The importance of large prey animals during the Pleistocene and the implications of their extinction on the use of dietary ethnographic analogies
Introduction
The higher extinction rate of large herbivores, relative to smaller herbivores, was considered to be a Late Pleistocene phenomenon (Koch and Barnosky, 2006). However, evidence for a much earlier decline in the richness and size of large herbivores in Africa is mounting (Faith et al., 2019: Smith et al., 2018, Potts et al., 2018, Bibi et al., 2017).
The result of this decline, as the title of Faith et al. (2019) paper makes clear, is that “Early hominins evolved within non-analog ecosystems.” In order to appreciate the significance of the decline to humans, we describe the unique importance of large prey animals to humans during the Pleistocene. We then examine the effect of the new findings on the validity of past estimates of the human trophic level, which are used intensively in explanations of human evolution and behavior and which rely heavily on presumed analogies between the recent ethnographic record and the Paleolithic.
The presence of large and very large animals in many archaeological sites throughout the Pleistocene is well-accepted. However, their value as a source of food during the Pleistocene has been contested.
Speaking directly to the subject of this paper, Speth (2010) comes out against the paradigm of the preference of large prey as a source of food with a set of arguments that rely in part on ethnographic sources. James O'Connell writes in the foreword to the book (Speth, 2010:viii): “ethnographic research further shows that hunters truly concerned with feeding their wives and children would generally do better by targeting a broader range of prey, including small game and plant foods.”
In studying the importance of large prey to humans in the Paleolithic, we identified two additional factors, besides the reliance on the ethnographic record, that hinder the full appreciation of the importance of large prey animals during the Paleolithic. The first is the potential for relative underrepresentation of large animals, and especially megaherbivores, in zooarchaeological assemblages. The second is the lack of research as to why large prey would be so uniquely valuable to the survival of humans.
In Ben-Dor and Barkai (In press), we investigated the potential for the underrepresentation of large animals in archaeological sites by analyzing an actualistic data of 60 hunts of the Hadza (Bunn et al., 1988, O'Connell et al., 1990). We found a substantial underrepresentation of the largest animals when using the standard Minimum Number of Individuals (MNI) and the Number of Identified Species (NISP) abundance indexes. The underrepresentation was substantially corrected with the use of MNI (but not NISP) based biomass index that takes into account the animals' weight. We also found a higher potential for the underrepresentation of megaherbivores because of a higher probability for the complete nonrepresentation of megaherbivores' hunts in the MNI. The underrepresentation of megaherbivores at Paleolithic sites was also advocated by other researchers using different methods of analysis (Bocherens et al., 2005, Mackie, 2019, Morin et al., 2016, Sinet-Mathiot et al., 2019).
In the present paper, we will try to explain how bioenergetic considerations, coupled with higher fat content, made large animals critical to human survival during the Pleistocene.
The different preservation potential of plants and animals remains in the archaeological record may have been the reason for the intensive use of analogies with the ethnographic record as a source for estimates of the Paleolithic human trophic level as it is represented in the Plant:Animal-sourced food ratio (Pla-AniR) (Cordain et al., 2000, Eaton and Konner, 1985, Konner and Eaton, 2010, Kuipers et al., 2012, Kuipers et al., 2010, Lee, 1968, Marlowe, 2005, Stahl et al., 1984, Ströhle and Hahn, 2011). For example, Lee (1968:43) hypothesized, based on his observations of the Ju/’hoansi, and data from 57 other hunter-gatherers (HG) groups, that HG would emphasize the more reliable food source, namely plants and consume a Pla-AniR of 65%:35%. Most of the ethnography based Pla-AniR estimates find a relatively large plant portion of the diet, especially in lower latitude regions like Africa.
Estimates of the human trophic level during the Paleolithic form the basis for many hypotheses and interpretations regarding human evolution and behavior (e.g., Dominguez-Rodrigo et al., 2014a, Hawkes and Coxworth, 2013, Hockett, 2012, Kaplan et al., 2007, Leonard et al., 2007, Mann, 2000, Aiello and Wheeler, 1995, Foley et al., 1991, Speth, 1989, Milton, 1987). Cordain et al.’s (2000) paper, which is based exclusively on an analysis of the ethnographic record, was cited in over 800 articles, many of which discuss various aspects of human behavior during the Paleolithic. Among examples of the use of ethnography-based estimates of human trophic levels in the evolutionary context, one can count competing explanations for humans' extended longevity – the 'grandmother hypothesis' (Hawkes and Coxworth, 2013) and the 'embodied capital hypothesis' (Kaplan et al., 2009) – both of which are based on different assumptions of the relative dietary importance of gathered plant food versus hunted animal food during human evolution. At times, the plant-dominated diet of a single recent hunter-gatherers (HG) group, such as that of the Hadza of Tanzania, is carried across over two million years and several Homo taxonomic units to represent an analog for humans' diet of early Homo in predictions of human evolution, albeit with some words of caution (e.g., Hawkes, 2016, Schnorr et al., 2014). The research may also center around the question of a 'generalist' vs. 'specialist' direction in human evolution (Ungar et al., 2006, Wood, 1992). In such cases, the actual variability of recent HG's Pla-AniR can be argued as being analogous to Paleolithic human conditions (Marlowe, 2005). Pla-AniR estimates also serve in hypotheses about an ideal, evolutionarily compliant, contemporary diet (Cordain et al., 2005, Eaton, 2006).
Following the description of the importance of large prey to Paleolithic humans, we review the prey size decline, globally, and in Africa. Turning to an examination of the validity of ethnographic analogies in predicting human trophic level during the Paleolithic, we review the general reliability of the Pla-AniR ethnographic record and that of two much-cited HG groups, the Hadza and the Ju/'hoansi. We assess to what extent their technology and high plant consumption could be explained by our model as an adaptation to the decline in prey size.
Section snippets
Acquisition of large prey animals during the Pleistocene
Recent analyses of the archaeozoological and paleontological East African record portray H. erectus as a habitual hunter of large prey (Domínguez-Rodrigo and Pickering, 2017, Roach et al., 2018). Preference for large prey animals during the Pleistocene is a conventional interpretation of archaeological assemblages (Bunn and Ezzo, 1993, Surovell and Waguespack, 2009). Large animals, including elephants, are a common feature in African early Pleistocene sites (Bunn and Ezzo, 1993,
Why humans preferred to acquire large prey?
We propose that four factors made large prey, and especially megaherbivores, a primal target of human predation. They are their relatively high fat content, high energetic return, the relatively simple technology employed in their acquisition, and their high biomass density in the landscape. All these aspects may be inter-related. For example, high biomass density and relatively less complex methods of acquisition may be the cause of their higher energetic return. Not tending to escape, which
Changes in the availability of large animals and their ecological implications – Worldwide and Africa
Having recognized the importance of large prey animals to the economy of past hunter-gatherers, we study the dynamics of their availability during the Pleistocene and until the recent ethnographically described past. It is also essential to study the implications of the decline in megaherbivores abundance on the relative environmental availability of plant and animal-sourced foods.
Faith et al. (2019) document, continuously, throughout the Pleistocene, a substantial decline in Africa of size 5
Ethnographic – Paleolithic analogies of Pla-AniR
The use of comparative analogies in archaeology is the subject of a considerable body of literature, reflecting its importance as well as its controversiality (see references in Lyman and O'Brien, 2001). A consensus may be forming that the validity of the analogies should be evaluated on a context by context basis (Currie, 2016). There is, however, a little argument about the basic requirements for a valid analogy as they were already well-phrased by Ascher (1961). Summarizing the opinions of
The association between large prey extinctions and plant food consumption
Our model assigns a dual role for large prey in human survival: relative bioenergetic efficiency and provision of fat to help overcome a physiological constraint on the metabolism of protein. The ceiling on protein consumption explains why HG would be expected to increase the consumption of plants with the decline of the fatter large animals, even though medium and even smaller size animals still provide a higher net energetic return than plants (see Section 3.2). This situation can be
Conclusion
Humans' evolution, like that of any animal, was heavily influenced by the availability of their dietary resources. We have explained a marked reliance of humans on the acquisition of large animals during most of the Pleistocene by the nutritional and bioenergetic implications of the large animals' high fat content, high energetic return, superior biomass density, and the relative incomlexity of their acquisition, compared to the acquisition of smaller fleeting prey. We consequently interpreted
Credit authorship contribution statement
Miki Ben-Dor: Conceptualization, Writing - review & editing, Writing - original draft, Data curation. Ran Barkai: Conceptualization, Writing - review & editing.
Declaration of Competing Interest
The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.
Acknowledgements
We hold a deep sense of gratitude to the editors and the anonymous reviewers, and especially to one of them, who relentlessly led to us to markedly revise a more modest original manuscript to arrive at the present, hopefully much improved one.
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