Functional and phylogenetic approaches reveal the evolution of diversity in a hyper diverse biota

Ant communities in tropical forests may be governed by varying assembly mechanisms, depending on the particular habitat investigated. We compared phylogenetic diversity and structure across two forest biomes (dry and humid) and two vertical layers (arboreal and terricolous) in ant communities in Madagascar, and assessed the influence of invasive species on this community structure. We estimated phylogenetic signal and correlated evolution for habitat and several functional traits and tested for conservatism in relevant functional and habitat traits. Ancestral states were reconstructed to illuminate the evolution of habitat traits. All analyses utilized phylogenies estimated from newly generated data from three nuclear markers for 290 Malagasy ant taxa. Dry forests, although lower in species richness, were found to support equally high lineage diversity as humid forests. In contrast, phylogenetic diversity was much lower in arboreal than in terricolous communities. We observed significant phylogenetic clustering in the combined humid forest and in the arboreal–humid, arboreal–dry and terricolous–humid communities, whereas the combined dry forest community was overdispersed. Among ant communities in Madagascar, overdispersion and competition therefore may be more prevalent in dry forest, and habitat filtering may be more dominant in humid forest. Excluding invasive ant species had little overall effect on community structure. All investigated traits showed low to intermediate conservatism; strong support for correlated evolution was found for increased eye size and an arboreal lifestyle. Habitat transitions from humid to dry and from terricolous to arboreal occurred more frequently, and ancestors of most lineages were predicted to be terricolous or humid-forest adapted. We conclude that most Malagasy ant clades first colonized humid forests and subsequently transitioned into dry forests, indicating that previous hypotheses on the evolution of Madagascar's hyperdiverse biota may not apply to ants and other arthropods.