Scott A. Blumenthal

Dietary changes of large herbivores in the Turkana Basin, Kenya from 4 to 1 Ma

Significance Stable carbon isotopes give diet information for both modern and fossil mammals and can be used to classify diets as C4 grazers, C3–C4 mixed, or C3 browsers. We show that diets of some major African herbivore lineages have significantly changed over the past 4 million years by comparing fossils from the Turkana Basin in Kenya with modern mammals from East and Central Africa. Some fossil assemblages have no modern analogues in East and Central Africa, suggesting different ecological functions for some mammals in the past as compared with their modern counterparts. The development of modern tropical grassland ecosystems are products of the coevolution of both grasses and herbivores. A large stable isotope dataset from East and Central Africa from ca. 30 regional collection sites that range from forest to grassland shows that most extant East and Central African large herbivore taxa have diets dominated by C4 grazing or C3 browsing. Comparison with the fossil record shows that faunal assemblages from ca. 4.1–2.35 Ma in the Turkana Basin had a greater diversity of C3–C4 mixed feeding taxa than is presently found in modern East and Central African environments. In contrast, the period from 2.35 to 1.0 Ma had more C4-grazing taxa, especially nonruminant C4-grazing taxa, than are found in modern environments in East and Central Africa. Many nonbovid C4 grazers became extinct in Africa, notably the suid Notochoerus, the hipparion equid Eurygnathohippus, the giraffid Sivatherium, and the elephantid Elephas. Other important nonruminant C4-grazing taxa switched to browsing, including suids in the lineage Kolpochoerus-Hylochoerus and the elephant Loxodonta. Many modern herbivore taxa in Africa have diets that differ significantly from their fossil relatives. Elephants and tragelaphin bovids are two groups often used for paleoecological insight, yet their fossil diets were very different from their modern closest relatives; therefore, their taxonomic presence in a fossil assemblage does not indicate they had a similar ecological function in the past as they do at present. Overall, we find ecological assemblages of C3-browsing, C3–C4-mixed feeding, and C4-grazing taxa in the Turkana Basin fossil record that are different from any modern ecosystem in East or Central Africa.

Detecting intraannual dietary variability in wild mountain gorillas by stable isotope analysis of feces

We use stable isotope ratios in feces of wild mountain gorillas (Gorilla beringei) to test the hypothesis that diet shifts within a single year, as measured by dry mass intake, can be recovered. Isotopic separation of staple foods indicates that intraannual changes in the isotopic composition of feces reflect shifts in diet. Fruits are isotopically distinct compared with other staple foods, and peaks in fecal δ13C values are interpreted as periods of increased fruit feeding. Bayesian mixing model results demonstrate that, although the timing of these diet shifts match observational data, the modeled increase in proportional fruit feeding does not capture the full shift. Variation in the isotopic and nutritional composition of gorilla foods is largely independent, highlighting the difficulty for estimating nutritional intake with stable isotopes. Our results demonstrate the potential value of fecal sampling for quantifying short-term, intraindividual dietary variability in primates and other animals with high temporal resolution even when the diet is composed of C3 plants.

Paleoenvironmental context of the Middle Stone Age record from Karungu, Lake Victoria Basin, Kenya, and its implications for human and faunal dispersals in East Africa.

The opening and closing of the equatorial East African forest belt during the Quaternary is thought to have influenced the biogeographic histories of early modern humans and fauna, although precise details are scarce due to a lack of archaeological and paleontological records associated with paleoenvironmental data. With this in mind, we provide a description and paleoenvironmental reconstruction of the Late Pleistocene Middle Stone Age (MSA) artifact- and fossil-bearing sediments from Karungu, located along the shores of Lake Victoria in western Kenya. Artifacts recovered from surveys and controlled excavations are typologically MSA and include points, blades, and Levallois flakes and cores, as well as obsidian flakes similar in geochemical composition to documented sources near Lake Naivasha (250 km east). A combination of sedimentological, paleontological, and stable isotopic evidence indicates a semi-arid environment characterized by seasonal precipitation and the dominance of C4 grasslands, likely associated with a substantial reduction in Lake Victoria. The well-preserved fossil assemblage indicates that these conditions are associated with the convergence of historically allopatric ungulates from north and south of the equator, in agreement with predictions from genetic observations. Analysis of the East African MSA record reveals previously unrecognized north-south variation in assemblage composition that is consistent with episodes of population fragmentation during phases of limited dispersal potential. The grassland-associated MSA assemblages from Karungu and nearby Rusinga Island are characterized by a combination of artifact types that is more typical of northern sites. This may reflect the dispersal of behavioral repertoires-and perhaps human populations-during a paleoenvironmental phase dominated by grasslands.

Aridity and hominin environments

Significance Oxygen isotopes in modern and fossil mammals can provide information on climate. In this study, we provide a new record of aridity experienced by early hominins in Africa. We show that past climates were similar to the climate in eastern Africa today, and that early hominins experienced highly variable climates over time. Unexpectedly, our findings suggest that the long-term expansion of grasses and grazing herbivores since the Pliocene, a major ecological transformation thought to drive aspects of hominin evolution, was not coincident with aridification in northern Kenya. This finding raises the possibility that some aspects of hominin environmental variability might have been uncoupled from aridity, and may instead be related to other factors, such as rainfall seasonality or ecological interactions among plants and mammals. Aridification is often considered a major driver of long-term ecological change and hominin evolution in eastern Africa during the Plio-Pleistocene; however, this hypothesis remains inadequately tested owing to difficulties in reconstructing terrestrial paleoclimate. We present a revised aridity index for quantifying water deficit (WD) in terrestrial environments using tooth enamel δ18O values, and use this approach to address paleoaridity over the past 4.4 million years in eastern Africa. We find no long-term trend in WD, consistent with other terrestrial climate indicators in the Omo-Turkana Basin, and no relationship between paleoaridity and herbivore paleodiet structure among fossil collections meeting the criteria for WD estimation. Thus, we suggest that changes in the abundance of C4 grass and grazing herbivores in eastern Africa during the Pliocene and Pleistocene may have been decoupled from aridity. As in modern African ecosystems, other factors, such as rainfall seasonality or ecological interactions among plants and mammals, may be important for understanding the evolution of C4 grass- and grazer-dominated biomes.

Stable isotopic variation in tropical forest plants for applications in primatology.

Stable isotope analysis is a promising tool for investigating primate ecology although nuanced ecological applications remain challenging, in part due to the complex nature of isotopic variability in plant‐animal systems. The aim of this study is to investigate sources of carbon and nitrogen isotopic variation at the base of primate food webs that reflect aspects of primate ecology. The majority of primates inhabit tropical forest ecosystems, which are dominated by C3 vegetation. We used stable isotope ratios in plants from Kibale National Park, Uganda, a well‐studied closed‐canopy tropical forest, to investigate sources of isotopic variation among C3 plants related to canopy stratification, leaf age, and plant part. Unpredictably, our results demonstrate that vertical stratification within the canopy does not explain carbon or nitrogen isotopic variation in leaves. Leaf age can be a significant source of isotopic variation, although the direction and magnitude of this difference is not consistent across tree species. Some plant parts are clearly differentiated in carbon and nitrogen isotopic composition, particularly leaves compared to non‐photosynthetic parts such as reproductive parts and woody stem parts. Overall, variation in the isotopic composition of floral communities, plant species, and plant parts demonstrates that stable isotope studies must include analysis of local plant species and parts consumed by the primates under study from within the study area. Am. J. Primatol. 78:1041–1054, 2016.

Stable carbon, oxygen, and nitrogen, isotope analysis of plants from a South Asian tropical forest: Implications for primatology

Stable isotope analysis of primate tissues in tropical forest contexts is an increasingly popular means of obtaining information about niche distinctions among sympatric species, including preferences in feeding height, forest canopy density, plant parts, and trophism. However, issues of equifinality mean that feeding height, canopy density, as well as the plant parts and plant species consumed, may produce similar or confounding effects. With a few exceptions, researchers have so far relied largely on general principles and/or limited plant data from the study area as references for deducing the predominant drivers of primate isotope variation. Here, we explore variation in the stable carbon (δ13C), nitrogen (δ15N), and oxygen (δ18O) isotope ratios of 288 plant samples identified as important to the three primate species from the Polonnaruwa Nature Sanctuary, Sri Lanka, relative to plant part, season, and canopy height. Our results show that plant part and height have the greatest effect on the δ13C and δ18O measurements of plants of immediate relevance to the primates, Macaca sinica, Semnopithecus priam thersites, and Trachypithecus vetulus, living in this monsoonal tropical forest. We find no influence of plant part, height or season on the δ15N of measured plants. While the plant part effect is particularly pronounced in δ13C between fruits and leaves, differential feeding height, and plant taxonomy influence plant δ13C and δ18O differences in addition to plant organ. Given that species composition in different regions and forest types will differ, the results urge caution in extrapolating general isotopic trends without substantial local baselines studies.

Hippopotamus (H. amphibius) diet change indicates herbaceous plant encroachment following megaherbivore population collapse.

Megaherbivores (>1000 kg) are critical for ecosystem health and function, but face population collapse and extinction globally. The future of these megaherbivore-impoverished ecosystems is difficult to predict, though many studies have demonstrated increasing representation of C3 woody plants. These studies rely on direct observational data, however, and tools for assessing decadal-scale changes in African ecology without observation are lacking. We use isotopic records of historical common hippopotamus (Hippopotamus amphibius) canines to quantify herbaceous vegetation change in Queen Elizabeth National Park, Uganda following a period of civil unrest and poaching. This poaching event led to population collapse of two threatened African megaherbivore species: hippopotamus and African elephants (Loxodonta africana). Serial carbon isotope ratios (δ13C) in canine enamel from individuals that lived between 1960–2000 indicated substantial increases in C3 herbaceous plants in their diet (<20% C3 in the 1960s to 30–45% C3 in the 80s and 90s), supported by other observational and ecological data. These data indicate megaherbivore loss results in succession of both woody and herbaceous C3 vegetation and further reaching effects, such as decreased grazing capacity and herbivore biodiversity in the area. Given multiple lines of evidence, these individuals appear to accurately capture herbaceous vegetation change in Mweya.

Diets of mammalian fossil fauna from Kanapoi, northwestern Kenya

Carbon isotope ratios of mammalian teeth from the Kanapoi site in northern Kenya are interpreted in the context of C3 and C4 derived resources to investigate the paleoecology of Australopithecus anamensis. δ13C values of large mammals, when compared at the taxon level, show an ecosystem that is strongly biased towards mixed feeders and browsers. However, sufficient C4 resources were present such that some C4 dominated grazers were also present in the large mammal fauna. Analyses of micromammals shows that their diets were C3 dominated or C3-C4 mixed. Carbon isotope studies of primates shows that the major primate tribes-Colobini, Papioini, Hominini-all made some use of C4 resources in their respective diets; the Hominini had a higher fraction of C3 diet resources than the other primate tribes represented in the fossil record.