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date: 20 October 2018

Archaeozoology: Methods

Summary and Keywords

Archaeozoology is the study of animal remains, mainly bones and other hard parts, from archaeological sites. It contributes to a more complete understanding of various aspects of human life in the past. Ideally, archaeozoologists, like other specialists, should be involved in the entire process of an archaeological research project, from its design, to fieldwork and data collecting, to final reporting and publication. For efficient communication and fruitful collaboration, the archaeologists involved in this process need to understand the basics of archaeozoological methodology and the range of questions that the discipline can answer. Methods vary among archaeozoologists—not least with regard to quantification—and it is important to be aware of these differences and their possible impact on results when comparing data for different sites. While the actual analysis of animal remains is done by the archaeozoologists, preferably in circumstances where they have access to a comparative collection of recent animal skeletons, the excavation and collection of remains is often the responsibility of the archaeologists. Animal remains are affected by a host of taphonomic processes of loss that are beyond our control. To avoid additional loss of information at the fieldwork stage, appropriate methods are particularly important. The use of sieves with a mesh size no greater than 2 mm is essential in order not to miss the smaller, but no less informative, animal remains. Project leaders play an important role in providing good storage facilities for archaeozoological remains after excavation and after study. With the rapid development in analytical methods, it can be extremely interesting to return to previously studied remains and sample them.

Keywords: vertebrate animals, bones, teeth, taphonomy, archaeozoology, methods, paleo-economy, paleo-ecology, later Holocene, Africa

The Study of Animal Bones and Teeth from African Historical Sites

Archaeozoology, or zooarchaeology, is the study of all kinds of physical remains or traces of animals from archaeological contexts. Here the focus is on bones and teeth of vertebrates (fish, amphibians, reptiles, birds, and mammals). Together with mollusk and bird egg shells, which are also hard and sometimes well preserved, these are the most commonly found archaeological animal remains. More exceptionally, soft parts are also preserved, such as feathers, hair, horn, or even skin. Traces left by animals include gnawing marks, footprints, feces, and burrows. Archaeozoologists may be specialized in a certain animal group, region, period, or method. Although archaeozoological research may sometimes emphasize zoological aspects, this article primarily focuses on how archaeozoologists contribute to the reconstruction of the life of humans in the past, of their natural environment, and of the interaction between the two.

The preservation of animal remains and the reasons for their presence in archaeological deposits are discussed first. Some basic methods are explained of archaeozoological analysis of bones and teeth, including methods of collection and recovery, which are often the responsibility of the field archaeologists. Finally, the diverse ways in which archaeozoological research contributes to the archaeology of historical periods in Africa are discussed. The goal is to give non-specialists an understanding of the principles, methods, and research potential of archaeozoology, and to stimulate communication and collaboration with faunal specialists. These specialists, ideally, should be involved in the entire process of the archaeological research project, from its design, to fieldwork and data collecting, to final reporting and publication. For non-zooarchaeologists making use of published faunal datasets, it is also important to have some knowledge of taphonomy and zooarchaeological methods, and of how these may affect results.

Taphonomy

In an archaeozoological context, the term taphonomy refers to the study of all events between the time of death of an animal and the discovery of its remains by archaeologists, encompassing how and why the animal ended up in the archaeological deposits, and the processes that resulted in the preservation of (part of) its remains.

Preservation

Except in special circumstances, for animal remains to be preserved, they should be buried relatively quickly after deposition because on the ground surface, various processes such as sun, rain, and trampling by humans or animals will quickly lead to their destruction. Once the remains are in the ground, acidity and soil water management are probably the main factors influencing preservation.1 Bones and teeth consist of an organic (collagen) and an inorganic (hydroxyapatite) fraction, and it is mainly thanks to the latter that they can be preserved. Tooth enamel and dentine have higher percentages of inorganic material than do bones (96–99.5% and 75–80% vs. 65%, respectively) and therefore have the best chances of surviving in archaeological deposits.2 Acidic soils are detrimental to the inorganic fraction. Because such soils prevail in large parts of equatorial Africa, bones have been preserved particularly poorly there. Soil acidity explains, for example, why no bones at all have been retrieved from recent excavations at Nok sites in central Nigeria (c. 1000 bce–500 ce).3 Faunal remains from Central Africa are mainly restricted to cave sites or to particular structures, such as pits, in fairly recent sites, where localized preservation conditions are better.4 Intermittently wet and dry conditions are favorable for the survival of microorganisms that break down the organic fraction of bones and teeth, and thus also negatively affect the preservation of bone. Very dry conditions can lead to exceptionally good organic preservation, as witnessed in the deserts of Egypt, where there are examples of archaeological animal remains with hair, skin, and horn still adhering.5 Waterlogged conditions also lead to good preservation, as found at the historical site of Bolondo in the Central Congo Basin (c. 1500 ce).6 In general, the better preservation conditions are found mostly in the arid and semi-arid areas of Africa. However, although bones macroscopically look well preserved, most of the collagen will often be degraded, hampering aDNA and other analysis.

In addition to the burial circumstances, the nature of the animal material itself determines its preservation chances. Tooth enamel preserves better than either dentine or bone, and thicker mammal bone preserves better than thin bird bone. Often an overrepresentation of elements that are more resistant to destruction—such as teeth and small, compact bones, including carpals and tarsals—can be noted.7 This so-called differential preservation also needs to be taken into account when researchers are looking at numbers of remains by animal taxon, as the remains of some taxa will preserve better than those of others. In fish, for example, skulls of most catfish (Siluriformes) preserve particularly well compared with bones of other fish.8

Taphonomic Groups

Animal remains preserved in archaeological sites can be grouped according to the reasons they ended up there. These groupings are referred to as taphonomic groups.9 It is important to distinguish between remains that are contemporary with human habitation and those that are older or younger. The group termed intrusives consists of individuals that were not brought to the site intentionally by its human inhabitants. Any surface will likely have some animal remains lying on it when humans first start to use it, and these remains can become incorporated into the archaeological layers. Animal remains may also be reworked into the archaeological deposit from older (geological) deposits. Remains of more recent animals (termed late intrusives) are often from burrowing taxa that died inside burrows they made in archaeological deposits. But even remains of animals that date from the time around the human occupation of a site may be intrusive. This group, called pene-contemporaneous intrusives, typically consists of mice and rats, as well as other small creatures that could have lived and died on the site without much human interference.

Although sites can yield a diverse group of intrusive animal taxa, remains of animals that did end up at the site through deliberate human actions are usually more numerous. The most important taphonomic group is probably the food waste. Another group is the artisanal refuse, which includes any worked animal remains, waste from working activities, and remains of animals killed for their skin or fur. Worked animal remains are usually treated as artifacts, but these should nevertheless be made available to the faunal specialist, who can often determine the raw material (bone, tooth/ivory, other) as well as the taxon, or at least the size class and the larger group of animals (e.g., mammal, bird). There are also such things as pseudo-artifacts, bones that look worked to a non-specialist, but which are, actually, entirely natural—another reason to have the faunal specialist review all of the bones deemed from worked animals. A further taphonomic group is the carcasses. These are animals that were not eaten after their death, and that usually served either as a companion animal or as a provider of power while alive. Animals that were not eaten were often dumped outside of settlements, which significantly reduced their chances of being preserved and found during archaeological excavations. This explains the relative lack of dromedary (Camelus dromedarius), horse (Equus caballus), and donkey (Equus asinus) from historical African archaeological contexts in areas where these animals must have occurred.10 A final taphonomic group is the ritual remains, meaning animals offered or consumed in a ritual or religious context.

Zooarchaeologists use the find context, the cultural context (e.g., which animals were considered edible), and traces on the remains to determine to which taphonomic group the remains are likely to belong. For at least some of the remains, the taphonomic status remains unclear. All archaeozoologists reflect on the reasons for the presence of the animal remains they are studying, as these have important implications for the interpretation of the data, but some will discuss this more explicitly in their reports than others.

Methods

Excavation, Collection, and Selection

It is advisable that archaeozoologists, as well as other specialists, be involved in the design of archaeological projects, the planning of fieldwork, and the selection of contexts to excavate. This helps specialists to answer their own research questions and it allows them to better contribute to the general goals of the project. In certain circumstances, and depending on their own habits and training, faunal specialists may be on-site during excavation. However, that is not usually the case, and the field archaeologists are responsible for the excavation and collection of archaeological animal remains. When animal skeletons in articulation are encountered on a site, meaning that the bones are still in the same anatomical position as in a live animal (often animal burials, but not restricted to that) (figure 1), a faunal specialist is better called in to record in situ, as this will allow gathering maximal information. The same applies for any special contexts with animal bones.

Archaeozoology: MethodsClick to view larger

Figure 1. Example of complete animal skeletons found in articulation: Two cats (Felis catus) from a pet cemetery at Berenike (Egypt, late 1st–early 2nd century ce). Most animal remains that archaeozoologists study are disarticulated, fragmented, and found within a mixture of remains of different species (see figure 5). Photo courtesy of Marta Osypińska.

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Figure 2. Schematic summary of the taphonomic history of animal remains, from living animals to the studied animal bone assemblage. Each step in the scheme results in a loss of information. Adapted from Simon Davis. The Archaeology of Animal Bones (London, UK: Routledge, 1987), 22.

Each step in the taphonomic process leads to a loss of information (figure 2), and most of these steps cannot be controlled. However, loss due to recovery methods can be controlled. Usually larger bones are handpicked from the excavation trenches. The key is to collect not only these larger bones, which are usually easy to see, but also the small to very small bones, as these are equally informative. This includes bones of small-bodied taxa (a typical example is fish) and small bones and fragments of bones of larger taxa. To this end, all remaining anthropogenic sediment should be screened, with or without the aid of water, depending on the soil type and what is possible logistically. For reasons of efficiency, it is helpful to coordinate archaeozoological sampling with archaeo-botanical and other types of sampling.

The collection methods used will significantly affect the list of identified animal taxa and elements, with an underrepresentation of smaller taxa and bones in case of a lack of (fine) screening. Zooarchaeological specialists can judge how fine the screening should be for a good balance between results and time invested, as this differs between sites and contexts. Exclusive use of screens with mesh aperture larger than 2–4 mm is not advised, and if such relatively large mesh sizes are used for some of the recovery, subsamples should be sieved on a finer mesh.11 Subsample volumes must be noted to enable extrapolation to the total deposit. The sieve residues usually also contain various remains, such as large sediment particles, that are not of interpretive use, and it may be very time consuming to separate out the bones and other useful finds. Sorting should be done with care and, if possible, in lab circumstances, rather than directly from the sieve in the field, because it is more difficult to see the bones in field conditions, and there is usually an element of time pressure.

Washing faunal material is not advised because it can damage bones and may adversely affect certain analytical methods. Decisions on any cleaning that may be required is better left to the zooarchaeologist. Ideally, remains should be dried (slowly) prior to packaging. If this is not possible and the remains are still wet, they should be placed in bags or containers that are left open or are permeable to moisture. The remains must be protected from damage, with smaller, fragile bones packed separately from more bulky remains. All packing should be clearly, durably, and comprehensively labeled. It is important to give the specialists all details on find numbers, collection methods, contexts, soil volumes, etc.

It is not always possible or even advisable to study all of the animal remains collected during an archaeological excavation project, and a selection should then be made in consultation between the excavator and the archaeozoologist. Selection can be by type of context, by date, or by animal group (fish, mammals, birds, etc.), and it can be preceded by an initial assessment of all of the faunal material. Excavators should not triage the animal remains themselves, either for size or for what may seem more “diagnostic” to them. It is also better to leave any sorting into animal groups to the specialist, or at least to coordinate strategies with them.

Identifications

Before attempting identification, archaeozoologists sometimes find it necessary to clean animal remains. They usually do this using a soft, dry brush. Bones with (recent) breaks may be glued back together to help identification or to allow the taking of (additional) measurements. Identification includes the determination of the type of material; the skeletal element; which portion of the element, in cases where bones are incomplete; the laterality (left or right); and the taxon. Although identifying the material is usually straightforward, it can also be difficult, for example, when distinguishing between bone and ivory or between different types of ivory.12 Ivory refers to any mammal teeth that are large enough to be worked; in Africa this refers mainly canines and incisors of elephant (Loxodonta africana), hippo (Hippopotamus amphibius), and (wild) pig (Suidae).

Identification of the animal taxon is done as precisely as possible by the faunal specialist. The most precise level of identification that can be reached is normally the species level, but identification may also be restricted to genus or family, or to an even higher taxonomic level (figure 3). For each level, there are scientific names and conventions for how to write these. Scientific names are mandatory in archaeozoological reports to meet international standards. They also prevent confusion, particularly when work is translated into different languages. However, common names are usually added, as they are easier to understand for non-specialists. Archaeozoologists record uncertainties in their identifications by adding “cf.,” “?,” and other qualifiers.

Archaeozoology: MethodsClick to view larger

Figure 3. Taxonomic levels and scientific nomenclature illustrated for wolf and its domesticated form, dog. Note that there are two systems for naming domesticated animals, although it has been argued that the second system should be adopted by all researchers.13

Ideally, identification should be done in lab facilities housing a reference collection of (recent) skeletons of the animal taxa that can be expected at the site. Archaeological faunal remains are identified through direct comparison with these reference skeletons. Due to local conditions in Africa and export constraints, it is not always possible to do identifications using comparative collections. Even for experienced researchers the lack of such collections can make it very difficult to work and to make identifications with confidence. Archaeozoologists should mention in their reports which comparative collections they have used, if any, because this can help colleagues to evaluate the reliability of the identifications. Pictures and drawings of bones, as well as modern aids, such as 3D scans, are helpful, but they cannot replace actual reference skeletons. Criteria for the identification of element and taxon are shape and size of the remains, although size may vary within a single species (see later). Functionally similar elements look more similar the more closely animals are related (figure 4). Some types of bones are more diagnostic than others, meaning that they show distinguishing traits that allow identifications to species. Ribs and vertebrae are typically not diagnostic. When bones are depicted in reports, they are usually oriented proximal side up, and the view will be mentioned (figure 4). While describing the parts of bones present, archaeozoologists can use simple, quick descriptions or refer to more detailed, standardized systems.14

Archaeozoology: MethodsClick to view larger

Figure 4. Left Femurof horse (Equus), cattle (Bos), and sheep (Ovis). Cranial view. Proximal side is up, distal side is down. Note the similarities among these bones, despite the size and shape differences that allow for species identification. Adapted from Elisabeth Schmid, Atlas of Animal Bones: For Prehistorians, Archaeologists, and Quaternary Geologists. Knochenatlas: Für Prähistoriker, Archäologen, und Quartärgeologen (New York, NY: Elsevier, 1972), 111.

General bone atlases exist for African mammals, containing drawings of the most common bones of the most frequently encountered species.15 To distinguish among similar species, more detailed identification keys are available in many cases. Specific to Africa is the distinction between different species of antelopes.16 Often species identification is not possible, and they are grouped according to size classes. In historical contexts, in addition to antelopes, domesticated bovids (sheep, goat, and cattle [Bos sp.]) are also to be expected. A common problem in archaeozoology is the distinction between sheep (Ovis ammon) and goat (Capra hircus).17 The distinction between wild and domestic bovids is also not always straightforward; for example, the distinction between cattle and buffalo (Syncerus caffer) can be ambiguous.18 As with the antelopes, size categories of unidentified bovids are often used. A very well known classification system of African bovids by size class (Class I–IV, from small to large, based on body weight) is that of C. K. Brain; As with the antelopes, in such cases size categories of unidentified bovids are often used. A very well known classification system of African bovids by size class (Class I-IV, from small to large, based on body weight) is that of Brain,19 others have used similar systems that are tailored to the regions and periods they are working in.

Archaeozoology: MethodsClick to view larger

Figure 5. Animal remains from one archaeological unit at the Iron Age site of Oursi village (Burkina Faso, ce 0–1400). Typically about a quarter of the bones are identifiable, at least to order level. Photo by author.

All faunal collections contain remains of which no more can be said with respect to taxon than that they are of mammal, fish, etc. Sometimes even identification to class may not be possible with traditional methods. When archaeozoologists call remains “identifiable,” they may not always be referring to the same taxonomic level, everything identified at least to class, to order, and so on. Some researchers divide unidentifiable fragments into size classes, type of bone, etc.20 Other details that may be recorded for unidentifiable remains are state of preservation and modifications (see later), as well measurements of their length and/or thickness.21 The identification rate—that is, the percentage of the remains that could be identified—depends on various factors, including collection method(s), state of preservation, the presence of similar taxa, the experience of the researcher, and the available reference collection (figure 5). Low identification rates are not necessarily bad; they may mean that the animal remains were collected rigorously, including all the small, unidentifiable fragments.

Quantification

Treatment of archaeozoological data should, by preference, be quantitative, but this is not statistically justifiable when the assemblages are very small. Quantification of the relative frequency of the taxa identified (figure 6) is probably the aspect of archaeozoological methodology that shows the most variation among researchers.22 Most common and most straightforward is probably counting numbers of remains, known as number of identified specimens (shortened to NISP).

Archaeozoology: MethodsClick to view larger

Figure 6. Typical graphical summary of relative abundance of animal taxa, here presented both by NISP and by MNI. The indication of the total numbers, on which the graphs are based, are necessary to estimate the representativeness of the proportions. The taxa in the graph are the major groups of tetrapod fauna at Fukuchani (FK) and Unguja Ukuu (UU) (Zanzibar, second half 1st millennium ce). Source: Mary E. Prendergast, Eréndira M. Quintana Morales, Alison Crowther, Mark C. Horton, and Nicole L. Boivin, “Dietary Diversity on the Swahili Coast: The Fauna from Two Zanzibar Trading Locales,” International Journal of Osteoarchaeology 27, no. 4 (2017): 621–637.

To reconstruct meat weight, NISPs are sometimes multiplied by average live weight by taxon.23 There are some issues with the NISP method, for example, that a single skeletal element that has broken into several fragments may be counted several times. This is a problem particularly with jaws, which may break into multiple bone fragments and loose teeth and tooth fragments. Such problems can be resolved by estimating minimum numbers of elements (MNE) or minimum number of individual specimens (MNISP).24 The other common quantification method is minimum number of individuals (MNI), in which the minimum number of individual animals necessary to explain the remains present is calculated. This is done by context, usually taking laterality, portion present, and sometimes also size and age into account. Although the basic principles are simple, there is no standardized way of calculating MNI. There are advantages and disadvantages to both the MNI and NISP method. When comparing data from different sites or contexts, it is important to ascertain that quantification was done in a similar way. NISP is often preferred for comparative purposes because calculations are simpler and done in a more uniform way, and are also more commonly available. For contexts such as burials, in which clearly complete individuals or parts of complete individuals are present, calculating MNI is definitely recommended. Another common quantification method is to weigh animal remains. The premise of this method is that bone weight is a better reflection of the food represented by the remains than are bone counts. In any case, it is very important to realize that none of these three main quantification methods has a direct correlation with how much each taxon contributed to the diet (certainly not in terms of the different caloric values of their meat or other parts eaten), nor with how many of these animals actually lived at or near the site.

Ageing and Sexing

Data on the age and sex of the animals represented are collected, as they inform us about domestic herd management and hunting and fishing strategies. The biological age of fish and reptiles can be inferred from their size, as these animals keep on growing throughout their life at documented rates. In mammals, as long as long bones are still growing, their epiphyses remain unfused. For domestic mammals and some wild species, the fusion state can be used for age determination, because the approximate age at which each epiphysis fuses to the corresponding diaphysis is known.25 For fetuses of domestic species, the lengths of long bones without articulations allow for estimation of the age at death.26 Tooth eruption and tooth wear also provide an indication for age at death. Especially for domesticated species, aging systems based on observations on the (mandibular) dentition of modern animals of known biological age are well developed.27 Although age estimates based on teeth are preferred because they provide greater precision, such estimates are mainly useful where large numbers of relatively complete jaws are preserved and mortality profiles can be constructed. Such profiles, in theory, will look different depending on whether livestock was kept mainly for meat, milk, or other secondary products.28 Unfortunately, in sub-Saharan Africa, it is rare to find large numbers of jaws preserved.29 Moreover, because the methods for age identifications of domesticates were developed on modern individuals raised in places other than sub-Saharan Africa, there may be some deviations, because the growth rate of animals as well as their rate of tooth replacement vary with the breed and living conditions.

The sexing of certain archaeological animal remains is possible based on traits that are unique to one of the two sexes, for example baculae (penis bones) in male carnivores and certain male primates and spurs in male chickens (Gallus domesticus). Sexing is also possible based on sexual dimorphism, meaning that there are morphological differences in the same skeletal parts of two sexes of one species. This is, for example, the case for the pelvic bone of mammals, due to the fact that only the females carry young. Size can also allow the distinction between two sexes of one species. In mammals, males are usually larger than females, while in birds it is the other way around.

Measurements and Size Reconstructions

Measuring bird and mammal bones is done in a very standardized way, mainly with callipers, and generally with a precision of 1 or 0.1 mm.30 Measurements are also taken on the bones of some fish. Because of the greater variety in morphology of fish skeletons, measurements need to be defined separately for different taxonomic groups of fish.31 Measurements can be useful for species identification—for example, to separate between different species of antelopes, to determine sex, or to reconstruct types (breeds) of a certain domesticated species.32 Size changes in wild mammals can also be linked to climatic fluctuations of temperature (cf. Bergman’s rule) and of precipitation.33 To look at size variation through time and space for a certain species, measurements on a particular skeletal element can be used. When not enough data on the same skeletal part are available, there are methods that allow for the combining of measurements on different elements. The most commonly used of these is the Logarithmic Size Index (LSI), which is based on comparisons of archaeological measurements with that of a selected reference individual of the same species.34 For domesticated species, withers heights can be reconstructed by multiplying the lengths of long bones by specific factors. However, in contexts with poor preservation, such length measurements are usually not obtainable. Moreover, the relationships between long bone measurements and withers height vary between types, and the factors mediating this variation have not been defined for African breeds.35

In fish, measurements can be used as a means to calculate body length, expressed either as standard length (SL), that is, the distance from the tip of the snout to the beginning of the tail, or total length (TL), provided that equation curves based on a large number of comparative specimens are available.36 Another way to estimate fish size is by direct comparison of archaeological bones with those of recent specimens of known size. Such estimates will be more reliable the more fish of different sizes are represented in the reference material used. Fish will usually be attributed to size classes, for example, of 10 cm (0–10 cm, 10–20 cm, 20–30 cm, etc.).

Description of Modifications and Pathologies

During faunal analysis, archaeozoologists describe the appearance of the animal remains and the diverse modifications that can be seen on them. These modifications can be either anthropogenic or natural and can inform on the various processes that the remains have undergone after the death of the animal, both before and after burial. The detailed recording of such modifications is more commonly done for prehistoric than for historical assemblages.37 Depending on the researcher and on the type of assemblage, modifications are recorded for all remains or just for the identifiable ones.

To describe the state of preservation of bone remains, archaeozoologists often use the standard system designed by Behrensmeyer, with its so-called weathering stages, ranging from 0 to 5 for larger mammals, ranging between very well preserved bone without cracks or flakes (0) and very poorly preserved bone, crumbling down in splinters (5).38 Better preservation increases the chances of identification.

A type of natural modification that can often be seen even with the naked eye is traces of animal gnawing, and these can often be attributed to a specific group of animals. Rodents gnaw bones and other things to sharpen their continuously growing incisors. The incisors of these animals leave distinctive traces consisting of a repetition of two parallel lines. Marks of chewing and breaking by domestic dogs (Canis familiaris) or other carnivores also leave distinctive traces. These animals can also swallow small bones or bone fragments, which they vomit up or pass out with their feces. Once attacked by stomach juices, such fragments may show varying degrees of etching or may disappear entirely.39 Finds of carnivore gnawing marks, etching, and feces can indicate the presence of dogs at or near the site, even where the bones of these animals are missing.

Burning or charring may be anthropogenic or natural. Burning causes alterations in bone color indicative of the temperature the bones were exposed to, with brown (charred) at the lower end and grey-white (calcined) at the upper end of the scale.40 It can be difficult to distinguish between (dark) brown colors that are the consequence of exposure to fire, on the one hand, or of diagenetic processes, on the other; and the white color that is the result of burning can potentially be confused with whitish coloration resulting from sun bleaching. Anthropogenic burning can have different possible reasons. During roasting, the exposed bone ends will usually be burned, while the meat-adhering parts are only burned when roasted to beyond the point where the flesh is still edible. Burning may also be a consequence of deliberate burning of refuse, which results in particularly large concentrations of burned bones.41 Often burning is accidental, due to bones coming in contact with fire at some point after their deposition.

Another important group of anthropogenic marks are left by tools. The main types are cut-marks (shallow), chop-marks (deeper), and marks indicating that the bone has been chopped right through. Cut-marks and chop-marks mainly result from cutting away meat or removing skin with knife-like implements, while through-and-through chops usually result from dividing up carcasses with non-knife tools, such as axes or cleavers.42 It is possible to distinguish between marks left by iron tools and marks left by stone tools using microscopy.43 The types of marks, their number, and their position on the bones are described by archaeozoologists. Some give graphical summaries of the places on the skeleton where marks were found and the frequencies in which these marks were recorded, as this shows how animals were processed (skinning, dividing up of the carcass, and removal of meat). Archaeologists tend to overemphasize the importance of marks left by tools as evidence for the consumption of animals. Such marks can be necessary to firmly prove the consumption of species that are less commonly accepted as being edible, such as dogs. However, the mere fact that animal skeletons are found in settlement contexts, in a disarticulated state, and in a mixture of bones of different taxa, points to consumption.44

Traces of pathologies or trauma are sometimes also recorded. These can tell us more about how animals were treated by humans. The use of animals for power, for example, can leave marks on their skeletons. The presence of a range of pathologies on their skeletons allowed for the conclusion that the donkeys buried in an early pharaonic tomb in Abydos (Egypt, c. 3000 bce), which were not yet morphometrically distinguishable from the wild form, had been used for pack transport.45 For cattle bones, a standardized method exists to record draught-related pathologies.46

Sampling for Analytical Methods

Apart from radiocarbon dating, diverse newer biochemical analytical methods can be applied to animal bones and teeth. Such methods developed quickly over the past decades and have made it possible to address a wide range of new questions or to test existing models with new types of evidence. Stable isotope analysis, mainly of the elements carbon, nitrogen, oxygen, and strontium, allows reconstruction of animal diet, the climate and environment animals were living in, herd management and breeding strategies, (seasonal) movements, and geographic provenance.47 Ancient DNA (aDNA) research can be applied on remains of domesticated animals, provided that organic fractions (collagen in bone and teeth) are preserved, to study their (introduction) history in a certain area, as well as their population movements.48 In fact, aDNA analysis aimed at making such reconstructions has prompted a discussion on the correctness of identifications of archaeological animal bones as belonging to domesticates.49 An aDNA bulk-bone metabarcoding (BBM) approach to faunal identification can be helpful. In that case, DNA from unidentified bones is extracted and sequenced in parallel, resulting in a list of taxa present in the sample.50 Zooarchaeology through mass spectrometry (ZooMS) can, based on the analyses of specific collagenous and non-collagenous proteins in bones and teeth, also allow species-level identifications where traditional criteria cannot be used, or where more traditional species identification needs to be verified.51 ZooMs is much cheaper than aDNA analysis, and less time-consuming, as proteins are easier to extract and have better chance of preservation.52 One drawback of most of these analytical methods is that they are usually destructive, although in many cases the sample size is very small and the same sample can potentially be used for multiple techniques.

Because further methodological developments can be expected, it is advisable to retain faunal remains that have been studied by archaeozoologists and to store them in appropriate conditions to allow for new methods to be applied on them. At the moment, none of the analytical methods can replace traditional archaeozoological methods, and the two remain complementary. Moreover, for the selection or sampling of bones for analytical methods, an archaeozoological specialist is still needed, for example because certain species or certain skeletal elements are to be targeted.

Databases

Archaeozoologists enter the data they collect into a computer spreadsheet or database; most no longer record on paper. There are initiatives for common databases, where data are entered in a standardized way, which should allow for easier data exchange (e.g., Ossobook), but to our knowledge no such system is used yet in Africa.53 There have also been some attempts to create online common databases for Africanist archaeozoologists, but at the moment nothing is up and running. A recent book gives distribution maps of larger mammals in Africa through time, based on a large compilation of archaeozoological data.54 A similar but older book exists for southern Africa.55

Research Themes

Although research questions addressed by archaeozoologists vary according to the specific region and period they are dealing with, they can be grouped into a few general, recurrent, and often interrelated themes. In what follows, some of the major themes relevant to the later parts of human history are discussed. Here, too, archaeozoologists do not operate in isolation but, rather, together with archaeologists and other specialists.

Diet

Archaeozoologists usually attempt to reconstruct past human diet, at least as far as the animal component is concerned, and archaeozoological reports will often have the word diet in their titles.56 However, because of diverse problems of preservation, quantification of results, etc., it is extremely difficult to estimate how much of the different types of meat and other animal food items were consumed, not just relative to each other, but also, and perhaps more importantly, relative to plant foods. Where possible, nowadays, stable isotope analysis of human remains is increasingly carried out for dietary reconstruction.57 This type of analysis allows archaeozoologists to fill the gap in determining the relative importance of major food groups, such as plant food versus meat, or (marine) fish versus mammals. However, unlike traditional archaeozoological studies, stable isotope analysis does not allow us to determine which particular taxa were consumed, thus, the two methods for dietary reconstruction remain complementary. Despite the problems inherent in quantifying dietary importance, regional comparisons of quantitative archaeozoological data can reveal interesting patterns.58 In West Africa, fish bones are more common at sites near larger water basins, although this does not allow archaeozoologists to say how much fish was consumed exactly.59 While the consumption of meat and body fat will leave traces in the form of bones, the use of milk and blood will not leave comparable evidence. In theory, mortality profiles will differ in domesticated animals when they were used for milk, but, as indicated above, African sites rarely yield sufficient data to construct such profiles. However, organic residue analysis of pottery from the Central Sahara has shown that milk was used in Africa from at least the late 6th millennium bce.60 Related to diet and the consumption of animals, archaeozoologists also attempt to reconstruct the ways animals were butchered, their carcasses divided, and their meat prepared. For example, it has been suggested that a reduction in the proportion of burned bones at the site of Siouré in the Senegal Valley in the course of its occupation (mainly 1st millennium ce) is linked to increased boiling instead of roasting of meat.61

Subsistence Strategies

Diet is inevitably linked to the subsistence activities people carried out. Were they herding, fishing, hunting, or a combination of these; did they obtain food through contacts with other people engaged in these activities? A major theme in African archaeology is the introduction of food production and the arrival of the first domesticated animals.62 Very generally speaking, domesticates appear later in time the farther away one moves from northeastern Africa, because the earliest introductions were through the Levant. The continued exploitation of wild animal resources by hunting and fishing, after the arrival of domesticates, is characteristic for certain parts of Africa and appears to relate to the difficulties of keeping domesticated taxa (mainly due to disease risks), rather than to the richness in wild resources. This seems to be the case, for example, in woodland areas, such as most of Ghana, as well as in wetlands.63 Although skeletal distribution can allow for a distinction to be made between consumption sites (containing mainly meat-bearing bones) and production sites (containing more bones that would have had little adhering meat), the distinction is usually not straightforward. High proportions of vertebrae of Nile perch (Lates niloticus) at the site of Garumele (northern Niger, 1500–1800 ce) may indicate that fish were processed elsewhere and that only the flesh-bearing parts were transported to the site.64 Another example of a consumption site comes from Old Kingdom Egypt (3rd millennium ce), where, based on the species distribution and the age and sex of the animals, it has been suggested that the workmen and their overseers building the pyramids at Giza were being provisioned with food.65

Cultural Behavior

Although the environment played a major role in subsistence strategies, many choices were also cultural, particularly when societies became more complex. In some parts of West Africa, such as the Inland Niger Delta of Mali, different ethnic groups have different specializations (fishing, farming, or herding), and researchers have attempted to extrapolate these back in time, based on faunal evidence, among other methods.66 Recent osteometric studies combined with isotopic analysis of domestic animals from the archaeological site of Jenné-Jeno (Mali, c. 250 bce–1400 ce) suggest that subsistence strategies were much more diversified than previously thought and that ethnically linked subsistence specialization emerged only relatively recently.67 Which animals were considered edible and which ones were not was often culturally determined. Dogs are a typical example of animals that were eaten by some ethnic groups, but whose consumption was considered gruesome by others. In northern Burkina Faso, there is clear evidence for dog eating for the period between 1000 and 1400 ce, at a time of influences from northern African peoples, Berbers, and Islamic Arabs, through trans-Saharan trade.68 North African Berbers were known for eating dogs, while Islam considered dogs impure. Animal burials and finds of animals in ritual contexts also testify to the symbolic role of animals, usually specific species. Chickens, in particular, were probably popular in ritual spheres. However, some sacrificial practices may have left few archaeological traces, because sacrificed animals were not buried and therefore quickly taken away by predatory animals.69

Interregional Connections and Trade

Most domesticated animals are not native to Africa, and their distribution is revealing for our understanding of population movements and interregional contacts. While the first domesticates—namely, cattle, sheep, goat, and dog—arrived relatively early in the Holocene, other taxa and types arrived later. Bone finds from Ethiopia, dated at the latest to c. 600 bce, for example, indicate that chickens were introduced to the Horn of Africa through the maritime Red Sea trade.70 Genetic studies of modern African cattle show that zebu cattle were probably introduced through the same route.71 The presence of zebu is harder to prove from archaeological bone remains because they are difficult to distinguish from other types of cattle with traditional methods. Several commensal animal species, including black rat (Rattus rattus), also arrived from Asia via the East African coast, where their remains are found at archaeological sites from at least the second half of the 1st millennium ce.72 Another major route for the spread of domesticates was through trans-Saharan trade between coastal northern African and sub-Saharan Africa, a probable route for the spread of chicken, horse, donkey, and dromedary. While chickens were relatively common because they were eaten, the latter three were much rarer in the archaeological record.73 European contacts resulted in the introduction of new taxa, such as pig (Sus domesticus) and rabbit (Oryctolagus cuniculus), but Europeans also brought with them sheep, goat, cattle, and other species that were already present in Africa.74 Bones of wild animals can also prove interregional contacts. Marine fish at Aksum, in inland Ethiopia (1st millennium ce), for example, testify to trade connections with ports on the Red Sea.75 Ivory is a typical example of an animal product traded over long distances. Hippo as well as elephant ivory is found across most of Africa. Stable isotope analysis provides opportunities to provenance this ivory.76

Human–Environment Interaction

Apart from human dietary reconstructions, probably the most important category of interpretations of archaeozoological data is the reconstruction of the former environment, and archaeozoological reports typically contain a section on the paleo-environment.77 Reconstructions are based on the habitat requirements of the wild animal taxa represented in the bone assemblages, taking into account, of course, that (parts of) animals may have been brought in from farther away. Moreover, in mammals, the larger the animal, the larger its living range. Nevertheless, based on shifts in species composition, archaeozoological studies have successfully shown local environmental changes linked to (global) climate change, to which humans had to adapt their survival strategies.78 After the mid-Holocene, adaptations to more arid climates in large parts of Africa appear often to have coincided with an increase in the use of domesticated resources.79 The relative importance of the major domesticated animal species can also be used to make general inferences on the environment. Cattle need sufficient good grazing and water, while sheep and goat are usually less demanding, goat being the more hardy of the two.80 The composition of the domestic livestock herds is determined by both the environment and the mobility pattern of the people keeping them. Sedentary groups of the Sahel in Burkina Faso keep mainly sheep and goat, while cattle are more common among mobile groups.81 The same domesticated species will also look different depending on the environment. For example, long-legged sheep are common in arid areas, while dwarf goat and cattle are common in humid environments; this can also be shown archaeologically.82 Seasonal variation in the occurrence and abundance of natural resources affects human activities and settlement patterns. Fish, for example, are a resource that is typically easier to catch in certain parts of the year.83

It is clear that climate and environment had a major influence on human lifeways. Conversely, humans have also had an impact on their environment. Human predation can lead, for example, to smaller sized individuals of a certain animal taxon, and it has been argued that a size reduction of fish in the Senegal River valley during the 1st millennium ce, as inferred from archaeozoological data from the sites of Cubalel and Siouré, was due to overfishing.84 Populations of a certain species may also have disappeared or been locally reduced due to human pressure. A clear reduction in numbers of bones of kob (Kobus kob), a species of antelope, through time observed at the site of Mege (northeastern Nigeria, c. 550 bce–late-20th century ce) may be read as a consequence of overhunting.85 Kob is quite vulnerable to human exploitation due to its territorial behavior; it continues to live in an area after humans settle there. A reduction in wild game due to intensive hunting in the Iron Age of Southern Africa caused people to rely more on domesticated cattle.86

Discussion of the Literature

In the first half of the 20th century ce, data on archaeological animal remains were, at best, published as appendices to archaeological reports, which were often not more than a list of the taxa identified. Collection usually happened selectively, and remains were not retained after study, making it impossible for other researchers to verify the identifications.87 Since approximately the 1970s, the collection and study of archaeological animal remains from Africa has improved, coinciding with worldwide developments in the discipline and founding of the International Council of Archaeozoology, in 1976. Since the 1970s and 1980s, the range of issues addressed by archaeozoologists has widened, from dietary and behavioral–ecological aspects, to the social role of animals in human societies.88 But, though the research potential of archaeozoological remains has been proven, even today, animal remains from African archaeological sites are sometimes not well collected and/or not studied. There are geographic differences, for example, with particularly large numbers of good studies from southern Africa.89 There are also differences relating to the age of the sites. Traditionally, animal remains have received more attention from pre-historians than from historical archaeologists, a situation that continues to some extent today. This is mainly due to the fact that the former usually don’t have as many sources of information available to them. As a consequence, they use these sources to the maximum extent. However, it should be clear that archaeozoological research can yield new and relevant information for the reconstruction of human lifeways at any point in the past. In fact, the combined use of written sources and archaeozoological evidence can be mutually reinforcing.

Archaeozoologists working in a commercial setting in some countries outside of Africa have guidelines for good practice.90 Archaeozoologists working in Africa could benefit also from setting up some kind of guide of good practice. Good practice actually starts in the field, with the choice of the area to be excavated, as well as the excavation and collection methods. The direction that archaeozoology is taking globally is clearly linked to new and improving analytical methods, such as aDNA and stable isotope analysis, but these are combined with traditional methods for the moment. Another trend, perhaps a bit less strong in Africa, is the meta-analysis of large datasets, involving multiple sites, which has become easier with improved computer technology.91 While some parts of archaeozoological methodology, such as bone measurements, are standardized, others are not. Data exchange and compilations of large datasets would benefit from more standardization. However, not all aspects are as easy to standardize or to agree upon among researchers.92

One theme in African archaeozoology of the later Holocene that continues to dominate much of the research is the history of domesticated animals, including when they appeared and the routes through which they were spread. This is a domain in which genetic studies are now playing a crucial role, based not only on aDNA, but also on the DNA of living African domesticated animals. Most domesticated animal species were introduced from outside of Africa. Since the 1980s, there has been debate on a possible local African domestication of cattle, and this has not yet been completely resolved.93 Today, the constraining role of disease on the spread of domesticated animals over the African continent is receiving renewed attention.94 A more general archaeological research theme to which archaeozoologists in Africa have often contributed is the rise of complex societies.95 In Africa, archaeozoological research seems to be more frequently linked with ethno-archaeological research than it is in some other parts of the world, because of the persistence of traditional livelihoods. Through analogy, this allows us to better interpret the archaeological record, for example, concerning butchery and hide working practices.96 Moreover, it sheds light on aspects that are nearly impossible to understand based on pure archaeological evidence, such as gender issues in food preparation and consumption, or the ritual meaning of certain animal species.97 Beyond this, research topics are varied, depending on space and time, as well as on the overarching goals of archaeological research projects. However, diet, trade connections, ritual, and other cultural behaviors are recurrent themes.

Primary Sources

The most basic primary source of archaeozoological data is newly excavated, collected, and studied animal remains. In view of the crucial importance of good comparative collections for the study of animal remains, it is useful to know where such collections can be found. Inside Africa, a few places exist: for instance, the Ditsong National Museum of Natural History (Pretoria and Johannesburg, South Africa), the National Museums of Kenya (Nairobi), and the IFAN Museum of African Arts (Dakar, Senegal). Outside Africa, the Natural History Museum (London, UK), the Royal Belgian Institute of Natural Sciences (Brussels), the Royal Museum for Central Africa (Tervuren, Belgium), the Muséum national d’Histoire naturelle (Paris, France), and the Smithsonian Institution National Museum of Natural History (Washington, DC, USA) have good reference collections of African fauna. Some of the institutions mentioned also house collections of animal remains from African archaeological sites, but generally archaeological bones are spread over various storage facilities for antiquity museums, excavations houses, university departments, etc. Many archaeozoological reports mention where the studied bones are stored. Where this is not the case, or when there are no faunal reports, the search for the actual bones will be more difficult and will likely involve contacting archaeologists and archaeozoologists.

A good starting point when looking for published archaeozoological data, especially for a certain African country, is the newly published Atlas of Mammal Distribution through Africa from the LGM (18 Ka) to Modern Times, by Hélène Jousse. It contains references to the body of archaeozoological reports published until 2009. Most raw data are typically to be found in PhD theses, in site monographs, and more in online appendices to articles in scholarly journals. Examples of journals that have published archaeozoological data from Africa are African Archaeological Review; Azania: Archaeological Research in Africa; Journal of African Archaeology; International Journal of Osteoarchaeology; and Quaternary International. Attempts to set up common databases and online repositories have not been successful so far. The international archaeozoological community has a very active online discussion list: ZOOARCH Home Page, where a wide range of questions are asked and answered and conferences and job postings are announced. An online archive of this list is helpful when looking for information and the latest views on a certain topic.98

Further Reading

Albarella, Umberto, Mauro Rizzetto, Hannah Russ, Kim Vickers, and Sarah Viner-Daniels. The Oxford Handbook of Zooarchaeology. Oxford, UK: Oxford University Press, 2017.Find this resource:

Blench, Roger M., and Kevin C. MacDonald. The Origins and Development of African Livestock: Archaeology, Genetics, Linguistics, and Ethnography. London, UK: UCL Press, 2000.Find this resource:

Davis, Simon. The Archaeology of Animal Bones. London, UK: Routledge, 1987.Find this resource:

Driesch, Angela von den. A Guide to the Measurement of Animal Bones from Archaeological Sites, vol. 1. Peabody Museum Bulletin. Cambridge, MA: Harvard Peabody Museum Press, 1976.Find this resource:

Jousse, Hélène. Atlas of Mammal Distribution through Africa from the LGM (18 Ka) to Modern Times: The Zooarchaeological Record. Oxford, UK: Archaeopress, 2017.Find this resource:

O’Connor, Terry. The Archaeology of Animal Bones. Stroud, UK: Sutton Publishing, 2000.Find this resource:

Plug, Ina. What Bone Is That? A Guide to the Identification of Southern African Mammal Bones. Cape Town, South Africa: Rosslyn Press, 2014.Find this resource:

Reitz, Elizabeth J., and Elisabeth S. Wing. Zooarchaeology (2nd ed.). Cambridge, UK: Cambridge University Press, 2008.Find this resource:

Schmid, Elisabeth. Atlas of Animal Bones: For Prehistorians, Archaeologists, and Quaternary Geologists. Knochenatlas: Für Prähistoriker, Archäologen, und Quartärgeologen. New York, NY: Elsevier, 1972.Find this resource:

Sykes, Naomi. Beastly Questions: Animal Answers to Archaeological Issues. London, UK: Bloomsbury Academic, 2014.Find this resource:

Walker, Rikki. A Guide to the Post-Cranial Bones of East African Mammals. Norwich, UK: Hylochoerus Press, 1985.Find this resource:

Worley, Fay, and Polydora Baker. Animal Bones and Archaeology: Guidelines for Best Practice. Swindon, UK: English Heritage, 2014.Find this resource:

Notes:

(1.) Gill Campbell, Lisa Moffett, and Vanessa Straker, Environmental Archaeology: A Guide to the Theory and Practice of Methods, from Sampling and Recovery to Post-Excavation (2nd ed.; Swindon, UK: English Heritage, 2011).

(2.) Elizabeth J. Reitz, and Elisabeth S. Wing, Zooarchaeology (2nd ed.; Cambridge, UK: Cambridge University Press, 2008), 34 and Table 3.3.

(3.) Peter Breunig and Nicole Rupp, “An Outline of Recent Studies on the Nigerian Nok Culture,” Journal of African Archaeology 14 (2016): 237–256.

(4.) Wim Van Neer, “Domestic Animals from Archaeological Sites in Central and West-Central Africa,” in The Origins and Development of African Livestock: Archaeology, Genetics, and Ethnography, ed. Roger M. Blench and Kevin C. MacDonald (London, UK: UCL Press, 2000), 163–190.

(5.) Veerle Linseele, Wim Van Neer, Harco Willems, and Bart Vanthuyne, “An Unusual Cattle Burial at Dayr Al-Barshā (Middle Egypt),” in Archaeozoology of the Near East 9, eds. Marjan Mashkour and Mark Beech (Oxford, UK: Oxbow Books, 2017), 353–377.

(6.) Hans-Peter Wotzka, Studien zur Archäologie des Zentralafrikanischen Regenwaldes: Die Keramik des Inneren Zaïre-Beckens und Ihre Stellung im Kontext der Bantu-Expansion (Cologne, Germany: Heinrich-Barth-Institut, 1995), 369–381.

(7.) For example, Veerle Linseele, Archaeofaunal Remains from the Past 4000 Years in Sahelian West Africa: Domestic Livestock, Subsistence Strategies and Environmental Changes (Oxford, UK: Archaeopress, 2007).

(8.) Wim Van Neer, “Fishing in the Senegal River during the Iron Age: The Evidence from the Habitation Mounds of Cubalel and Siouré,” in Animals and People: Archaeozoological Papers in Honour of Ina Plug, eds. Shaw Badenhorst, Peter Mitchell, and Jonathan C. Driver (Oxford, UK: Archaeopress, 2008), 117–130.

(9.) Achilles Gautier, “Taphonomic Groups: How and Why?” Archaeozoologia 1 (1987): 47–52.

(10.) Linseele, Archaeofaunal Remains.

(11.) Reitz and Wing, Zooarchaeology, 147–150; and Campbell, Moffett, and Straker, Environmental Archaeology.

(12.) Edgard O. Espinoza and Mary-Jacque Mann, Identification Guide for Ivory and Ivory Substitute (Baltimore, MD: The World Wildlife Fund & the Conservation Foundation, 1991).

(13.) Anthea W. Gentry, Juliet Clutton-Brock, and Colin P. Groves, “The Naming of Wild Animal Species and Their Domestic Derivatives,” Journal of Archaeological Science 31 (2004): 644–651.

(14.) For example, Keith Dobney and Kevin Rielly, “A Method for Recording Archaeological Animal Bones: The Use of Diagnostic Zones,” Circaea 5 (1988): 79–96.

(15.) Rikki Walker, A Guide to the Post-Cranial Bones of East African Mammals (Norwich, UK: Hylochoerus Press, 1985); and Ina Plug, What Bone Is That? A Guide to the Identification of Southern African Mammal Bones (Cape Town, South Africa: Rosslyn Press, 2014).

(16.) Wim Van Neer, Contribution to the Archaeozoology of Central Africa (Tervuren, Belgium: Royal Museum for Central Africa, 1989); Joris Peters, Osteomorphology and Osteometry of the Appendicular Skeleton of Grant’s Gazelle, Gazella granti (Brooke, 1872), Bohor Reedbuck, Redunca redunca (Pallas, 1767) and Bushbuck, Tragelaphus scriptus (Pallas, 1766) (Ghent, Belgium: Laboratorium voor Paleontologie, Ghent University, 1986); and Joris Peters, Wim Van Neer, and Ina Plug, Comparative Postcranial Osteology of Hartebeest (Alcelaphus buselaphus), Scimitar Oryx (Oryx dammah) and Addax (Addax nasomaculatus) with Notes on the Osteometry of Gemsbok (Oryx gazella) and Arabian Oryx (Oryx leucoryx) (Tervuren, Belgium: Royal Museum of Central Africa, 1997).

(17.) Melinda A. Zeder and Heather A. Lapham, “Assessing the Reliability of Criteria Used to Identify Postcranial Bones in Sheep, Ovis, and Goats, Capra,” Journal of Archaeological Science 37 (2010): 2887–2905; Melinda A. Zeder and Suzanne E. Pilaar, “Assessing the Reliability of Criteria Used to Identify Mandibles and Mandibular Teeth in Sheep, Ovis, and Goats, Capra,” Journal of Archaeological Science 37 (2010): 225–242, and references therein.

(18.) Joris Peters, Osteomorphology and Osteometry of the Appendicular Skeleton of African Buffalo, Syncerus caffer (Sparrman, 1779) and Cattle, Bos primigenius f. taurus (Bojanus, 1827) (Ghent, Belgium: Ghent University, 1986).

(19.) C. K. Brain, “Some Suggested Procedures in the Analysis of Bone Accumulations from Southern African Quaternary Sites,” Annals of the Transvaal Museum 29 (1974): 1–8.

(20.) For example, Kristina Douglass et al., “Multi-Analytical Approach to Zooarchaeological Assemblages Elucidates Late Holocene Coastal Lifeways in Southwest Madagascar,” Quaternary International 471 (2018): 111–131.

(21.) Plug, What Bone Is That? 11.

(22.) On frequency of the taxa, see Reitz and Wing, Zooarchaeology, 202–213.

(23.) Wim Van Neer, “Food Security in Western and Central Africa During the Late Holocene: The Role of Domestic Stock Keeping, Hunting, and Fishing,” in Droughts, Food, and Culture: Ecological Change and Food Security in Africa’s Later Prehistory, ed. Fekri A. Hassan (New York, NY: Kluwer Academic/Plenum, 2002), 251–274, and see also discussion on quantification therein.

(24.) On estimating numbers of elements (MNE), R. Lee Lyman, Vertebrate Taphonomy (Cambridge, UK: Cambridge University Press, 1994), 102–104. On estimating numbers of specimens (MNISP), see Kevin C. MacDonald and Rachel H. MacDonald, “Mammalian, Avian, and Reptilian Remains,” in The Search for Trakrur: Archaeological Excavations and Reconnaissance Along the Middle Senegal Valley, ed. Roderick J. McIntosh, Susan K. McIntosh, and Hamady Bocoum (New Haven, CT: Yale University Department of Anthropology and Yale Peabody Museum of Natural History, 2016), 311–334.

(25.) I. A. Silver, “The Ageing of Domestic Animals,” in Science in Archaeology, ed. Don Brothwell and Eric Higgs (London, UK: Thames and Hudson, 1963), 250–268; Karl-Heinz Habermehl, Die Altersbestimmung bei Haus- und Labortieren (Hamburg, Germany: Verlag Paul Parey, 1975); Karl-Heinz Habermehl, Die Altersbestimmung bei Wild- und Pelztieren (Hamburg, Germany: Verlag Paul Parey, 1985); Melinda A. Zeder, “Reconciling Rates of Long Bone Fusion and Tooth Eruption and Wear in Sheep (Ovis) and Goat (Capra),” in Recent Advances in Ageing and Sexing Animal Bones: Proceedings of the 9th Conference of the International Council of Archaeozoology, Durham, August 2002, ed. Deborah Ruscillo (Oxford, UK: Oxbow Books, 2006), 87–118.

(26.) Habermehl, Altersbestimmung Haus- und Labortieren.

(27.) Silver, “Ageing of Domestic Animals”; Sebastian Payne, “Kill-off Patterns in Sheep and Goats: The Mandibles from Asvan Kale,” Anatolian Studies 23 (1973): 281–303; and Zeder, “Reconciling Rates of Long Bone Fusion.”

(28.) Payne, “Kill-off Patterns in Sheep and Goats”; and Jean-Denis Vigne and Daniel Helmer, “Was Milk a ‘Secondary Product’ in the Old World Neolithisation Process? Its Role in the Domestication of Cattle, Sheep, and Goats,” Anthropozoologica 42 (2007): 9–40.

(29.) But see, for example, MacDonald and MacDonald, “Mammalian, Avian, and Reptilian Remains.”

(30.) Angela von den Driesch, A Guide to the Measurement of Animal Bones from Archaeological Sites, vol. 1, Peabody Museum Bulletin (Cambridge, MA: Harvard Peabody Museum Press, 1976).

(31.) For example, Van Neer, Contribution à l’Ostéométrie, for Nile perch.

(32.) For example, Kevin C. MacDonald and Rachel H. MacDonald, “The Origins and Development of Domesticated Animals in Arid West-Africa,” in The Origins and Development of African Livestock: Archaeology, Genetics, Linguistics and Ethnography, ed. Roger M. Blench and Kevin C. MacDonald (London, UK: UCL Press, 2000), 127–162.

(33.) Richard G. Klein and Kathryn Cruz-Uribe “Size Variation in the Rock Hyrax (Procavia capensis) and Late Quaternary Climatic Change in South Africa,” Quaternary Research 46 (1996): 193–207.

(34.) Sensu Richard H. Meadow, “The Use of Size Index Scaling Techniques for Research on Archaeozoological Collections from the Middle East,” in Historia Animalium ex Ossibus: Beiträge zur Paläoanatomie, Archäologie, Ägyptologie, Ethnology und Geschichte der Tiermedezin: Festschrift für A. von den Driesch zum 65. Geburtstag, ed. Cornelia Becker, Henriette Manhart, Joris Peters, and Jorg Schibler (Rahden, Germany: Verlag Marie Leidorf, 1999), 285–300.

(35.) Summarized in Angela von den Driesch and Joachim Boessneck, “Kritische Anmerkungen zur Widerristhöhenberechnung aus Längenmassen Vor- und Frühgeschichtlicher Tierknochen,” Säugetierkundige Mitteilungen 22 (1974): 325–348.

(36.) For example, Van Neer, Contribution à l’Ostéométrie, for Nile perch.

(37.) For example, Yolanda Fernández-Jalvo and Peter Andrews, Atlas of Taphonomic Identifications (Dordrecht, Netherlands: Springer, 2016).

(38.) Anna K. Behrensmeyer, “Taphonomic and Ecologic Information from Bone Weathering,” Paleobiology 4 (1978): 150–162.

(39.) Sebastian Payne and Patrick J. Munson, “Ruby and How Many Squirrels? The Destruction of Bones by Dogs,” in Palaeobiological Investigations: Research Design, Methods, and Data Analysis, eds. N. R. J. Fieller, D. D. Gilbertson, and N. G. A. Ralph (Oxford, UK: British Archaeological Reports, 1985), 31–40.

(40.) See summary in Lyman, Vertebrate Taphonomy, Fig. 9.9.

(41.) Harco Willems et al., “An Industrial Site at Al-Shaykh Sacīd/Wādī Zabayda,” Ägypten und Levante 19 (2009): 293–331.

(42.) Terry O’Connor, The Archaeology of Animal Bones (Stroud, UK: Sutton Publishing, 2000), 45–46.

(43.) Haskel Greenfield, “The Origins of Metallurgy: Distinguishing Stone from Metal Cut-Marks on Bones from Archaeological Sites,” Journal of Archaeological Science 26 (1999): 797–808.

(44.) Veerle Linseele, “Cultural Identity and the Consumption of Dogs in Western Africa,” in Behaviour Behind Bones: The Zooarchaeology of Ritual, Religion, Status and Identity, ed. Sharon Jones O’Day, Wim Van Neer, and Anton Ervynck (Oxford, UK: Oxbow Books, 2003), 318–326.

(45.) Stine Rossel et al., “Domestication of the Donkey: Timing, Processes, and Indicators,” Proceedings of the National Academy of Science USA 105 (2008): 3715–3720.

(46.) László Bartosiewicz, Wim Van Neer, and An Lentacker, Draught Cattle: Their Osteological Identification and History (Tervuren, Belgium: Royal Museum for Central Africa, 1997).

(47.) Emma Loftus, Patrick Roberts, and Julia A. Lee-Thorp, “An Isotopic Generation: Four Decades of Stable Isotope Analysis in African Archaeology,” Azania: Archaeological Research in Africa 51 (2016): 88–114.

(48.) Diane Gifford-Gonzalez and Olivier Hanotte, “Domesticating Animals in Africa: Implications of Genetic and Archaeological Findings,” Journal of World Prehistory 24 (2011): 1–23.

(49.) K. Ann Horsburgh, Jayson Orton, and Richard G. Klein, “Beware the Springbok in Sheep’s Clothing: How Secure Are the Faunal Identifications upon which We Build Our Models?” African Archaeological Review 33 (2016): 353–361; K. Ann Horsburgh, “A Reply to Plug 2017: Science Requires Self-Correction,” Azania: Archaeological Research in Africa 53 (2017): 114–118; and Ina Plug, “Reply to Horsburgh et al. 2016: ‘Revisiting the Kalahari Debate in the Highlands’,” Azania: Archaeological Research in Africa 53 (2017): 98–113.

(50.) Alicia Grealy et al., “Tropical Ancient DNA from Bulk Archaeological Fish Bone Reveals the Subsistence Practices of a Historic Coastal Community in Southwest Madagascar,” Journal of Archaeological Science 75 (2016): 82–88.

(51.) Michael Buckley and Caroline Wadsworth, “Proteome Degradation in Ancient Bone: Diagenesis and Phylogenetic Potential,” Palaeogeography, Palaeoclimatology, Palaeoecology 416 (2014): 69–79; Nienke L. Van Doorn, “Zooarchaeology by Mass Spectrometry (Zooms),” in Encyclopedia of Global Archaeology, ed. Claire Smith (New York, NY: Springer, 2014), 7998–8000.

(52.) Caroline Wadsworth et al. “Comparing ancient DNA survival and proteome content in 69 archaeological cattle tooth and bone samples from multiple European sites,” Journal of Proteomics 158 (2017): 1–8.

(53.) Daniel Kaltenthaler et al. 2017. Ossobook, Version 5.6.2, Ludwig Maximilian University of Munich.

(54.) Hélène Jousse, Atlas of Mammal Distribution through Africa from the LGM (18 Ka) to Modern Times: The Zooarchaeological Record (Oxford, UK: Archaeopress, 2017).

(55.) Ina Plug and Shaw Badenhorst, Distribution of Macromammals in Southern Africa over the Past 30,000 Years as Reflected in Animal Remains from Archaeological Sites (Pretoria, South Africa: Transvaal Museum, 2001).

(56.) Nadja Pöllath and Anna-Katharina Rieger, “Insights in Diet and Economy of the Eastern Marmarica. Faunal Remains from Greco-Roman Sites in North-Western Egypt (Abar-El-Kanayis, Wadi Umm El-Ashdan and Wadi Qasaba),” Mitteilungen des Deutschen Archäologischen Instituts, Abteilung Kairo 67 (2013): 163–180.

(57.) Loftus, Roberts, and Lee-Thorp, “An Isotopic Generation.”

(58.) Such as the ones shown by Van Neer, “Food Security in West and Central Africa during the Late Holocene.”

(59.) Veerle Linseele, “The Exploitation of Aquatic Resources in Holocene West Africa,” in The Oxford Handbook of Zooarchaeology, ed. Umbert Albarella, Mauro Rizzetto, Hannah Russ, Kim Vickers, and Sarah Viner-Daniels (Oxford, UK: Oxford University Press, 2017), 439–451.

(60.) Julie Dunne, Savino di Lernia, Marek Chłodnicki, Farid Kherbouche, and Richard P. Evershed, “Timing and Pace of Dairying Inception and Animal Husbandry Practices across Holocene North Africa,” Quaternary International 471 (2018): 147–159.

(61.) MacDonald and MacDonald, “Mammalian, Avian, and Reptilian Remains.”

(62.) Blench and MacDonald, Origins and Development of African Livestock; Richard T. Chia and A. Catherine D’Andrea, Food Production in the Forest Zone of West Africa: Archaeological and Historical Perspectives; Freda Nkirote M’Mbogori, Farming and Herding in Eastern Africa: Archaeological and Historical Perspectives; Paul Lane and Anna Shoemaker, Interdisciplinary Perspectives on Precolonial Sub-Saharan African Farming and Herding Communities, Oxford Research Encyclopedias, African History.

(63.) For example, Achilles Gautier and Wim Van Neer, “The Continuous Exploitation of Wild Animal Resources in the Archaeozoological Record of Ghana,” Journal of African Archaeology 3 (2005): 195–212; Josette Rivallain and Wim Van Neer, “Les Fouilles de Koyom (Sud du Tchad). Etude du Matériel Archéologique et Faunique,” L’Anthropologie 87 (1983): 221–239; and Josette Rivallain and Wim Van Neer, “Inventaire du Matériel Archéologique et Faunique de Koyom, Sud du Tchad,” L’Anthropologie 88 (1984): 441–448.

(64.) Veerle Linseele and Anne Haour, “Animal Remains from Medieval Garumele (Niger),” Journal of African Archaeology 8 (2010): 167–184.

(65.) Richard Redding, “Status and Diet at the Workers’ Town, Giza, Egypt,” in Anthropological Approaches to Zooarchaeology: Colonialism, Complexity and Animal Transformations, eds. Douglas Campana, Pamela Crabtree, Susan deFrance, Justin S. Lev-Tov, and Alice M. Choyke (Oxford, UK: Oxbow Books, 2010), 65–75.

(66.) Kevin C. MacDonald and Wim Van Neer, “Specialised Fishing Peoples in the Later Holocene of the Méma Region (Mali),” in Fish Exploitation in the Past, Proceedings of the 7th Meeting of the ICAZ Fish Remains Working Group, ed. Wim Van Neer (Tervuren, Belgium: Royal Museum for Central Africa, 1994), 243–251.

(67.) Abigail Chipps Stone, “Finding the Ephemeral: Herding Strategies and Socio-Economic Organization in an Urban West African Context,” Quaternary International 471 (2018): 160–174.

(68.) Linseele, “Cultural Identity and the Consumption of Dogs.”

(69.) Timothy Insoll, “Talensi Animal Sacrifice and Its Archaeological Implications,” World Archaeology 42 (2010): 231–244.

(70.) Helena S. Woldekiros and A. Catherine D’Andrea, “Early Evidence for Domestic Chickens (Gallus gallus domesticus) in the Horn of Africa,” International Journal of Osteoarchaeology 27, no. 3 (2017): 329–341.

(71.) Olivier Hanotte et al., “African Pastoralism: Genetic Imprints of Origins and Migrations,” Science 296 (2002): 336–339.

(72.) Nicole Boivin, Alison Crowther, Richard Helm, and Dorian Q. Fuller, “East Africa and Madagascar in the Indian Ocean World,” Journal of World Prehistory 26 (2013): 213–281; Mary E. Prendergast et al., “Dietary Diversity on the Swahili Coast: The Fauna from Two Zanzibar Trading Locales,” International Journal of Osteoarchaeology 27 (2017): 621–637.

(73.) Stephen Dueppen, “Evidence for Chickens at Iron Age Kirikongo (c. AD 100–1450), Burkina Faso,” Antiquity 85 (2011): 142–157; and MacDonald and MacDonald, “Origins and Development of Domesticated Animals.”

(74.) Adam Heinrich, “Faunal Analysis and the Development of the Meat Industry in the 17th and 18th Centuries,” in Historical Archaeology in South Africa: Material Culture of the Dutch East India Company at the Cape, ed. Carmel Shrire (Walnut Creek, CA: Left Coast Press, 2014), 65–100.

(75.) Louis Chaix, “The Fauna from the Uno/Bu Excavations at Bieta Giyorgis (Aksum) in Tigray, Northern Ethiopia: Campaigns 1995–2003; Pre-Aksumite, 700–400 BC to Late Aksumite, AD 800–1200,” Journal of African Archaeology 11 (2013): 211–241.

(77.) Sharon E. Nicholson, “Climatology: Methods,” The Oxford Research Encyclopedias, African History.

(78.) Nadja Pöllath, “Surviving in a Profoundly Changing Landscape: The Mid-Holocene Archaeofaunal Record from Abu Tabari (NW-Sudan),” in People and Animals in Holocene Africa: Recent Advances in Archaeozoology, eds. Hélène Jousse and Joséphine Lesur (Frankfurt-am-Main, Germany: Africa Magna Verlag, 2011), 59–74.

(79.) Hélène Jousse, “What Is the Impact of Holocene Climatic Changes on Human Societies? Analysis of West African Neolithic Populations Dietary Customs,” Quaternary International 151 (2006): 63–73.

(80.) Gudrun Dahl and Anders Hjort, Having Herds: Pastoral Herd Growth and Household Economy (Stockholm, Sweden: Liber Tryck, 1976).

(81.) Linseele, Archaeofaunal Remains from the Past.

(82.) Stephen A. Dueppen, “Cattle in the West African Savanna: Evidence from 1st Millennium ce Kirikongo, Burkina Faso,” Journal of Archaeological Science 39 (2012): 92–101.

(83.) Wim Van Neer, “Evolution of Prehistoric Fishing in the Egyptian Nile Valley,” Journal of African Archaeology 2 (2004): 251–269; and Ina Plug, Peter Mitchell, and Geoff Bailey, “Late Holocene Fishing Strategies in Southern Africa as Seen from Likoaeng, Highland Lesotho,” Journal of Archaeological Science 37 (2010): 3111–3123.

(84.) Wim Van Neer, “Fishing in the Senegal River during the Iron Age: The Evidence from the Habitation Mounds of Cubalel and Siouré,” in Animals and People: Archaeozoological Papers in Honour of Ina Plug, eds. Shaw Badenhorst, Peter Mitchell, and Jonathan C. Driver (Oxford, UK: Archaeopress, 2008), 117–130.

(85.) Linseele, Archaeofaunal Remains from the Past.

(86.) Shaw Badenhorst, “Intensive Hunting During the Iron Age of Southern Africa,” Environmental Archaeology 20 (2015): 41–51.

(87.) Joris Peters, “A Revision of the Faunal Remains from Two Central Sudanese Sites: Khartoum Hospital and Esh Shaheinab,” in Archaeozoologia: Mélanges: Publiés à lʼoccasion du 5e Congrès International dʼarchéozoologie, Bordeaux – Août 1986, ed. Pierre Ducos (Grenoble, France: La Pensée Sauvage Éditions, 1986), 11–35.

(88.) Naomi Sykes, Beastly Questions: Animal Answers to Archaeological Issues (London, UK: Bloomsbury Academic, 2014).

(89.) Annie R. Antonites, Karin Scott, and Evin Grody, “New Directions in South African Archaeozoology of the Last 2,000 years,” African Archaeological Review 33 (2016): 345–351.

(90.) Fay Worley and Polydora Baker, Animal Bones and Archaeology: Guidelines for Best Practice (Swindon, UK: English Heritage, 2014), for the UK.

(91.) Gerrit L. Dusseldorp, “Faunal Assemblage Structure Suggests a Limited Impact of the Introduction of Domestic Stock on Later Stone Age Subsistence Economies in South Africa,” African Archaeological Review 33 (2016): 363–383.

(92.) Gifford-Gonzalez and Hanotte, “Domesticating Animals”; and Diane Gifford-Gonzalez and Olivier Hanotte, “Domesticating Animals in Africa,” in The Oxford Handbook of African Archaeology, eds. Peter J. Mitchell and Paul Lane (Oxford, UK: Oxford University Press, 2013).

(93.) Michael Brass, “Early North African Cattle Domestication and Its Ecological Setting: A Reassessment,” Journal of World Prehistory 31, no. 1 (2018): 81–115.

(94.) Peter J. Mitchell, “The Constraining Role of Disease on the Spread of Domestic Mammals in Sub-Saharan Africa: A Review,” Quaternary International 471 (2018): 95–110.

(95.) Kevin C. MacDonald, “Socio-Economic Diversity and the Origins of Cultural Complexity along the Middle Niger (2000 BC to AD 300)” (PhD diss., University of Cambridge, 1994).

(96.) Joséphine Lesur, John W. Arthur, Kathryn Weedman Arthur, and Matthew C. Curtis, “Hide and Meat among Boreda Hideworkers: Ethnoarchaeozoology of Consumption and Craft Practices in Gamo (Southwest Ethiopia),” Quaternary International 471 (2018): 81–94; see Mark McGranaghan, “Ethnographic Analogy in Archaeology: Methodological Insights from Southern Africa,” in The Oxford Research Encyclopedia of African History (New York, NY: Oxford University Press, October 2017); and Constance Smith, “Anthropological and Ethnographic Methods and Sources,” in The Oxford Research Encyclopedia of African History (New York, NY: Oxford University Press, 2017).

(97.) Elizabeth R. Arnold and Diane Lyons, “Ethnozooarchaeology of Butchering Practices in the Mahas Region, Sudan,” in Ethnozooarchaeology: The Past and Present of Human–Animal Relationships, eds. Umberto Albarella and Angela Trentacoste (Oxford, UK: Oxbow Books, 2011), 105–119; and Simone Brunton, Shaw Badenhorst, and Maria H. Schoeman, “Ritual Fauna from Ratho Kroonkop: A Second Millennium AD Rain Control Site in the Shashe-Limpopo Confluence Area of South Africa,” Azania: Archaeological Research in Africa 48 (2013): 111–132.

(98.) Jousse, Atlas of Mammal Distribution.