Miocene ungulates and terrestrial primary productivity: where have all the browsers gone?

Progressive changes are observed in both the composition of mammal faunas and vegetation during the Miocene epoch [24-5 mega-annum (Ma)]. These changes are usually interpreted as a response to climatic changes. In the traditional view, forests or woodlands gradually gave way to more open habitats, with grazing (grass-eating) ungulate (hoofed) mammal species replacing the browsing (leafy-vegetation-eating) species as grasslands expanded. However, data from fossil assemblages in the Great Plains region of North America show that this faunal change was not a one-for-one replacement of browsers by grazers, as usually thought. Typical late early Miocene (17 Ma) fossil communities included extraordinarily high numbers of browsing ungulate species, comprising a fauna that cannot be directly analogized with any present-day community. Both maximum species richness of all ungulates and the proportion of browsers declined steadily in ungulate communities through the middle Miocene, to levels comparable to those of the present by the late Miocene. The resulting dramatic, cumulative loss of browsing species constitutes one of the strongest faunal signals of the late Tertiary (but was not a single "event"). We suggest that the early Miocene browser-rich communities may reflect higher levels of primary productivity in Miocene vegetation, compared with equivalent present-day vegetation types. The observed decline in species richness may represent a gradual decline in primary productivity, which would be consistent with one current hypothesis of a mid-Miocene decrease in atmospheric CO2 concentrations from higher mid-Cenozoic values.