Did the Middle Stone Age make any difference for hunters?
Reading a meta-analysis of faunal data by Geoff Smith and coworkers that concludes that all Middle Pleistocene African peoples hunted the same prey animals.
If there is one generalization that we can make from the African record, with all its imperfections, it is that large cutting tools like handaxes were more common in assemblages from the earlier Middle Pleistocene than later. Those are defined as Acheulean. A second generalization is that Levallois-flaked points and evidence of hafting are more common in the later Middle Pleistocene than earlier. Across Africa, assemblages based on prepared core techniques like Levallois have been defined as Middle Stone Age.
The change in technology from Acheulean to Middle Stone Age in Africa has been portrayed as a major event in human prehistory. But was it really?
Paleoanthropologists bake a lot of assumptions into their thinking about technological change in the Pleistocene record. The biggest is the assumption of optimality, the idea that toolmakers made choices based on limited information to maximize returns and minimize costs. When two societies in the same place used different technical strategies—like handaxes versus prepared core flaking—the optimality assumption leads to the prediction that either the returns were different, the costs were different, or the information was different.
Returns and costs may be testable by looking at what the people ate. With hunting, that comes down to the choice of prey animals and hunting strategies. The costs of hunting depends on prey abundance, local landscape and habitat type, prey species defense strategies, and competition from other hunters—whether carnivore or hominin. Butchered animals bones in archaeological contexts are what we have to tell this story.
Did Acheulean hominins hunt differently than MSA hominins? You might think that MSA hafted spear points might make some difference in how ancient hominins hunted, scavenged, and ate animals. The biggest-ever review of this question was published in 2019, by Geoff Smith and coworkers. The work drew data from sites across Africa from 800,000 to 130,000 years ago. Looking at sites from Morocco across to Ethiopia and down through Kenya and Tanzania to South Africa, Smith and coworkers compiled faunal lists and other information from a varied array of more than 40 sites, some of which had faunal evidence from multiple time periods.
They conclude that what seems like a big change in technology made no difference at all to which animals hominins hunted.
The currently available faunal data do not support a broadening of the hominin dietary niche during the Middle Pleistocene. While smaller-sized bovids, such as gazelles, grysbok, southern reedbuck, and springbok, are preserved throughout the Middle Pleistocene sample, these species never illustrate significant increases. Similarly, dangerous game (e.g., Cape buffalo and long-horned buffalo) are recorded in small numbers throughout the faunal dataset. Increased proportions of dangerous game within MSA assemblages have been argued to reflect improvements in projectile technology, which provides the ability to hunt game at greater distances (Klein et al., 2007). While the proportion of these species increases from the early (0.03%) to late Middle Pleistocene (4%), they never predominate. Results from this metastudy suggest that such a change neither occurred on a broad scale during the Middle Pleistocene nor in parallel with the appearance of MSA technologies.
Big and small prey animals
The data are pretty clear about two things. First, the faunal evidence differs a lot between open air and cave sites. Cave assemblages have almost none of the very large prey species like elephants, hippos, and buffalo. Open air sites have these almost as often as smaller prey species.
What’s going on? These results reflect two biases: one caused by prey transport costs, and the other caused by researchers. Hominins did not move the bones of elephants and hippos to caves, and they rarely moved large bovid remains. The giant bones of these animals are hard to carry, so nobody carried them. They mostly disarticulated, butchered and ate such prey at or near the kill site. Small antelopes and other smaller prey could be carried whole or dismembered at a kill site with meaty parts carried off to a safer location—sometimes a cave or rockshelter.
The excavation bias is that archaeologists have been more likely to investigate open air sites that have obvious preservation of large mammal remains along with stone artifacts. Cave and rockshelter sites are likely to be excavated even without such obvious faunal evidence.
Both of these are known biases in zooarchaeology. Still, when you see data that illustrate the extreme difference of cave and open air faunal assemblages, they remind you just how much the hominin behavioral record is biased by the places that archaeologists prefer to dig. These biases matter. There are many more MSA cave sites, and in the African record included by Smith and coworkers, there is only one single Acheulean cave site! A large fraction of hominin foraging behavior is entirely missing from the record.
There really is no difference between Acheulean and MSA faunal size distributions other than the greater number of MSA sites that are caves.
Carcass access and processing intensity
Another way that sites can differ in faunal exploitation is how each individual animal was processed. At some sites, carnivores may have been able to get carcasses from hominin hunters, or hominins may have scavenged carcasses that were killed by carnivores. At other sites, hominins may have excluded carnivores almost entirely. This suggests a degree of control over their environments.
Dividing the sites into earlier and later Middle Pleistocene, Smith and coworkers found that the earlier sites were more likely to show carnivore traces on the prey animals. Later sites have more cutmarks and evidence for transport of selected body parts. However, that greater intensity of hominin involvement is confounded with the bias toward cave sites in the later Middle Pleistocene. Hominins that transported small prey animals to caves were less likely to have carnivore involvement; hominins that left cutmarks on elephant bones in open air sites were unlikely to keep carnivores from leaving their mark.
There is too little data on mortality profiles across these sites for Smith and coworkers to do a comparison of selectivity of prime age adults in prey species. Still, there are many Oldowan-era sites that have a clear bias toward selection of prime age adults by hominins, at least for some prey size classes. It is not probably going to be very informative to look at more detailed aspects of prey selection without first knowing that the hominin contribution is much higher than carnivore contribution to faunal assemblages.
Who were the hominins?
An enormous hole in this kind of project is the lack of knowledge of which hominins made the artifacts and interacted with the faunal remains. To their credit, Smith and coworkers are agnostic about which species of hominins are represented by the behavioral patterns across these many sites:
Currently, associated hominin fossils are limited and do not allow for direct correlations between various African Middle Pleistocene hominin species (e.g., Homo heidelbergensis, Homo rhodesiensis, Homo naledi and Homo sapiens) and a specific lithic technology or subsistence strategy. In general, this period is recognized by a mosaic of lithic entities and hominin species and the broad scale trends discussed here are complementary with more local and regional scale variation in terms of exact timings and causal mechanisms of behavioral change and continuity.
That’s an appropriately cautious statement. We do not know which hominin species made the artifact assemblages sampled at most Middle Pleistocene sites. The species or populations listed by Smith and coworkers—Homo heidelbergensis, Homo rhodesiensis, Homo naledi and Homo sapiens—are just an archaeologist’s sketch of the groups that may have been present across the continent during more than 600,000 years of time. I personally wouldn’t use all these names, but the point is that there were many populations with deep divergences across this time period in Africa. All of them were tool users and ate hunted animals. All of them were cultural.
It has been common over the last decade for archaeologists to assume that MSA assemblages were made by Homo sapiens in the narrow sense, meaning modern humans and their immediate ancestors. This has been a big component of the idea that modern humans had a “pan-African” origin.
But some of the earliest remains that almost all researchers attribute to H. sapiens are those from the Herto Member of in the Middle Awash region of Ethiopia, commonly known as the Herto hominins. Desmond Clark and coworkers (2003) described the stone artifact assemblages from the Herto Member as late Acheulean. By contrast, the Kabwe hominin skull—type fossil of H. rhodesiensis and not recognized by any researchers as a modern human—was found in a cave that had an artifact assemblage classified by Smith and collaborators as Sangoan or MSA. None of these finds are very solid evidence of archaeological association with fossil hominins. In the Kabwe case, the association was not recorded at the time of discovery and may just be a grab bag of material from a cave deposit; in the Herto case the artifacts and hominin fossils do not come from the same localities. But both cases help to remind us that the handful of known associations are not adequate to show which population or group may have made which artifacts.
With that caution in place, it is interesting that the faunal assemblages tell no story at all. There is no pattern over time across faunal assemblages in prey size selection. There is possibly an increase over time in cutmarks and decrease in carnivore involvement, but those cannot be tied to any particular hominin group.
I think that this pattern is striking evidence that earlier Middle Pleistocene hominins had achieved the ability to efficiently hunt prey species in the proportions that made ecological sense for them. The fact that later hominins selected the same prey sizes suggests that this aspect of hominin hunting may have been near equilibrium regardless of the precise toolkits they used. If you ask me who was doing the hunting, I’d say it was every hominin that existed in Africa across this time period. I think it is likely that they all had similar niches with respect to carnivory.
Note: I did a bit of a refresh on this post in April, 2026, to discuss the evidence with a bit more background.
References
Clark, J. D., Beyene, Y., WoldeGabriel, G., Hart, W. K., Renne, P. R., Gilbert, H., Defleur, A., Suwa, G., Katoh, S., Ludwig, K. R., Boisserie, J.-R., Asfaw, B., & White, T. D. (2003). Stratigraphic, chronological and behavioural contexts of Pleistocene Homo sapiens from Middle Awash, Ethiopia. Nature, 423(6941), 747–752. https://doi.org/10.1038/nature01670
Grün, R., Pike, A., McDermott, F., Eggins, S., Mortimer, G., Aubert, M., Kinsley, L., Joannes-Boyau, R., Rumsey, M., Denys, C., Brink, J., Clark, T., & Stringer, C. (2020). Dating the skull from Broken Hill, Zambia, and its position in human evolution. Nature, 580(7803), Article 7803. https://doi.org/10.1038/s41586-020-2165-4
Smith, G. M., Ruebens, K., Gaudzinski-Windheuser, S., & Steele, T. E. (2019). Subsistence strategies throughout the African Middle Pleistocene: Faunal evidence for behavioral change and continuity across the Earlier to Middle Stone Age transition. Journal of Human Evolution, 127, 1–20. https://doi.org/10.1016/j.jhevol.2018.11.011