An analysis of the Cordylus Polyzonus complex (Reptilia : Cordylidae) in the South-Western Cape
Thesis (MSc (Botany and Zoology))--University of Stellenbosch, 1990.
Mouton and Oelofsen (1988) suggested that melanistic cordylid populations in the southwestern Cape represent relict, cold-adapted populations which evolved under adverse climatic conditions during the Last Glacial Period, 18000-16000 BP. The first section of this study was undertaken to test their model for the evolution of melanism in the cordylid species, Cordylus polyzonus. For this purpose, geographic character variation among populations of Cordylus polyzonus in the region south of 32°15' latitude and west of 19015' longitude was investigated. Variation in 122 external morphological characters was analysed in 306 specimens from 93 localities. Although no inter-locality variation was observed in most of the characters, a high degree of concordancy was observed in the geographical variation of three headshield characters. These three headshield characters are considered non-adaptive and can accordingly be regarded as good indicators of genealogical relationship. Since they varied together with other adaptive characters such as melanism and body size, it can be concluded that geographically isolated melanistic populations of Cordylus polyzonus represent relicts of a once larger melanistic population. The data underscore the Mouton-Oelofsen hypothesis for the evolution of melanistic cordylid taxa in the south-western Cape. A further corrolary of the Mouton-Oelofsen hypothesis, is that melanistic cordylids are presently restricted to cool enclaves. This assumption was evaluated in the second section of this study by analysing prevailing climatic conditions associated with the occurrence of extant melanistic populations of the genera Cordylus and Pseudocordylus in the southwestern Cape. Climatic data were obtained from 123 weather stations in the study-area. A close correspondence was found between the distribution of melanistic populations along the westcoast and the primary upwelling zones of the southern Benguela Current. Lower mean daily temperatures and a high incidence of advective sea fog on the adjacent coastal regions are direct effects of these cold upwelled waters. Likewise, montane melanistic populations, occurring at relatively high altitudes along the extreme western borders of the Cape Fold Mountains, also experience lower mean daily temperatures and a high incidence of orographic fog and cloud cover. Melanistic cordylid populations therefore generally have to 4. contend with relatively exposed environmental conditions of low temperature and limited solar radiation due to the filtering effect of fog and cloud cover. The fact that all the melanistic taxa in this region occur as small isolated populations limited to cool enclaves, suggests that they are presently in a contracted state under strong environmental pressure. It is postulated that palaeoclimatic conditions very similar to climatic conditions presently prevailing in these enclaves were experienced over the entire western coastal region, at least as far as the Orange River, during the Last Glacial Maximum when melanism probably evolved. Our results therefore corroborate the Mouton-Oelofsen hypothesis that isolated melanistic cordylid populations represent cold-adapted relicts. One would accordingly expect melanistic populations to posses some enhanced ability to absorb infrared radiant heat to cope with these conditions of limited sunshine and lower temperatures. In section three of this study, differences in dorsal skin reflectivity between melanistic and turquoise ~. polyzonus populations were quantified. Furthermore, the physiological ability of this species to change body colour was investigated, as well as ontogenetic colour change. Significant differences in dorsal skin reflectivity existed between melanistic and turquoise specimens in the 500-1300 mjl spectral range, demonstrating the greater heat-absorbing capacity of melanistic populations. The data underscore the MoutonOelofsen model that melanistic populations are "cold-adapted" relicts. Furthermore, ontogenetic colour change is a real phenomenon in the melanistic variation of C. polyzonus. In this respect it is unique among the melanistic cordylid taxa in that ontogenetic colour change seemingly does not occur in the other forms. Experiments to investigate short term colour change in response to different temperatures regimes, produced no conclusive results. On the other hand, seasonal colour changes could be demonstrated for both melanistic and turquoise specimens, indicating that this species has the physiological capacity to change colour. It is, however, believed that geographic colour variation in ~. polyzonus cannot be attributed to this capacity, but is rather the result of selection over time. This view is underscored by the results of section one demonstrating that, apart from colour and body size, melanistic and turquoise forms also differ in certain non-adaptive traits; suggesting that they do not belong to the same primary gene pool.