A selection of extinct African megaherbivores
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Scarce emergence of new species drove the decline of African megaherbivores | Museo Nacional de Ciencias Naturales
Africa’s elephants, rhinoceroses and hippopotamuses are surviving representatives of a much richer assemblage of giant herbivores that once included such animals as
Deinotherium, an elephant relative with downward-curving tusks, and the pig-like anthracotheres, relatives of modern hippopotamuses. These animals were not merely conspicuous members of their ecosystems. As “ecosystem engineers”, megaherbivores—here defined as herbivorous mammals weighing at least one tonne—alter vegetation, disperse seeds, open pathways, create water holes and redistribute nutrients.
Why so much of this diversity disappeared has usually been approached as a question of extinction. Large animals reproduce slowly, require extensive habitats and comparatively large quantities of food, so it seems intuitively reasonable to assume that they must have been especially vulnerable when climates and environments changed. Human hunting has also figured prominently in explanations of the much more recent global disappearance of large mammals.
But the diversity of any group depends upon two processes, not one. Extinction removes species, while speciation produces new ones. Even a lineage with a comparatively modest extinction rate will decline if its species disappear faster than evolutionary divergence can replace them. Conversely, a group can withstand relatively frequent extinctions if new species continue to arise sufficiently rapidly.
A new study
published open access in Nature Communications. has now examined both sides of that evolutionary balance. Juan L. Cantalapiedra and colleagues analysed 3,327 African fossil occurrences representing 396 species of herbivorous mammals. Using neural-network models, they reconstructed changes in speciation and extinction over the past 23 million years while considering body size, tooth-crown height, evolutionary relationships and environmental change. Their results challenge the apparently obvious assumption that Africa’s largest herbivores declined because they were inherently more prone to extinction.
In fact, the models indicate that extinction rates were generally somewhat lower among the largest species. High-crowned teeth, which better withstand abrasive diets, were also associated with a reduced risk of extinction. The underlying problem for megaherbivores was that their already low speciation rates could not compensate for the species that were lost.
The imbalance developed over millions of years. As Africa became increasingly arid from about 7.2 million years ago, both speciation and extinction initially accelerated as changing vegetation and expanding open habitats reorganised its ungulate communities. Speciation then levelled off after about 3.6 million years, while extinction continued to rise and increased sharply with the beginning of the Pleistocene, about 2.58 million years ago. During the driest intervals, smaller herbivores generated new species at substantially higher rates, whereas speciation among megaherbivores was suppressed still further. The resulting decline was already well under way long before humans possessed the technology needed to hunt such enormous animals systematically.
This does not mean that humans played no part in the most recent losses, nor that climate alone explains every extinction. Rather, it places recent events within a much longer evolutionary history in which changing climate, declining ecosystem productivity, extinction and unequal rates of speciation gradually transformed the African fauna.