Ecology and morphology of the Kalahari tent tortoise, Psammobates oculifer, in a semi-arid environment

Philosophiae Doctor - PhD (Biodiversity and Conservation Biology)

Southern Africa harbours one-third of the world's Testudinid species, many of which inhabit arid or semi-arid areas, but ecological information on these species is scant. I studied the habitat, morphology and ecology of Kalahari tent tortoises over 13 months in semi-arid Savanna at Benfontein farm, Northern Cape Province, South Africa. In order to allow continuous monitoring of individuals, I attached radiotransmitters to males and females, split equally between two habitats, sites E (east) and W (west), with apparent differences in vegetation structure. Results of the study were based on data obtained from 27 telemetered tortoises and 161 individuals encountered opportunistically. Female Kalahari tent tortoises were larger than males and the sex ratio did not differ from 1:1. Based on person-hours to capture tortoises, the population appeared to have a low density, with more time required to capture a juvenile (35 hours) than an adult (10-11 hours). The frequency distribution of body size ranges was indicative of recruitment. Relative age, based on annuli counts, suggested that males were younger than females, perhaps because males as the smaller sex are more predation-prone than females. Linear relationships between annuli counts and shell volume indicated that, after reaching sexual maturity, female body size increased faster in volume than did male body size, possibly because a larger volume may enhance female reproductive success. Body condition differed between sites, sexes and among seasons. The hot and dry summer may account for low summer body condition, whereas vegetation differences and size effects, respectively, may account for the low body condition of tortoises in site W and in males. Site E was sandy with grasses, particularly Schmidtia pappophoroides, being the prevalent growth form. This habitat resembled a Savanna vegetation type Schmidtia pappophoroides – Acacia erioloba described for a neighbouring reserve. Site W was stonier, dominated by shrubs, and was reminiscent of Northern Upper Karoo vegetation (NKu3). Neither site resembled Kimberley Thornveld (SVk4), the designated vegetation type of the area. Differences in substrate and grazing intensity may have contributed to site vegetation differences. Rainfall had an important influence on seasonal vegetation. Short grass abundance correlated with rainfall and annual plants sprouted after spring rain. Refuge use changed according to season and sex. Males selected denser refuges than females did, perhaps because males were smaller and more vulnerable to predation and solar heat. Tortoises selected sparse, short grass as refuges in cool months, probably to maximise basking whilst remaining in protective cover. During hot periods, mammal burrows were preferred to vegetation as refugia. The smaller males spent more time in cover than females, which may be related to predator avoidance or thermoregulation. Females spent more time basking than males, perhaps due to their larger size and to facilitate reproductive processes. Tortoises did not brumate, but through a combination of basking, and orientation relative to the sun in their refuges, managed to attain body temperatures that allowed small bouts of activity. Body temperature for active tortoises was similar among seasons, and was higher for more specialised active behaviours, such as feeding and socialising, than for walking. Increased activity by males in spring could relate to mating behaviour while females were more active in autumn, when they foraged more than males, perhaps due to the high cost of seasonal reproductive requirements. Males displaced further per day than did females, but home range estimates did not differ between sexes. Annual home range estimates varied substantially among individuals: 0.7–306 ha for minimum convex polygons and 0.7–181 ha for 95% fixed kernel estimates. The ability to cover large areas would assist tortoises in finding resources, e.g., food, in an area where resource distribution may be patchy. Differences among seasonal home ranges and movements probably reflect seasonal climatic change; activity areas shrinking when temperatures were extreme. In order to assess the effects of a semi-arid environment on the morphology of P. oculifer, I compared its morphology to that of its ‘cool-adapted’ sister taxon Psammobates geometricus, using live and museum specimens. Both P. oculifer and P. geometricus are sexually dimorphic and differences between the two species could indicate environmental or sexual selection effects, or a combination of the two. The shorter bridge length, which allowed more leg space, and wider front feet in P. oculifer cohorts probably represent traits for manoeuvring in a sandy habitat, while wider heads in P. oculifer possibly relate to interspecific differences in diet. The flatter shell in female P. oculifer, relative to P. geometricus, may represent a trade-off between space for reproductive structures, e.g., eggs, and the need to fit into small refuges, e.g., mammal burrows. Male P. oculifer had wider shells, more space around their hind legs, and wider hind feet than P. geometricus males had, all characteristics which may assist males to fight and mate in a sandy environment.

South Africa