Seasonal variation in the thermal biology of the rock hyrax (Procavia capensis)
Animals in the Southern African sub-region are faced with unpredictable seasonal rainfall patterns and unpredictably low resource availability due to the influence of the El Nino Southern Oscillation System. This has led to conservative energetic traits in animals that offset the costs of maintaining homeostasis in the unpredictable environments they inhabit. One of these animals is the rock hyrax (Procavia capensis). Past research has looked at the thermoregulation of rock hyrax in the laboratory. Results indicated that rock hyrax had labile body temperatures that reached lethal levels at ambient temperatures above 39°C. Laboratory studies separate endogenous thermoregulation from behavioural thermoregulation and do not reflect the overall thermoregulatory potential of the rock hyrax in maintaining body temperatures. This study looked at the thermoregulation of rock hyrax in their natural environment. Body temperatures were measured in the field using Thermochron iButtons inserted into the intraperitonial cavity of the animals. Behavioural observations were also recorded on subgroups and individually marked animals. Rock hyraxes were exposed to large fluctuations in ambient temperatures and food availability during the course of this study. In winter, ambient temperatures ranged between 5-25°C and in summer between 18-42°C. Our results show that rock hyrax seasonally and daily altered both their physiological and behavioural thermoregulation to control body temperature efficiently. The physiological alterations observed in rock hyrax differed between winter and summer. During winter, when food availability was low, rock hyrax maintained body temperatures at a lowered level relative to summer. Body temperatures fluctuated to a greater extent during winter as a result of reduced body temperatures at night and increased body temperatures due to basking during the diurnal hours. During summer, rock hyrax displayed high body temperatures, which reached hyperthermic levels. This enabled rock hyrax to forage during midday hours since heat loads could be easily dissipated through passive conduction in the cooler rock crevices. Rock hyrax employed different behavioural patterns in winter and summer. It is proposed that rock hyraxes are unable to meet energetic demands on a low quality and patchy food resource under low ambient temperatures. Predation is also a cost to foraging and becomes increasingly important when rock hyraxes have to move large distances away from crevices to find food. The most frequent behaviour recorded in rock hyrax during winter was basking. Basking enabled rock hyrax to maintain body temperatures with very little thermoregulatory cost. Differences were also recorded in behavioural patterns in rock hyrax of different sizes. Juveniles, due to their small body size and high energetic demands foraged more frequently than adult rock hyrax. During summer, rock hyrax spent considerable time in the rock crevices. This was because ambient temperatures exceeded the upper limit of the thermoneutral zone of the rock hyrax throughout most of the day. Rock hyrax therefore escaped the excessive temperatures by utilizing the cooler rock crevices, which remained at temperatures within the rock hyraxes thermoneutral zone. The most frequent behaviour recorded in rock hyrax aboveground was foraging. Since forage was abundant around the rock crevices during summer, predation risk was less of a factor influencing foraging behaviour. Since basking is an essential component of the rock hyraxes thermal biology during winter we examined basking behaviour in more detail. Basking in the morning was not used to increase body temperatures from hypothermic levels as otherwise thought. Instead, it was used to maintain body temperatures at low ambient temperatures by altering posture, orientation to the sun and basking bout lengths. During the early morning, when heating rates were greatest, rock hyrax orientated their bodies exposing the greatest surface area to solar radiation. During midday, reduced basking bout lengths and the reduction of surface areas exposed to the sun reduced the heat loads during the hottest parts of the day. Rock hyrax appeared to utilize the warm rock surfaces during the late afternoon when the sun was setting to maintain body temperatures before entering the crevices for the night. The combination of physiological and behavioural thermoregulation therefore enables rock hyrax to maintain homeostasis with very little energetic costs in an environment that displays variability in both ambient temperatures and resource availability.