What can hippopotamus isotopes tell us about past distributions of C4 grassy biomes on Madagascar?

Abstract

Societal Impact StatementToday, expansive C4 grassy biomes exist across central, western, and northern Madagascar. Some researchers have argued that the island's now‐extinct pygmy hippopotamuses belonged to a megaherbivore grazing guild that maintained these grasslands prior to human arrival. However, the chemistry of hippo bones indicates that C4 grasses were only a minor part of hippo diet. This, in turn, suggests that C4 grasses were present but not widespread when hippos were alive and that grasses expanded only after Malagasy people shifted from hunting and foraging to agropastoralism approximately 1000 years ago. These results have important implications for environmental reconstructions and biodiversity management.Summary Extinct hippopotamuses (Hippopotamus spp.) were part of Madagascar's megaherbivore guild. Stable carbon (δ13C) and nitrogen (δ15N) isotopes in radiometrically dated bone collagen track spatial and temporal variation in diet and habitat. If hippos helped maintain C4 grassy biomes, then they should have regularly consumed C4 grasses, which have high δ13C values. However, if expansive C4 grassy biomes are anthropogenic, then forests would have been more extensive in the past, and hippos would have predominantly consumed C3 plants with low δ13C values. Nitrogen isotopes can clarify foraging habitat (moist or dry). We assessed δ13C and δ15N values for hippos from different ecoregions of Madagascar and compared these with data for extinct herbivorous lemurs from the same ecoregions. We further explored the effects of wet/dry transitions on isotopic trends for hippos from the central highlands and spiny thicket ecoregions. Carbon isotopes suggest (1) limited C4 consumption by hippos in the central highlands, dry deciduous forest, and succulent woodland ecoregions; and (2) moderate consumption of C4 resources in the spiny thicket. Nitrogen data indicate that hippos foraged in wetter habitats than sympatric lemurs in all regions. Malagasy hippos did not regularly graze C4 grasses in dry, open habitats, even in regions blanketed by C4 grassy biomes today. Malagasy grasses are adapted to grazing and fire, but these are likely ancient adaptations that accompanied grasses when they initially spread to Madagascar. C4 grassy biomes were spatially limited in extent in the past and only expanded after the Late Holocene introduction of domesticated ungulates.