the face of rapid biodiversity declines [1]. Yet the resources for conservation are often limited and shared amongst many competing priorities [2]. As a consequence, even basic monitoring surveys are absent within most protected areas [3]. Although a range of wildlife monitoring methods exist, considerable focused survey effort is often required to yield accurate and precise estimates [4]. This makes monitoring difficult to sustain or replicate, limiting access to the data required for evidence-based conservation decisions. Citizen-scientists have been proposed as an important complement to the finite resources available for basic monitoring within protected areas [5]; however, the full potential of this approach has yet to be realised. Wildlife tourists and guides are especially focussed on encountering and photographing fauna and flora, yet the data collected in these efforts is rarely harnessed for conservation monitoring within protected areas. A detailed understanding of photographic tourism's potential role in wildlife monitoring has been lacking, but is essential for the development of new tools to harness the data being collected through tourism. Here, we demonstrate that tourist-contributed data can aid wildlife monitoring in protected areas by providing population estimates of large carnivores comparable to those from traditional survey methods. Our approach could capitalize upon the immense number of wildlife photographs being taken daily as part of the global > 30-billion USD, wildlife-based tourism industry.

Energetics and evasion dynamics of large predators and prey: pumas vs. hounds.

Quantification of fine-scale movement, performance, and energetics of hunting by large carnivores is critical for understanding the physiological underpinnings of trophic interactions. This is particularly challenging for wide-ranging terrestrial canid and felid predators, which can each affect ecosystem structure through distinct hunting modes. To compare free-ranging pursuit and escape performance from group-hunting and solitary predators in unprecedented detail, we calibrated and deployed accelerometer-GPS collars during predator-prey chase sequences using packs of hound dogs (Canis lupus familiaris, 26 kg, n = 4-5 per chase) pursuing simultaneously instrumented solitary pumas (Puma concolor, 60 kg, n = 2). We then reconstructed chase paths, speed and turning angle profiles, and energy demands for hounds and pumas to examine performance and physiological constraints associated with cursorial and cryptic hunting modes, respectively. Interaction dynamics revealed how pumas successfully utilized terrain (e.g., fleeing up steep, wooded hillsides) as well as evasive maneuvers (e.g., jumping into trees, running in figure-8 patterns) to increase their escape distance from the overall faster hounds (avg. 2.3× faster). These adaptive strategies were essential to evasion in light of the mean 1.6× higher mass-specific energetic costs of the chase for pumas compared to hounds (mean: 0.76 vs. 1.29 kJ kg-1 min-1, respectively). On an instantaneous basis, escapes were more costly for pumas, requiring exercise at ≥90% of predicted [Formula: see text] and consuming as much energy per minute as approximately 5 min of active hunting. Our results demonstrate the marked investment of energy for evasion by a large, solitary carnivore and the advantage of dynamic maneuvers to postpone being overtaken by group-hunting canids.

Movement, resting, and attack behaviors of wild pumas are revealed by tri-axial accelerometer measurements.

BACKGROUND: Accelerometers are useful tools for biologists seeking to gain a deeper understanding of the daily behavior of cryptic species. We describe how we used GPS and tri-axial accelerometer (sampling at 64 Hz) collars to monitor behaviors of free-ranging pumas (Puma concolor), which are difficult or impossible to observe in the wild. We attached collars to twelve pumas in the Santa Cruz Mountains, CA from 2010-2012. By implementing Random Forest models, we classified behaviors in wild pumas based on training data from observations and measurements of captive puma behavior. RESULTS: We applied these models to accelerometer data collected from wild pumas and identified mobile and non-mobile behaviors in captive animals with an accuracy rate greater than 96%. Accuracy remained above 95% even after downsampling our accelerometer data to 16 Hz. We were further able to predict low-acceleration movement behavior (e.g. walking) and high-acceleration movement behavior (e.g. running) with 93.8% and 92% accuracy, respectively. We had difficulty predicting non-movement behaviors such as feeding and grooming due to the small size of our training dataset. Lastly, we used model-predicted and field-verified predation events to quantify acceleration characteristics of puma attacks on large prey. CONCLUSION: These results demonstrate that accelerometers are useful tools for classifying the behaviors of cryptic medium and large-sized terrestrial mammals in their natural habitats and can help scientists gain deeper insight into their fine-scale behavioral patterns. We also show how accelerometer measurements can provide novel insights on the energetics and predation behavior of wild animals. Lastly we discuss the conservation implications of identifying these behavioral patterns in free-ranging species as natural and anthropogenic landscape features influence animal energy allocation and habitat use.