Escarpment Retreat Drives Diversification of Eastern Madagascar through Allopatric Speciation

Madagascar, as a biodiversity hotspot on Earth, exhibits a high level of endemism as observed from the prevalent vicariant speciation of terrestrial mammals, amphibians, and flora. Species richness of the island is uneven, with the highest species richness and endemism found on the steep great escarpment of the eastern margin. The unevenness is further observed within the escarpment region in that phylogenic turnover shows both latitudinal and altitudinal variations. Madagascar has remained almost tectonically inactive since the last rifting with Seychelles-India in the late Cretaceous. The high diversity and endemism of Madagascar challenge the conventional notion of uplift-driven speciation, which argues that speciation is driven by the formation of diverse habitat types from tectonic uplift.

Although the fundamental topographic framework of Madagascar has been in place since the late Cretaceous, it is modified in the Cenozoic by multiple processes including island-wide mantle-driven dynamic uplift, erosion-driven landward retreat of the escarpment at the eastern margin, localized volcanic and faulting activities. Our topographic reconstruction reveals that the dominant correlation is between the escarpment and species richness. To investigate the causal mechanisms of the diversity at the eastern escarpment, we constructed landscape evolution models, tracing the dynamics of habitable land surface patches throughout model simulations.

We investigated two distinct landscape scenarios: an escarpment retreat model simulating river incision into a pre-existing plateau with negligible tectonic uplift, and a tectonic uplift model featuring spatially and temporally constant uplift with river incision into the resulting mountain range. The steady-state topographic height of the tectonic uplift model is calibrated to match the plateau elevation of the escarpment model to ensure the same number of habitat types between models. The landscape of a great escarpment is highly dynamic and the heterogenous retreat of the escarpment and the water divide makes the geographically isolated drainage basins expand landward at different rates during the retreat process. Within the escarpment region, habitat patches dynamically appear, disappear, fragment, or merge at a frequency that scales with the retreat rate. In contrast, the tectonic uplift model only exhibits similar dynamic landscape change during the transient phase with habitat patches stabilizing spatially and temporally once a steady state topography was achieved.

The models predict that escarpment retreat fosters habitat patch dynamics such that patches isolate, or reconnect with a frequency on the order of a million years, appropriate for allopatric speciation. The habitat patch dynamics are a consequence of processes of catchment expansion, river captures, isolation of highland remnants, and formation of topographic barriers during the retreat. We conclude that the spatially heterogeneous but temporally steady retreat of the Madagascar escarpment since rifting has sustained allopatric speciation over evolutionary timescales resulting in the observed high diversity and its spatial pattern of eastern Madagascar.