African elephant genetics: enigmas and anomalies

During the last two decades, our understanding of the genetics of African elephant populations has greatly increased. Strong evidence, both morphological and genetic, supports recognition of two African elephant species: the savanna elephant (Loxodonta africana) and the forest elephant (L. cyclotis). Among elephantids, phylogeographic patterns for mitochondrial DNA are highly incongruent with those detected using nuclear DNA markers, and this incongruence is almost certainly due to strongly male-biased geneflow in elephants. As our understanding of elephant population genetics has grown, a number of observations may be considered enigmatic or anomalous. Here, several of these are discussed. (i) There are a number of within-species morphological differences purported to exist among elephants in different geographic regions, which would be difficult to reconcile with the low genetic differentiation among populations. (ii) Forest elephants have a higher effective population size than savanna elephants, with nuclear genetic markers much more diverse in the forest elephants than savanna elephants, yet this finding would need to be reconciled with the life history of the two species. (iii) The savanna and forest elephants hybridize and produce fertile offspring, yet full genome analysis of individuals distant from the hybrid zone suggests that gene flow has been effectively sterilized for atleast ∼500,000 years. (iv) There are unexplored potential ramifications of the unusual mito-nuclear patterns among elephants. These questions are considered in light of highmale and low female dispersal in elephants, higher variance of reproductive success among males than females, and of habitat changes driven by glacial cycles and human activity.