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date: 09 May 2021

Faunal Analysis in African Archaeologyfree

  • Jessica C. ThompsonJessica C. ThompsonYale University


Faunal analysis (or zooarchaeology) in African archaeology is the identification, analysis, and interpretation of the remains of animal bones recovered from archaeological sites in Africa. Its methods and theoretical underpinnings derive from archaeology, paleontology, biology, and geochemistry, and they extend across all faunal categories. Much of the work in African faunal analysis concerns large-bodied mammalian taxa, but the approach encompasses analysis of fish, shellfish, birds, reptiles, and indeed all animal remains found in association with archaeological sites. The diversity of research encompassed within faunal analysis is also especially high in Africa, where the earliest reported archaeological site is far older than the earliest archaeological site outside of Africa. The extra time depth affords the African archaeological record a wide arena of research questions that are answerable using faunal data. Major themes in African faunal analysis include the origins of unique components of human diet and hunting ability, reconstruction of the transition from hunting and gathering to food production, and analysis of the historical use of animals in trade, exchange, and social status.

Overview of Faunal Analysis in African Archaeology

Faunal analysis (or zooarchaeology) in African archaeology is the identification, analysis, and interpretation of the remains of animal bones recovered from archaeological sites in Africa. It is a core approach in investigations of the African past and is used at any site where information is needed from preserved faunal remains. Faunal analysis is inherently multidisciplinary; its methods and theoretical underpinnings derive from archaeology, paleontology, biology, and geochemistry. A typical faunal assemblage may contain remains from birds, small and large mammals, reptiles, fish, and mollusks. An analyst must be familiar enough with all these groups to separate them, although they may then specialize in one particular taxonomic category (e.g., fish or large mammals). Within their specialization, they must still be familiar with a wide range of possible species that may be present. This differs from bioarchaeology, which is the study of human remains from archaeological contexts, and therefore has a focus on a single species.

Many of the major issues in African faunal analysis concern large-bodied mammalian taxa, as these taxa are most commonly associated with human hunting and pastoral behaviors. These animals are therefore often considered to have the greatest likelihood of informing about past human subsistence, and they are often the most abundant remains at sites. However, work on other faunal categories has steadily increased as researchers investigate the role of other taxa in ancient human economic and social behavior. Furthermore, animal remains that were not directly accumulated by people can offer insight into site formation processes, paleoenvironments, and other useful contextual information.

Africa has the longest archaeological record, and therefore the diversity of archaeological questions that can be addressed with faunal data is greater than what is possible elsewhere. The earliest reported archaeological site in the world is at Lomekwi 3 in Kenya, dating to 3.3 million years ago [Ma] (Harmand et al. 2015). This is far older than the earliest archaeological site outside of Africa, claimed at either 2.4 Ma in India (Gaillard et al. 2016) or 2.1 Ma in China (Zhu et al. 2018), resulting in nearly a million years of additional time in Africa in which faunal analysis can contribute important information about past behavior and ecology. Major themes in African faunal analysis therefore include the origins of unique components of human ancestral diet and hunting ability, in addition to questions common in later time periods. The latter include analyses of early human diet, reconstruction of the transition from hunting and gathering to food production, and evaluation of the historical role of animals in trade, exchange, and social status.

Fundamental Methods in Zooarchaeology

Typically, a faunal analyst will collect data on a range of attributes from the specimens they study. The most fundamental data are what taxa are present (species abundances), what anatomical parts are represented (skeletal element abundances), and what alterations have been made to the remains (fragmentation, fossilization, burning, modification on the surfaces, etc.) (Gifford-Gonzalez 2018). A “faunal assemblage” refers to the animal remains under analysis that derive from a specific location. For example, an assemblage may reference all the remains from a given site, a given layer within a site, or some other coherent set. The most basic description of a faunal assemblage is the NSP, or Number of Specimens. The NISP, or Number of Identified Specimens, is either equal to or a subset of the NSP. The NISP differs from the NSP because what is identified is specific to different questions or collections. For example, the NISP for large mammals in an assemblage could refer to all the specimens identified as derived from large mammals, all the specimens from large mammals that could be identified to skeletal part, or all the specimens from large mammals that could be identified to a certain taxonomic level. Therefore, it is important to define at the start of a study what is considered an “identified specimen.” The ratio of the NSP to NISP can also be a useful and expedient measure of fragmentation, as the more heavily fragmented an assemblage, the higher the number of specimens (NSP), but the lower the number that can be identified (NISP).

NSP and NISP are simple counts, but other measures of faunal abundance are derived through more complex means. The MNE, or Minimum Number of Elements, is the minimum number of a specific skeletal element that is represented in an assemblage. This can be just the general element (e.g., the MNE for femur) or a more specific category (e.g., the MNE for the right femur). Criteria such as body size, ontogenetic age, and provenance may be used to calculate the MNE, as long as these are not specimens that can reasonably be expected to have derived from the same element and the same individual. For example, a left femur from a small adult animal, one from a large juvenile animal, one from a large adult animal, and one from a large adult animal dated to a different time period cannot all come from the same individual. The MNE in this example would be four for left femur. By extension, the MNI (Minimum Number of Individuals) would also be four. If, for the same assemblage, the MNE for right radius is calculated as five, then the MNI would rise to five, while the MNE for left femur would remain at four.

In his seminal ethnoarchaeological work with the Nunamiut in Alaska, Binford (1978) observed that different body parts of a carcass have different nutritional values, and that these translate into butchery and transport decisions on the part of hunters. This, then, has implications for which skeletal parts ultimately become a part of the archaeological record. By assigning a value to these skeletal parts, it is possible to infer how prehistoric hunters selected body parts rich in fat, meat, or other resources to be butchered off-site or brought back to a central place. From this concept, additional zooarchaeological measures can be derived. MAU, the Minimum Animal Units, is the MNE divided by the number of times that element is represented in a skeleton (e.g., the MAU derived from an MNE of six for femur would be three, because the femur occurs twice in the body). The percentage of MAU can then be used to standardize how frequently a given element is represented by dividing the MAU for an element by the MAU for the highest represented element, and multiplying by 100. For example, if the highest MAU for all skeletal parts is three, then the %MAU for femur in the sample above would be 100 percent. This is useful because then the representation of different skeletal parts can be compared against other meaningful measures, such as the Food Utility Index (FUI) (Faith and Gordon 2007), which measures the caloric returns that could have been derived from them when they were still in a fleshed state. In principle, an optimally behaving hunter would maximize their caloric returns by preferentially investing effort in processing and transporting the skeletal parts with the highest caloric values, or FUI.

In addition to analyses of which skeletal parts are present, information about hunting and transport decisions can also be gleaned through analysis of which taxonomic groups are present in an assemblage, and in what abundances. A high diversity of taxa may indicate a broader diet breadth, suggesting that hunters took prey from a range of different habitats and did not specialize in particular taxa. High ubiquity across sites or between temporally constrained layers at the same site may indicate that there were certain staple taxa that were always taken. This could potentially speak to either human decision making or environmental constraints on prey availability, both of which are of interest in reconstructing past behavior. There are a number of ways to quantify diversity and ubiquity in zooarchaeological assemblages, and many of these are drawn explicitly from paleoecological method and theory (Lyman 2008).

The interpretation of abundances of either skeletal parts or taxonomic groups demonstrates how all types of zooarchaeological data must be analyzed and evaluated within a theoretical framework that links these traces to past human and non-human behaviors. Theoretical approaches that have traditionally been applied to primary zooarchaeological data are Optimal Foraging Theory and Middle Range Theory, which connect the present to the past via processes observable in the modern day. These are useful frameworks because faunal data often reflect human subsistence, which lies at the core of human economic systems. However, interpretations derived from zooarchaeological data may also be deployed within theoretical frameworks that more directly address the social and political aspects of human economies (Russell 2011).

Taphonomy and Actualism

Faunal analysis in African archaeology has been especially instrumental in the development of taphonomic method and theory. Taphonomy is the study of what happens to an organism’s remains after death and includes processes that can severely impact what parts survive and ultimately become part of the fossil record. Common taphonomic processes include human butchery, carnivore consumption and scattering of the remains, burial and decomposition, and post-depositional fragmentation or alteration through the actions of trampling animals, wind, water, and microorganisms (Fernandez-Jalvo and Andrews 2016). All of these leave traces on the bones in the form of bone surface modifications (e.g., cut marks from knives, percussion marks from breaking open bones for marrow, tooth marks from carnivores biting the bones), breakage, and burning patterns. They also influence what bones or bone portions are preserved (Marean et al. 1992) and how identifiable a fragment of bone can be to the analyst (Merritt and Davis 2017).

Taphonomy has had a variable role in faunal analysis since its first sporadic application in the 1950s (James and Thompson 2015). In the early part of the 20th century, faunal analysis mainly focused on the production of a list of species that were found in archaeological assemblages. In African contexts, zooarchaeology saw much of its growth at the same time and in the same contexts where the significance of taphonomy was starting to be realized. Thus, many of the key early studies in African zooarchaeology were also taphonomic studies. For example, C. K. Brain (1981) drew extensively on his work observing modern taphonomic processes in the 1960s with the aim of unraveling the processes that had led to the formation of hominin-bearing sites in what later became the World Heritage-designated Cradle of Humankind near Pretoria, South Africa. In his work, Brain challenged the idea that early hominins such as Australopithecus or Paranthropus had killed and accumulated the animals represented in the fossil deposits. Instead, he argued, most bones—including hominin bones—derived from the kills of contemporaneous predators.

Lewis Binford also produced a work in the same year (Binford 1981) that used taphonomy to challenge prevailing views about African faunal assemblages. He embedded this work in Middle Range Theory, which deploys the concept of uniformitarianism in the interpretation of archaeological sites. Uniformitarian assumptions, as in geology, are that processes happening in the modern day are the same as those that happened in the past. Because these processes leave traces that may preserve in the archaeological record, one can observe modern phenomena to identify the traces unique to them. In so doing, analysts can use preserved traces to reconstruct what happened during a time that is not directly observable. For example, a carnivore may be observed to bite a bone and leave behind distinctive tooth marks on the bone surface. When similar marks are found in the fossil record, an interpretation can be more confidently made that the bone was subjected to carnivore modification in the past.

These middle-range linkages are often made through actualistic research, or observations of present-day phenomena, that is designed to connect an agent, effector, and actor to a given trace (Gifford-Gonzales 1991). Experiments involve the direct manipulation of variables, while naturalistic observations rely on passively observing processes as they occur in a natural setting (Pobiner 2015). This work continues to be foundational to zooarchaeologists working especially in the earliest time periods, where traces of past behavior may be subtle and both hominins and other processes likely contributed to the formation of a faunal assemblage. However, analysts have begun to scrutinize two underlying assumptions of Middle Range Theory in zooarchaeology: (1) that behavioral and biomechanical processes have remained the same, or at least comparable and knowable, over the entire span of human prehistory; and (2) that traces in the archaeological record have a single, discernible cause. Thus, many of the debates that have dominated the zooarchaeology of human origins center around the problem of equifinality, where multiple different causal chains can produce the same archaeological trace (Domínguez-Rodrigo, Egeland, and Pickering 2007).

Early Hominin Diets

At the earliest end of the time spectrum, researchers use faunal analysis in African archaeology to seek to understand the origins of major changes in hominin subsistence (Thompson et al. 2019). Humans and some human ancestors are the only primates to consume animals of the same or larger body size than themselves, and this change in diet facilitated a number of other key changes in human biological evolution, such as increased brain and body size around 1.8 Ma (Antón, Potts, and Aiello 2014). As our ancestors moved into a more carnivorous niche and came into greater competition with other predators, they fundamentally shifted the way they interacted with other organisms in their ancestral environments (Faurby, Daniele, Lars, and Alexandre, 2020). Because bones and teeth preserve better than other plant or animal parts, these interactions are more likely to leave traces in the zooarchaeological record than elsewhere.

The earliest zooarchaeological evidence for stone tool-assisted butchery of large mammals derives from the DIK-55 site at Dikika, in the Afar region of Ethiopia (McPherron et al. 2010). The evidence consists of two specimens of fossil bone, dated to ~3.4 Ma, that bear traces consistent with the application of a sharp-edged stone tool. If hominins were using stone tools to access large mammal resources as early as 3.4 Ma, then this implies a very different kind of ecology from that typically inferred for Australopithecus or Kenyanthropus, the two hominin genera from this time period. It also means that archaeologists should be consistently collecting and examining broken fragments of potentially butchered fossil bone from the landscapes that contain early hominin fossils, even if flaked stone tools themselves are rare or absent. This is a very different strategy from that which has been historically employed, which is one in which the most complete—and therefore taxonomically identifiable—elements have been collected. It also shifts the emphasis on finding the earliest traces of hominin behavior away from stone tools and squarely into the realm of zooarchaeology. However, the DIK-55 traces have been contested as being caused by other agency, such as the hooves of ungulates trampling them in a rocky substrate (Domínguez-Rodrigo, Pickering, and Bunn 2011) or the sharp edges of crocodile teeth (Sahle, El Zaatari, and White 2017). If either scenario is true, then this modifies the significance of zooarchaeology at these early time periods from one of empirical urgency to one of theoretical possibility.

The considerable controversy over the DIK-55 specimens illustrates the power of equifinality to shape interpretations of early hominin behavior, justify methodological decisions, and undermine support for major narratives of human origins. It also highlights theoretical and methodological problems that have lain somewhat dormant in zooarchaeology. For example, most bone surface modifications on fossils (e.g., stone tool cut marks, carnivore tooth marks, hammerstone percussion marks, or ungulate trampling marks) are identified by an analyst through the application of expert knowledge. An expert learns the criteria that apparently separate different kinds of marks from one another, typically through working with modern actualistic assemblages, and then applies those criteria to the fossil record to make a diagnosis. That diagnosis is often binary, that is, a mark “is” or “is not” a cut mark. Newer approaches seek to remove the analyst from such a direct role in the decision-making process through machine learning (Domínguez-Rodrigo and Baquedano 2018) or by assigning a probability of agency to a given mark rather than a firm diagnosis (Harris et al. 2017). Additional methods of dealing with equifinality include the development of more objective means of measuring mark attributes (Pante et al. 2017), including 3D shape attributes (Otárola-Castillo et al. 2018).

The nature of the DIK-55 controversy in many ways echoes another ongoing debate, deemed the “hunting versus scavenging” debate (Domínguez-Rodrigo 2002). This debate began with the taphonomic challenges posed by Binford (1981), who drew from his ethnoarchaeological work to observe that bones are rarely left unmodified and undisturbed after they are discarded. He applied this logic to early hominin sites such as Olduvai Gorge, Tanzania, where “living floors” of bones and stone artifacts had been reported in association with hominin remains. He argued that a better explanation was that these early sites were palimpsests of hominin and carnivore activity, and that most of the bone accumulations were the result of hominins scavenging after carnivore kills. The interpretation was challenged by Bunn and Kroll (1986), who used both cut marks and skeletal part abundances at the FLK “Zinjanthropus” site at Olduvai Gorge, Tanzania, to argue that meat-bearing bones were both abundant and frequently bore cut marks at the site. Therefore, hominins likely had primary access.

This result was later contested by Blumenschine (1988), who used actualistic data to show that when carnivores had first access to carcasses, they left many tooth marks as they broke open bones. Hominins that scavenged after the carnivores would not leave many marks because most of the nutrients were already gone and further processing would not be useful. He argued that data from the FLK “Zinj” site showed a pattern consistent with carnivores having first access. This was further challenged on the basis of equifinality by Domínguez-Rodrigo and Barba (2006), who proposed that many of the purported carnivore tooth marks were actually fungal and bacterial etchings. Thus, the frequency of carnivore tooth marks became lowered relative to hominin marks, implying the reverse scenario: hominins had first access. The debate has continued, with a fresh analysis of FLK “Zinj” indicating that there is abundant evidence for primary hominin access, but that it is especially apparent on smaller-bodied mammals (Parkinson 2018). This adds additional nuance to information about early hominin subsistence strategies and corroborates data from similar-aged sites in Kenya (Ferraro et al. 2013).

Origins of Modern Subsistence and Social Behavior

In addition to being the home of our lineage, Africa also contains the earliest sites produced by members of our own species, Homo sapiens. Faunal analysis has been deployed extensively as a way to understand two key aspects of sites dating between ~500 thousand and 50 thousand years ago (ka): what environments were like at the time of early modern human evolution, and when our species first achieved its ecological dominance. Modern hunter-gatherers use a number of complex technologies and social behaviors in their daily foraging and hunting tasks, and faunal analysis is useful for understanding when these behaviors first emerged. Dietary shifts may also have been instrumental in driving some of this technological change, especially as hunting became more important in our lineage (O’Driscoll and Thompson 2018).

Some of the earliest efforts to understand subsistence at the origin of our species included the use of taxonomic lists to identify which animals were regularly hunted and which were not. Klein (1976) suggested that early humans who were anatomically modern nonetheless did not have fully modern hunting abilities, and thus the faunal assemblage from Klasies River Mouth, a key site along the South African coast dating between about 130 thousand and 44 thousand years ago, contained a paucity of the most dangerous animals that existed on the landscape (specifically, bushpig and buffalo). Binford (1984) contested this interpretation with an even more extreme perspective, arguing on the basis of a lack of meat-bearing bones that early humans were only the most marginal of scavengers. In spite of his early work on taphonomy, he based this interpretation on the fundamental assumption that the abundances of different kinds of faunal remains are a direct reflection of the behavior of the accumulator. He recognized that many different actors may actually accumulate faunal remains (in this case carnivores and humans), but there was little work at the time that showed how assemblages were subsequently modified from their original depositional states.

It was not until more than a decade later that understanding density-mediated attrition (or destruction) would become critical for interpreting zooarchaeological assemblages (Lam et al. 2003). This concept recognizes that certain anatomical parts will preserve more commonly at archaeological sites than others as a function of their bone mineral density, even if the entire animal was originally present in the deposits. Thus, neither species abundances nor skeletal element abundances should be taken as an unfiltered view of what was once present at the site. Long bone shaft fragments, teeth, and foot elements are particularly resistant to density-mediated attrition, but long bone shafts are less taxonomically informative than the long bone ends. Therefore, abundances of different anatomical parts will vary depending on if they are calculated based on shafts or on ends. If most calculations such as the MNE or its derivatives are done on ends, which are rarely preserved, then head and foot elements will appear to be more common. It was based on this assumption, the low %MAU of high-utility elements, that Binford inferred people had scavenged from carcasses where the meat-bearing parts had already been consumed by carnivores. However, at Klasies River Mouth, most shaft fragments from meat-bearing long bones had been discarded, thus leaving the false impression that the assemblage contained mostly head and foot parts and not meat-bearing parts (Marean and Kim 1998).

Although the concept of density-mediated attrition is often explicitly applied to mammalian remains, differential preservation can also affect shells or other faunal components, thus causing entire aspects of assemblages to be missing or altered from their original composition. At Die Kelders Cave 1, also on the South African coast, massive dissolution of shells, only detectable at a microscopic level, removed entire faunal categories from the site and caused slumping of the overlying sediments (Goldberg 2000). Research bias has also served to render certain faunal components “invisible.” For example, Blombos Cave, an important early modern human site on the southern African coast with deposits dating between 125 and 75 ka, contains high quantities of tortoise and small mammal remains relative to large mammal remains. However, with research focused mainly on early hunting abilities, these components had been largely ignored (Thompson and Henshilwood 2014). If a gendered division of labor existed among these early foragers in a similar way to its prevalence among modern groups, then this focus on large hunted mammal remains places undue emphasis on the subsistence activities of men rather than the diverse and cooperative strategies that are in place with modern people.

There is now broad recognition that early Homo sapiens sites in Africa contain faunal assemblages that attest to fully capable hunting and foraging abilities. In some cases, it is the smallest and least dangerous animals that have become most informative. For example, at Sibudu Cave on the eastern coast of South Africa, abundant small antelope may have been captured through nets, snares, and other complex technologies (Wadley 2010). At Die Kelders Cave 1, patterns in cut marks on small mammals suggest that they were exploited not only for meat, but also for their skins (Armstrong 2016). Along both the northern (Chakroun et al. 2017) and southern (Marean 2016) coasts of Africa, marine shell from archaeological sites attests to widespread incorporation of coastal ecotones into early human diets during the Late Pleistocene, which started at 128 ka. This foraging behavior is not present in earlier hominins and can require significant scheduling and ecological mapping in order to be a successful strategy. Thus, faunal research in this time period has shifted away from understanding early hunting behavior and toward identifying those aspects of an assemblage that can inform about innovation, within-group agency, between-group interactions, and diversity in ancient subsistence.

This emerging trend has been aided by increased work on the social aspects of early human behavior that can be revealed through faunal analysis. After about 40,000 years ago, faunal remains were often modified into tools or ornaments. The African continent contains most of the oldest evidence for this kind of innovation in organic technology. This has been used to document increased complexity in early technological and social systems and suggests continuity across large timescales (d’Errico et al. 2012). Bone harpoons at Katanda, in the Democratic Republic of Congo, have been dated to ~95,000 years ago (Yellen et al. 1995), and bones sharpened into knives occur in the northern African record by 90,000 years ago (Bouzouggar et al. 2018). In southern Africa, bone technology also includes points, which indicates an elaboration of hunting technology by at least 75,000 years ago (d’Errico, Backwell, and Wadley 2012).

Faunal remains that have been decorated or shaped into ornaments also offer one of the best opportunities for preservation of evidence for early symbolic behavior. Perforated Nassarius shell beads have been found in both the northern and southern African records, sometimes with evidence for deliberate color selection or modification (Bar-Yosef Mayer 2015). Also in South Africa, incised ostrich eggshell fragments dated to about 60,000 years ago suggest that functional objects such as water containers may have also held symbolic value early in their history of use (Texier et al. 2013). Deliberately shaped beads made from ostrich eggshell or land snail do not occur until later in the record, although directly dated ostrich eggshell beads and bead preforms from multiple sites in eastern Africa date to at least 50,000 years ago (Miller and Willoughby 2014). Because this is very close to the limits of radiocarbon dating, ostrich eggshell beads could in principle be as old in eastern Africa as engraved ostrich eggshell is in southern Africa. However, because beads are very small objects, it is difficult to detect if they have been mixed into sediments from different time periods. In addition to dating them, ongoing work is needed to elucidate how they were made and what taphonomic factors can alter them. This is a rich area of research, as different approaches to bead manufacture may offer insight into cultural differences between human groups over space and time.

Resource Intensification

Analyses of African faunal assemblages dating to the end of the Pleistocene (~10.5 ka) and early Holocene most frequently deal with the evolution and diversification of hunter-gatherer economies. Across Africa, hunter-gatherers were beginning to make more intensive use of the same resources their ancestors had already been exploiting. In the realm of faunal analysis, these resources commonly took the form of aquatic foods, such as fish, that were present but uncommon during earlier time periods (Prendergast and Beyin 2018). In some cases, these changes translated to major accumulations of middens where a single taxon or series of taxa dominate. The processes underlying these transformations are often lumped under the general rubric of “intensification” (Morgan 2015).

The concept of intensification can be painted using a wide brush, but it is useful for examining how later hunters and gatherers dealt with the changing abundances of resources that came with major environmental shifts such as the Last Glacial Maximum ~24 ka or the end of the Ice Ages ~11.5 ka. Most researchers place at least some of the responsibility for Holocene intensification on the recursive impacts of population expansion. Shell middens began to grow in size and quantity especially along the southern and northern coasts of Africa, which share a similar Mediterranean-type environment. In South Africa, shells from these middens are on average smaller than shells found in deposits from earlier time periods, suggesting that increased numbers of foragers had been placing more intense predation pressure on these sedentary resources (Klein and Steele 2013). Similarly, tortoise sizes also decreased between the transition from the Middle to Later Stone Ages, c. ~24 ka, implying increased harvesting of individuals before they grew to their largest potential sizes (Henshilwood et al. 2001).

Along with this inferred increase in population size, Holocene hunter-gatherers in many areas reduced their mobility and began to intensively use resources in restricted areas. The South African coast has some of the earliest global evidence for shellfish consumption, going back into the Middle Pleistocene, but large shell middens do not begin to accumulate until the Holocene (Kyriacou et al. 2015). The largest date to within the last three thousand years, and dietary and provenance isotopes of associated skeletons show that these were hunter-gatherers who maintained separate geographic territories (Sealy 2016). Faunal data are a critical part of this interpretation. For example, at Lambert’s Bay on the western coast of South Africa, there is a notable shift in marine shell species and their abundances relative to terrestrial fauna, as hunter-gatherers during the “mega-midden phase” began to focus even more on marine resources and small antelope and less on large mobile game (Jerardino 2010).

Faunal data from other regions provide an important line of evidence that corroborates a trend of increasingly localized subsistence strategies over the course of the Holocene and more intensive and repeated occupation of the same sites. For example, at Rifle Range in Somalia, foragers began to decrease their emphasis on large, mobile herding mammals as they moved into the Holocene and instead acquired most of their hunted resources from small antelope (Jones, Brandt, and Marshall 2018). These trends of increased population and intensification around key geographic features is particularly pronounced in parts of Africa with highly productive lacustrine resources, such as Lake Turkana and Lake Victoria in East Africa (Prendergast and Beyin 2018), or Paleolake Gobero in what is now the Sahara Desert (Sereno et al. 2008). This increased focus on intensifying subsistence around productive resource patches, such as lakes, rivers, and inselbergs, had a profound effect on intersocial relationships, and it is in the context of more sedentary hunter-gatherers where the first substantial evidence for interpersonal violence appears in the bioarchaeological record (Lahr et al. 2016). As with earlier time periods, ornaments made from faunal remains may be informative about the nature of social relationships, either through data on technological and stylistic approaches or by using provenance isotopes to track their ancient movements across the larger landscape.

In some cases, Early Holocene forager-hunter-fishers created substantial shell middens around lakes and river systems, and their clear reliance on aquatic resources led early researchers to describe them and their Late Pleistocene counterparts under the general cultural umbrella of the “African aqualithic” (Sutton 1977). This concept was an explicit attempt to explain the transition to food production in Africa via a pathway unique from other parts of the world, but universal across the African continent (Holl 2005). The concept of an aqualithic has since been replaced with a more nuanced set of research that emphasizes the local roots of phenomena such as intensive fishing economies rather than framing them as simple precursors to food production. Thus, since the first comprehensive zooarchaeological analysis of prehistoric fishing in East Africa conducted by Stewart (1989), faunal analysis has been a central element to critical examination of the concept of an African aqualithic.

In some cases, however, intensification can be interpreted as a preadaptation that facilitated later changes that would occur with food production. For example, in the early Holocene of North Africa, wild caprines (Barbary sheep) appear to have been the specialized hunting targets. By ~8500–7500 bp, they were being corralled in caves in the Libyan Sahara, accumulating dung and potentially kept as a reserve food source for controlled slaughter (di Lernia 2001). Similarly, regions of northern and eastern Africa that had early evidence of intensification were also some of the first to host interactions between hunter-gatherers and pastoralists. This was usually preceded by several thousand years of hunting and gathering economies that already incorporated a moderate degree of delayed return (Dale and Ashley 2010). Some of these economies, such as that associated with the Early Holocene Kansyore pottery tradition around Lake Victoria or the Capsian of the Mahgreb, used ceramics but not domesticated fauna. This combination of faunal resource intensification, experience with delayed-return subsistence, and use of technological elements such as ceramics may have predisposed certain groups to the adoption of animal husbandry when it spread into the region (Mulazzani et al. 2016).

Origins and Spread of Pastoralism

In the later part of the Holocene, the African continent witnessed a major change in human subsistence and land use patterns with the rise and expansion of food production. However, unlike in most other parts of the world, African food production began with livestock management and did not appear as a package with or after agriculture (Gifford-Gonzalez and Hanotte 2013). Faunal analysis has played a pivotal role in debates about the origins and spread of pastoralism, using lines of evidence such as the morphology of animal bones, isotopes from animals and soils where they were found, DNA from modern animals, and ancient DNA.

There is continued ambiguity surrounding whether cattle domestication occurred independently within the confines of Africa or if it spread from the Near East via the Nile Valley. The earliest morphologically distinctive cattle bones were recovered in the 1970s from Nabta Playa and Bir Kiseiba in the Egyptian Sahara, and genetic divergence estimates from modern cattle suggest an independent origin of African and Near Eastern populations (Hanotte et al. 2002). However, there is a large gap in archaeological data between this putative early domestication in North Africa and what is known as the Pastoral Neolithic of eastern Africa, which spread across the Lake Victoria Basin from ~5000–1200bp (Grillo et al. 2018).

By ~3000 bp, a mosaic of subsistence strategies had developed in the Lake Victoria Basin, including specialized pastoralism. At this point, there was an apparent pause in the spread of cattle south of this region until after ~2000bp. Gifford-Gonzalez (2001) proposed that there could have been an ecological barrier posed by the brushy habitat that houses tsetse fly, which carries trypanosomiasis (sleeping sickness). As this disease can be communicated to livestock, while wild game have developed more natural resistance, it could pose a significant problem to herders seeking to take their livestock south. However, analysis of prehistoric soil isotopes shows that there were large tracts of grasslands around Lake Victoria that could have facilitated the spread of pastoralism because the brush comprising prime tsetse fly habitat was not as dense as previously inferred (Chritz et al. 2015). This is a problem to which ancient DNA of early livestock could potentially contribute, as it could pinpoint the timing of the emergence of advantageous alleles that living cattle breeds exhibit with respect to a number of potential pathogens—including trypanosomes (Kim et al. 2017).

Once pastoralism did spread further south of the Lake Victoria Basin, it merged with other innovations in subsistence and technology, namely, agriculture and metalworking. This accompanied a massive reconfiguration of human populations known as the Bantu expansion, so-named because of the shared language family to which most living populations in tropical and southern Africa belong. This cultural “package” reached the southeastern tip of South Africa by ~1300 years ago. Modern and ancient DNA as well as artifactual and human skeletal evidence all indicate that this expansion was both cultural and biological in nature, involving the physical movement of people with a West African origin. Faunal analysis helps to not only identify which remains were found at Iron Age and historical sites, but also to investigate the degree to which wild fauna may have continued to play a role in subsistence even within an economy that was reliant on food production (Badenhorst 2015).

With most of the emphasis on cattle domestication, additional layers of complexity exist regarding whether bovine and caprine domestication took place together or at different times and places (di Lernia 2013). Caprines (sheep and goats) are the first domesticates to have arrived in southwestern Africa, sparking substantial debate about whether the process was predominately driven by cultural diffusion (Jerardino et al. 2014) or by the migration of people into the region, bringing their livestock with them (Smith 2014). Faunal data have been integral to tracking the route of early domesticates along the southwestern coast of Africa, for example, at the site of Leopard Cave in Namibia, where remains that are morphologically consistent with sheep have been directly dated to ~2200 cal bp (Pleurdeau et al. 2012). However, some purported early sheep remains from other parts of southern Africa have recently been challenged as such through advances in ancient DNA (Horsburgh, Orton, and Klein 2016). This has led to heated methodological debates about the utility of conventional morphology-based zooarchaeology versus the use of more recently developed molecular methods (Scott and Plug 2016).

In addition to documenting the origins and spread of both bovines and caprines, there remains much work to be done on understanding the use of secondary products, such as milk. In most cases, this appears to lag behind the initial domestication of bovines by at least 1000 years (Dunne et al. 2018). In some regions, milk, rather than meat, may have been one of the primary drivers of pastoralism (Lombard and Parsons 2015). Lines of evidence include rock art depictions of cattle with full udders, ceramic vessels with residues of milk products, and zooarchaeological data showing differential growth of males and females (implying castration), and the timing of slaughter of male versus female animals. Genetic data from modern people also show the strongest selection for lactase persistence among populations that live in regions with the longest zooarchaeological evidence for pastoralism, but is also present in mixed-ancestry Khoe groups in southern Africa (Breton et al. 2014). This offers strong evidence that the spread of pastoralism into southwestern Africa involved the migration of people with their livestock into the region, and that its rapid dissemination was facilitated by both cultural and biological processes.

Zooarchaeologists who specialize in the African Neolithic are increasingly also becoming engaged with multidisciplinary work that reveals ways in which a long history of pastoralism in Africa has had a lasting impact on environments, and even climates. In the Sahel region south of the Sahara, pastoralism has been implicated as causing changes in vegetation patterns that led to local shifts in precipitation and increasing desertification (Wright 2017). In the grasslands of eastern Africa, the practice of kraaling animals in concentrated places has led to the development of nutrient-rich patches that in turn structure wild plant and animal communities (Marshall et al. 2018). Thus, pastoralism has been an embedded part of the fabric of African ecosystems for many thousands of years, as revealed by a combination of faunal data and paleoclimatic and paleoenvironmental data derived from other proxies. It remains today a key element of African economies, and there is potential to learn from archaeological data how it can best be managed in a world where—as in the past—human populations and their environmental contexts rarely remained stable and unchanging.

Historical Economies and Networks

Much can be revealed about ancient trade, exchange, and identity based on what people ate in the past. This is a particularly important area of faunal research in Late Iron Age and historical contexts, where preservation is typically better than at older sites and there is a more diverse array of material culture and contemporaneous accounts upon which to draw. However, even in these cases, there is an under-recognized place for taphonomy. For example, the Castle of Good Hope in Cape Town, South Africa, is an historical fort built in the 17th century by the Dutch East India Company. The faunal assemblage from its “Granary” location was originally interpreted as the material residue of the meals of enslaved people based on what was called the “slave pattern” of faunal representation. In this pattern, meat-poor bones, bones from older animals, and heavily fragmented remains seemed to indicate access to mainly undesirable parts of animals (Hall 1999). A reanalysis by Heinrich (2012) took a much more explicit taphonomic approach and found that a better explanation for the fragmentation and skeletal part abundance patterns was post-depositional alteration of the assemblage. There were in fact many meaty bones, and some faunal components, such as bones from wild ungulates, that were more consistent with the diets of high-status individuals in the compound. Upon closer inspection of both site formation and historical documentation, interpretation of the Granary assemblage changed from one in which it exclusively contained the remains of slave diets to one in which it was the mixed residue of multiple social elements within the fort.

Osteometric and isotopic work has also been used to reveal nuances in faunal assemblage formation and meaning. At Jenné-jeno in Mali’s Inland Niger Delta, which was occupied from ~250 bce to 1400ce, subsistence specialization has been proposed as part of an explanation for the appearance of clusters of urban centers without the presence of an obvious single seat of authority (McIntosh 2005). Under this model, subsistence specialization and complex trade relationships would provide the buffer necessary for the maintenance of elaborate skill specializations in a highly variable environment. However, data from the faunal remains show more diverse subsistence patterns within each site than would be expected (Stone 2018). The majority of livestock were small and local rather than partitioned into local sheep herding and large, transhumant cattle-herding specialists. When larger cattle and more variation in provenance began to appear, it was at the end of the tenure of Jenné-jeno rather than being an essential part of its rise and organization.

Other aspects of the economy are apparent in the use of faunal remains that were not the immediate products of subsistence. At Diouboye, a medieval settlement along the Falémé River in eastern Senegal in western Africa, zooarchaeological data show the hunting of wild animals that have thick skins or ivory, and carnivores (Dueppen and Gokee 2014). The underlying economy of leather and animal skin production is rendered visible through faunal analysis and shows which places emphasized hunting for secondary products rather than mainly for subsistence. Similarly, at early farming sites in KwaZulu Natal in South Africa, secondary ivory products were confirmed to derive from elephants, even though non-ivory hippopotamus remains had been found at the sites (Coutu et al. 2016). Moreover, geochemical sourcing of the ivory showed that a range of localities formed the sources of elephant ivory and that these were specific to each site. Thus, interior sources of ivory ultimately destined for the Indian Ocean trade could be identified as early as the 7th to 10th centuries ce.

Faunal remains have long been used as indicators of cultural separation between people, be it via social hierarchy within the same community (as in the example with slave diets) or via separation between urban centers or even greater regional areas. In some cases, the degree of reliance on wild versus domesticated fauna can be an indicator of social and trade relationships as much as it is an indicator of economic organization. Substantial dietary diversity at two sites in Zanzibar that date to the later 1st millennium ce show that not all communities made a clear and decisive shift to rely exclusively on domesticated fauna. The smaller site, Fukuchani, shows substantial economic dependency on wild fauna, while the larger site, Unguja Ukuu, emphasizes domesticates such as caprines (Prendergast et al. 2017). Sites such as these also contain early remains of non-native fauna, such as black rats, chickens, and cats. This speaks to broader patterns of trade and exchange across the greater Indian Ocean region. However, neither site shows reliance on deep-sea species of fish that would indicate a larger maritime adaptation.

The concept of “maritimity” along the Swahili Coast is an emerging area of debate that draws heavily on zooarchaeological data. The degree to which communities engaged with oceanic resources carries with it implications about the extent to which occupants of the Swahili Coast were directly involved in Indian Ocean networks or instead were more passive agents in the societal changes wrought by these networks. It is also an important concept for understanding the degree of social and economic connection between coastal and interior communities. Fleischer et al. (2015) describe the advent of a fully maritime adaptation after the 1st millennium ce, including deep-sea fishing, sailing, increased permanency of occupation along coastlines, and construction of architectural features specifically designed to aid in maritime activities. Countering this, Ichumbaki (2015) notes that people along the Swahili coast were strongly engaged with marine resources back into the Pleistocene and that therefore the definition of maritimity should be much broader and consider the local roots of later expansions into this niche. Similarly, Kusimba and Walz (2018) critique the notion that maritime adaptations of the Swahili Coast emerged with little or no influence from the interior. Much of this debate depends on faunal data sets, which empirically show the extent to which populations relied on resources from the ocean and how economic and social relationships were negotiated between the interior and the coast.

The spread of introduced fauna into various parts of Africa has similar implications for the extent and timing of broader regional interactions. As with evaluating evidence for the spread of pastoralism, it is an issue where relatively recent molecular techniques and more traditional morphological evaluations of faunal remains are beginning to intersect. For example, one of the most economically important introduced taxa is the domestic chicken, which has its earliest morphological date in Ethiopia between 820 and 595 cal bce (Woldekiros and D’Andrea 2017). Morphological and molecular analysis of chicken remains from East Africa support a scenario of independent introductions of this taxon into different parts of Africa and at different times (Prendergast et al. 2017). This empowers faunal data to play a role in both hypothesis formation and hypothesis testing with respect to major questions about ancient human interactions within Africa.

Summary of Major Issues in African Faunal Analysis

Faunal analysis in African archaeology represents a challenging and diverse landscape of approaches that are structured around a series of fundamental problems in African archaeology. In many cases, the theoretical and methodological frameworks are applied in ways that reflect the quality of data preservation. For example, in earlier time periods, there is strong emphasis on taphonomy, site formation processes, and quantification of skeletal parts. These are basic components of the zooarchaeological toolkit, but have become especially salient in the context of very ancient sites because, in some cases, there is uncertainty as to the degree of hominin involvement in their formation—or if they are even archaeological sites at all. However, this treatment of taphonomy as mainly having relevance to older time periods is one that is changing, as evidenced by new insights that have been gleaned from taphonomic analysis of faunal assemblages found at historical sites.

Similarly, use of fauna to inform about social relationships is no longer restricted to historical or near-historical periods. There is much emerging research that deals with this problem in hunter-gatherer contexts and takes a long-term view of human–animal relationships as a process rather than a series of distinctive stages. An important element to all zooarchaeology is the establishment of taxonomic and anatomical part lists, as these can reveal changes in prey choice, hunting or collecting strategies, or—later in time—the development of pastoral economies. However, the development of such lists is nearly as fraught as the reconstruction of early site formation processes. Some sites have heavily fragmented or poorly preserved fauna that can be difficult to identify, which can result in very low sample sizes of identifiable remains. There is a fresh realization that taphonomic processes can affect some categories of fauna differentially, thus causing some to appear to be more abundant than others. In other cases, there is contention over the accuracy of identifications themselves. Advances in molecular approaches are beginning to play a stronger role in resolving the problem of identification.

Zooarchaeology by Mass Spectrometry (ZooMS) is a collagen fingerprinting technique that can characterize the basic taxonomic category of a given bone, shell, or tooth sample at relatively low cost (Buckley 2018). In addition to assisting with identification of heavily fragmented remains, it has been applied to the identification of organic ornaments, such as ivory from archaeological contexts, in order to elucidate ancient patterns of trade, exchange, social hierarchies, and symbolic incorporation of different species into utilitarian objects (Bradfield et al. 2019). Along with ancient DNA analysis, it has also been used to confirm the identifications of introduced taxa before they are directly dated or to revisit sites that are key anchor points in the reconstruction of the ancient movement of domesticates. These approaches build on a long-established tradition of using faunal remains as sources for isotopic proxies of ancient environment and diet, but they remain limited to more recent time periods where organic preservation is still viable.

In some aspects, such as its emphasis on taphonomy, African zooarchaeology has made significant contributions to global zooarchaeology. In others, such as the movement of early domesticates—introduced and native—much basic work remains. As with other aspects of African archaeology, there has been relatively little research done across a vast continent that houses the longest record of human occupation. There simply needs to be more primary empirical work before the nuances of major issues in African zooarchaeology become further resolved. The fundamental shape of major shifts in human diet, economic systems, and social networks is now established, but the details remain fuzzy. Poor age constraint, uneven geographic coverage, and poor molecular preservation are all obstacles that remain to be overcome, and they all must begin with more basic research. Because faunal remains are a common component of any archaeological site with organic preservation, zooarchaeology is well-positioned to lead the way in filling these gaps.

Further Reading

  • Binford, L. R. 1981. Bones: Ancient Men and Modern Myths. New York: Academic Press.
  • Brain, C. K. 1981. The Hunters or the Hunted? An Introduction to African Cave Taphonomy. Chicago: University of Chicago Press.
  • Gifford-Gonzales, D. 1991. “Bones Are Not Enough: Analogues, Knowledge, and Interpretive Strategies in Zooarchaeology.” Journal of Anthropological Archaeology 10: 215–254.
  • Gifford-Gonzalez, Diane. 2018. An Introduction to Zooarchaeology. Cham: Springer International.
  • Giovas, C. M., and M. J. LeFebvre, eds. 2018. Zooarchaeology in Practice: Case Studies in Methodology and Interpretation in Archaeofaunal Analysis. Cham: Springer International.
  • Horsburgh, K. Ann, Jayson Orton, and Richard G. Klein. 2016. “Beware the Springbok in Sheep’s Clothing: How Secure Are the Faunal Identifications Upon Which We Build Our Models?” African Archaeological Review 33 (4): 353–361.
  • James, Emma C., and Jessica C. Thompson. 2015. “On Bad Terms: Problems and Solutions within Zooarchaeological Bone Surface Modification Studies.” Environmental Archaeology 20 (1): 89–103.
  • Kusimba, Chapurukha M., and Jonathan R. Walz. 2018. “When Did the Swahili Become Maritime?: A Reply to Fleisher et al. (2015), and to the Resurgence of Maritime Myopia in the Archaeology of the East African Coast.” American Anthropologist 120 (3): 429–443.
  • Lyman, R. L. 2008. Quantitative Paleozoology. Cambridge, U.K.: Cambridge University Press.
  • Marean, Curtis W. 2016. “The Transition to Foraging for Dense and Predictable Resources and Its Impact on the Evolution of Modern Humans.” Philosophical Transactions of the Royal Society B 371 (1698): 20150239.
  • Marshall, Fiona, Rachel E. B. Reid, Steven Goldstein, Michael Storozum, Andrew Wreschnig, Lorraine Hu, Purity Kiura, Ruth Shahack-Gross, and Stanley H. Ambrose. 2018. “Ancient Herders Enriched and Restructured African Grasslands.” Nature 561: 387–390.
  • Mitchell, Peter, and Paul Lane. 2013. The Oxford Handbook of African Archaeology. Oxford: Oxford University Press.
  • Morgan, Christopher. 2015. “Is It Intensification Yet? Current Archaeological Perspectives on the Evolution of Hunter-Gatherer Economies.” Journal of Archaeological Research 23 (2): 163–213.
  • Russell, N. 2011. Social Zooarchaeology: Humans and Animals in Prehistory. Cambridge, U.K.: Cambridge University Press.
  • Stahl, A. 2005. African Archaeology: A Critical Introduction, edited by A. Stahl. Oxford: Blackwell.


  • Antón, Susan C., Richard Potts, and Leslie C. Aiello. 2014. “Evolution of Early Homo: An Integrated Biological Perspective.” Science 345 (6192): 1236828.
  • Armstrong, Aaron. 2016. “Small Mammal Utilization by Middle Stone Age Humans at Die Kelders Cave 1 and Pinnacle Point Site 5–6, Western Cape Province, South Africa.” Journal of Human Evolution 101: 17–44.
  • Badenhorst, Shaw. 2015. “Intensive Hunting during the Iron Age of Southern Africa.” Environmental Archaeology 20 (1): 41–51.
  • Bar-Yosef Mayer, Daniella E. 2015. “Nassarius Shells: Preferred Beads of the Paleolithic.” Quaternary International 390: 79–84.
  • Binford, L. R. 1978. Nunamiut Ethnoarchaeology. New York: Academic Press.
  • Binford, L. R. 1981. Bones: Ancient Men and Modern Myths. New York: Academic Press.
  • Binford, L. R. 1984. The Faunal Remains from Klasies River Mouth. New York: Academic Press.
  • Blumenschine, R. J. 1988. “An Experimental Model of the Timing of Hominid and Carnivore Influence on Archaeological Bone Assemblages.” Journal of Archaeological Science 15: 483–502.
  • Bouzouggar, Abdeljalil, Louise T. Humphrey, Nick Barton, Simon A. Parfitt, Laine Clark Balzan, Jean-Luc Schwenninger, Mohammed Abdeljalil El Hajraoui, Roland Nespoulet, and Silvia M. Bello. 2018. “90,000 Year-Old Specialized Bone Technology in the Aterian Middle Stone Age of North Africa.” PloS One 13 (10): e0202021.
  • Bradfield, Justin, Tim Forssman, Luke Spindler, and Annie R. Antonites. 2019. “Identifying the Animal Species Used to Manufacture Bone Arrowheads in South Africa.” Archaeological and Anthropological Sciences 11 (6): 2419–2434.
  • Brain, C. K. 1981. The Hunters or the Hunted? An Introduction to African Cave Taphonomy. Chicago: University of Chicago Press.
  • Breton, Gwenna, Carina M. Schlebusch, Marlize Lombard, Per Sjödin, Himla Soodyall, and Mattias Jakobsson. 2014. “Lactase Persistence Alleles Reveal Partial East African Ancestry of Southern African Khoe Pastoralists.” Current Biology 24 (8): 852–858.
  • Buckley, M. 2018. “Zooarchaeology by Mass Spectrometry (Zooms) Collagen Fingerprinting for the Species Identification of Archaeological Bone Fragments.” In Zooarchaeology in Practice: Case Studies in Methodology and Interpretation in Archaeofaunal Analysis, edited by C. M. Giovas and M. J. LeFebvre, 227–247. Cham, Switzerland: Springer International.
  • Bunn, H. T., and E. M. Kroll. 1986. “Systematic Butchery by Plio-Pleistocene Hominids at Olduvai Gorge, Tanzania.” Current Anthropology 27: 431–452.
  • Chakroun, Amel, Driss Chahid, Larbi Boudad, Emilie Campmas, Arnaud Lenoble, Roland Nespoulet, and Mohamed Abdeljalil El Hajraoui. 2017. “The Pleistocene of Rabat (Morocco): Mollusks, Coastal Environments and Human Behavior.” African Archaeological Review 34 (4): 493–510.
  • Chritz, Kendra L., Fiona B. Marshall, M. Esperanza Zagal, Francis Kirera, and Thure E. Cerling. 2015. “Environments and Trypanosomiasis Risks for Early Herders in the Later Holocene of the Lake Victoria Basin, Kenya.” Proceedings of the National Academy of Sciences USA 112 (12): 3674–3679.
  • Coutu, Ashley N., Gavin Whitelaw, Petrus le Roux, and Judith Sealy. 2016. “Earliest Evidence for the Ivory Trade in Southern Africa: Isotopic and ZooMS Analysis of Seventh–Tenth Century AD Ivory from Kwazulu-Natal.” African Archaeological Review 33 (4): 411–435.
  • Dale, Darla, and Ceri Z. Ashley. 2010. “Holocene Hunter-Fisher-Gatherer Communities: New Perspectives on Kansyore Using Communities of Western Kenya.” Azania: Archaeological Research in Africa 45 (1): 24–48.
  • d’Errico, Francesco, Lucinda Backwell, Paola Villa, Ilaria Degano, Jeannette J. Lucejko, Marion K. Bamford, Thomas F. G. Higham, Maria Perla Colombini, and Peter B. Beaumont. 2012. “Early Evidence of San Material Culture Represented by Organic Artifacts from Border Cave, South Africa.” Proceedings of the National Academy of Sciences USA 109 (33): 13214–13219.
  • d’Errico, Francesco, Lucinda R. Backwell, and Lyn Wadley. 2012. “Identifying Regional Variability in Middle Stone Age Bone Technology: The Case of Sibudu Cave.” Journal of Archaeological Science 39 (7): 2479–2495.
  • di Lernia, S. 2001. “Dismantling Dung: Delayed Use of Food Resources among Early Holocene Foragers of the Libyan Sahara.” Journal of Anthropological Archaeology 20: 408–441.
  • di Lernia, S. 2013. “The Emergence and Spread of Herding in Northern Africa: A Critical Reappraisal.” Chap. 36. In The Oxford Handbook of African Archaeology, edited by Peter Mitchell and Paul Lane, 527–540. Oxford: Oxford University Press.
  • Domínguez-Rodrigo, M., C. P. Egeland, and T. R. Pickering. 2007. “Equifinality in Carnivore Tooth Marks and the Extended Concept of Archaeological Palimpsests: Implications for Models of Passive Scavenging in Early Hominids.” In Breathing Life into Fossils: Taphonomic Studies in Honor of C. K. (Bob) Brain, edited by T. R. Pickering, K. Schick, and N. Toth, 255–267. Gosport, IN: Stone Age Institute Press.
  • Domínguez-Rodrigo, M. 2002. “Hunting and Scavenging by Early Humans: The State of the Debate.” Journal of World Prehistory 16: 1–54.
  • Domínguez-Rodrigo, M., and R. Barba. 2006. “New Estimates of Tooth Mark and Percussion Mark Frequencies at the Flk Zinj Site: The Carnivore-Hominid-Carnivore Hypothesis Falsified.” Journal of Human Evolution 50: 170–194.
  • Domínguez-Rodrigo, Manuel, and Enrique Baquedano. 2018. “Distinguishing Butchery Cut Marks from Crocodile Bite Marks through Machine Learning Methods.” Scientific Reports 8 (1): 5786.
  • Domínguez-Rodrigo, Manuel, Travis Rayne Pickering, and Henry T. Bunn. 2011. “Reply to Mcpherron Et Al.: Doubting Dikika Is About Data, Not Paradigms.” Proceedings of the National Academy of Sciences USA 108 (21): E117-E17.
  • Dueppen, Stephen A., and Cameron Gokee. 2014. “Hunting on the Margins of Medieval West African States: A Preliminary Study of the Zooarchaeological Record at Diouboye, Senegal.” Azania: Archaeological Research in Africa 49 (3): 354–385.
  • Dunne, J., S. di Lernia, M. Chłodnicki, F. Kherbouche, and R. P. Evershed. 2018. “Timing and Pace of Dairying Inception and Animal Husbandry Practices across Holocene North Africa.” Quaternary International 471: 147–159.
  • Faith, J. T., and A. D. Gordon. 2007.“Skeletal Element Abundances in Archaeofaunal Assemblages: Economic Utility, Sample Size, and Assessment of Carcass Transport Strategies.” Journal of Archaeological Science 34: 872–882.
  • Faurby, S., Daniele Silvestro, Lars Werdelin, and Alexandre Antonelli. (2020). Brain expansion in early hominins predicts carnivore extinctions in East Africa. Ecology Letters 23: 537–544.
  • Fernández-Jalvo, Yolanda, and Peter Andrews. 2016. Atlas of Taphonomic Identifications: 1001+ Images of Fossil and Recent Mammal Bone Modification. Dordrecht, The Netherlands: Springer.
  • Ferraro, Joseph V., Thomas W. Plummer, Briana L. Pobiner, James S. Oliver, Laura C. Bishop, David R. Braun, Peter W. Ditchfield, et al. 2013. “Earliest Archaeological Evidence of Persistent Hominin Carnivory.” PLoS One 8 (4): e62174.
  • Fleisher, Jeffrey, Paul Lane, Adria LaViolette, Mark Horton, Edward Pollard, Eréndira Quintana Morales, Thomas Vernet, Annalisa Christie, and Stephanie Wynne-Jones. 2015. “When Did the Swahili Become Maritime?” American Anthropologist 117 (1): 100–115.
  • Gaillard, Claire, Mukesh Singh, Anne Dambricourt Malassé, Vipnesh Bhardwaj, Baldev Karir, Amandeep Kaur, Surinder Pal, et al. 2016. “The Lithic Industries on the Fossiliferous Outcrops of the Late Pliocene Masol Formation, Siwalik Frontal Range, Northwestern India (Punjab).” Comptes Rendus Palevol 15 (3): 341–357.
  • Gifford-Gonzales, D. 1991. “Bones Are Not Enough: Analogues, Knowledge, and Interpretive Strategies in Zooarchaeology.” Journal of Anthropological Archaeology 10: 215–254.
  • Gifford-Gonzalez, D. 2001. “Animal Disease Challenges to the Emergence of Pastoralism in Sub-Saharan Africa.” African Archaeological Review 17: 95–139.
  • Gifford-Gonzalez, D., and O. Hanotte. 2013. “Domesticating Animals in Africa.” In The Oxford Handbook of African Archaeology, edited by Peter Mitchell and Paul Lane, chap. 34, 491–506. Oxford: Oxford University Press.
  • Gifford-Gonzalez, Diane. 2018. An Introduction to Zooarchaeology. Cham, Switzerland: Springer International.
  • Goldberg, P. 2000. “Micromorphology and Site Formation at Die Kelders Cave 1, South Africa.” Journal of Human Evolution 38: 43–90.
  • Grillo, Katherine M., Mary E. Prendergast, Daniel A. Contreras, Tom Fitton, Agness O. Gidna, Steven T. Goldstein, Matthew C. Knisley, Michelle C. Langley, and Audax Z. P. Mabulla. 2018. “Pastoral Neolithic Settlement at Luxmanda, Tanzania.” Journal of Field Archaeology 43 (2): 102–120.
  • Hall, M. 1999. “Subaltern Voices? Finding the Spaces between Things and Words.” In Historical Archaeology: Back from the Edge, edited by Pedro Paulo, Paulo Funair, Martin Hall, and Sian Jones, 193–203. New York: Routledge.
  • Hanotte, Olivier, Daniel G. Bradley, Joel W. Ochieng, Yasmin Verjee, Emmeline W. Hill, and J. Edward Rege. 2002. “African Pastoralism: Genetic Imprints of Origins and Migrations.” Science 296 (5566): 336–339.
  • Harmand, Sonia, Jason E. Lewis, Craig S. Feibel, Christopher J. Lepre, Sandrine Prat, Arnaud Lenoble, Xavier Boes, et al. 2015. “3.3-Million-Year-Old Stone Tools from Lomekwi 3, West Turkana, Kenya.” Nature 521 (7552): 310–315.
  • Harris, Jacob A., Curtis W. Marean, Kiona Ogle, and Jessica Thompson. 2017. “The Trajectory of Bone Surface Modification Studies in Paleoanthropology and a New Bayesian Solution to the Identification Controversy.” Journal of Human Evolution 110: 69–81.
  • Heinrich, Adam. 2012. “Some Comments on the Archaeology of Slave Diets and the Importance of Taphonomy to Historical Faunal Analyses.” Journal of African Diaspora Archaeology and Heritage 1 (1): 9–40.
  • Henshilwood, Christopher S., Judith C. Sealy, Royden Yates, Kathryn Cruz-Uribe, Paul Goldberg, Frederick E. Grine, Richard G. Klein, et al. 2001. “Blombos Cave, Southern Cape, South Africa: Preliminary Report on the 1992–1999 Excavations of the Middle Stone Age Levels.” Journal of Archaeological Science 28 (4): 421–448.
  • Holl, A. F. C. 2005. “Holocene ‘Aquatic’ Adaptations in North Tropical Africa.” In African Archaeology: A Critical Introduction, edited by A. Stahl, chap. 7, 174–186. Oxford: Blackwell.
  • Horsburgh, K. Ann, Jayson Orton, and Richard G. Klein. 2016. “Beware the Springbok in Sheep’s Clothing: How Secure Are the Faunal Identifications Upon Which We Build Our Models?” African Archaeological Review 33 (4): 353–361.
  • Ichumbaki, Elgidius B. 2017. “When Did the Swahili Become Maritime? A Reply to Jeffrey Fleisher et al. (2015).” In Sea Ports and Sea Power: African Maritime Cultural Landscapes, edited by Lynn Harris, 1–11. Cham, Switzerland: Springer International.
  • James, Emma C., and Jessica C. Thompson. 2015. “On Bad Terms: Problems and Solutions within Zooarchaeological Bone Surface Modification Studies.” Environmental Archaeology 20 (1): 89–103.
  • Jerardino, Antonieta. 2010. “Large Shell Middens in Lamberts Bay, South Africa: A Case of Hunter–Gatherer Resource Intensification.” Journal of Archaeological Science 37 (9): 2291–2302.
  • Jerardino, Antonieta, Joaquim Fort, Neus Isern, and Bernardo Rondelli. 2014. “Cultural Diffusion Was the Main Driving Mechanism of the Neolithic Transition in Southern Africa.” PLoS One 9 (12): e113672.
  • Jones, Mica B., Steven A. Brandt, and Fiona Marshall. 2018. “Hunter-Gatherer Reliance on Inselbergs, Big Game, and Dwarf Antelope at the Rifle Range Site, Buur Hakaba, Southern Somalia ∼20,000− 5,000 BP.” Quaternary International 471: 55–65.
  • Kim, Jaemin, Olivier Hanotte, Okeyo Ally Mwai, Tadelle Dessie, Salim Bashir, Boubacar Diallo, Morris Agaba, et al. 2017. “The Genome Landscape of Indigenous African Cattle.” Genome Biology 18 (1): 34.
  • Klein, R. G. 1976. “The Mammalian Fauna of the Klasies River Mouth Sites, Southern Cape Province, South Africa.” South African Archaeological Bulletin 31: 75–98.
  • Klein, Richard G., and Teresa E. Steele. 2013. “Archaeological Shellfish Size and Later Human Evolution in Africa.” Proceedings of the National Academy of Sciences USA 110 (27): 10910–10915.
  • Kusimba, Chapurukha M., and Jonathan R. Walz. 2018. “When Did the Swahili Become Maritime?: A Reply to Fleisher et al. (2015), and to the Resurgence of Maritime Myopia in the Archaeology of the East African Coast.” American Anthropologist 120 (3): 429–443.
  • Kyriacou, K., J. E. Parkington, M. Will, A. W. Kandel, and N. J. Conard. 2015. “Middle and Later Stone Age Shellfish Exploitation Strategies and Coastal Foraging at Hoedjiespunt and Lynch Point, Saldanha Bay, South Africa.” Journal of Archaeological Science 57: 197–206.
  • Lahr, M. Mirazón, F. Rivera, R. K. Power, A. Mounier, B. Copsey, F. Crivellaro, J. E. Edung, et al. 2016. “Inter-Group Violence among Early Holocene Hunter-Gatherers of West Turkana, Kenya.” Nature 529 (7586): 394–398.
  • Lam, Y. M., O. M. Pearson, Curtis W. Marean, and Xingbin Chen. 2003. “Bone Density Studies in Zooarchaeology.” Journal of Archaeological Science 30 (12): 1701–1708.
  • Lombard, Marlize, and Isabelle Parsons. 2015. “Milk Not Meat: The Role of Milk Amongst the Khoe Peoples of Southern Africa.” Journal of African Archaeology 13 (2): 149–166.
  • Lyman, R. L. 2008. Quantitative Paleozoology. New York: Cambridge University Press.
  • Marean, C. W., and S. Y. Kim. 1998. “Mousterian Large-Mammal Remains from Kobeh Cave: Behavioral Implications for Neanderthals and Early Modern Humans.” Current Anthropology 39 (S): S79–S113.
  • Marean, C. W., L. M. Spencer, R. J. Blumenschine, and S. D. Capaldo. 1992. “Captive Hyena Bone Choice and Destruction, the Schlepp Effect, and Olduvai Archaeofaunas.” Journal of Archaeological Science 19: 101–121.
  • Marean, Curtis W. 2016. “The Transition to Foraging for Dense and Predictable Resources and Its Impact on the Evolution of Modern Humans.” Philosophical Transactions of the Royal Society B 371 (1698): 20150239.
  • Marshall, Fiona, Rachel E. B. Reid, Steven Goldstein, Michael Storozum, Andrew Wreschnig, Lorraine Hu, Purity Kiura, Ruth Shahack-Gross, and Stanley H. Ambrose. 2018. “Ancient Herders Enriched and Restructured African Grasslands.” Nature 561: 387–390.
  • McIntosh, Roderick J. 2005. Ancient Middle Niger: Urbanism and the Self-Organizing Landscape. Cambridge, U.K.: Cambridge University Press.
  • McPherron, Shannon P., Zeresenay Alemseged, Curtis W. Marean, Jonathan G. Wynn, Denne Reed, Denis Geraads, Rene Bobe, and Hamdallah A. Bearat. 2010. “Evidence for Stone-Tool-Assisted Consumption of Animal Tissues before 3.39 Million Years Ago at Dikika, Ethiopia.” Nature 466 (7308): 857–860.
  • Merritt, Stephen R., and Kellyn M. Davis. 2017. “Diagnostic Properties of Hammerstone-Broken Long Bone Fragments, Specimen Identifiability, and Early Stone Age Butchered Assemblage Interpretation.” Journal of Archaeological Science 85: 114–123.
  • Miller, Jennifer Midori, and Pamela Rae Willoughby. 2014. “Radiometrically Dated Ostrich Eggshell Beads from the Middle and Later Stone Age of Magubike Rockshelter, Southern Tanzania.” Journal of Human Evolution 74: 118–122.
  • Morgan, Christopher. 2015. “Is It Intensification Yet? Current Archaeological Perspectives on the Evolution of Hunter-Gatherer Economies.” Journal of Archaeological Research 23 (2): 163–213.
  • Mulazzani, Simone, Lotfi Belhouchet, Laure Salanova, Nabiha Aouadi, Yosra Dridi, Wassel Eddargach, Jacob Morales, et al. 2016. “The Emergence of the Neolithic in North Africa: A New Model for the Eastern Maghreb.” Quaternary International 410: 123–143.
  • O’Driscoll, Corey A., and Jessica C. Thompson. 2018. “The Origins and Early Elaboration of Projectile Technology.” Evolutionary Anthropology: Issues, News, and Reviews 27 (1): 30–45.
  • Otárola-Castillo, E., M. Torquato, H. Hawkins, E. James, J. Harris, C.W. Marean, S.P. McPherron, and J.C. Thompson. 2018. “Differentiating between Cutting Actions on Bone Using 3d Geometric Morphometrics and Bayesian Analyses with Implications to Human Evolution.” Journal of Archaeological Science 89: 56–67.
  • Pante, Michael C., Matthew V. Muttart, Trevor L. Keevil, Robert J. Blumenschine, Jackson K. Njau, and Stephen R. Merritt. 2017. “A New High-Resolution 3-D Quantitative Method for Identifying Bone Surface Modifications with Implications for the Early Stone Age Archaeological Record.” Journal of Human Evolution 102: 1–11.
  • Parkinson, Jennifer A. 2018. “Revisiting the Hunting-Versus-Scavenging Debate at Flk Zinj: A Gis Spatial Analysis of Bone Surface Modifications Produced by Hominins and Carnivores in the Flk 22 Assemblage, Olduvai Gorge, Tanzania.” Paleogeography, Paleoclimatology, Paleoecology 511: 29–51.
  • Pleurdeau, David, Emma Imalwa, Florent Détroit, Joséphine Lesur, Anzel Veldman, Jean-Jacques Bahain, and Eugène Marais. 2012. “‘Of Sheep and Men’: Earliest Direct Evidence of Caprine Domestication in Southern Africa at Leopard Cave (Erongo, Namibia).” PloS One 7 (7): e40340.
  • Pobiner, Briana L. 2015. “New Actualistic Data on the Ecology and Energetics of Hominin Scavenging Opportunities.” Journal of Human Evolution 80: 1–16.
  • Prendergast, Mary E., and Amanuel Beyin. 2018. “Fishing in a Fluctuating Landscape: Terminal Pleistocene and Early Holocene Subsistence Strategies in the Lake Turkana Basin, Kenya.” Quaternary International 471: 203–218.
  • Prendergast, Mary E., Michael Buckley, Alison Crowther, Laurent Frantz, Heidi Eager, Ophélie Lebrasseur, Rainer Hutterer, et al. 2017. “Reconstructing Asian Faunal Introductions to Eastern Africa from Multi-Proxy Biomolecular and Archaeological Datasets.” PloS One 12 (8): e0182565.
  • Prendergast, Mary E., E. M. Quintana Morales, A. Crowther, M. C. Horton, and N. L. Boivin. 2017. “Dietary Diversity on the Swahili Coast: The Fauna from Two Zanzibar Trading Locales.” International Journal of Osteoarchaeology 27 (4): 621–637.
  • Russell, N. 2011. Social Zooarchaeology: Humans and Animals in Prehistory. Cambridge, U.K.: Cambridge University Press.
  • Sahle, Yonatan, Sireen El Zaatari, and Tim D. White. 2017. “Hominid Butchers and Biting Crocodiles in the African Plio–Pleistocene.” Proceedings of the National Academy of Sciences USA 114 (50): 13164–13169.
  • Scott, Karin, and Ina Plug. 2016. “Osteomorphology and Osteometry Versus aDNA in Taxonomic Identification of Fragmentary Sheep and Sheep/Goat Bones from Archaeological Deposits: Blydefontein Shelter, Karoo, South Africa.” Southern African Humanities 28: 61–79.
  • Sealy, J. C. 2016. “Intensification, Diet, and Group Boundaries among Later Stone Age Coastal Hunter-Gatherers Along the Western and Southern Coasts of South Africa.” In The Oxford Handbook of the Archaeology of Diet, edited by Lee-Thorp Julia, M. Anne Katzenberg, and Sealy Judith. Oxford: Oxford University Press.
  • Sereno, Paul C., Elena A. A. Garcea, Hélène Jousse, Christopher M. Stojanowski, Jean-François Saliège, Abdoulaye Maga, Oumarou A. Ide, et al. 2008. “Lakeside Cemeteries in the Sahara: 5000 Years of Holocene Population and Environmental Change.” PloS One 3 (8): e2995.
  • Smith, Andrew. 2014. The Origins of Herding in Southern Africa: Debating the “Neolithic” Model. Saarbrücken, Germany: LAP Lambert Academic Publishing.
  • Stewart, Kathlyn. 1989. Fishing Sites of North and East Africa in the Late Pleistocene and Holocene: Environmental Change and Human Adaptation: British Archaeological Reports, vol. 521. Oxford: British Archaeological Reports.
  • Stone, Abigail Chipps. 2018. “Finding the Ephemeral: Herding Strategies and Socio-Economic Organization in an Urban West African Context.” Quaternary International 471: 160–174.
  • Sutton, J. E. G. 1977. “The African Aqualithic.” Antiquity 51 (201): 25–34.
  • Texier, Pierre-Jean, Guillaume Porraz, John Parkington, Jean-Philippe Rigaud, Cedric Poggenpoel, and Chantal Tribolo. 2013. “The Context, Form and Significance of the MSA Engraved Ostrich Eggshell Collection from Diepkloof Rock Shelter, Western Cape, South Africa.” Journal of Archaeological Science 40 (9): 3412–3431.
  • Thompson, J. C., S. Carvalho, C. W. Marean, and Z. Alemseged. 2019. “Origins of the Human Predatory Pattern: The Transition to Large Animal Exploitation by Early Hominins.” Current Anthropology 60 (1): 1–23.
  • Thompson, Jessica C., and Christopher S. Henshilwood. 2014. “Nutritional Values of Tortoises Relative to Ungulates from the Middle Stone Age Levels at Blombos Cave, South Africa: Implications for Foraging and Social Behaviour.” Journal of Human Evolution 67: 33–47.
  • Wadley, Lyn. 2010. “Were Snares and Traps Used in the Middle Stone Age and Does It Matter? A Review and a Case Study from Sibudu, South Africa.” Journal of Human Evolution 58 (2): 179–192.
  • Woldekiros, H. S., and A. C. D’Andrea. 2017. “Early Evidence for Domestic Chickens (Gallus Gallus Domesticus) in the Horn of Africa.” International Journal of Osteoarchaeology 27 (3): 329–341.
  • Wright, David K. 2017. “Humans as Agents in the Termination of the African Humid Period.” Frontiers in Earth Science 5: 4.
  • Yellen, J. E., A. S. Brooks, E. Cornelissen, M. J. Mehlman, and K. Stewart. 1995. “A Middle Stone Age Worked Bone Industry from Katanda, Upper Semliki Valley, Zaire.” Science 268: 553–556.
  • Zhu, Zhaoyu, Robin Dennell, Weiwen Huang, Yi Wu, Shifan Qiu, Shixia Yang, Zhiguo Rao, et al. 2018. “Hominin Occupation of the Chinese Loess Plateau since About 2.1 Million Years Ago.” Nature 559 (7715): 608–612.