Taxonomy, phylogeny and biogeography of seed-harvesting ants in the Tetramorium solidum-group (Hymenoptera: Formicidae)
Thesis (MSc)--Stellenbosch University, 2013.
Ants emerged during the Cretaceous Period more than 100 million years ago (Grimaldi & Engel 2005; Perrichot et al. 2008) and diversified independently on different landmasses following the Early Cretaceous fragmentation of Pangaea (Grimaldi & Agosti 2000). Today, ants represent one of the most ecologically successful groups globally (Wilson 1987), and their diversity far exceeds that of other social insects (Hölldobler & Wilson 1990). Currently 21 subfamilies, 305 genera and 12465 recorded ant species exist (Bolton 2012). Ants occur in almost all terrestrial habitats (Brown 2000) and are not uniformly distributed across regions, biomes and continents. As with all other taxa, historical abiotic factors have played a major role in their diversification and biogeographic structure (Tolley et al. 2006), including climatic changes (Holldobler & Wilson 1990) and the necessity to adapt to new environmental conditions (Tolley et al. 2006). Many global areas still lack data on regional biodiversity and the historical processes that may have shaped it. This is particularly so for southern Africa where the ant fauna is still relatively poorly known (Robertson 2000) and where there have been no published biogeographic analyses for any ant taxon. There is a highly endemic ant fauna that has evolved in the dry regions of southern Africa, which shows unique adaptations to the extreme thermal niches of these regions. For example, ants in the genus Ocymyrmex have adapted to forage on barren ground at temperatures as high as 67 ºC, when most insects and other arthropods are unable to survive (Marsh 1985). Ocymyrmex have long legs and usually hold the head, thorax and gaster high above the substrate, which helps with body temperature regulation. These ants are most active at the hottest time of the day and they move very rapidly. For example, O. barbiger has been observed running at speeds of 23 m/min (Marsh, 1985). In some ant species such extreme temperatures may limit foraging activity (Ward, 2007) but, in the case of Ocymyrmex, adaptations to this extreme thermal niche allowed exploitation of resources (dead and heat stressed arthropods). Moreover, these ants have subterranean nests that are ideal to escape environmental extremes and to move their brood deeply under the ground (Marsh, 1985). The distribution of some ant taxa in these dry regions closely reflects borders between particular biomes. For instance, in South Africa the ponerine ant, Pachycondyla hottentota, and the sugar ant, Camponotus storeatus, follow the borders of the Nama and Succulent Karoo biomes respectively (Robertson 2000). In the light of this background the overall objective of this study is to revise the taxonomy and determine the historical drivers of diversification in an important group of arid-adapted ants, the Tetramorium solidum-group, which is mainly restricted to dry semi-desert areas in southern Africa. Members of the Tetramorium solidum-group are commonly encountered seed harvesters yet despite their apparent ecological importance, their taxonomy and biogeography remains largely unresolved. Thirty years elapsed since the last revision of the group by Bolton (1980) and since, ant surveys have considerably expanded the material available for study and provide an opportunity for a fresh perspective on this group. The specific aims of this project were twofold: 1) to revise the species of the Tetramorium solidum-group; and 2) to explore the phylogenetic relationships among members of the group and the bioclimatic factors restricting their current distributional ranges in southern Africa. My study is timely, as to date no attempt has been made to unravel the history of arid-adapted ant diversification in southern Africa. Such information is much needed for identifying areas of high species diversity and endemism (Azuma et al. 2006) and prioritizing areas for conservation (Faith 1992). The thesis consists of three chapters. In chapter one I use morphological characters to review the current taxonomy of the Tetramorium solidum-group. From these data I have compiled an updated identification key for the group and described five new species. In chapter two I have constructed a fossil-calibrated dated phylogeny for Tetramorium solidum-group species in order to explore the historical processes that may have contributed in structuring the current distribution patterns and endemism within and among the species of the group. In this chapter I also explore bioclimatic factors that may restrict the current distribution of members on the Tetramorium solidum-group, using correlative bioclimatic modeling approaches. Chapter three provides a brief and overall conclusion to the study.