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ABSTRACT Modern African ungulates navigate seasonal variation in resource availability through diet‐switching (primarily mixed‐feeders) and/or migrating (primarily grass grazers). These ecological generalisations are well‐documented today, but the extent to which they apply to the non‐analog ecosystems of the Pleistocene are unclear. Drawing from serially‐sampled stable isotope measurements from 18 Kenyan large herbivore species from the Last Glacial Period (LGP), we evaluate how diet, diet‐switching, and migration compare to observations from present‐day settings. We find a higher grazing signal in most LGP species and a greater magnitude of diet‐switching than in the present. Additionally, we find that the relationships between grass intake, migration, diet‐switching, and body size during the LGP were unlike those observed today. This establishes a revised paleoecology of LGP herbivore communities and highlights that LGP herbivores were behaviourally non‐analog. Our results imply that ecological observations from present‐day settings offer an incomplete perspective of herbivore‐environment interactions. 

For many animals, migration is an important strategy for navigating seasonal bottlenecks in resource availability. In the savannas of eastern Africa, herds of grazing animals, including blue wildebeest (Connochaetes taurinus), Thomson's gazelle (Eudorcas thomsonii), and plains zebra (Equus quagga), travel hundreds of kilometers annually tracking suitable forage and water. However, we know nearly nothing about migration among the extinct species that often dominated Late Pleistocene communities. Using serially sampled 87Sr/86Sr and δ13C, we characterize the prehistoric movement and diet of the enigmatic wildebeest Rusingoryx atopocranion from two localities (Karungu and Rusinga Island) in the Lake Victoria Basin of western Kenya. We find clear evidence for migration in all four individuals studied, with three 87Sr/86Sr series demonstrating high-amplitude fluctuations and all falling outside the modeled isoscape 87Sr/86Sr ranges of the fossil localities from which they were recovered. This suggests that R. atopocranion exhibited migratory behavior comparable to that of its closest living relatives in the genus Connochaetes. Additionally, individuals show seasonally-variable δ13C, with a higher browse intake than modern and fossil eastern African alcelaphins indicating behavioral differences among extinct taxa otherwise unrecognized by comparison with extant related species. That this species was highly migratory aligns with its morphology matching that of an open grassland migrant: it had open-adapted postcranial morphology along with a unique cranial structure convergent with lambeosaurine dinosaurs for calling long distances. We further hypothesize that its migratory behavior may be linked to its extinction, as R. atopocranion disappears from the Lake Victoria Basin fossil sequence coincident with the refilling of Lake Victoria sometime after 36 ka, potentially impeding its past migratory routes. This study characterizes migration in an extinct eastern African species for the first time and shapes our ecological understanding of this unique bovid and the ecosystems in which Middle Stone Age humans lived. 

Abstract Though herbivore grass dependence has been shown to increase with body size across herbivore species, it is unclear whether this relationship holds at the community level. Here we evaluate whether grass consumption scales positively with body size within African large mammalian herbivore communities and how this relationship varies with environmental context. We used stable carbon isotope and community occurrence data to investigate how grass dependence scales with body size within 23 savanna herbivore communities throughout eastern and central Africa. We found that dietary grass fraction increased with body size for the majority of herbivore communities considered, especially when complete community data were available. However, the slope of this relationship varied, and rainfall seasonality and elephant presence were key drivers of the variation—grass dependence increased less strongly with body size where rainfall was more seasonal and where elephants were present. We found also that the dependence of the herbivore community as a whole on grass peaked at intermediate woody cover. Intraspecific diet variation contributed to these community‐level patterns: common hippopotamus (Hippopotamus amphibius) and giraffe (Giraffa camelopardalis) ate less grass where rainfall was more seasonal, whereas Cape buffalo (Syncerus caffer) and savanna elephant (Loxodonta africana) grass consumption were parabolically related to woody cover. Our results indicate that general rules appear to govern herbivore community assembly, though some aspects of herbivore foraging behavior depend upon local environmental context.