Fluctuating climate and dietary innovation drove ratcheted evolution of proboscidean dental traits.

Identification of the selective forces that shaped adaptive phenotypes generally relies on current habitat and function, but these may differ from the context in which adaptations arose. Moreover, the fixation of adaptive change in a fluctuating environment and the mechanisms of long-term trends are still poorly understood, as is the role of behaviour in triggering these processes. Time series of fossils can provide evidence on these questions, but examples of individual lineages with adequate fossil and proxy data over extended periods are rare. Here, we present new data on proboscidean dental evolution in East Africa over the past 26 million years, tracking temporal patterns of morphological change in relation to proxy evidence of diet, vegetation and climate (aridity). We show that behavioural experimentation in diet is correlated with environmental context, and that major adaptive change in dental traits followed the changes in diet and environment but only after acquisition of functional innovations in the masticatory system. We partition traits by selective agent, showing that the acquisition of high, multiridged molars was primarily a response to an increase in open, arid environments with high dust accumulation, whereas enamel folding was more associated with the amount of grass in the diet. We further show that long-term trends in these features proceeded in a ratchet-like mode, alternating between directional change at times of high selective pressure and stasis when the selective regime reversed. This provides an explanation for morphology adapted to more extreme conditions than current usage (Liem's Paradox). Our study illustrates how, in fossil series with adequate stratigraphic control and proxy data, environmental and behavioural factors can be mapped on to time series of morphological change, illuminating the mode of acquisition of an adaptive complex.