A thesis submitted to the Faculty of Science, University of the Witwatersrand, Johannesburg, in fulfilment of the requirements for the degree of Doctor of Philosophy, 2017 / Under current and previous global climate change, environments are changing and have changed at a rapid rate. Species with the potential to undergo adaptive radiation are likely to survive environmental change. The genus Rhabdomys is widespread in southern Africa, occurring along the east-west rainfall gradient in South Africa. Rhabdomys may have undergone adaptive radiations in the past, which may have resulted in the current suite of species in various habitats of different aridity. Some Rhabdomys species also occur in sympatry in some locations in South Africa. The aim of my study was to investigate adaptive variation in Rhabdomys by studying the behaviour of 5 populations, representing 3 Rhabdomys species, across South Africa. Using selected taxa, my approach was, firstly, to describe variation in two traits, personality and spatial cogntion, well known for showing environmentally-linked (i.e. adaptive) variation. Secondly, I manipulated the development of exploratory and anxiety behaviour to assess the limits of the adaptive variation (i.e. test the nature of the reaction norm of the characters measured). I first established the taxon-level personality of 4 taxa (2 sympatric) in 5 standard behavioural tests. Generally, the semi-desert living R. pumilio was the boldest together, surprisingly, with R. d. dilectus occurring in grasslands of central South Africa, contradicting previously published results. Comparatively, R. bechuanae from central South Africa and R. dilectus from far north-eastern South Africa, also occurring in grasslands were less bold, even though R. bechuanae is sympatric with R. dilectus in central South Africa. My data indicate adaptive variation at the extreme populations and possibly character displacement in the sympatric populations. In the next chapter, I investigated whether early rearing environment shapes exploratory behaviour and anxiety responses of R. pumilio and R. bechuanae. I predicted that using an interspecies cross-fostering protocol would reveal a gene x environment interaction on behaviour, so that fostered offspring would display an intermediate behaviour phenotype compared to their non-fostered siblings. I showed that a novel rearing environment mostly did not influence the adult behaviour of cross-fostered inidividuals. This indicates genetic constraints on exploratory behaviour and anxiety responses. Next, I tested whether physical rearing environment shapes exploratory behaviour and anxiety responses. I reared semidesert R. pumilio, sympatric R. bechuanae and R. dilectus and allopatric R. bechuanae under either no cover or high cover for 2 generations. The taxa were mostly similar and altering the phyical housing condition did not alter behaviour, but there were small differences between the taxa in exploratory behaviour. In the final experimental chapter, I established whether the environment predicts the spatial cognition in semi-desert R. pumilio, sympatric R. bechuanae and R. dilectus and an allopatric population of R. dilectus from far north-eastern South Africa. The populations showed very similar performance in a modified Barnes maze, indicating a possible phylogenetic constraint on spatial cognition. Overall, my study suggests that there is adaptive variation in personality but not spatial cognition. In contrast to previous studies in the genus, alterations to the social and physical environments failed to separate out genetic and environmental effects (i.e. reaction norm) that would potentially provide the mechanisms for adaptive variation within and between species. The similarity in spatial cognition between taxa and similar responses to environmental modification indicate phylogenetic constraints on traits that were predicted to vary geographically. / XL2018

The striped-mouse, Rhabdomys pumilio, is widely distributed throughout southern Africa within a variety of habitats and rainfall regimes. It is found at sea level in the Eastern and Western Cape regions and at altitudes above 2700 m in the Drakensberg mountains. The attraction of R.pumilio to cultivated land and crops has resulted in extensive damage to plants and cultivated crops. A study of the genetic variation between populations of R.pumilio from different regions of southern Africa was undertaken by protein electrophoresis and randomly amplified polymorphic DNA using the polymerase chain reaction (PCR-RAPD). A cytogenetic study was also undertaken. The mean heterozygosity (H=0.074) for R.pumilio was more than twice that estimated for mammals (H=0.036) while the mean percent polymorphism (P=16.1%) was only slightly higher than the mean percent polymorphism obtained for mammals (P=14.7%). The highest heterozygosities were recorded in the Potchefstroom (H=0 .145) and Zimbabwe (H=0 .118) samples and the lowest mean heterozygosity was recorded in the peninsular Western Cape (H=0. 032). A mean Fst value of 0.459 was obtained, suggesting a high degree of genetic differentiation between the samples of R.pumilio but the negative Fis (-0.01) value emphasized that R.pumilio retained an outbreeding population structure. The similarity coefficient between the samples of R.pumilio using PCR-RAPD's ranged between 0.471 and 0.853 and substantiated the argument for genetic divergence between the samples of R.pumilio. An isolation by distance model for the population genetic structure of R.pumilio was supported by the allozymes (r=0.58, p<0.00l) and PCR-RAPD's (0.75, p<0.00l). Temperature and rainfall also had an influence on the allelic frequency distribution of certain loci of R.pumilio. Rogers (1972) genetic similarity varied between 0.796 and 0.988 while the values for Nei's (1978) unbiased genetic distance varied between 0.000 and 0.189 for the different samples of R.pumilio. Subgrouping of the KwaZulu-Natal samples, the peninsular Western Cape and Eastern Cape samples of R.pumilio was evident with the allozymes. With the PCR-RAPD' s the Zimbabwe sample showed the least similarity to the other samples with a KwaZulu-Natal/Potchefstroom subgroup separating from the less well defined Eastern Cape and Western Cape subgroup. Cytogenetic studies of specimens of R.pumilio from some of the localities in southern Africa revealed a chromosomal number of 2n=48 , while the Potchefstroom and Zimbabwe specimens displayed a chromosomal number of 2n=46. Homology in G-and C-banding was recorded. The allozymes, PCR-RAPD's and chromosomal studies suggested subspecies status for the Zimbabwe population of R.pumilio. The Potchefstroom sample displayed a greater genetic similarity to the remaining South African samples of R.pumilio than the Zimbabwe samples and therefore could not be considered for subspecies status. Although the South African samples of R.pumilio displayed a certain degree of genetic divergence, it was insufficient to warrant subspecies status although evolution in this direction was suggested. / Thesis (Ph.D.)-University of Natal, Durban, 1999.

Rainfall is the most important factor limiting primary production, and hence food availability, in ecosystems. This study tests the thesis that, under conditions where rainfall is unpredictable and food supply varies spatially and temporally from year-to-year, physiological energy-saving mechanisms should evolve in animals for which migration is not an option to avoid periods of low habitat productivity. Apart from the normally erratic rainfall patterns characteristic of arid regions, unpredictable seasonal rainfall may also occur in mesic zones as a consequence of climatic phenomena such as the El Niño Southern Oscillations (ENSO). Using two hypothetical models, this study attempts to determine whether the respective evolutionary history of both aridity and ENSO can be detected in the various phenotypic physiological traits of Rhabdomys pumilio from across southern Africa. Model 1 predicts that a gradient of conservatism of physiological traits related to metabolism, will be detectable along an aridity gradient. Model 2 predicts an increase in the conservatism of similar traits correlated with both the aridity gradient and the gradient of negative rainfall anomalies that are associated with the ENSO. Measurements of circadian rhythms of body temperature and oxygen consumption of individuals of R. pumilio collected from six sites across southern Africa showed that the geographic variation in the physiological traits and proximate responses of this species was explained better by Model 2 than by Model 1. Furthermore, for some traits and proximate responses of this species it was shown that, although the stresses associated with unpredictable rainfall occur less frequently in non-desert ENSO-zones, they are nevertheless sufficient to mimic the selection pressures of rainfall unpredictability in arid zones. This study also highlighted the low basal rates of metabolism and the other conservative physiological traits of this species as a whole. The basal metabolic rate of 0.819 mlO₂g/h and the minimum wet thermal conductance of 0.110 mlO₂/g/h/ºC are lower than that predicted by allometry. The circadian amplitude of body temperature of 2.52°C and the circadian amplitude of oxygen consumption of 1.39 mlO₂/g/h were higher than those predicted by allometric equations. These deviations from predicted values were attributed to the adaptedness of this species to the physiological energy stresses associated with unpredictable resource availability in southern Africa. / Thesis (M.Sc.)-University of Natal, Pietermaritzburg, 1998.