"Nutcracker Man" debunked

This week, Thure Cerling and colleagues report in PNAS [1] carbon stable isotope data from 24 specimens of Australopithecus boisei. This is a huge sample as fossil hominins go, and they give a very consistent picture about the diet of this most robust of the australopithecines. These 24 individuals got between 61 and 91 percent of their carbon from grasses.

My 2005 explainer on stable isotope chemistry and early hominin diets fills in the details about carbon-12, carbon-13 and their relationship to 3- and 4-carbon photosynthetic cycles. The salient aspect of the comparisons involving A. boisei here is that C₄ plants, mostly grasses, incorporate relatively more carbon-13 than do other plants, and herbivores assimilate this carbon-13 into their bones and teeth.

The high ratio of grass-derived carbon in A. boisei is fundamentally different from all living and fossil apes, and it is far higher than the values found for other early hominins. The only other primate that comes close is the fossil giant gelada Theropithecus oswaldi, a savanna-living species.

What were these extinct species really eating? Was grass the food? For living geladas, grass consumption includes seeds -- a fact that led Clifford Jolly to suggest that early hominins might also have specialized on seeds [2]. Of course, humans today also specialize on grass seeds. We call them grains, eat them in bread and drink them in soda. And beer.

But what about A. boisei? The large, thick-enameled premolars and molars, with their low cusps, seem well suited to grinding small hard objects and resisting the resulting wear. But Cerling and colleagues devote a good chunk of their discussion to the description of molar wear in A. boisei and other early hominins. Their argument is that the teeth of A. boisei show no signs of "hard object" feeding:

Of perhaps greater moment than its potential specific simila- rities, the microwear of P. boisei molars, which shows remarkable uniformity over time from about 2.3 Ma to about < 1.4 Ma (9, 24), stands in stark contrast to the wear fabrics exhibited by primate hard-object consumers. Indeed, there is no evidence beyond the anecdotal [e.g., the broken left first permanent molar crown in the KNM-ER 729 P. boisei mandible (8) and the observation that a couple of P. boisei molars show antemortem enamel chipping (25)] that these food items were hard.

These observations are not new, but putting them together with the evidence of grass consumption makes it pretty clear that seed eating was not a predominant source of dietary carbon. The "Nutcracker Man" sobriquet, applied to A. boisei because of its powerful jaw mechanics, must be false. No significant hard object feeding, very low dietary carbon from trees and non-grassy (or sedgy) plants.

Instead, Cerling and colleagues propose that both A. boisei and other early hominins wore their teeth on the, well, grassy parts of grass.

P. boisei cheek teeth display notable gradients of gross wear, resulting in large, deeply excavated dentine exposures, and in this regard, they are similar to other australopith species (e.g., A. afarensis and A. africanus) that also possess low tooth cusps with thick enamel. Thus, like other australopiths, P. boisei undoubtedly had a diet that consisted of foods with abrasive qualities—the gross wear is as likely due to repetitive loading of phytolith-rich tough foods as exogenous grit. Thus, either grass or sedge consumption and/or exogenous grit might well have contributed to P. boisei’s notable wear gradient.

And:

Recent dental microwear studies suggest that the mechanical properties of A. afarensis (and A. anamensis) diets were nearly identical to those of P. boisei (9, 24, 40–42). If this is so, could it be that the australopith masticatory package represents an adaptation to C4 resources such as grasses or sedges? The similarity in dental microwear fabrics among the eastern African australopiths, all of which lack any evidence for hard-object food consumption (9, 24, 40–42), is consistent with the notion that their craniodental morphology could reflect “repetitive loading” rather than hard-object consumption (7, 8, 43).

Grit might get in from eating underground parts like rhizomes. Phytoliths are small, hard silicate structures in the green parts of plants, including the stems and leaves of grass.

Last year I wrote about carbon isotope analysis of two specimens of Australopithecus boisei, the famous OH 5 "Zinj" specimen, and the Peninj mandible. Both specimens show evidence of a high consumption of grass-derived carbon -- estimated at 77% and 81% grass-derived carbon, respectively. Those levels are characteristic of grazing animals. Cerling and colleagues show that these values are right in the middle of the range among specimens of A. boisei that cover a half million years in Kenya and Tanzania.

In the paper reporting the carbon stable isotopes of OH 5 and Peninj, van der Merwe and colleagues [3] suggested that A. boisei may have relied on papyrus as a staple. The culms and rhizomes of papyrus both have substantial nutritional content but are very fibrous and require much chewing and spitting out fiber at intervals. The hypothesis would imply that A. boisei relied on these foodstuffs for the majority of its calories.

Cerling and colleagues do not mention papyrus, and take a much more direct approach on grass-eating. But they do report data on oxygen stable isotopes from the specimens that may be relevant to the ecological context of grass (or sedge) consumption. Oxygen isotopes in bone and teeth reflect the pattern of water consumption by an animal. Oxygen-16 evaporates and transpires preferentially from leaves, so an animal living in an arid environment that gets most of its water from plants will be relatively enriched for the heavier oxygen-18. An animal that depends on drinking water from lakes or rivers will tend to have lower oxygen-18. A. boisei is almost as low in oxygen-18 composition as hippopotamus, suggesting they were strongly dependent on water sources.

A highly water-dependent grass-eating A. boisei is a very different picture of the biology of this robust species. The South African robust species, A robustus, is very different in this regard. These two species are often lumped together, but this is unfair in many ways to their distinctive anatomical patterns. Knowing that their dietary adaptations were very distinct, we should be more inclined to focus on the details where they differ.

Bottom line: A. boisei represents a highly distinctive dietary pattern, not present in any living ape, that no longer exists. At least the giant gelada, T. oswaldi, may also have exploited similar resources. Some grass resources, including papyrus corms and rhizomes, have high caloric and nutritional value, but require adaptations to deal with the fibrous content.

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