Rib remodelling changes with body size in fossil hippopotamuses from Cyprus and Greece

Abstract

Large species that are isolated for thousands of years on islands often evolve extreme degrees of dwarfism. Very little is known about physiological processes that accompany such extreme transitions in extinct dwarf species. We tested whether physiological cycles of bone maintenance (remodelling) in dwarf adult hippopotamuses correlate with insularity-driven body mass shifts that may occur due to variables such as ecological release from predation pressure and change in access to resources. We hypothesised that hippopotamuses with the smallest body size should show higher values of osteocyte lacunae, proxies for osteoblast proliferation during cycles of remodelling, when compared to relatively larger dwarf forms, as well as much larger mainland common hippopotamuses. We examined 20 ribs from three extinct Pleistocene Hippopotamus species spanning a gradient in body size: H. minor (~132 kg, Cyprus), H. creutzburgi (~398 kg, Crete), and H. antiquus (~3200 kg, mainland Greece). Ribs were selected because they reflect bone metabolic rates that are not completely clouded by factors such as biomechanics. Densities of osteocyte lacunae (Ot.Dn) were examined in 864 individual secondary osteons observed in histology sections. We found the highest average Ot.Dn in the H. minor ribs, intermediate Ot.Dn in the H. creutzburgi ribs, and the lowest Ot.Dn in the H. antiquus ribs. It appears that Ot.Dn distinctly separated these three species, possibly signifying a gradient in bone remodelling such that bone tissue optimises maintenance in the face of insularity-driven reduction of body size. We discuss hippopotamus rib bone microstructure and the utility of Ot.Dn in palaeontological analyses for elucidating intricate biological processes occurring in bone of insular fossil mammals.