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File: Lecture 7 Coastal Migration and Aquatic Resources
Populations of H. erectus were the first human emigrants from Africa. Traveling a route that must have been much the same as that hypothesized for moderns, Lecture 6, they reached the islands of Indonesia (review the map in lecture 5) where fossil crania have been found at Trinil, Ngandong, Sangiran, and Perning near the city of Mojokerto. A group of investigators led by O. Frank Huffman and Yahdi Zaim have just undertaken a major multi-disciplinary project to properly describe the Mojokerto site where the partial skull of a juvenile was found. In the latest publication of this project the authors describe a paleolandscape similar to a modern landscape from Mojokerto pictured above.
The preliminary analysis of the fauna and flora in the beds at the site indicate an environment very much similar to that pictured above where mangrove vegetation, grassland and montane forest provided a variety of resources including small and large bovids, Asian elephants, deer, and turtles.
Bantengs (wild oxen) would have come to gaze on the grassy plains between the water and the montane forest inhabited by large cats, monkeys, and pigs. The oxen shared their grazing ground with muntjaks, antelope, rhinos and hippos, some of which foraged on smaller shrubs. Preliminary analysis of fossil teeth from the site indicate that the grasses on these grazing grounds were primarily C4 tropical grasses. The analysis of phytoliths in the sediments reveal two types of grasses, open-land taxa from the subfamilies Panicoidae and Arundinoidae and the temperate climate subfamily Pooideae which probably drifted in from montane habitats. Notice from the opening photograph in this lecture the habitat diversity within a short distance from shore line to ridge line. Early humans might have varied their foraging strategies with the seasons depending upon which resources were most abundant in each habitat. Notice also that the presence of large herbivores near the shore doesn’t rule out big game hunting in habitats characterized by aquatic resources that women and children could collect.
At other locations in Java where crania of H. erectus have been found, a succession of dry and wet climatic regimes can be documented in the fossil pollen record. The stratigraphy of the Sangiran dome, near where Sangiran 17 was found, documents a record from 2.6 M. A to 0.2 M. A. (Semah et al. 2003 ). The sediments at the Plio-Pleistocene boundary indicate the retreat of the seas and emergent land bridges between mainland Southeast Asia and the Indonesian islands. At all times, the habitat of East Java was characterized by high diversity with the hominid fossils dating to 800,000 years associated with pollen documenting a drier climate but tropical rain forest taxa still present. “In the area outside Sangiran, an Indonesian-French team has worked in the Southern Mountains of Java documenting a long sequence of occupation and adaptation to wet tropical environments from the late middle Pleistocene to the Holocene (Semah et al. 2003 p. 161).” The second wave of human emigrants from Africa, H. sapiens, surely passed along these shores as they reached southern Australia by at least 40,000 years ago. The use of coastal, marine, and estuary resources by sapiens in Southeast Asia has been little considered as most of the focus in Asian archaeology has been on H. erectus in China and later populations transitioning from hunting-gathering strategies to farming. Erlandson (2001) attributes this neglect of coastal and aquatic adaptations to two factors, the widespread perception that hominids did not adapt to aquatic habitats before around 15,000 years ago and the obsessive attention to the idea that male-dominated big game hunting explained the origin of tool use, the formation of the nuclear family, and in its most recent incarnation the evolution of large brains. Erlandson labels one set of ideas about aquatic resources the “Gates of Hell” model as theorists proposed that humans only began to use lower quality aquatic resources when forced into marginal habitats by declining returns from hunting and/or by density dependent population growth, that is when they were forced from more desirable habitats. In the next section we are going to ask if foraging returns derived from one habitat can be used to guide inferences about returns in another time and place. This issue is especially relevant to the claims that evidence for the use of aquatic resources is evidence for habitat stress and decline of higher ranked resources. Last resort scenarios ignore the evidence for the nutritional value of aquatic resources, the cultural complexity of societies subsisting on those resources, high human population densities in some of the most widely exploited riverine and coastal habitats, and the emerging archaeological data suggesting early adoption of aquatic resources. Instead proponents, of the last resort hypothesis, cite the small size of many of the marine resources such as mollusks, crabs, sea urchins, barnacles, and shrimp while ignoring the giant clams available on the reefs of the southern oceans, the abundance of individuals in extensive mollusk beds that characterize rocky coastlines north and south, the large number of stranded fish that can be acquired in drying pools, and nesting sea turtles as a source of eggs and meat. Since many of the aquatic resources are sessile and predictable in time and space little search and no pursuit is involved in their harvest and interactions with other predators are rare. Further shell fish gathering requires no prior tool production, maintenance or preparation. Shellfish in particular are characterized by low variance, high density, and ease of collection, the very qualities that meet the daily nutritional requirements of growing children who can gather many of these resources for themselves. Erlandson cites a study by Jones and Richman (1995) showing that “mussel beds produce one of the highest rates of biomass production on earth (Erlandson 2001 p 294).” The supposed low quality of marine resources is derived from an estimate by Bailey (1978) “that 156,800 cockles were required to provide the caloric yield of one red deer (Erlandson 2001 p. 294).” Erlandson questions the accuracy of these estimates and later notes the analysis by Lindstrom (1996) of returns from the Truckee River fishery that are higher than return rates calculated by Simms (1987) for terrestrial Great Basin habitats. Shell fish are low calorie foods so would rank low in foraging models based on net energetic return but they have high nutritional value as sources of protein, calcium, and omega fatty acids. Ordinarily I would not post information from commercial sources but the chart comes from a USDA handbook and is a useful comparison of beef and shellfish. Later in the semester we will review arguments over the value of marine resources in the diet when we consider the research from the Okinawan centenarian study. When you read the assigned pages in Erlandson pay attention to arguments over aquatic resources, the ambiguity of the archaeological record, the theoretical prejudices of the investigators, the nature of the resource in question, and the problem of changes in sea level with high interglacial levels destroying whatever archaeological evidence there might have been from periods of lower seas. Remember from Walter et al. (2000), Lecture 6, that the Pleistocene reef terrace at Eritrea was preserved by tectonic activity that uplifted a portion of the former shallow water reef. Notice, from Table 1 page 306 Erlandson (2001), that the sites with evidence of use of aquatic strategies span the time from 2.3M to 16.5K, a considerable time frame for the occupation of “marginal” habitats. Instead of regarding these habitats and strategies as marginal, new evidence and new hypotheses call for serious consideration of aquatic resources as central to the evolution of human dietary strategies. Often the most obvious fact of human subsistence is unmentioned because it is so taken for granted. Erlandson calls our attention to the fact that freshwater for drinking is the “single most important aquatic resource for humans (Erlandson 2001 p. 293).” Even as our earliest ancestors exploited the resources of the woodlands and grasslands, they must have stayed close to sources of water. The important archaeological sites of Africa are all in riverine or lacustrine environments. It is hard to imagine wild foragers ignoring the resources in and and around water sources especially when those resources could be captured and processed with very simple tools. Where simple tools were not adequate to the job, early sapiens were capable of making more sophisticated tools as evidenced by the hafted bone point technology from Katanda. Although I have emphasized coastal strategies in this lecture we know that early anatomically modern humans did move into other ecological niches as shown by the occupation of Niah Cave in Borneo. The Deep Skull, tentatively dated to the time period between 43,000 – 40,000 B.P., has been found in late Pleistocene sediments of the cave. Niah cave is one of a series of caverns in sheer-sided limestone walls that rise nearly 400 m above the lowland rain Though the evidence from Niah is scanty, starch grains from palm pith and deep-rooted species of yams have been identified from the site. At other sites in Melanesia, starch grains of elephant yams, taro, ginger, and swamp taro have been identified. Some of these taros require cooking, drying or leaching to remove toxic calcium oxalate crystals but the pith of certain palms can be eating raw. These palms have high energetic yields but some processing costs as the palms have to be felled and the fiber pounded or otherwise extracted from the trunk. Evidence from sites in Southeast Asia and the evidence we will discuss in Lecture 9 from the Ache demonstrates that forest environments provided rich habitats for human foragers. Recall from Lecture 6 that the islands of Indonesia were accessible by land during glacial periods when water was locked up in northern hemisphere ice shields.
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File: Lecture 8 Optimal Foraging Theory
The lectures to date have focused on the evolution of Plio-Pleistocene diets inferred from the archaeological evidence and from paleoecology informed by modern food preferences through an implicit assumption that the environment is populated with nutritious plants and animals to be captured by individual hominids ranging over that landscape. Most of you know from other classes and from your own diets that foragers are selective in their choices. The simple coincidence of resource and forager in the same habitat tells us little about the diets of the forager. To be fair we have looked at circumstantial evidence for use of some resources such as cut marks on bones, opened and cracked shells, processed fish bodies, special tools for capturing specific animals, and isotope ratios but all of that evidence provides only a coarse grained picture of early diets. Archaeologists interested in refining their assessments of past foraging strategies have applied models from optimal foraging theory (OFT) to help them understand the past. These models were originally developed by biologists interested in explaining the specific resource choices they observed in their study populations. They were subsequently used by evolutionary anthropologists to study diet choices in modern foragers subsisting on wild resources. Most of you know that the faculty and students from the University of Utah departments of biology and anthropology were very active in developing and applying these models.
How can the information from OFT be reconciled with the Grandmother Hypothesis for the evolution of human life histories from O’Connell et al. 1999 (Lecture 5)? The fundamental premise of OFT is sound as we expect natural selection to favor individuals that allocate energy to tasks that promote somatic growth, maintenance and reproduction. In any habitat those genotypes better at allocation decisions are expected to leave the most descendants, that is have the highest fitness, yet the expectations of the diet breadth models are often violated in real populations. All models assume a generic forager but the early work with the Ache demonstrated that men routinely ignore the highest ranked resource in the forest, palm starch, in favor of hunting. This falsification of the model was as important as the confirmation of most of the predictions for other resources because it raised the question of why men hunt which led through many populations and many papers to important ideas about the role of costly signaling in male-male competition and linked sexual selection theory and foraging theory. But sexual selection on males is not the only factor determining diet choice as risk and uncertainty influence forager choices. The idea that the value of a resource item is determined by the costs of search and handling changed the way investigators ranked prey items in the foragers environment. Many resources are energy dense but have high search and handling costs. It is not the absolute value, to a forager, of the resource but the net energy gain to the consumer after the energetic costs of pursuit and handling are debited. In most investigations of actual foragers net energy gains are calculated by debiting post encounter costs against the currency of the gain, usually calculated in caloric return per unit of time spent in pursuit, capture and handling.In the simple version of the optimal diet model resources are included in the diet based on the tradeoff between handling upon encounter and the option of continuing to search for other items. The decision is based on the mean rate of return for that particular habitat and the suite of resources within it. Be aware that post encounter returns are always much larger than returns reported with search time included unless search time is nil. The inclusion of processing costs in the equation reflects the net gain to the forager from choosing any resource and influences forager preferences for certain resources over others. Richard Lee’s failure to include processing time in the cost of mongongo nuts seriously under estimated the contribution of women’s work to family energy budgets and led to the characterization of the Dobe !Kung as the the original affluent society, meeting their subsistence needs with few hours of work. I have assigned two short readings, Sih and Milton (1985) and Hawkes and O’Connell (1985) debating the the use of OFT to study human resource choices and specifically the ranking of mongongo nuts in !Kung diets. These two papers will give you a sense of the early arguments over the application of OFT models to human foragers and the value and problem of simplifying assumptions. As you have already read in Erlandson, the failure to report search time for resources leads to the over estimation of returns from resources such as collared peccaries and makes comparisons of post encounter returns of resources, such as red deer and mussels from two very different habitats, meaningless. Note in all of the lectures that follow I have tried to maintain the notation, calories, cal/hr, Cal/hr, or Kcal/hr used in the assigned papers. The notation is confusing, but the link provides a simple explanation of the differences. For our purposes we assume that investigators are referring to Kcal when they use cal, Cal or calories to indicate the energetic gains from any resource.
Winterhalder, Lu and Tucker (1999 p. 302-303) distinguish between risk and uncertainty by defining risk as “unpredictable variation in the outcome of a behavior, with consequences for an organism’s fitness or utility.” Further “with risk the probability distribution of outcomes is in some sense known to the organism, but stochasticity makes any particular outcome unpredictable.” “Uncertainty refers to incomplete knowledge of outcome probabilities. Uncertainty can be overcome by acquiring information about an environment; risk cannot.” They make the point that risk as used in behavioral ecology and economics does not “mean exposure to danger.” The authors point out that simple foraging models assume the forager always experiences the average conditions of its habitat where decisions have predictable outcomes. This assumption is central in all debates over the role of big game hunting in the evolution of the human diet. When we examine Hadza foraging strategies we will consider the high variance in game capture and contrast that to figures for the average amount of meat in the diet. In contrast to deterministic models, risk sensitive models assume organisms adjust to a range of possibilities arrayed along a probabilistic distribution. In addition risk-sensitive analysis specifies the relationship between the outcome and its fitness value resulting in a sigmoid curve. To the left of the inflection point of the curve value rises with increasing resources but at an accelerating marginal rate when the resources are scarce and at a decelerating marginal rate when they are abundant. In simple language, with too little food “added increments have high value and with too much, added increments count for little (Winterhalder et al. 1999 p 304).” The work of Curaco and his colleagues, with yellow-eyed juncos, showed that the utility function for the birds takes the sigmoid form. There is no reason to suspect that hominids are any less responsive to variable outcomes. If you are interested in further exploration of these ideas the figures presented in the Winterhalder et al. (1999) paper will give you a much better sense of the issues. I have posted the paper under also recommended for those of you interested in a more detailed explanation. Since value is measured as a rate, usually in calories gained per unit time, the sigmoid value function is time sensitive and this sensitivity varies with respect to the physiology of the organism and the character of the resource. Think back to Lecture 1 and the link to the Moran Eye center Web vision site with the graphic demonstrating the daily rebuilding of the pigment containing discs of the rods and cones. Stores of vitamin A must be sufficient to support the daily demands of this process. Winterhalder et al. (1999) make the same point about water acquisition. The preference for a reward of k liters/hr and equal probabilities of 0 or 2k liters/hr will depend upon whether the result will be suffered for hours, days or weeks before choice and outcome are reiterated. Responses of the organism to risk are determined by design by natural selection over the long time frame and the physiological state of the organism over shorter time frames. In general females are expected to be more risk adverse than males and organisms in negative energy balance are predicted to be more risk prone. The latter prediction was confirmed in experiments with yellow-eyed juncos whose foraging choices were observed under different temperature regimes that influenced energy balance. Given two foraging options with constant or variable rewards, birds in negative energy balance favored the high variance reward, presumably in the attempt to gain a reward sufficient to change their energy balance. The excitement generated by these results was tempered by a number of experiments that failed to replicate the results. Winterhalder et al. (1999 p. 323) conclude; “These studies suggest that the expected energy budget rule may apply only rarely to hominids, nonhominid primates, and modern humans which are omnivorous and relatively large species.” They might have qualified their statement to include only adult humans as anthropologists and archaeologists, with few exceptions (cf. O’Connell et al. 1999), have ignored the consequences of short periods of negative energy balance on the growth and development of neonates, infants, and juveniles. Conversely, resistance to short-term perturbations in energy balance may be an advantage of large body size that contributed to the success and expansion of H. erectus. We need to be careful here not to attribute the evolution of large body size to a resistance to short term negative energy balance. Such resistance can be a consequence of large body size but the causal arrow on the evolution of large body size runs from mortality risks in the environment of the species. Following Charnov’s (1993) mammalian life history model, large body size is a consequence of more time to grow larger because lower average adult mortality, from extrinsic causes, favors delaying the change from somatic to reproductive allocation. Organisms that experience low average adult mortality rates, grow longer, mature later and have longer lifespans. These are the derived traits that distinguish us from our nearest primate relatives. These arguments were reviewed by O’Connell et al. (1999).
The last assigned reading for this lecture is a paper by Elston and Zeanah in which they use models from behavioral ecology to understand the Holocene transition evident in the archaeological record of the Great Basin. Most of you know that our department has been in the forefront of extending foraging theory to hunter-gatherers but you may not be aware of the innovative ways the models are being applied to solve longstanding problems in Great Basin archaeology. The paper by Elston and Zeanah is a good example of this work as well as providing a link, in this course, from ancestral populations known only from the archaeological record and the next set of papers from more recent foragers. The opening graphic of this lecture, from the work of Edward S. Curtis, is intended to emphasize the importance of Great Basin studies to applications of optimal foraging theory to human foragers and to highlight the importance of ethnographic analogs to this work. The record of human occupation in the basin can be divided into two distinct time frames labeled Prearchaic for the period from 10,500-8000 BP and Archaic for later occupation. The early Holocene (EH) climate experienced by Prearchaic populations was cooler and wetter than climates after about 7800 BP. “The relatively cool, even EH climate, abundant surface water and complex steppe vegetation created productive habitats for a rich biota of fish, waterfowl and mammals (Elston and Zeanah 2002 p. 107).” The tool assemblages from this time period have an array of projectile points and flaked tools indicating a hunting economy but the coprolites recovered from the sites indicate a diet that included many of the resources of the lakes and marshes plus seeds and small animals, quite unlike a diet predicted from the graph of energy yield posted above. Basin occupants in this time period had access to a variety of resources within short travel distances as basins with lakes and marshes were separated by ridges with brushy steppes and juniper woodland from mid-elevation to ridge tops providing high-quality habitat for larger game animals. Moreover the scatter, size and character of the archaeological sites from this time period indicate low population density and high mobility of foraging groups. An attempt to use the diet breadth model to explain the evidence for the addition of seeds to the diet failed as the model developed by Simms predicted that women should have by-passed seed under all conditions, including the absence of large game, a prediction contrary to the clear evidence for seed use. A second model analyzed the distribution of soil types in the basin to develop a patch choice model. The simulations based on the distribution of lowland and upland habitats predicted that women should bypass seeds in favor of other plant resources and small game and men should have targeted only large and medium size game. This model fails to explain the evidence for small animal consumption found in the coprolites. Finally a risk sensitivity analysis returned the prediction that risk prone foragers should favor the high variance option “preferentially pursuing large game over smaller game and foraging in habitats where encounters with large game were most likely (Elston and Zeanah 2002 p. 115)” while risk sensitive foragers should prefer lower-ranked resources characterized by low variance in acquisition. From this model, Pinson (1999, cited in Elston and Zeanah 2002) hypothesized that Prearchaic foragers avoided starvation risk by pursuing small game. That is they were risk adverse. Elston and Zeanah (2002) note the following problems in Pinson’s analysis; 1) the model ignores the possibility that Prearchaic foragers might have entered basins where foraging returns did not meet daily requirements at which point the risk of starvation should have prompted the risk prone strategy of choosing the high variance option of large mammal hunting, 2) if Prearchaic foragers were risk adverse they would be expected to forgo migration from known basin habitats to new, unknown locations, and 3) if EH environments were as patchy as predicted by Pinson, the Prearchaic archaeological record should more nearly correspond to the Archaic record when climate deterioration reduced the density of high ranked prey. Increasing search time would have been a consequence under both conditions. Zeanah et al. (1999) used some of the techniques from Pinson’s analysis of Carson Desert foraging in simulations of optimal foraging in Railroad Valley but in the latter analysis “more detailed regional palaeoenvironmental data, better information on the productive capacity of modern soil types and improved GIS (Geographic Information System) capabilities permitted a more finely tuned reconstruction of prehistoric foraging landscapes than was feasible in the Carson Desert (Elston and Zeanah (2002 p. 117).” The Railroad Valley simulations considered male and female foragers randomly encountering resources across spatial and seasonal variation. From the early Holocene (EH)to middle Holocene (MH) hunting returns for men diminished by as much as 75% in all seasons. By contrast women’s returns varied much less across the transition, increasing somewhat in autumn as pinyons entered the ecosystem. As habitats dried out in the MH, wetland resources disappeared as options for women allowing lower ranked small seeds to enter the diet in late winter-early spring. The authors conclude that the highly productive ecosystems of the EH and the low population numbers encouraged high mobility that allowed men and women to forage in the most productive habitats for their resource choices. As the climate changed and women’s wet land resources declined, Great Basin foragers faced the risk of starvation in certain seasons. In the face of this risk women collected small seeds in seasons when they were abundant and stored them for the short season. Seed caches changed the pattern of mobility for both men and women, encouraged residential bases, lower investment in tools for large game hunting and more investment in tools such as grinding stones for seed processing. Bright, Ugan and Hunsaker (2002) predicted increasing investment in processing tools to reduce handling cost as lower ranked resources enter the diet in response to a decline in the density of higher ranked resources. The change in grinding stone technology, through time, in the Little Boulder Basin area of north-central Nevada supports that prediction. These innovative approaches linking optimal foraging theory, resource changes through time, and changing investments in technology increase our confidence that ancestral diet choices can be studied and understood. In lecture 9 we will look at how foraging models, applied to resource choices in modern hunter-gathers, have improved our understanding of human diet choices.
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File: Lecture 9 The Ache: Broadleaf Evergreen Foragers
The Ache are foragers of the subtropical, broadleaf, evergreen forest of Paraguay. I chose this example because Ache foraging has been intensively studied by the Utah Ache Project and as a comparison to semiarid African strategies. The warm, wet summer regime of the Ache forests are similar to habitats widely distributed from South America to Indonesia between 20 degrees north and south of the equator in regions with annual average precipitation varying from 1600 mm to 2000 mm. Temperature regimes in these habitats are characterized by warm summers and warm to cool winters with January maximums in the Ache region of around 40 degrees centigrade and July minimums of about -3 degrees centigrade. Early anatomically modern humans moving south toward Australia and north into China, Viet Nam, and Thailand must have encountered these habitats and foraged in them. The early Pleistocene archaeological record of upland Southeast Asia is little explored but the Ache can serve as a reasonable analog of human foraging strategies in humid subtropical ecosystems. Keep in mind that the distribution of these ecosystems, through time, was determined by northern hemisphere glacial cycles. On the Koppen climate classification map pictured below the Ache habitat is shown in green, warm to hot wet summer, cool dry winter.
The required reading for this lecture, Hawkes, Hill, and O’Connell (1982) is one of the earliest reports of the use of OFT to analyze resource choices of humans foraging on wild food. Even though the preliminary data in this paper has been revised and corrected in subsequent papers, I have assigned this early paper because it provides a brief history of the arguments, in anthropology, over human diets and because of the broad picture it provides of Ache foraging. The northern Ache were full-time hunter-gatherers until mid-1970 but now live at settlements sponsored by a Catholic mission. At the time the research was conducted by the University of Utah Ache project, they still engaged in foraging expeditions and lived on wild resources during their time away from camp. Hawkes et al. (1982,) contrast the views of Richard Lee, developed from his field experience with the !Kung, to those of Marvin Harris developed in a series of debates with Napoleon Chagnon over the causes of violence among the Yanomamo. Harris argued that the Yanomamo fought over scarce protein resources while Lee’s tally of foods eaten daily in !Kung camps convinced him that plant foods were more important than meat. Harris argued that meat was nutritionally superior to plants and would only be replaced by plants when large game animals had been depleted by hunting pressure. The important themes that would subsequently dominate debate over foraging strategies in modern hunter-gatherers are set out in Hawkes et al. (1982 p. 380). “Lee argues that plant foods are favored because they are abundant, reliable, and readily located, and therefore more efficiently exploited than are animal foods. Plants are said to be low-risk/high-return resources, while animals are high-risk/low-return resources. Animals are taken in spite of the inefficiencies involved because of the taste appeal of the meat and the prestige that accrues to successful hunters.” Hawkes et al. (1982) use two models from OFT, the optimal diet model and the patch choice model, to predict the resources the Ache should gather if they are maximizing net energy return. You were introduced to the optimal diet model by Elston and Zenah (2002), lecture 8, so you should know that the forager is assumed to make a decision about whether to take a resource upon encounter or to continue searching. Resources are ranked based on the return in calories over the post encounter processing times including both pursuit, for mobile prey, and processing to turn the resource into food. The forager is predicted to take only those resources that give a return rate equal to or higher than the average rate for resource in the optimal diet. Recall the Hollings Curve which showed that search and handling time are distinct elements of the problem, the search time is determined by the density of prey on the landscape but the net returns from that prey are determined by the handling time. It is the net return, after handling, that determines the rank of the prey in the diet. In the simple form of the model high ranked prey should always be taken when encountered. Hawkes, Hill and O’Connell make a very important point on page 388: “Note that the resource rankings of this model say nothing about the quantitative importance of a resource to optimal foragers. High-ranked items may be so rarely encountered that they represent only a very small portion of the diet; low-ranked items in the optimal set may be encountered with sufficient frequency to contribute the bulk.” Keep this in mind when we get to the disputes over the importance of various foods for contemporary human nutrition. Three additional points, 1) when the cost of search is added to the cost of any resource, its value to the forager, in absolute terms, may decline, 2) all optimal diet models are specific to a habitat and time period, and 3) the optimal diet model assumes a fine-grained environment where resources are encountered at random. Foraging trips with the Ache showed that the environment is coarse-grained or patchy with many resources clumped. On foraging trips the Ache often stop to harvest in some resource patches but not others. The authors suggest that the “distribution of tools” (Hawkes et al. 1982 p. 391) might account for the decision but discard that suggestion because the Ache nearly always take oranges and honey, even though both are harvested with the use of axes, but often pass palm fiber. The patch choice model predicts that foragers use patches that produce the best return when travel time, search within the patch, and handling time are considered. Notice that return figures for patchy resources include the time for searching within the patch. If both animals and oranges are considered as patches the average energetic gain return for hunting, including search is about 1115 Cal. per hunter-hour while the return for oranges is 4438 Cal per forager-hour and the return for honey is 3231 Cal/hr. Palm larvae has a similarly high return rate of 1849 Cal per forager-hour but involves the risk of high variability across logs. Returns calculated for fishing patches were similarly variable, ranging from > 2000 Cal per forager-hour to about 733 Cal per forager hour leading to two questions, when should foragers exploit palms and when to take fish. The average return rate in calories per hour of sixteen resources taken by the Ache are listed in Table 3, page 389. Notice that collared peccaries are a very high ranked resource but no search time is factored into the estimate of return.
Subsequent analyses of the data from five years of research on Ache resource choices reported in Hill et al. (1987) highlighted both the value and problem of using optimal foraging theory for analysis of human foraging choices. First detailed records showed that a theory for generic foragers failed to capture the different strategies of males and females. Among the Ache, men achieve significantly higher returns, 1253 Cal/hr, than women, 1087 Cal/hr. These figures are averaged over all resources taken by Ache men and women and include both search and handling costs. Before processing the rates are more nearly the same, 1339 Cal/hr for men and 1221 Cal/hr for women. The differences in rates indicate the high cost of processing palm fiber. Men often by pass palms but when they do take them they gain higher rates because they process the fiber faster than women. Consistent with the predictions of OFT models only one of the 26 resources taken by the Ache gave returns, after encounter, lower than the overall mean. That resource was palm larvae, mostly taken by women but sometimes also collected by men. Contrary to the predictions of the theory that foragers should preferentially target high return resources, returns from adult male hunting, 1349 Cal/hr, were lower than the caloric return, 2630 Cal/hr that could be obtained from palm starch and hearts (Hill et al. 1986). The observations that men gain higher returns than women, on average, changes if returns are compared for days when camp is not moved. On days when the group is moving women carry babies, pets, household belongings, and any meat the men catch. Men keep hunting after turning over meat they have already caught. Because of the opportunity cost of processing, most Ache processing work is completed in camp at the end of the day. When the Ache women are moving they pass many resources they might otherwise gather so their return rates on camp move days is very low. By contrast when they stay in one camp for several days, the women achieve a return on gathering of 2804 calories per hour compared to a return rate for men, on these days, of 1344. These figures include all time spent in food acquisition and processing. These results raised the question of why men hunt and suggested two alternative hypotheses, 1) caloric return per unit of time is not the only currency by which foraging strategies should be judged and 2) fitness payoffs for hunting are broader than meeting subsistence requirements. The first hypothesis proposes, in agreement with Harris, that meat is more highly valued than plant food because of high protein and lipid content. You can evaluate this hypothesis by recalling the human RDA for lipids and protein in the diet as reported by Conklin-Brittain et al., lecture 2 and the values of lipids and proteins that can be acquired from plant foods. When you open the link search on RDA to compare values in chimpanzee diets to RDA for humans. Also have a look at the figures and ask if annual averages reported in the charts are misleading. Monthly mean lipid levels in the diets are much more variable than monthly mean protein levels. Even though the annual mean lipid level in the chimpanzee diet seems sufficient to meet the RDA for humans the high variance may not provide adequate daily lipid consumption. The second hypothesis, suggested but not fully developed in this paper, is that men opt for a high variance strategy that promises a large payoff when successful. The payoffs for hunting include not only the calories, proteins, and lipids gained from the meat but mating benefits for good hunters. This last payoff is more fully developed in the papers on Hadza hunting. The work with the Ache demonstrated that human foragers differ, by age and sex, in the amount of time they spend foraging, the resources they take, and the amount of time they spend processing. The resources a forager takes on any day is not only a function of the costs of pursuit and handling but of the foragers state and the time frame over which the subsistence needs exist. Lower ranked fruit that can easily be gathered and eaten while moving will often be collected to satisfy short term needs while large game and palm starch are taken to satisfy needs over several days. In all cases the ranking of resources is habitat specific and even in the relatively stable habitat of the lowland deciduous forest can vary by season. The Ache only hunt and dig armadillos in the late wet season when the animals are fat and the average return is 3948 cal/hr compared to a return of 1220 cal/hr in the early wet season. The emphasis on highly ranked resources might leave the impression that the Ache diet is very narrow but the list of wild plants and animals taken by the Ache suggests otherwise. Have a look at Table 2 from Hill et al. (1987), below to get some idea of the variety in the diet. Remember larva are the only resource that reduces the average return, over all resources included in the diet. The optimal diet model predicts that the Ache should never take bamboo larva but ethnographic observations indicate larva are often collected. Do the Ache take larva because they value the high fat content of larva? This question is still the subject of debate so no clear answer but exceptions to the predictions of OFT do not diminish the usefulness of the model. Instead they open new windows into human foraging strategies.
The Ache are exceptional, among studied hunter-gatherers, in the number of calories they consume daily, 3610, and in the amount of meat they eat, 80% of calories from meat. Notice in Table 2, the difference in return rates depending upon the included costs. The return for White-tipped peccary is 5323 calories/hour if time spent tracking is debited but as high as 8755 calories per hour if only post encounter rates are considered. Similarly Ache men obtain a much higher return for 9-banded Armadillos they encounter on the surface compared to those they dig up. The variation in these rates makes it nearly impossible to compare return rates across foragers if the ethnographers don’t clearly distinguish between the methods of calculating rates. Keep these problems in mind as we survey foraging strategies across habitats.
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File: Lecture 10 Semiarid African Strategies
In this lecture we focus on the foraging strategies of the !Kung and Hadza hunter-gatherers of the semiarid region of Africa. The work of Lorna and John Marshall and Richard Lee made the Kalahari San the archetype for hunter-gatherers and the evolution of human foraging strategies. Their work in Southern Africa was complemented by the early work of James Woodburn among the Hadza of Tanzania and the detailed work of James O’Connell and Kristin Hawkes among the same group. As a consequence of these efforts we know more about human foraging in semiarid environments than in almost any other habitat. You viewed the map of present climatic regimes in the last lecture. Go back and have a look at the distribution of semiarid lands in Africa, the bright yellow and three shades of orange areas. The !Kung San live in the Kalahari desert of southwestern Botswana and eastern Namibia and the Hadza occupy the area around Lake Eyasi just south of the equator in Tanzania. Both areas are characterized by seasonal rainfall and shortages of surface water in the dry season. In these semiarid, seasonal rainfall climates, precipitation is highly variable from year to year and across the region in any one rainy season. In the Kalahari, over eleven years, precipitation varied from a low of 252.1 mm in 1962 to high of 788.9 mm the next year. Temperatures in the Kalahari vary from a high of 40 degrees centigrade in November-December to a low of nearly -10 degrees centigrade in July and August. The climatic regime of the two regions is very similar but the topography couldn’t be more different. These topographic differences have important implications for differences in foraging strategies of !Kung and Hadza children. In the lecture on children’s strategies we will consider those differences. This is a long lecture but the reading assignment is not excessive and I promise you a short Lecture 11.
The widely read early work and films from Lorna and John Marshall and later the careful measures of !Kung foraging returns by Richard Lee established the !Kung as archetypal human hunter-gatherers even though their habitat represents a very small percent of the habitats occupied by early H. sapiens. The notion that early humans evolved in an ecological niche similar to those occupied by the !Kung and Hadza focused the attention of researchers on big game hunting as the principle dietary strategy of humans to the exclusion of other foraging strategies. You were introduced to this problem in the reading from Erlandson (2001), lecture 7. Review it here so that you can put the dietary strategies of semi-arid lands in the context of variable human responses to the opportunities presented over the long migration from Africa. Once Richard Lee measured the actual food brought into a !Kung camp, in the Dobe area of the Kalahari, he came to the conclusion that plant foods represented the principal component of the diet. The Dobe !Kung men do hunt but the returns are highly variable. In spite of the variable nature of returns from meat, it is a highly desired resource. When a large game animal comes into camp everyone eats meat but on the many days when there is no meat in camp, men, women and children do not go hungry. They enjoy other foods gathered from the sandy plains of their homeland. I have assigned chapter 2 of Jiro Tanaka’s 1980 report on the !Kung San of the central Kalahari. You can download the reading from Marriott Course reserve. Tanaka’s field work was accomplished in the sixteen months he spent in the field between 1966 and 1968, sixteen months from April 1971 to August 1972 and two more months in 1974. In his introduction to San, Hunter-Gatherers of the Kalahari, Tanaka makes the point that desert is a misnomer for this region as it is an expansive, monotonous, grassland dotted with open woodland trees and shrubs characterized by a summer rain season from December to March. Tanaka gives you a study of diet in an area where there are no mongongo nuts, the most important dietary staple in the area where Richard Lee worked. Richard Lee made the San famous through his measurements of the number of hours spent gathering food. Since he didn’t count processing times his estimates left the impression that the !Kung could satisfy their dietary needs in no more than 2-3 days per week. As you know from reading the Hawkes and O’Connell (1982) response to Sih and Milton (1982) Lecture 8, mongongo nuts have particularly high processing costs. The principal foods of the central Kalahari San are listed in Table 8. Even though Tanaka did not use the framework of OFT to study resources gathered, the first eleven foods listed in Table 8 constitute the important resources in the diet. Notice the importance of melons and roots, that store water, for a people who have no access to surface water through much of the year. Tanaka attributes the ability of the San to survive in this very habitat to the technology of the digging stick which allows humans to access deeply buried tubers that cannot be gathered by any other primate, including savannah baboons who live on shallow rooted species in other habitats but need access to surface water so are unable to colonize the central Kalahari. Four of the important resources are shown in the next table. I was unable to find a good picture of Bauhinia beans. The shrub is a beautiful ornamental of the nursery trade so all of the online images feature the flowers and not the pods and beans.
Consistent with OFT, Tanaka (p 59) reports that beans of Bauhinia petersiana are the “central food item for several months” making “up nearly the whole of the San’s diet” in months when they are ripe and water is available. “Even though this is the period of greatest abundance and variety of food types during the whole year, people often totally ignore other foods.” This latter is somewhat surprising as the beans must have high carrying and processing costs since the inedible pod accounts for 75% of the whole weight so approximately 5 kg of pods and beans are carried back to camp to yield 1 kg of edible beans. Table 12 in Tanaka gives nutrient analysis of the important plant foods and Table 13 provides an estimate of the daily caloric intake per person. The figures are not intended to be totaled but to show what might be gained if 5 kg of Kan melon were eaten each day. On page 74, Tanaka gives an estimate of 2,000 kcal for the per capita yield of food taken during the study period and finds that return consistent with the caloric needs of individuals of the size and weight of adults in the population. Compare the protein content of the dried beans of T. esculentum and B. petersiana and dried Terfezia to the protein composition in animal foods, Table 12 and Table 14, Tanaka. We can’t be sure the units reported are the same but if we assume g/100 grams in each case the plant foods are comparable in protein gain to the animal foods. T. esculentum, marama bean, is being developed as a commercial crop in arid regions of Australia and Texas as it “is an excellent source of good quality protein and compares well with other protein foods including soybeans. Its oil is rich in mono- and di-unsaturated fatty acids and contains no cholesterol. It is also a good source of calcium, iron, zinc, phosphate, magnesium, B vitamins and folate (quoted from the web abstract). ” The number of game animals taken during the study period is shown in Table 11. Giraffes are the largest animals hunted by the San but they are rarely captured in the very dry habitat of the Central Kalahari. More commonly captured are the gemsbok, eland and kudu each weighing 200 to 300 kg. When the amount of meat taken is averaged over all camps, Tanaka estimates that a group of 50 San kill and share approximately 5,600 kg of game in a year or about 112 kg per person or .30 kg per day. From his description we know that meat comes into camp rarely so averages are hardly adequate to estimate the amount of meat eaten. When a large animal is captured people eat nothing but meat until the animal is finished and then go without meat for many days. On page 67, Tanaka describes the sharing pattern for game, large game animals weighing 100 to 300 kg are shared among all the members of the camp, smaller animals such as duiker and steenbok are distributed over a smaller circle of families and still smaller game, birds, springhares and hares are “usually consumed within the family.”
The !Kung Bushmen who live in the Dobe area of the Kalahari were studied by Richard Lee from October 1963 to January 1965. Dobe is in the northwest corner of the Kalahari where there are some permanent waterholes. During the dry season, May to October, groups cluster around the permanent water holes, moving out each morning to collect resources within about a 6 km radius of the camp. In the summer rain season between December and April they scatter across the landscape visiting the pools of water that collect from December through April. The population is more widely dispersed at this time of year. In the season of the rains the women gather fruits, berries, melons, and leafy greens. In the dry season they gather roots, bulbs and resins. Remember from Lecture 2, that chimpanzees add resins to their diets in the dry season. Mongongo nuts are gathered year around as the nuts that fall to the ground in one season are preserved by the very hard shell surrounding the kernel. Richard Lee (1968 p 33) makes the point that “food is a constant but distance required to reach food is a variable, it is short in the summer, fall and early winter and reaches its maximum in the spring” that is before the rains begin. Recall the debate from the Sih-Milton/Hawkes-O’Connell exchange, assigned in lecture 8, over the high processing cost of mongongo nuts. That cost, when only time is considered, needs to be reviewed in the context of other resources that might be gathered in each season and against the background of high variability in game capture. It is almost certain than no dry season resources provide a higher return, otherwise the women would not pay the opportunity cost of processing mongongo. Lee reports the nuts account for 50% of the vegetable diet by weight with the average daily consumption of 300 nuts yielding about 1260 calories and 56 grams of protein. Lee (1968 p 33) estimates that 7.5 ounces of nuts “contains the caloric equivalent of 2.5 pounds of cooked rice and the protein equivalent of 14 ounces of lean beef.”
The figures in this table, copied from Richard Lee (1968) What Hunters Do for a Living, or, How to Make Out on Scarce Resources In Man the Hunter eds. R. B. Lee and I De Vore. pp 30-48. Chicago: Aldine, show that mongongo nuts are an important source of both protein and calories in the diet of the Dobe !Kung. Even though vegetable foods comprise from 60-80 per cent of the total diet by weight, meat is still highly valued and widely sought. During the period charted in Table 5 !Kung hunters brought 410 lbs of meat into camp. Lee assumed that meat was shared equally by all consumers in camp, during that month between 23 and 40 individuals, distributing 410 lbs over an average of 31.8 persons per day over the 28 days between July 6 and August 2 gives an average of just over 7 ounces per person per day. In Table 5 Lee reports the weight in grams (230) which is just over 8 ounces. Beware of these average reports. Looking at the large animal prey pictured above this might be one medium sized animal. Just after the animal is brought in people eat much more meat per day than 8 ounces and when the carcass is fully consumed they go without meat for many days. However, since the !Kung hunt small game as well, the total figure may represent many small animals. San Bushmen may hunt singly or go out in groups but once an animal has been spotted and hit the hunters return to camp to rest. Since the poison used on the arrows is slow acting, the wounded animal will wonder over the landscape for some time before it dies. The !Kung men know they will have a long tracking job, the next day, before they find the dying animal. I have copied a few pages from Tanaka describing the hunt. The copy I have posted for you is missing the figures of hunting equipment but the description gives you a good sense of the difficulty of procuring a gemsbok that has only been hit in the leg. Since the amount of poison on an arrow is small and the poison is slow acting the wounded animal may wonder a long distance before it dies. Notice that the !Kung men of the central Kalahari also scavenge prey from other carnivores and take many small species. According the Tanaka’s description, the internal organs and the ribs are cooked and eaten at the spot of the kill and the bones are smashed to get at the marrow. The uneaten portions are cut up for transport back to camp. The San consume the entire animals except for hooves, horns, bones and stomach contents.
Can you reconcile the information in Table 2 with the graph in Figure 1? The Table shows that the Hadza are most successful at scavenging large game in the late dry season but the Figure shows that large herbivore biomass increases with annual rainfall. Two points should be taken from this observation 1) large game animals may have been relatively more abundant at the Plio/Pleistocene boundary when the genus Homo diverged from the australopithecines so more scavenging opportunities and more opportunities to eat meat, but 2) game animals, predators and humans are more dispersed over the landscape in the wet season and more concentrated around the water holes in the dry season. By now you should expect human diets to vary with the seasons. Recall Stewart’s hypothesis, from Lecture 5, for seasonal dependence on fish easily taken from drying ponds. No matter if fish or mammalian flesh provide the meat component of the diet when no meat comes into camp people still need to eat.
The women walk from 4 to 60 mins. to the tuber patch where they spend the day collecting. Within the patch they stop about mid-day to build a fire and cook some of the tubers they have collected. In the afternoon they continue collecting, stopping again to cook and eat before they gather up the tubers they are taking back to camp for the evening meal. About 39% of the tubers gathered are eaten in the field. During the wet season women add berries to their foraging schedule, spending about half their foraging time in berry patches. Women gain very high returns, measured in grams per hour in the berry patches but they spend about 34% of the foraging time traveling to the patches and 23% walking within the patch between bushes. I estimated the daily gain for berries based on 6 hours of foraging in the wet season at 26% of 6 hours times a gain of 4017 cal/hr to be 6268 cal. for a days work picking berries. In the dry season women spend 4 hours foraging for tubers. On those trips 38% of the time is spend digging for an average of 1.52 hours digging tubers that provide a return of 2000 kcal/hr or 3040 kcal/day. The foraging hours reported here are for child-bearing women. In general senior, post-cycling women forage 22-52 % longer than women of child-bearing age. When you consider the “take home calories” remember that the women and children over 6 that accompany them have been eating in the field so their need for daily subsistence from the take home “bag” is less than that for individuals who were not in the patch. You can see from these estimates that changes in the distance to tuber or berry patches could have considerable effects on the returns that women are able to gain from their daily foraging round. In general the travel distance to berry patches is higher than the distance to tubers because tubers are more densely distributed in the environment. These estimates are reported by Hawkes, O’Connell and Blurton Jones (1989) for observations October to November 1985 and March to April 1986 ( you can find the reference in O’Connell et al. 1999).
How many calories do foragers need? How many hours should they work for a net energetic gain? Was Marshall Sahlins right, needing little they meet their needs in a few hours of work and have many hours of leisure? If we remove our western lens and think about foraging through the lens of OFT we get a new perspective. The theory predicts that foragers should target resources that provide a net energetic return after the costs of pursuit and handling are considered. We have already learned that search time is not debited from the gross return, however all researchers know that walking time is energetically expensive. As a consequence of occupying a niche different from that of our nearest relatives we range more widely in the food quest but because we are bipeds we may be more efficient over large ranges. Leonard and Robertson (1997) modeled the energetic cost of walking for a generalized quaduped compared to a biped and found that male and female bipeds are more efficient over all speeds from 2.4 to 6.0 km/hr. They estimate that the evolution of obligate bipedality produced an energetic savings of 40 to 47% for male and female hominids compared to large bodied primates. Since modern hunter-gathers have an average range of 13.10 km/d compared to 1.77 km/d for large bodied apes an increase in locomotor efficiency is particularly important to occupation of the human foraging niche. However the cost of search is related not only to distance traveled but to other variables such as temperature, humidity, topography, and load carried. Nursing infants are a particularly important component of the female forager’s load which also includes food transported back to camp where other dependent children are waiting to be fed. Unlike chimpanzees, human males carry tools, tool stone and large portions of meat. Could it be that foragers allocate their efforts to maximize calorie gain while minimizing energetic expenditure. Are foraging strategies a tradeoff between energetic returns from the resources in their environment and the energetic cost of searching for, handling and processing those resources? Are there seasons of the year and periods of the day when it pays a forager to conserve energy by not foraging? In the extreme temperatures of the Kalahari resting in the shade during the hottest part of the day may be a very efficient strategy. Lee reports that Dobe !Kung women work an average of 2-3 days per week but Hawkes and O’Connell (1985) point out that Lee ignores the cost of processing mongongo nuts, 5 hours of cracking and pounding to produce 1 kg of nutmeat. Lee reports per capita consumption of 210 grams of mongongo nuts so a women could produce enough nuts for approximately 4 days of consumption if she only cracked for herself. However she cracks for her family dependents as well. If we compare the work effort of the foragers we have studied so far we find differences in their foraging patterns and the number of hours worked. Tanaka (Table 16) reports large variation in male work effort, hours out of camp searching for food, among the !Kung he studied, 9.35 to 4.15 hours but less variation in women’s effort; 4.25 to 1.50. The Ache move camp almost daily, walking during the day, processing food, cooking, visiting and resting during the evening and night. The !Kung gather resources during the morning hours, unless they are tracking game, and process food, rest in the shade, and visit during the mid-day and evening hours. The Hadza also begin gathering early, stopping at mid-day and sometimes gathering late in the afternoon but often spending no more than 6 hours in the field. Hadza men are more similar to Ache men in the habit of going out to hunt every day. Variation in work effort and foraging returns across habitats would make a good term paper topic.
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File: Lecture 11 Meriam Aquatic Foragers
Mer island is located in the Torres Strait on the northern end of the Great Barrier Reef 142 km southeast of Papua New Guinea. With this lecture we revisit the sub-humid tropics for a study of foraging strategies of a population surrounded by the sea. The island is fringed by reef, the light blue-green color center left and zigzagging toward the center of the photo to the right of the island. The foragers of Mer practice a subsistence strategy that includes marine foraging, growing of yams, manioc, and bananas as well as keeping of pigs and chickens and subsidies from the Australian government. The gardens and domesticated animals provide a small fraction of the diet as most carbohydrates are now purchased from a small store on the island and most of the meat in the diet comes from marine resources. Three distinct strategies are found in the marine niche; hand-line fishing and netting on the nearshore at high tide, shell fish collecting and spear fishing on the reef flat at low tide, and deep and shallow water fishing, lobster collecting and turtle hunting offshore, and hunting and collecting turtles in the nesting season. This lecture has two purposes; 1) to introduce sub-tropical marine collecting strategies, and 2) to consider an important body of ideas advanced to explain widely noted falsifications of the predictions of optimal foraging strategies. Those falsifications were first noted in the finding that Ache men rarely collect palm starch in spite of the high caloric returns provided by that resource (Hill et al. 1987). The evidence that men pursue resources characterized by high variance and wide sharing, once captured, was included in the discussion by Elston and Zenah (2002), developed by O’Connell et al. (1999) in the review of the grandmother hypothesis as an explanation for the evolution of the genus Homo, and evidenced in the hunting strategies of the Kalahari San Bushmen and the Hadza big game hunters. You were prepared for these ideas by the four hypotheses advanced by Mitani and Watts (2001) to explain hunting by chimpanzee males. By now you should suspect that male foraging is about more than satisfying personal nutritional needs. Everywhere humans forage, men and women target different resources. The resources taken by women are characterized by low day to day variance in acquisition rate, by small package size and by limited distribution within the family. By contrast men target resources characterized by high variance in acquisition, large package size, and distribution to all in the vicinity of the catch. The fact that men do not discriminate between their wives and offspring and the families of other men provides strong evidence that male strategies are not about parental investment or offspring provisioning. The argument that male hunting can best be explained as sexual selection for male-male competition is set out in the assigned papers for this lecture.
Notice that women allocate very little time to spearing, 9% of their time on the reef compared to 76% of the time shell fish collecting. Because of the high cost of transporting the shells of the very large clams they collect, women process as they collect. Women collect only three species, Hippopus hippopus, Tridacna spp. and Lambis lambis, the spider conch which is very much smaller than the conchs found in tropical Atlantic waters. The largest tridacna clams can reach 140 cm, over 4 feet on the longest dimension.
Women forage on the dry reef carrying a bucket, knife, hammer and a small spear. As they collect they cut out the meat from the shell and drop it in the bucket using the hammer to crack conch shells and the spear for balance, although they do stop to spear small fish or octopus trapped in the tide pools. When the man and woman return home from the reef he shares his fish with the neighbors and she cooks her share of the fish and the resources she has collected from the reef for the family dinner. Over a sample of 44 women, female forages brought in 1962 grams/per bout, on average, while spearfishermen (n = 36) brought in 356 grams. Study Table 1 (Bliege Bird, Smith and Bird 2001) where they report returns from the ebb tide reef flats according to activity. Reef collecting of large clams gives the highest return in calories, proteins and fat. Even though men and women spend the same time on the reef, their energetic gains are quite different. Figure 1, in this same reading, illustrates the gains achieved by forages targeting shellfish compared to gains from spearfishing. Pay attention to the kcal scale on each panel. Although the curves seem comparable the gains for spearfishing vary from 0 to 1700 kcal while those from shellfish collecting vary from 0 to 5500 kcal for just over 140 mins of foraging time. Notice also the high variance in returns on panel A. The extreme outlier of just under 1800 kcal in 25 minutes of time is not representative of the average gain reflected in the regression through the mean. The returns plotted in panel B cluster more tightly around the mean. This evidence lends support to the hypothesis that spearfishing is a costly signal of male talent. Keep this example in mind when we get to the lecture on children’s foraging and the argument that male foraging strategies explain the evolution of long juvenile periods in ancestral H. sapiens. If foragers process as they go, they leave no record of past meals. Think about how we derive information on ancient diets. The archaeological record consists of remains left at central processing spots or middens, kill sites, camp sites, and dumping sites. The record is over-weighted with resources with durable discard. When foragers process as they forage, inedible portions are scattered across the landscape. Absence from the archaeological record never implies absence from the diet of ancestral humans. You can begin to appreciate the importance of ethnographic evidence drawn from foragers who still collect wild foods. The power of the simple OFT models helped ethnographers understand the evidence they collected. Even where modern foragers practice a mixed subsistence, including cash purchases of food, a great deal has been learned about the energetic costs of wild foods from the use of OFT models to understand foraging strategies for wild resources. On their small, 2.8 by 1.7 km, bounded world the Meriam practice a seasonal round. As noted above, March to early Throughout the year men take turtles in the open water. There is a rich description of this type of hunting in the required reading for this lecture. You might be surprised that a reputation for a good hunter falls on the leader of the boat who directs the driver, makes decisions about pursuit and directs the chase rather than on the jumper who actually goes into the water and wrestles the turtle. Turtle hunting is open to all men between the ages of 16-47 but participation varies. “Thus 44.5% of the 90 Meriam males ages 16-47 hunted at least once in the study period, but the 3 most active participants (3.3% of males) were over 5 times more likely to engage in a turtle hunt than the average Meriam male in this age range (Bliege Bird et al. 2001 p 14).” These same 3 men were named as the best turtle hunters in a series of interviews with other island residents. Hunting in open water is more costly than hunting on the beaches. Leaders of open water hunts are older and more experienced while young men who are jumpers in open water hunts are active participants in nesting season captures perhaps as a way of working toward hunt leadership.
In all discussion of optimal foraging across environments and in the same environment across methods we need to exercise care to make certain we are comparing net returns calculated in the same way. If search costs are debited, as they are in Fig. 7, the returns will be much lower than returns reported as post encounter return rates, the method of calculating returns using the classical optimal foraging models.
One of the features that distinguishes humans from other primates may well be male-male competition in the arena of provisioning others. Proponents of costly signaling hypotheses to explain the evolution of men’s work propose that human males signal their quality as mates and allies and their danger as competitors by providing high variance, high return resources that involve skill in acquisition and some personal risk. The caloric returns from the two types of signaling explored in this population are vastly different. The return from spearfishing is insignificant over the total collected resources but turtle hunting in the open water provides a very high caloric return shared over many individuals in very public feasts. In the first case, the spearfishermen signal the personal physical qualities that make them successful hunters of small mobile prey; hand-eye coordination, patience, and endurance. Their successes on the reef provide few caloric benefits to observers, but provide useful information to potential competitors and allies. Those who attend to the signal receive honest information about the quality of the man because the task is too difficult to be accomplished by those of lesser talent. In the second case, turtle hunting in the open water, observers gain not only information but valuable calories and opportunities for communal feasting. Pay attention to the predictions Bliege Bird et al. (2001) derive from the costly signaling hypotheses and how they are tested. I have also assigned Hawkes and Bliege Bird because they review Zahavi’s handicap principle, link it to the analysis of chimpanzee hunting by Mitani and Watts (Lecture 2) and to Veblen’s classic analysis of conspicuous consumption. Finally, Hawkes and Bliege Bird (2002) argue that reciprocal altruism fails as an explanation of most human sharing patterns because pair-wise exchanges for large meat packages have not been demonstrated. These are important arguments and you should be clear about them before the second exam. If you are not familiar with modern arguments linking conspicuous consumption and sexual selection in humans, have a look at Geoffrey Miller’s prize winning essay Waste is Good. This essay should be especially interesting to those of you majoring in psychology or marketing.
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File: Lecture 12 Juvenile Foraging
Unlike other primate females, human mothers have overlapping dependents, a new child is born before the previous child becomes independent. A number of hypotheses have been developed to explain who feeds a women and her child when her foraging returns are constrained by child care. As illustrated in this photo and those in previous lectures, women take their nursing infants with them but here you see a !Kung women from the Dobe area carrying two of her dependents plus the food she has just gathered. In the early versions of the man the hunter hypotheses, investigators imagined that women with dependent children remained in a central place caring for children while men provisioned the family with large game. A more recent hypothesis links the evolution of long juvenile dependencies to food sharing between older female kin and young dependents (see O’Connell et al 1999). In environments of low extrinsic mortality young animals can risk devoting more time to personal growth, growing longer and larger before maturing. In the mosaic habitats of the Plio-Pleistocene boundary mothers and children were able to move into new habitats because long-lived grandmothers helped provision weaned infants. In turn the benefits of provisioning increased longevity genes in these populations, further reducing average adult mortalities and promoting later age of maturity. It is late age of maturity that provides long juvenile periods. The consequence of selection for longevity may well be long learning periods but as always we should be careful to separate cause and consequence in evolutionary puzzles. The authors assigned in this lecture review these arguments over and over. Be sure you understand the different hypotheses because they are all related to the occupation of new habitats, the provisioning of juveniles, the evolution of the human diet, and the evolution of longevity in our lineage. By now you have ample evidence, from ethnographic studies, demonstrating that the man the hunter scenario under-estimates the work of women who forage for themselves and their children even when diets are supplemented by widely shared resources provided by hunting men. The second proposition of the hunting scenario, that human children are dependent until they become adults is true for the !Kung and some South American subsistence horticulturists but quantitative studies of other foragers demonstrate that young children and juveniles provide some of their own resources. Differences across habitats and subsistence strategies are indicators that juvenile strategies might be structured by ecology and physical development and not by learned capabilities. The differences, across habitats, bear upon the modern version of the man the hunter proposal, the embodied capital hypothesis, which proposes that the long juvenile period in humans evolved in response to the value of a long period of learning for mastering adult foraging skills. Once individuals, males in this particular hypothesis, master hunting skills they are able to provision mates and juvenile dependents. The evidence that children can be effective foragers at a young age challenges this proposition. The various explanations for the evolution of our unique life histories are well rehearsed in the assigned readings for this lecture.
Hadza children as young as 5 years old forage near camp for resources to supplement their diets. In certain seasons, children are left at home when mothers, carrying only nursing infants, go to dig tubers. Detailed observations reported in Blurton Jones, Hawkes and O’Connell (1989) showed that infants older than 2 1/2 years rarely accompany mothers to tuber foraging patches. Children left in camp forage on tubers growing near the surface, the honey of stingless bees, and baobab fruits. The latter fruits have high processing costs as the pods need to be cracked and the pith pounded into a dry powder. This powder is moistened with water or eaten dry with water. The seeds can be cracked and the kernels eaten raw. In spite of high processing costs, children aged 5-10 years can gain 629 cal/h from baobab, while those 10-15 get about 1014 cal/h. However children do not allocate many minutes to foraging so it would be a mistake to imagine that Hadza children entirely support themselves. Honey is mostly taken by boys between the ages of 12-15 if one can get an axe to chop out the nest. Honey yields about 339 cal/h but boys who accompany women on foraging trips often acquire 650-1350 calories on these trips. Blurton Jones et al. (1989 p 380) estimate that “If children spent just 2 hours per day foraging for baobab, or (for older boys with access to an axe) honey, they would acquire around 800-1000 calories, almost half the calories they need.” In a subsequent field season, the same research team completed time allocation studies to determine how mothers change their own foraging patterns in response to opportunities presented by resources they can gather with their children (Hawkes, O’Connell and Blurton Jones 1995). Contrary to the expectation from optimal foraging theory that mothers choose foraging patches that optimize their own return rates, they found that women, in certain seasons, choose to forage in berry patches with their children instead of traveling to patches to collect tubers. Mothers maximize team foraging rates by focusing on resources that children can gather with adults. “If higher rates of food acquisition are advantageous to women because with higher rates they can feed children more, or feed more children, then when children are active foragers themselves a woman’s children will consume food at a higher rate if she chooses the strategy that maximizes the team rate she and her children earn collectively, even if the rate she earns herself is less than the maximum possible (Hawkes et al.1995 p 695 emphasis original).” The team rates achieved for mothers and children depend upon the number of children and the time allocated to foraging. The more children a mother has the higher the team rate. If the team forages for longer than 556 mins, berries are the best choice. If a women wants to maximize her own return rate collecting tubers she needs to forage for an additional 2.5 hours. Over the mean length of observed trips, women and children do best focusing on berries. The foraging strategies of mothers and kids might have been influenced by the scarcity of baobab fruit during the late dry season of 1988 when the time allocation studies were completed. In spite of children’s abilities to forage for themselves around the Hadza camp, they still depend upon mothers. The time allocation data show a significant difference between foraging time for children with co-resident mothers and those without. Notice on Table 2, copied from Hawkes et al. (1995), that an 8 year old girl with no mother or father in camp spent more time foraging than any other child. In a sample of 20 juveniles, aged 3.5 to 17 years old, 9 had no coresident mother in camp.
The caloric returns children achieve from their own foraging efforts depend upon the resources they target. The two types of tubers, Makalita and //ekwa, gathered by children yield 73 to 85 Cal/100 g and children over the ages of 7 and 8 can gather 598 and 314 g/hr, respectively, of these two resources gaining about 200 calories for 29 mins of digging Makalita. When children accompany mothers to the berry patch they can gain from 964 to 2,223 Cal/hr depending upon which berry, Salvadora persica or Cordia sp. they are picking. Children gain about 50-70% of the adult rate picking S. persica berries, Table 6 (Hawkes et al. 1995). Since Cordia berries were not fully ripe until the end of the observation period, rates by age were not available for the berries yielding the higher caloric return. There are two important take home points from the Hadza field work on children’s foraging. 1) Hadza children’s foraging choices are consistent with optimal foraging theory predictions, near camp children generally choose to forage on the tubers with the highest caloric return just as they do in the berry patch. Before Cordia berries ripened children picked berries giving a lower return but once the Cordia ripened they ignored S. persica berries even though they were still available. 2) Mothers adjust their foraging strategies to include resources their children can gather. In berry picking season mothers forego tuber picking, where they earn high returns, to travel long distances to berry patches in order to maximize the team rates they can gain with their children. The stashing rate shown in Table 6 is defined as the “total weight of fruit accumulated for transport per hour in the resource patch. They represent the minimum rate that would have been taken had foragers eaten none of the berries they picked (Hawkes et al. 1995 p 693).” Actual picking rates are a combination of eating and stashing rates. The last column is an attempt to measure the amount of time spent picking, the stashing rate and consumption rates are added and divided by the tin measured rate. The results show that women are the most steady pickers and men, boys and children the least effective. Men may have spent more time in the berry patch in this particular field season as “hunting was poor(only eight large animals taken over 43 days of direct observation)(Hawkes et al. 1995 p 689.).” Men pick at very high rates but they also consume at a high rate so their stashing rate is lower than that of women and girls.
The foraging efforts of Hadza youngsters stand in sharp contrast to those of Dobe !Kung children who do almost no foraging. Instead !Kung children contribute to their own subsistence by cracking mongongo nuts carried home from distant patches by their mothers. Hawkes, O’Connell and Blurton Jones (1995) attribute the difference to the scarcity, in the Kalahari, of near camp resources that children can gather and the high, dry season temperatures and lack of water that preclude taking children on long walks to the nut groves. Instead !Kung mothers and kids do better if toddlers and older youngsters stay in camp and crack nuts that mothers have carried home. Recall the topographic differences between Hadza country and the Kalahari with no prominent landmarks to guide children from foraging sites to home camps. Blurton Jones et al. (1989) suggested that the harsh dry season temperatures keep !Kung children from foraging with mothers and the danger of getting lost in the flat environment restrain children from foraging alone near camp.
At both of their field sites, Mer Island and the Western Desert of Australia, Douglas and Rebecca Bliege Bird conducted quantitative studies of juvenile foragers to determine if optimal foraging theory could explain choices made by children. At both sites they found that children take resources that give them high rates of caloric return as predicted by theory but their foraging returns are constrained because the rate at which they encounter resources is determined by size and walking speed rather than age or experience. Their evidence indicates that physical abilities rather than learned capabilities determine juvenile foraging strategies. On Mer Island, mixed-age groups of children, independent of adults, forage on the reef flats after school or on weekends. On these trips they collect some of the same resources collected by adults but they encounter them at a lower rate so their overall returns, in the patch, are not reduced by stopping to take smaller shellfish such as black lipped conch and small top-shell. Post encounter processing costs of the large shellfish are higher for children because they do not have the upper body strength to cut out and remove edible flesh from the largest shells. These differences, between adults and children in physical strength rather than skill, explain why children take resources ignored by adults. Further Bliege Bird and Bird, in Children on the reef (2002), suggest that children are very efficient at extracting meat from small shells because they can more easily reach in the small valve openings with their small fingers to pull out the meat. Mer children also fish, from the beach, a foraging strategy requiring some skill. Across the Torres Strait islands, Meriam kids are known for their skill in hand-line beach fishing. Children begin as young as five years old and quickly master the technique. After an initial steep rise in efficiency there is little effect of age on efficiency. However, at every age there are large differences in individual skill and efficiency such that most of the variation in this task is explained by individual differences with one 62 year old man out-ranking all others in return rate. Figure 1 taken from Bliege Bird and Bird (2002b) illustrates the scatter and the large disparity when a very good older fisherman (the open circle top right) is included in the analysis. Residuals of overall return rates are plotted on the y axis to remove the effects of varying sample numbers across foragers. In spite of the title, return rates are calculated from both small-hook and large-hook fishing methods. Men and children of both sexes choose large-hook fishing and women choose small-hook. One difference between children and adults is that children usually have fewer choices of line size. From the figure, we might conclude that only the most skilled children choose to hand-line fish but without knowing the proportion of children who choose to fish it would be impossible to say the sample is self selected by only the best children.
Martu (sometimes Mardu) children of Northwest Australia are often left at home when mothers hunt goanna lizards in the sand hill flats but these children are active forages in patches distinct from those targeted by their mothers. Bird and Bliege Bird (2005) argue that size determines the The analyses, of Martu children’s foraging success, was designed to tease apart the factors which influence foraging choices of juveniles. The regression plots show that height has a larger effect on return rates than age (compare Figs 6.2 and 6.3). The proponents of the embodied capital hypothesis for the evolution of human life histories propose that the long human juvenile period evolved to provide a long period of subsidized dependence during which children forego supporting themselves in favor of learning the foraging skills that are later cashed out as productive adults. In this scenario juvenile dependency is subsidized by paternal support of wives and children. Bird and Bliege Bird show youngsters can be effective foragers using the same tools and techniques as their mothers but foraging in different patches. Since within patch encounter rates constrain juvenile choices, size not experience makes a difference to children’s effectiveness. These results are important because they undercut the proposition that long juvenile periods, in humans, evolved in response to the value of learning adult foraging skills. Instead they support the proposition that long juvenile periods evolved in response to the benefits of growing larger before maturing. The question remains, why don’t children forage longer hours? The work effort of the 8 year old Hadza girl with no co-resident parents suggests children are capable of longer work hours. It is possible that children allocate energy to fogging effort while still conserving energy for growth. Humans are determinate growers which means we reach skeletal maturation and hence most of our adult height about the time of sexual maturity. Bliege Bird and Bird (2002a) suggest that foraging efficiency increases after puberty because both males and females have greater opportunity to enhance fitness gains from foraging efficiency. During the thousands of years spent subsisting on wild resources, males excelled in male-male competition through costly signaling of hunting ability and providing resources for public consumption. In this same niche, females promoted their own reproductive success by gathering low variance resources to support their dependent children and grandchildren. Foraging efficiency increases at puberty in response to natural selection on females and sexual selection on males.
In Growing Up Mikea, Tucker and Young (2005) review the various arguments for the long juvenile period in human life histories and report detailed time allocation studies of Mikea tuber foraging. They emphasize what you have learned earlier in this lecture, the cost of children varies across habitats. In the dry woodland forest of Madagascar, where the Mikea live there are few dangers for children foraging in the woodland, no poisonous snakes or large predators and plenty of shade to moderate temperatures. Even though the Mikea practice a mixed subsistence economy that includes subsistence farming and herding, trade, and cash labor they still collect wild resources. Figure 7.2 shows that children spend much more time in leisure activities than other age classes but by adolescence young people spend as much time working as do married adults. Notice also that adult males and females allocate nearly the same proportion of daylight hours to food production. Among children, boys allocate nearly twice the time to food production as do girls but these figures change when allocation to tuber foraging is measured. Adolescent girls devote more time than any other age group to this task (Figure 7.3). Still across all foragers net acquisition rates increase with age and men achieve significantly higher rates than women but Table 7.2 shows that daily returns for men are low because they spend 1/3 the time that women spend foraging for tubers. In the early dry season, Mikea children actively forage with adults and mothers, who suffer no decline in foraging efficiency when accompanied by children. Notice in Figure 7.8 that adult females forage in groups with children and adolescents more frequently than do adult males. In certain seasons adults and children forage in the same patch but when patches near home are depleted in the largest tubers adults range further to exploit new patches while children stay in the older patches harvesting smaller tubers that lie close to the surface. The ovy tuber, collected by the Mikea, grows in sandy soil so digging and collecting is relatively easy. The ease of digging ovy combined with its higher energy density produces a net return rate much higher than the tubers collected by the Hadza. Have a look at Table 7.2 and notice that children gain 505 to 537 kcal/hr gathering ovy but they gather only 35-41 minutes/day. The same question arises over and over for children, why don’t they devote more hours to foraging? Tucker and Young suggest that children are neither rate-maximizers or time-minimizers. Since they are provisioned by adults they are also not energy limited. Instead they forage for the physical and mental challenge and for the social opportunities afforded by mixed age play groups. However the ability of Hadza, Merriam, Martu, and Mikea children to collect resources for themselves suggests that they might be able to support themselves if adult provisioning failed. As illustrated from the detailed Hadza data, children often have no co-resident parents. In such cases orphans must rely on near-relatives, neighbors and their own ingenuity. We expect natural selection to favor strategies, in young mammals, that help them survive maternal accidents. To expect youngsters to forage on the same resources as adults is unreasonable given their size and physical strength. The studies reviewed in this lecture demonstrate that children can be capable foragers depending upon the opportunities in their environment but over all environments, given adult support, they choose to allocate more time to leisure.
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Journal of Archaeological Research, The Archaeology of Aquatic Adaptations: Paradigms for a New Millennium Jon M. Erlandson1 Although aquatic resources are often Pleistocene cultural complexity, most ignored the role of aquatic or maritime human history. When did aquatic first play a significant role in by (1) reviewing various theories on lems that prevent archaeologists from the archaeological record for evidence conclude that aquatic resources, accessible, have probably always been Evidence suggests, however, that seafaring) played a significantly expansion of anatomically modern humans significant expansion occurred more sophisticated seafaring, fishing, KEY WORDS: aquatic resources; human
The average molluscan flesh is humans apparently existed for oyster. In any event, shell (Meighan, 1969, p. 417). Central to the success of our cal range and astounding population 1 Department of Anthropology, oregon.uoregon.edu.
288 intelligence, adaptive flexibility, torical or evolutionary framework, opportunists, omnivores who thrived in both natural and cultural. On a planet life itself is dependent on water to fully adapted for at least 2.5 million modern anthropological theory has were not systematically used by humans 1968; Cohen, 1977; Osborn, 1977a,b; 1968; Yesner, 1987). As Bass (1972, p. and broken by rivers and streams or habitats listed by Gamble (1994, pp. cestors as they spread around the habitats are nowhere to be found. As Washburn and Lancaster (1968, many archaeologists still seem to During most of human history, barrier and the inability to cope absence of remains of fish, using boats. There is no evidence pre-agricultural period . . . for More recently, Yesner (1987, p. 285) that maritime resources were not record has attracted a general ways did not precede late Upper If such statements are accurate, colonizing much of Africa and Eurasia aid of floats, boats, or the they survive in such a wide range of a physical and psychological ancestors—problem solvers and keen aquatic resources when hundreds of vores do not? Why is there so little resources until postglacial times, long nization of island Southeast Asia and aquatic resources were virtually I believe the general perception aquatic environments during the last on the evolutionary study of aquatic and the development of boats and other peripheralize the significance of them to an essentially incidental role The Archaeology of Aquatic Adaptations agricultural societies and marginal role in a relatively brief trajectory departed from its natural into increasingly artificial modes of As Yesner (1987) noted, however, foods of “last resort” is out of suggest that maritime or aquatic populous, and culturally complex than (Birdsell, 1953; McCartney, 1975; of the most complex and artistically veloped in rich marine environments, Tlingit, Haida, Aleut, Koniag, etc.) relatively unproductive for human ported some of the most complex and archaeological evidence for the limited. This results in a fundamental resources (although often both diverse nomic foundation for relatively complex tions and elaborated material cultures. for problematic aspects of such models aquatic paradox has yet to be In this paper, I discuss some of the nature and antiquity of aquatic broader implications for our human subsistence and technology, and human economic intensification, and short summary of historical thought then discuss some epistemological, that currently prevent any real aquatic adaptations. I then review the resource use and maritime migrations and some approaches I see as aquatic adaptations as we embark on our A BRIEF HISTORY The study of coastal and other thropology and archaeology, one that the two fields. Despite this long bate about the nature of aquatic human societies, and the role they have 1975, 1978; Binford, 1968; Claassen, 290 1995a; Glassow and Wilcoxon, 1988; Osborn, 1977a; Parmalee and Klippel, and Stocker, 1983; Raymond, 1981; Lancaster, 1968; Wilson, 1981; Yesner, the “New Archaeology” of the 1960s coherent body of theory on the broader ions expressed on such matters were varied widely (see Clark, 1936, p. 140; as Clark (1936) and many others and widespread shell mounds with middens and relatively intensive an important component of the Bailey, 1978; Binford, 1968). As an thesis came into vogue in the 1960s and focused on more global approaches to tions to aquatic environments. In 1994, largely for heuristic viewpoints in this debate as “Garden (Erlandson, 1994, p. 273). Garden of aquatic habitats as veritable inexhaustible and easily harvested—was 1995a; Hewes, 1968; Morgan, 1877, p. 1962). On a global level, such scription of the role of the sea in . . . the path of our evolution sea. No other setting is as the tidal shore, presented the diversity and abundance of ecologic niche in which animal Similar statements linked to specific or to regional archaeological sequences poused by a number of authors. Such however, that archaeological records for such aquatic largesse dating back level, moreover, the accumulation of chronometric dating techniques made If aquatic resources were so the archaeological record for their After the 1960s, following the scholars explicitly asserted that to the hunting of large terrestrial exploitation (e.g., Bailey, 1978; The Archaeology of Aquatic Adaptations et al., 1971; Osborn, 1977a). These the prevailing view of the time that, dominated big-game hunting was the and technological evolution. Shellfish in such models as marginal or even costly to harvest or process, poor unreliable, or requiring high fact that collecting shellfish and work in most ethnographic societies man economies (Claassen, 1998, p. 175). fore, that the archaeological record aquatic resources and the relatively ing peoples. They argue that humans did aquatic resources until the the intensive harvest pressure of extinction of the Pleistocene (and is) often assumed to be evidence degradation. Osborn, the most ardent cestors “ignored” shellfish and (Osborn, 1977b, p. 301) and that the virtually universal. . . . marine resources are sification, in the case of their low protein content. A that is too costly for human dependent selection. This given the option, we should expect tion of the sea, at least in p. 177). In practice, relatively few such polarized schemes, and most is considerably more complex. model has heavily influenced the work have worked with or discussed coastal (e.g., Bailey, 1975; Binford, 1968; Hayden, 1981; Isaac, 1971; Kelly, 1996; To square such a dismal view of evidence that many coastal societies tion densities, sedentism, and cultural further explanation. Osborn (1977b) societies was exaggerated because their in density calculations, but he could sedentism and cultural complexity. 292 Northwest Coast societies was a result never adequately explained how they marginal environments. Yesner (1987) cated explanation for the coastal environments were relatively a combination of megafaunal tion, and the development of mature to bloom. Thus, he argued, humans did until relatively late in human history, aquatic habitats ultimately fostered ity typical of many Middle or Late model include significant variation in tinction or survival, the considerable widespread extinctions, and little were relatively unproductive prior to As we shall see, none of these basic paradox of supposedly low aquatic tions and cultural complexity, or for that aquatic adaptations developed viously believed. Nor do they explain identical data sets can come to such velopment of such basic aspects of however, we must first review some of of aquatic resources, then examine some comprehensive understanding of the AQUATIC Much has been said about the sources: shellfish, fish, sea and others. I do not review these be the subject of an entire paper. It to examine some of the divergent classes of aquatic resources. Also often collectively lumped as “small” that they are therefore less productive subsistence and their presence in for resource stress or economic parate viewpoints about the major classes of aquatic organisms not plants, etc.) may also be significant The Archaeology of Aquatic Adaptations
Curiously, perhaps the single freshwater for drinking, is seldom on water is so fundamental and so is significant, however, because the our ancestors to aquatic habitats for resource, drinking water determined especially in relatively arid regions. habitats, hominids would have spent a animals in such environments, including or scavengers that fed on aquatic Under these circumstances, it seems have prevented similar opportunistic of our earliest ancestors living along similarities between many of the lakes, rivers, estuaries, and marine learning curve would have been required habitats. The intensity of such aquatic of course, depending on the relative the other subsistence pursuits As noted above, the notion of a with aquatic habitats is also difficult ancestors now appear to have spread 1.7 million years ago. How did they if they were afraid of the water and simple rafts, boats, or other flotation
No class of aquatic resources has ogists than shellfish (e.g., Bailey, 1998; Erlandson, 1988, 1991; Glassow 1995; Meehan, 1977, 1982; Meighan, Osborn, 1977a; Parmalee and Klippel, 1987; Yesner, 1987). The generic term ety of aquatic invertebrates, dominated also including crabs, sea urchins, organisms. Although the size of ably, from large octopi or giant clams shellfish are relatively small 294 shellfish make up for in quantity and and sessile aggregations. While most plete animal proteins and some vitamins carbohydrates, and calories (see ten been portrayed by anthropologists indicate that mussel beds produce one on earth (Jones and Richman, 1995). Since at least the early 1900s, and seemingly innocuous creatures as for humans. . . . procuring the essentials of existence. In all parts of the gathering for food the shells to the lower classes of society (Uhle, 1907, p. 31). Some archaeologists bolstered such nutritional content of shellfish calculated, for example, that 156,800 yield of one red deer. Some of these ignored the fact that shellfish may source or that they were often a source that could be gathered by children, and the elderly (see Meehan, 1977). The fact that shellfish most ethnographic societies (Claassen, suggests that such comparisons of inappropriate. Some scholars have also argued low caloric content, and their were relatively laborious to process countered that they required little could provide highly reliable and the high failure rates of hunting some researchers extolled shellfish as others noted that a heavy reliance on poisoning” (Noli and Avery, 1988), on many land mammals (bison, rabbits, (Buchanan, 1988). While some criticized to periodic El Nino, storm, or red tide could sometimes be predicted and that agricultural products and other to floods, droughts, disease, and The Archaeology of Aquatic Adaptations Finally, though Osborn (1977a,b) historical accounts to support the foods, Moss (1993) clearly exposed the often inherent in such accounts. I was Tlingit friend, Richard Newton, who his people. Responding to questions remains in Tlingit village and camp acterized their dietary role as similar society (Moss, 1993, p. 643). When explained that the ideal Tlingit man encouraged laziness because they were primarily by women, but that they also by Tlingit men (Moss, 1993). Certain Tlingit men, in fact, because they were All this debate has had little and other small resources are lower and O’Connell, 1999; Renfrew and concluded, for instance, that “no one diet” of any ancient society. continue to view the appearance of as evidence for human demographic economic intensification (e.g., Cohen, postglacial florescence of shell world, therefore, has become notion that human economies were broad-spectrum revolution.
Similar debates have taken place (e.g., Butler, 1996; Clark, 1948; 1979; Lindstrom, 1996; Morgan, 1877; Literally thousands of different habitats, from the deep abyssal floors adults, these fish range in size from largely solitary and relatively rare, in concentrated schools numbering in moreover, are characterized by a only one or two species and even these have low species diversity but a The nutritional value of fish calorie content of various species. 296 are a relatively nutritious source of Watt and Merrill, 1975). Fish eggs, quantities, are also generally very are also highly digestible and than the meat of land mammals. High populations—especially certain fish with lower rates of disease and greater Opinions expressed about the Some accounts have portrayed fishing ning with Morgan’s idealized unlimited in supply, and the only kind 1877, p. 21). In contrast, in comparing Kelly (1996, p. 209) stated that . . . fish are different. Some but not bottom feeders. And fish oceanic fish. The forager can randomly. If there are no fish accepting this as likely. Kelly’s characterization, however, is including the extremely productive and halibut or cod banks, kelp beds, and Others have argued that fishing and high technological investments. (1979) suggested, however, that under could be extremely productive even habitats, the seasonal drying of ponds on mud flats where they can be easily or anoxic conditions can also lead to of dead fish are deposited on the Stocker (1983, p. 549) described an known as “anchovy beaching,” in themselves on the beach roughly four lamprey eels, grunion, and many others) predation, and such spawning runs are logistical planning required for mass storage. Even when more sophisticated these need not be especially elaborate tidal weirs, for instance, can greatly truly impressive yields. Before California’s marine fisheries, available in nearshore and estuarine The Archaeology of Aquatic Adaptations huge quantities of these small fish later consumption. Still, considering prior to the advent of the Upper nologies involving cordage, baskets, beyond the capabilities of hominids ern humans. And some fishing sophisticated boats, or elaborate weir able investment in materials, labor, communication skills that may have been ancestors. Despite such technological ies have modeled the productivity of tive terrestrial subsistence pursuits 1987). Some of these estimates are propriate technologies in potentially still informative, suggesting that the The most sophisticated analysis of Truckee River fishery in the western gests that fishing harvests using a higher than the return rates calculated Lindstrom’s projected return rates ods of fishing produced yields that terrestrial alternatives.
There has also been considerable nomic productivity of aquatic mammal whales, seals, sea lions, sirenians Clark, 1946, 1947; Colten and Arnold, and Jones, 1992; Jones and Hildebrandt, Workman and McCartney, 1998). Aquatic freshwater animals (hippopotami, spend varying amounts of time in the sometimes be taken on land. Some rized as clearly marine or freshwater: distances up rivers; seals live other European lakes (Reeves et al., salt- or freshwater habitats; and river may also spend time in brackish or Clearly most aquatic mammals are of the largest animals on earth, which 298 hunting also were relatively abundant marine mammals weigh well over 500 kg, 100,000 kg. Osborn (1977b) argued that relatively high in the food chain, productivity of the world’s oceans. no relevance, however, to maritime lived in proximity to biannual pinnipeds (see Haggarty et al., 1991). Like virtually all mammals, the relatively rich sources of nutrients, Heller and Scott, 1967; Osborn, 1977b; marine) mammals also have a thick layer them with insulation and human hunters fat deposits can also be rendered into or used in lamps as a source of heat baleen of many aquatic animals also variety of technologies (houses, boats, many societies that actively pursue also gain significant status, Such potentially lucrative costs and risks of procuring aquatic gerous and seasonal pursuit, especially hunting forays are often relatively may also require relatively complex and thy boats and related hunting gear that to produce and maintain. This is recorded among ethnographic marine population densities and had hunted sea negative effects on the distribution Hildebrandt, 1995; Lyman, 1995). Prior been taken relatively easily while islands or other isolated coastal and pinniped carcasses off the beach huge) subsistence dividends, with and Kinahan, 1983). Finally, the costs must be measured against the greater hunting, fishing, and transportation As with virtually all classes of siderable variability in the potential. This includes aspects of availability to humans, the methods tivity of various procurement The Archaeology of Aquatic Adaptations diversity, it should be no surprise ferent conclusions about the general In recent discussions, for instance, mals as either central or peripheral to along the Pacific Coast of North and Hildebrandt, 1995) proposed that to predation in rookeries, some seals rine hunters, with later technological intensification as human impacts on strategies changed. Colten and Arnold little evidence for an early focus on suggested that its general economic (see also Kent, 1989, p. 5; Workman and resolving such debates are problems sentative samples of sea mammal remains within the larger economies of human PROBLEMS IN Underlying such debates, but ambiguity of the archaeological record have been supported with data from conclusions were drawn from virtually it possible for researchers to reach ysis of the same body of data? The taphonomic, methodological, reconstructions of the history of divergence of opinions about the to a variety of problems with the the way individual archaeologists differences in the preconceptions of
In part, different opinions can tion for what constitutes a dietary or terms such as coastal, aquatic, McCartney, 1998). Definitions for instance, from those groups who procure the sea to those who go to sea in boats Recognizing the complexity and 300 (1975, p. 344) tried to bring some by defining five broad adaptive ified marine, maritime, and riverine. (1991) differentiated littoral from groups who went to sea to obtain much to operationalize the definition of (1980, p. 728) defined “fully their calories or protein from marine riverine or lacustrine peoples, but in quantify the dietary contribution of and trace element studies of human bone general aspects of ancient diets, but a photosynthetic pathways, etc.) continue At times, we must even confront versus terrestrial resource. How do we that may be caught in the ocean one scavenged from the shoreline the next? mus, crocodile, land otter, or many in aquatic habitats but may also be taken from terrestrial colonies aquatic or sea lions taken from onshore elk captured—as they were sometimes America—while swimming to or from paralyzed by the cold on the beach just such ambiguous cases are relatively mon than many of us recognize, and they drawn between aquatic and terrestrial lacustrine habitats. If such behaviors, moreover, how can we hope to cases in the archaeological record? Changing Sea Levels, Coastal Despite such ambiguities, the tory of aquatic adaptations lies in the tremendously over the past 2 million repeatedly obliterated the resource use is most likely to be Quaternary, exceeded only by Last Many scholars are rightfully hesitant were once widespread along submerged The Archaeology of Aquatic Adaptations is ample reason to believe the riously underrepresented (Kraft et al., years ago, world sea levels stood exposing broad coastal plains around dated as seas rose to their present times during the Plio-Pleistocene, in coastal geography around the world. Worldwide, only Africa and levels were last comparable to today. terglacial sea stand of 125,000–130,000 have destroyed most evidence for global sea levels have risen ated with lower shorelines has either or both. Even today, with sea level many important coastal sites (e.g., Grotta dei Moscerini, Daisy Cave) years ago are being destroyed by marine with earlier interglacials are present maxima represent just a small fraction that associated occupation sites (e.g., are localities such as those in North artifacts (hand axes, etc.) have been testifying to the destruction of (e.g., Bar-Yosef, 1994; Howe, 1967). Equally important for adaptations are the effects of sea localities. As sea levels rise or fall, changes; the environmental setting of Reconstructions at coastal sites with that the exploitation territories of entirely terrestrial during earlier and van Andel, 1980). The maximum last 20,000 years, for instance, have areas (e.g., northern Australia) to have moved less than about 10 km are with relatively early evidence for ahead). Reconstructing the because a cave or open site located on or more from the coast at various times Study of modern coastal than about 5 or 10 km from a home base Meehan, 1982). When they do hunt or 302 shellfish, fish, or sea mammals are In most situations, therefore, sites shoreline are unlikely to contain Distances of even 1 or 2 km can remains (Wing, 1977). During periods of the intensity of aquatic resource use on its proximity to coastal habitats. of the last 17,000 years, coastal sites evidence for an intensification of in local environments rather than a human subsistence (see Bailey, 1983a; Some may argue that the loss of amining the antiquity of the human use problems inhibit such comparisons. diversity of most freshwater habitats nities. Second, it is not clear if the habitats is any more representative. dynamic, and climatic, glacial, and sea on their structure and productivity. shoreline erosion can produce shorelines. In riverine systems, while depositional cycles can bury them preservation and visibility problems systems as they are in marine Differential Another problem lies in the of organic remains. As we all know, the the primary record of human use of archaeological sites. Acidic soils, for acids in neutral soils, commonly lead in archaeological sites. In by relatively sedentary peoples, the mitigate the effects of soil acidity or or bone. For the Paleolithic or believe humans were relatively mobile, have been insufficient to counteract or other sites located some distance aquatic food remains was limited by shells and bones exposed to dilute acid deteriorate faster than bones, probably The Archaeology of Aquatic Adaptations lower collagen or lipid content. At a Hidden Falls in southeast Alaska South Africa (Goldberg, 2000), while shells had either disintegrated In the case of Die Kelders, despite the site strata, decalcification of the site while bone fragments were Among animal bones alone, the mals are more likely to be preserved in and Chatters, 1994). There has been ative survivability of skeletal remains but differential bone density is a aquatic vertebrates generally have tible to chemical dissolution and some taxa (sharks, etc.), fish bones high surface area to volume or mass vulnerable to chemical deterioration. rays, sturgeon, lamprey eels, etc.) bony parts, and small bony fish (i.e., The bones of many aquatic mammals are to differential deterioration from Numerous studies clearly show ologists dramatically affect the blages. Studies of faunal recovery, for fish bone and shellfish remains in vated sediments through coarser (0.25 1994; Garson, 1980; Koloseike, 1968; evaluating the evidence for aquatic where researchers had limited interest systematically recovered, or fine-screen tigators now routinely collect faunal flotation, but others still rely on Because the importance of hunting been emphasized, there have sometimes of other faunal remains from Upper Paleolithic subsistence, in fact, large land mammals (e.g., Barker, 1974; that produced a variety of faunal Gibraltar, I was surprised to find a in the Gibraltar Museum, materials at Gorham’s Cave. For some reason, any of the site publications, even shellfish were all published (see 304 1951, 1964; Zeuner and Sutcliffe, Middle and Upper Paleolithic levels at Morocco (Howe, 1967; Howe and Movius, sites from the south coast of the duced seal, fish, and “a series of” 21). Although vertebrate remains were fied (Arambourg, 1967). Description of statement that a “number of mollusks chaeological deposits in the Mugharet J. Clench of the Museum of Comparative for our purposes came of this however” Such problems may have been due, could identify and analyze the remains however, of the lower priority such as shellfish and fish that were tant. In Howe’s synthesis of the for instance, the description of stone remains are relegated to 5 pages in an short and obscure paragraph. The Even if we can overcome such still confronts us. This is the on archaeology. The historical economies has been dealt with at length 1997). The remnants of this outdated decade after most scholars recognized the meat early hominids consumed and than recognized in earlier us that human dentition is eclectic diet (Scott and Turner, 1997, the long haul species are rarely well medicine and nutritional studies show human health, growth, and reproductive as our hominid ancestors spread around success was their ability to adapt to a their relatively eclectic and Nonetheless, many theoretical compare the yields of shellfishing or many hominids relied heavily on the relative productivity of gathering The Archaeology of Aquatic Adaptations yields in such cases and must have been predictions based on optimal foraging human societies as groups of generic divisions of labor in hunting and cultures. Even for Holocene peoples, hunting yields to predict dietary priate, since large-game hunting was and some other aquatic resources were and older individuals. There is little of shellfish gathering in human the work of women or commoners, to an and to biases in historical and (Claassen, 1991, pp. 278–279; Moss, Until anthropology transcends some cance of aquatic adaptations will structions of human evolution. These large land mammals were virtually resources for our hominid ancestors; the central force that shaped human velopments; (3) that the utilization of for demographic pressure or resource preserves a representative picture of ARCHAEOLOGICAL EVIDENCE FOR OF AQUATIC Given the nature and ubiquity of know so little about the history of opment of aquatic adaptations we are theoretical assumptions and blessed where faunal preservation is systematically recovered and completely synthesis must rely on an from sites preserved above modern sea dating between about 120,000 and 15,000 from the modern coast. Despite such problems, numerous source use have been listed over the and Waselkov (1987) or mentioned by Scott, 1986; Yesner, 1980). None of appeared, and additional data have In Tables I–III, I have compiled my 306 Table I. Some Early Old World Description Locality/site and Homo habilis Senga 5, Possible use of Semliki molluscs, and River, Zaire with Oldowan Olduvai Gorge, Possible use of Tanzania crocodiles, and molluscs. Homo erectus Olduvai Gorge, Possible use of Tanzania crocodiles, (hippo), molluscs, and Kao Pah Nam, Pile of Thailand against cave hearth and Holon, Israel Freshwater turtle and hippo Acheulian scavenged (?) Mas des caves, Seal remains Lunel-Viel, located ca. 10 France Mediterranean Archaic Homo sapiens
Hoxne, England Remains of fish, waterfowl Acheulian similar to cultural
Duinefontein 2, Sea bird South Africa remains in dominated by
Terra Amata, Shellfish and France associated coastal
Lazaret, France Marine shellfish context.
Ramandils, Marine shellfish France Middle food remains.
Kebibat, Rabat, Aterian shell Morocco coast, remains.
Presqu’ile du Aterian site on Canal, contains Berard, limpets. Algeria
Haua Fteah, Marine shellfish Cyrenaica, strata. Libya The Archaeology of Aquatic Adaptations
Description Locality/site and
Mugharet el ’Aliya, Marine Morocco remains in strata.
La Grotte Zouhrah, Aterian Rabat, Morocco shellfish crab), Homo
Grotte des Aterian shell Contrebandiers, Coast Morocco sapiens
Devil’s Tower, “Thick Gibraltar Mousterian heap” of
Gorham’s Cave, A variety of Gibraltar from several occupation
Grotta dei Diverse marine Moscerini, fragments), Latium, Italy and clams. suggest human shell tools.
Vanguard Cave, Mousterian strata Gibraltar evidence” by limpets,
Ras el-Kelb, Mousterian Lebanon pericoastal numbers of from various
Salzgitter- Freshwater fish Lebenstedt, associated Germany assemblage.
Grotta Breuil, Small numbers of Latium, Italy shells from probably not significant” Gruta da Figueira Marine shells Brava, Portugal Mousterian and other Anatomically Modern Humans (Homo Middle Stone Age Klasies River Mouth, South mammals, and Africa Middle Stone Age Boegoeberg II, South Africa numerous
Abdur, Eritrea Middle Stone Age Early Middle Herolds Bay Cave, South Africa with mussels shellfish, associated
308
Description Locality/site and
Katanda 9 and 16, Thousands of fish Semliki River, with MSA barbed Zaire points in
Die Kelders 1, Sea mammals, South Africa remains deposits; poorly
Hoodjies Punt, Open air MSA site South Africa shellfish, sea
Sea Harvest, South Open air MSA site Africa the use of and fish.
Blombos Cave, MSA shell midden
South Africa densities of limpets, etc.), formal bone
Willandra Lakes, Abundant shellfish Australia from numerous associated with mixed economy. Ksar ‘Akil, Numerous Lebanon shellfish Paleolithic goose(?), and
Mugharet el ‘Aliya, Marine shellfish Morocco undated Upper New Britain, Several early Melanesia middens, fish substantial sea colonization of Riparo Mochi, Early Aurignacian Liguria, Italy almost 5000 shell (MNI ca. ornaments made
Castonet Shelter, Greenland seal France in early Mandu Mandu Low density Rockshelter, crab, and fish Western site ca. 5 km Australia occupation. Leang Burung, Abundant Sulawesi in cave site. Gorham’s Cave, Numerous marine Gibraltar Early Upper sea bird, seal, Kilu Rockshelter, Shell midden with Solomon Islands, fauna; Melanesia required by maritime The Archaeology of Aquatic Adaptations
Locality/site Shuwikhat-1, Catfish and Upper Egypt fishing and Ishango 11 and 14, Abundant fish Semliki River, shellfish, Zaire bone points assemblages lacustrine Site 1017 (Khor Khormusan Musa), Egyptian numerous Nubia mixed Ohalo II, Jordan Thousands of Valley, Israel with house Sea of La Riera, Asturias, Upper Spain shellfish, Ballana (Site Halfan 8859), Egyptian of burned Nubia fish
Altamira Cave, Solutrean use Santander, Spain within a site
Balmori Cave, Upper Asturias, Spain hundreds of limpets. Coberizas Cave, Shellfish Spain bones in Small numbers Cueva Ambrosio,
Almeria, Spain vertebrae shell
of cave ca. Note. M = million years; K = thousand have produced possible evidence for the maritime activities. These lists, too, have compiled such data for years but parent evidence for aquatic resource proliferation of such sites, in fact, I more than 15,000 years old and New is no question that aquatic resources ter these times, and the different help compensate for the fact that the very different times. Even so, early list individually. Instead, I first 310 Table II. Some Early New World
Locality/site Monte Verde, Chile Pericoastal coastal Broken Mammoth, Abundant Alaska fish, economy. Tule Lake, Fish and California resource in
Lewisville, Texas Several Clovis freshwater remains Quebrada Jaguay, Faunal Peru shellfish, Pedra Pintada Freshwater fish, Cave, Brazil several levels. Marmes Use of Rockshelter, along with Washington Healy Lake, Alaska Possible Quebrada Seabird, fish, Tacahuay, Peru Kanaka Rapids site, Isotopic Idaho skeleton component Ring site, Peru Basal levels of dated to
Dalton Complex, Rodgers Shelter, Missouri source.
Daisy Cave, San Abalones,
Miguel Island, shells in
California Early shellfish, with shell 49-PET-408, Human skeletal southeast Alaska marine Island Hidden Falls, southeast Alaska maritime poorly
Cutler Ridge, Shell midden Florida remains, continental
California coast Numerous Early
middens on with
Sabine River site, Submerged Gulf Texas with burned Chuck Lake II, Island shell southeast Alaska remains Note. Kyr = thousand years. The Archaeology of Aquatic Adaptations Table III. Islands Locality Flores Southeast Possible evidence Asia crossing of Sunda to New Guinea and Oldest sites in Australia evidence for voyaging, crossings up
Crete, Greece Homo sapiens poorly calcareous were cemented 51,000 ± Crete sea crossings. Bismarck Shell middens, Archipelago, several sites Melanesia with voyages Sicily, Italy Aurignacian Mediterranean voyage. Ryukyu Islands, Human skeletal Japan Yamashita-cho Okinawa and voyages of ca. Kozushima Island, Upper Paleolithic Japan crossing 50 km obsidian. Melos Island, Travel across ca. Greece obtain Admiralty Islands, Settlement of Melanesia 200 km voyage. Cyprus Occupation of Akrotiri of Cyprus. Channel Islands, Boat and marine California Paleoindian of at least 10
Southeast Alaska Presence on and British lifestyle and Columbia Note. Kyr = thousand years. during various stages of human way. After reviewing such “direct” questions often remain about the nal remains found in such sites. for early aquatic adaptations, sites submerged on continental shelves 312 of pericoastal sites that indicate some resources. Old For the Lower Paleolithic, tence. Preservation problems are taphonomic issues related to the origin evidence for hominid activity are use of aquatic resources by hominids where the remains of a variety of stone tools between about 2.5 and 1.7 Greenwood and Todd, 1970; Harris et 1990; Stewart, 1994). Probably left these lacustrine sites record the minids, as well as the background noise researchers today believe the remains were accumulated primarily via predators or other natural causes. however, that some of the small mammals hunted by hominids. Several early Rift aquatic or amphibious animals, etc. (Leakey, 1971). Because many of settings, however, any clear minid activities is difficult to to be closely associated with hominid tively abundant in both cultural and p. 240) and Stewart (1994) have argued, fish, Clarias sp.) would have been aquatic settings and are unlikely to seems logical, especially for hominids based on opportunistic scavenging and For Homo erectus, a series of ciations of artifacts and aquatic or reported that the bones of catfish and sediments dated between about 1.1 and of crocodiles, aquatic turtles, and the case with much of the Olduvai difficulty in determining whether but she argued that a cultural origin fragmentary condition and close 1981; Roe, 1994; Stewart, 1994). In Bed The Archaeology of Aquatic Adaptations and 0.8 million years ago, Leakey of distinctive pits and furrows of salt by Homo erectus. Along the coastlines of Africa widespread evidence for Lower p. 214; Howe, 1967; Wulsin, 1941). Most however, and contain choppers, hand interglacial beach deposits. Although cupation of coastal plains, the precise pericoastal, inland?) of such What appears to be relatively Homo erectus in Southeast Asia comes stone cave in northern Thailand According to Fagan (1990, p. 120), shells were found piled against the cobble-ringed hearth, and the remains found. Evidence for aquatic resource use of archaic Homo sapiens after about whether this increase represents real better preservation and greater Paleolithic site of Hoxne in England, been found in what have been et al., 1993). Although the dating of tentative, much of the Clactonian interglacial period between about fauna are dominated by large land numerous specimens of freshwater fish and smaller numbers of otter and origin of the aquatic and other faunal has not been firmly established, but that the distributions of all of cuvieri material . . . and most of pattern as the larger bones, also might be food remains Also in England, excavations at the duced fish and freshwater mussel of the site (ca. 425,000 years (Singer 1986, p. 140)) and the cultural origin About 300,000 years ago, archaic along the Mediterranean coast of France 314 and other marine shells were found at tity are poorly documented. Other early comes from several North African Middle Fteah in Libya (Klein and Scott, 1986; Morocco (Howe, 1967), and several sites
Sbihi-Alaoui, 1979; Roche and Texier, southern Europe, Mousterian use of Monte Circeo (Stiner, 1994) and in France (Cleyet-Merle and Madelaine, (Garrod et al., 1928), Gorham’s Cave (Barton et al., 1999) in Gibraltar. At rine shells with flaked edges suggest between about 60,000 and 80,000 years Neandertals and other archaic Homo evidence for the exploitation of fish Madelaine, 1995), and the exceptional beach. Pinniped bones also are rare in scavenging of stranded animals or archaic Homo sapiens occupying the shellfish and other intertidal With the appearance of or AMH), beginning about 125,000 years aquatic resources increases if the Aterian sites of northwest or nearly modern Homo sapiens sapiens evidence for such associations may come Abdur in Eritrea along the Red Sea Stone Age (MSA) stone tools were found other aquatic fauna in strata dated to Walter et al., 2000). With the clear whether the stone artifacts were the faunal remains represent the food documented associations and evidence in South Africa dating between about Klasies River Mouth caves (Deacon and Wymer, 1982), Die Kelders cave (Marean 1974), the Sea Harvest and Hoodjies Herolds Bay Cave (Brink and Deacon, et al., 1999b), and Blombos Cave the earliest evidence for relatively found, including the relatively and flightless seabirds (i.e., coastal MSA localities (Klein and The Archaeology of Aquatic Adaptations where a significant number of large midden strata associated with bone and Sealy, 1997). Initially estimated to be old, sediments capping the MSA levels via thermoluminescence (TL) to 1999). The dearth of fish in most Klein and Cruz-Uribe (2000) to suggest intellectual or technological is possible, however, that the higher discouraged such activities, just as along the California coast during the Rick et al., in press). Along with the latter idea comes from the carefully the remains of numerous large Semliki River in Zaire (Brooks et al., to about 80,000 years ago, the Katanda complex composite fishing technologies significant expansion of aquatic Similar barbed bone points also fish bones at the Late Stone Age site in Zaire, in strata dated to about (Yellen, 1998). Fish bone is relatively African sites, including the White 1994; Yellen, 1998) where the lower years ago, and a series of Nile River RYBP. In coastal areas, little is known period because sea levels were deeply African coastlines were far removed shore (see van Andel, 1989). This same interval in southwest lowering sea levels and extensive southern and southwest Europe have and consumption. Shellfish densities of the Pleistocene (e.g., Straus, 1990; clear if this represents an growth, increased sedentism, changes in a combination of such processes (see Straus and Clark, 1983). Numerous in Europe and southwest Asia also have from marine shells or artistic Madelaine, 1995; Clottes and Courtin, numbers of marine shell ornaments, in Mediterranean coastlines more than 100 316 traveled to the coast seasonally or coasts. In southern Asia, there is only from this time period. In Indonesia, a Burung attests to the systematic (Glover, 1981). At Longrien, a long limited evidence for aquatic resource layer dated to about 30,000 RYBP. can be little doubt that maritime or since at least 50,000 years ago. The peopling of Australia and New from Southeast Asia to Sahul would have even during periods of much lower sea evidence for the use of freshwater fish now appears to have been settled by (Roberts et al., 1990) and 60,000 years freshwater shell middens from the been radiocarbon dated between 38,000 recently argued that some of these as 60,000 years ago. Although evidence Pleistocene Australia is lacking, limited amounts of marine shell from RYBP (e.g., Bowdler, 1990; Morse, 1988; Mandu Creek rockshelter, located only the Last Glacial, a low-density midden crab, fish, and terrestrial fauna be interpreted as evidence for limited level and shoreline reconstructions and density of marine resources and the The apparent abandonment of most of the and the fact that they were reoccupied levels, can be interpreted as evidence strongly influenced local settlement shell middens located on the Melanesian and the Solomons—islands that reach—have been dated between about 1989a,b; Wickler and Spriggs, 1988). occupations is attested to not just by but also by the abundance of marine deposits. The presence of such sites in and New Guinea, is due to the steep rapidly into deep water and changes in on the local shorelines and the coastal The Archaeology of Aquatic Adaptations New In the New World, most early comes from the Pacific Coast, where the lateral displacement of postglacial 1998). The earliest sites currently troversial pericoastal site of Monte reportedly contains evidence for (Dillehay, 1997). At the coastal site dates between about 11,600 and 10,900 whale remains were all found associated land mammals (Nu˜ ez et al., 1994). At n coast, Llagostera (1979) also economy including the use of a variety and 9400 RYBP. Along the south coast of Peru, component from Quebrada Jaguay, where have been found in strata dated between faunal remains at Quebrada Jaguay animals, but the presence of obsidian site may be just one aspect of a (see Richardson, 1998). Also located on as old (10,800–10,500 RYBP), is from the earliest occupation are pelican) and fish (anchoveta, anchovy) remains (Keefer et al., 1998). Of the basal stratum at Quebrada Tacahuay, A third site on the south coast, the have been occupied as early as 10,600 north coast of Peru, Richardson (1998) of the Amotape complex, where unifacial the remains of mangrove shellfish 10,000, 9200, and 9000 RYBP (see also shell middens of the Las Vegas complex RYBP (Richardson, 1998; Stothert, The meticulous work of Roosevelt at Pedra Pintada cave in Brazil dated ter fish were an important component relatively eclectic and focused on Along the Pacific Coast of North maritime sites currently come primarily the California coast, Daisy Cave stone artifacts and a low-density shell 318 and other shellfish remains dated to That humans were on California’s has long been suggested by Orr’s 14 C woman) skeleton to ca. 10,000 RYBP suggests that Arlington Woman actually (Johnson et al., 2000), but a precise Channel Islands have been separated Pleistocene, these two sites boats during the terminal Pleistocene capabilities. Along the California dated between about 9,700 and 8,000 Moss, 1996; Jones, 1991). One of the where stratified shell midden deposits contain abundant shellfish and fish bird remains, numerous bone fishing made from sea grass (Connolly et al., Along the coastlines of northern are only two shell middens reliably Rockshelter on the northern California gon coast (Erlandson, 1994; Erlandson and Erlandson, 1995). The dearth of Pacific Coast now appears to be subsidence earthquakes along the monly associated with tsunamis and Minor and Grant, 1996). In British early coastal sites dated between about mented (Carlson, 1998; Erlandson and Moss, 1998), including portions of a PET-408 (On-Your-Knees Cave) on Prince 9,200 RYBP (Dixon, 1999, p. 118). The consistent with a diet comprised almost manufactured from a land mammal rib has been dated to about 10,300 RYBP site may have been occupied even to the estimated age of a basalt flake deposit located on the continental Columbia (Fedje and Christensen, 1999), regarded as very preliminary. Adjacent to the generally broader Gulf of Mexico and Atlantic coasts, common. On the Louisiana coast, where been prograding for millennia, Gagliano The Archaeology of Aquatic Adaptations with an 11,000-year-old archaeological near the intersection of a creek and the Louisiana and Texas border, Dunbar shell midden dated to about 8,500 RYBP fish bone. Along the Atlantic Coast of North about 8,000 years are extremely rare. stance, the Clovis-age shoreline is 150 km offshore (see Dunbar et al., line changes have been dramatic in most than about 5,000 years old are highly middens have been found. One exception located adjacent to a narrow stretch of shoreline changes associated with This important site, dated to as much reportedly has produced the remains of and shellfish (Dunbar, 1997). Although interior Paleoindian ized big-game hunters, there is of early sites. These include the where two well-stratified terminal one dating between about 11,800 and 10,300 and 9,600 RYBP (Yesner, 1996). early components. Identifiable (>60%) by aquatic birds (swan, and small land mammals (wapiti, bison, is dominated by the remains of large mammals, 10% waterfowl, and smaller remains. Another Paleoindian component is the Lewisville Clovis site located where archaeological deposits array of plant and animal remains. Of remains of freshwater mussel and snail ered were the remains of box turtle, egg shells, raccoon, snake, etc. (Story cated along the Brazos River, turtles and a few fish remains. A about 10,000 and 9,500 RYBP, also ing the remains of many freshwater along with a double human burial marine shells Oliva sayana and Neritina Another early North American site is a 320 on the shore of Tule Lake in northern a hearth dated to 11,450 ± 340 terfowl, and mammal bones. At Marmes mussels and salmon bones are reported and 11,000 RYBP. Shellfish Globally, the growing number of shellfish, fish, sea birds, sea aquatic resources were used relatively H. erectus, and H. sapiens. Due to a omy, recovery, and interpretation of it is difficult to evaluate how economies. Moreover, lists like those that must be addressed before we can resources was both early and animals and other noncultural processes aquatic animal remains in early sites press). Although recent taphonomic and predators transport bones into possibility that animals and not humans bones, or sea mammal remains into early After visiting several did not initially question whether the in the site deposits (other than a have been deposited by anything other analysis of faunal remains from the considered a host of possible sources tural mechanisms for the accumulation shown, however, that a wide range of coyotes, badgers, cats, and a variety vertebrates (seals, fish, birds, etc.) landforms (e.g., Erlandson and Moss, in 1999b). Caves and rockshelters, in of mammals and birds that hunt or carcasses or skeletal remains at site nal remains left by hominids. the nature of terrestrial and aquatic sites to determine whether the activity contributed significantly to the Unfortunately, such careful difficult or impossible to evaluate The Archaeology of Aquatic Adaptations the sites listed in Tables I and II. remains in early sites, the evidence localities may need to be reassessed. shells, for instance, but my in the cave deposits. Gorham’s Cave of birds, including seagulls and others (Erlandson and Moss, in press). Without to cultural activities, we cannot be Neandertal and Upper Paleolithic cave present, similar questions can be sites listed above, as well as the fish, and shellfish remains were Howe, 1967). In the New World, similar Pleistocene or Early Holocene “shell Channel Islands (e.g., Erlandson, 1994, Erlandson and Moss, in press). For other sites, the Middle Stone nent among them (but see Klein et al., with aquatic resource use seems much Devil’s Tower, for instance, Garrod a “large heap” of shells associated (1994, pp. 181–184) found that a burned, suggesting that they too were Old World examples include many of the Lakes in Australia and the Pleistocene Wickler and Spriggs, 1988). In the New about the predominantly cultural origin open air middens along the Pacific Mammoth, and Lewisville also seem The Distribution Even allowing for such remains—often even more serious for early sites—a significant number of systematic early aquatic resource use spatial and temporal distribution of amples, is of special interest. Yesner that such sites are exceptional and are ally high marine productivity. Thus rare examples of relatively intensive otherwise dominated by terrestrial 322 This association holds for some the evidence for early marine resource in Italy, Lebanon, Libya, and Algeria 1967; Stiner, 1994), where marine standards. My own comparison of the different conclusion. While a number of upwelling (Peru, California, Gibraltar, distribution of coastal sites to an atlas of the world’s oceans intensive marine upwelling, exceptional (zooplankton) productivity, sea or vulcanism, marine habitat, or found in areas of coral reefs, (by 8,000–10,000 years ago). They are boreal forests, savanna, chaparral, and contemporary interior sites contain game. I found only one trait that seems bathymetry. From California to Florida all the early sites are located along topography drops off rapidly. The shallow continental shelves generally for marine resource use, regardless of is due to the simple fact, clearly Parkington, 1981; Shackleton et al., modern coastlines were far removed from 250,000 years and more. Studies of shown that the skeletal remains of residential sites more than about 10 km 1982), except for those that have shorelines are steep, however, sites be found within the foraging radius of sites located along shallow continental had access to marine resources for the and shorelines approached the modern This general bathymetric which steep shorelines are associated resource use, while shallow shelves stronger predictor of the location of other aquatic or terrestrial traits I explain some puzzling anomalies. It are much more common along the The Archaeology of Aquatic Adaptations shallow Atlantic Coast of the United coast, all of which is characterized by the earliest coastal sites are bathymetry (Richardson, 1998). Finally, Florida coast, where beaches were as ago, the modern shoreline has produced than about 5,000 years old, except for where the Cutler Ridge site contains foraging dated to about 9,600 RYBP. The correlation between steep sites also seems to contradict two adaptations, (1) that steep bathymetry, tidal and nearshore habitats, reduces and renders them relatively adaptations only developed in the development of relatively broad, analysis of the distribution of early habitats are more productive than itime adaptations were more widespread archaeological record for the antiquity biased in most parts of the world. The Further support for this early development of seafaring in for Pleistocene maritime voyaging in date to the Holocene. For decades, the have made substantial migrations by marginalized maritime adaptations in relatively unsophisticated must have crossed rivers and other and through Eurasia. Prior to 1980, were a very recent addition to human 1976; Johnstone, 1980, p. xv). Due to earliest use of boats—as opposed to to cross small water barriers while depending primarily on indirect (Table III). Except for the and other peoples in the last 5,000 of not-too-distant islands that have in recent geological times, criteria 324 Archaeologists have long argued that Homo erectus was capable of making of the Straits of Gibraltar from and Harris, 1998; Rolland, 1998). As isolation of Neandertals in western it seems increasingly unlikely that routinely cross the potentially Mediterranean, there is limited but exploration by Neandertals (Cherry, may indicate that Homo erectus reached 700,000–800,000 years ago (Morwood et Bednarik (1998) and Bednarik et al. seafaring and maritime adaptations date however, there is little evidence for by Homo erectus or archaic Homo more likely to have been relatively Evidence for Pleistocene more compelling and more widespread, a number of unequivocal and substantial in press; Irwin, 1992). Evidence for aging comes primarily from eastern ages in excess of 20–200 km have now 50,000 and 15,000 years ago. The proof the Pleistocene requires a fundamental maritime voyaging was once thought to the 1970s, terminal Pleistocene of obsidian from the Mediterranean mainland Greece dated to as early as of seafaring was extended with the by 20,000 years ago (Lampert, 1971), a ago (Bowler et al., 1970), 40,000 years as 50,000–60,000 years ago (Roberts of the route chosen, colonization of separate sea crossings, including 1992). As a result, the colonization of liest evidence for planned maritime of the earliest evidence for world (Davidson and Noble, 1992). For a time, two puzzling facts colonization of Australia may have been times Australian Aborigines reportedly making substantial sea crossings about their ability to travel through The Archaeology of Aquatic Adaptations the rest of the world, Australia also direct evidence for maritime majority of such sites were less than With the discovery in the late Bismarck Archipelago and the Solomon et al., 1989a,b; Wickler and Spriggs, maritime voyaging in the peopling of of these islands, now dated to at least p. 185), added several significant reach Australia and New Guinea. More tively impoverished terrestrial flora the marine shellfish, fish, and other Melanesian evidence also suggests that significantly between about 35,000 and ment of New Guinea, New Britain, and 100 km, colonization of Buka in the quired a minimum sea voyage of 140 km By 15,000 years ago, moreover, the Admiralty group, which required an 90 km of which would have been Further evidence for Pleistocene Japan itself was connected to the Asian level, so its settlement did not gued, however, that new blade and 30,000 years ago when Japan was maritime contacts. This idea may be beneath a charcoal-rich stratum in about 32,100 RYBP (Matsu’ura, 1996, about 15,000 and 26,000 RYBP also have caves on Okinawa and the smaller which stretches southward from Japan Miyako Island, human remains found ated with charcoal dated to about the bathymetry of the Ryukyu Islands, quired to reach Okinawa from Japan, Reaching Miyako Island, from either longer voyages of up to 150 km. In that by about 21,000 RYBP, maritime tain obsidian from Kozushima Island Similar to Australia, despite oldest shell middens in Japan date to p. 224; see also Aikens and Higuchi, coastal sites have been submerged by 326 The evidence for Pleistocene it places competent mariners in the Pacific at a date early enough to have Americas (Engelbrecht and Seyfert, the Kurile Islands stretch to the Peninsula and the southern shores of Pleistocene seafaring capabilities of tocene seafarers in Japan, and the appear to have had the capabilities to Whether they made such a journey is resolve the issue—like so many adaptations—lies largely unstudied the North Pacific. Other Evidence for Two other sources of data need to uation of the antiquity of aquatic merged or “drowned” terrestrial limited evidence for marine or ination of either topic is beyond the to consider such evidence at all. Submerged Scholars have long known that tinental shelves around the world (see may fundamentally bias our tions (e.g., Emery and Edwards, 1966; Richardson, 1981; Shepard, 1964). What raises fundamental problems. We must be after all, and simply assuming that lines around the world and that coastal the human story leaves many scholars other hand, assuming that the clear evidence to the contrary is examine submerged coastal landscapes shell middens or other evidence for Unfortunately, this is not as limited so far. During marine sites would have been accompanied by destroying countless coastal sites The Archaeology of Aquatic Adaptations have redeposited most older sites into on wave-cut platforms such as the Lower World marine terraces. Along the the world, intact submerged sites would where local landforms are protected by are cemented or sealed under subsidence of sites into the intertidal settings, where wave energy and potential for the preservation of 1983, 1998). Even in these settings, has been accomplished on submerged Masters and Flemming, 1983). Due to moreover, the work that has been done comparatively close to shore and in have risen up to 150 m in the past prevented effective undersea 10,000–12,000 years old. Nonetheless, impressive numbers and the number of sites is rapidly In a recent summary, Flemming (1998, p. human occupation sites (SHOS) have been world, about 100 of which are older hand axes found in sediments underlying off the South African coast and creek bank submerged 18 m below sea of France (Flemming, 1998, pp. dating to the middle and early Holocene age also have been recovered from the et al., 1992; Fedje and Christenson, Stright, 1990). In a number of evidence for aquatic adaptations have feature remarkable preservation, as burials, canoes, canoe paddles, Holocene sites such as Tybrind Vig in Yam off the coast of Israel (Raban, One of the earliest submerged is the Upper Paleolithic Cosquer Cave coast of France (Clottes et al., 1992; partly submerged limestone cavern, the level. A narrow and gradually rising opening into a large cavern, only parts engraved or painted motifs have been cavern, and radiocarbon dates indicate 328 two periods about 27,000 and 18,500 Paleolithic artistic representations of Cosquer they make up about 12% of the seals, the great auk (Pinguinus the cave appears to have been located height of the Last Glacial, these to the artists. As Clottes and Courtin Paleolithic skeletons excavated in the 150 km to the east were associated with testifying to the symbolic importance of the area. Clearly, submerged terrestrial right conditions. The questions that sites represent the proverbial tip of earlier coastal sites lie offshore in found and sampled to help unravel some role of the sea in human history.
Like Cosquer Cave and the sites around the world that are located that lack dense accumulations of to linkages to aquatic habitats. These Table I, which at various times in located some distance from the coast. paleogeographic reconstructions, these densities of marine food remains, the objects (shell beads, baler shells, site abandonment. For a number of other further from aquatic habitats, such small amounts of aquatic food remains resources were relatively unimportant, residential periods on the coast, of closer to the coast, or some arguments has been made for interior marine shell ornaments are found far some economic or ritualized use of argued depending on the theoretical In this regard, I find the eral pericoastal sites (Mandu Mandu, old contain small numbers of shells, The Archaeology of Aquatic Adaptations between interior and coastal groups know for certain whether these shells Australians occasionally visited the ited the interior, or that they groups residing separately in coastal Australia, which we now know was have so few early coastal sites? Did don its coastlines in favor of more for tens of thousands of years? If so, systematic use of shellfish and fish years ago? Why do we see further Melanesia by 35,000–40,000 years ago? pericoastal sites in Australia are the by coastally adapted Pleistocene or destroyed by rising postglacial
To some, none of the individual a compelling argument for the early the theoretical and methodological knowledge about world sea levels and of lines of archaeological evidence, I the veracity of the current consensus that there is only limited evidence for to the end of the Pleistocene. This is appears to have been colonized near the were much lower than they are today. that this pattern accurately reflects To understand the history of aquatic region, we must first determine if the result from actual changes in human or taphonomic forces, or the recovery themselves. Unfortunately, except for recently (i.e., Polynesia), such rarely been done in a manner that A New It is possible, however, to theory about aquatic adaptations and ical data from the New World. If 330 smaller and less productive than reflects demographic pressure, the antiquity of coastal adaptations in case. Traditional theory, including ranking models, suggests that until mographic pressure to reduce the aquatic resources would not be pling of two vast continents with scapes, especially by small grated through an interior route into evidence for marine resource use or aquatic resources are not necessarily evidence for their use, at least in use compare favorably to those To adequately evaluate the first settled the Americas and whether of the full range of early adaptations, cannot answer either question with any analysis of the Peruvian archaeological inferior in the arid western slopes of the Peruvian evidence, he believed sus coastal occupation equal to more region, well over 12,000 years. This the earliest occupants of the region their population densities had trial environment. Thus the Andean on earth—was characterized as an In retrospect, we now know Erlandson, 1988; Perlman, 1980; Quilter he assumed the Andean archaeological disproved claim that tectonic uplift of level rise since the last glacial. clock began with initial human years ago, based on claims by MacNeish (made from the same rock as the cave (Flea) Cave. Even today, with the accept the dubious evidence for 1999, pp. 100–101). Today, the earliest widely logical site is Monte Verde (Dillehay, RYBP. Monte Verde is located in a While there is evidence that big game idence for a relatively eclectic The Archaeology of Aquatic Adaptations played a significant role. The cupants had links to the coast. Recent occupation of the Andean coast to et al., 1998; Richardson, 1998; brada Jaguay, shows interior links the opposite end of a seasonal we accept that Monte Verde is one of likely) and assume that its occupants resources themselves (which seems less earliest coastal versus interior This might be enough time for maritime focus, but it seems highly 11,000 years ago was due to demographic In North America, the situation for early coastal settlement and use of presence of people on California’s (Erlandson et al., 1996; Johnson et difficult to account for as a response tive and diverse terrestrial resources of the Channel Islands, moreover, seem small fish, plant foods, and dence that they were big-game hunters or massive elephant seals. Rather, they relied on a variety of resources, and a variety of small fish. The very this early date suggests that shellfish, ranked, highly regarded, and highly of North America, the evidence for as widespread, but in these areas the artifacts on the continental shelves of the coastal archaeological record. To me, the available data suggest were an integral part of many early New cance has been underemphasized in archaeological sites does not deterioration, population pressure, am not suggesting—as many have done restrial resources—that aquatic arguing that the use of aquatic stress and economic intensification. and the Old World, the factors that will be used, when, and by whom are this complexity, the diversity of 332 by our ancestors, and the flexibility global or universal statements about justice to the diversity and complexity record. SUMMARY AND I began this paper by suggesting nature of aquatic adaptations have lacustrine societies, relegating them aquatic resources are marginal and that recently renders their study issues addressed by archaeologists: the appearance of anatomically modern development of agriculture, the rise of a variety of taphonomic processes, lems, and data gaps raise serious systematic aquatic adaptations Specifically, I suggested that 1. postglacial sea level rise has about 10,000 years along which tions would logically be 2. differential preservation, selectively underemphasized archaeological sites around 3. traditional models of role of hunting in general, in many ancient societies; 4. normative cultural ecological man societies as aggregations diverse people (men, women, etc.) who often were engaged 5. prior to the development of minids relied to a significant increased the relative required no specialized 6. our hominid ancestors have usual environmental eclectic omnivores, an dinary success in colonizing on earth. The Archaeology of Aquatic Adaptations With these issues in mind, I use in archaeological sites around the earlier than about 15,000 years ago and old. I concluded that a variety of from determining when aquatic and how important they were in the of the earliest archaeological erectus in Africa contain the remains fish, crocodiles, molluscs, and with late Homo erectus or early archaic distribution of fish and other aquatic closely with artifacts and other faunal remains (aquatic and terrestrial) and to prove. Less equivocal evidence for archaic Homo sapiens also is found at that these hominids occupied coastal doubt that aquatic resources were used the intensity of such use remains Paleolithic subsistence remains largely Hominids clearly crossed aquatic through much of Eurasia, indicating necessarily the physical or the appearance of anatomically modern sources may have been limited largely of fish, birds, mammals, and other cialized technologies. Only with the do we find the evidence for a more of marine or aquatic resources. Not incides with the first evidence for transitional technologies, including stone blade and geometric or earliest widespread evidence for the relatively sophisticated boats. In the technologies at Klasies River Mouth localities in South Africa, for riety of shellfish, marine mammals, 2000), although some or all of the rather than hunted (Binford, 1984). the earliest evidence for complex harpoons from Katanda associated with 1995). From Blombos Cave in South marine fishing, probably dating to 334 1997). And from the Boegeberg 2 shell relatively intensive MSA use of Dated to about 90,000 years ago sapiens sapiens found outside of coastal Israel—suggesting that early of Africa by this time. Although have moved into most of Europe for evidence suggests that they spread into From there, within just a few voyages through island Southeast Asia By about 30,000–35,000 years ago, western Melanesia and the Ryukyu Although the current state of our earliest subsistence strategies that of marine or other aquatic resources anatomically modern humans. When such the exploitation of a range of stable resource base would have significantly to the development of to the reproductive success of Homo matic demographic and geographic regard, it is worth noting that current of anatomically modern humans allow multiregional models for this in press). Wherever anatomically modern carried with them a penchant for art complex problem-solving and 1992; Klein, 1998; Mellars, 1998). ing played a more significant role expansion, the geographic spread, and The available archaeological Hell and Garden of Eden models, with Aristotle’s “golden mean”—somewhere gorical statements to the contrary, we were first widely used by our hominid man evolution. However, it makes no ignored aquatic resources for more than was a significant hominid pursuit, in found in shallow water or on the shore ably abundant and available without has been some intensification of it also seems likely that our ancestors situationally whenever and wherever it The Archaeology of Aquatic Adaptations resources sometimes compare favorably raises significant questions about the as anthropological indicators of the economies (see Bailey, 1978). If shell economies, is the broad spectrum We cannot afford to ignore the exploitation of many types of marine requires relatively complex hook-and-line) that currently appear to minids other than anatomically modern aquatic habitats are extremely ied terrestrial habitats, and that ronments that they defy broad 1980). Given the nearly endless bility of aquatic versus terrestrial the antiquity and intensity of aquatic and time. Today, rather than searching aquatic settings, we should be working that currently inhibit our diversity of aquatic adaptations. More of dichotomized debates about whether of Eden or Gates of Hell. Once we through space and time, as well as the vironments that result from juxtaposing terrestrial habitats, we can focus on environments that took place as our subsistence strategies on both land and innumerable adaptive responses possible cal, demographic, and sociopolitical universal laws of aquatic adaptations As we move into the twenty-first models and polarized arguments that about the evolution of aquatic has suggested, “it is time to put to fish, sea mammal, or coastal forager. made great strides in understanding the adaptations in human history. As our numerous issues yet to be resolved and be studied. To truly understand the history, however, a number of issues Perhaps the most pressing are and the search for ancient sites sea. It is time to extend the search 336 of shorelines around the world and into dating to the crucial period between new technologies, offshore along steeply dipping shorelines of the Cave and submerged caves off Gibraltar been identified in limestone bedrock to preserve evidence for early coastal sea levels and coastal erosion excavation work might also be shorelines of the Japanese archipelago, antecedents of the Jomon peoples may where such sites are located and when resources played a significant role in such adaptations were. Critical to evaluating the idea moved rapidly out of Africa along the and beyond is a search for early shell eas associated with Last Interglacial and the islands of Southeast Asia. coastal sites elsewhere in the world, are focused along coastal stretches where lateral shoreline movements minimized. Also needed are renewed the remains of aquatic resources, with techniques, including fine screen of the origin of aquatic and other actualistic studies to help distinguish versus natural origin, work that will been done for terrestrial fauna in Ultimately, we need more data regions to better document the and time. In the twenty-first century, adaptations should focus on documenting resources played in human history as world, from sea to shining sea.
This paper is dedicated to the recognized the limits of what we could of aquatic adaptations. I would also The Archaeology of Aquatic Adaptations Alan Osborn, Geoff Bailey, and David views stimulated tremendous progress in tations. For freely sharing ideas and I thank Virginia Butler, James Dunbar, Glassow, Nina Jablonski, Antoinette Greg Nelson, John Parkington, Geoffrey Richardson, Torben Rick, Anna Vellanoweth, Larry Wilcoxon, John Stephen Nash, and an anonymous reviewer helped me revise an earlier draft. I am Price for inviting me to write this sistance, and for their patience. Most Madonna Moss—my wife, colleague, and my coastal journeys (physical and comments on an earlier version of this the opinions expressed in this paper.
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8-3 Thinking outside the box: a new perspective on diet breadth and sexual division of labor in the Prearchaic Great Basin Robert G. Elston and David W. Zeanah Abstract The archaeological record of the technology and mobility of Prearchaic foragers, suggesting a hunting-oriented imply a generalized, broad-spectrum work for understanding this discrepancy from the ethnographic present. ancient diets without an understanding reviews recent studies using behavioral behavior in a PHT resource landscape. explanation for the diversity of strategies for men and women, rather that Prearchaic hunter-gatherers the mobility and technological pro les Keywords Great Basin; Pleistocene/Holocene Introduction The North American Great Basin (Fig. 1) documenting the ecological they lived. Informed by ethnographic has demonstrated the existence of much of the Holocene (Jennings 1957, |
As rivers flowed off the highlands they formed deltas characterized by mangrove swamps with several species of trees including Nypa fruticans palms and the sugar palm Arenga, with flowers that produce a sugary juice, and shrubs of Passiflora with edible passion fruit, and edible climbing ferns, Stenochlaena palustris. The trunks of the Nypa are submerged so the fruit, which is edible when it is immature, would have been accessible to terrestrial hominids. The delta facies contain shells of edible mollusks including oyster shells but at low density and scattered distribution. Crocodile and turtle fossils have been reported from the same sediments at other places in the Perning district. To date only one fish fossil has been identified. The photos of bantengs and nypa palm were taken from a web site of photographs of Ujung Kulon National Park but unfortunately the web site is no long available. 
forest. The present climate in the vicinity of the cave is tropical super wet with mean annual rainfall greater than 3000 mm and rare or very short seasonal dry periods. The Pleistocene climate may have been more seasonal but changes in rainfall and temperature regimes at this site, through time, have not been resolved. At the time of Pleistocene occupation, the cave may have been about 30 km from the sea. Floral and faunal elements recovered from the late Pleistocene levels suggest that “humans were foraging in a mosaic of closed forest, scrub, grassland, swamp, and freshwater lakes and rivers (Crangrook 2000:83, cited in Barton 2005:57).” Barton, along with others interested in drawing as much information as possible from the archaeological record, has been involved in the development of a new line of evidence from archaeological assemblages, the identification of starch grains through the comparison of grains found in sediments and on tools with grains from modern starches.
Unlike the !Kung, the Hadza focus exclusively on large game animals. Over 256 days of observation in the years from 1985 through 1989, the Hadza killed or scavenged 72 large animals for an average of one animal every 3.6 days. But considering there were 6 hunters in camp during the wet season and 10 in the dry season, individual hunter success rates were not high. The average of 1 large animal every 4 days seems to guarantee that Hadza individuals eat meat every day but averages obscure the high variance in acquisition. The researchers calculate the chance of failure for any one hunter on any given day as 97%. They compare this to a success rate for the Dobe !Kung of one animal every four man-days and for the Ache, 2-3 successful days out of every 4. In the case of both the !Kung and Ache the success rate reported here includes all the small game added to the diet. Each Hadza hunter maximizes his mean rate of return by focusing on large game but at the cost of high variance, the probability of failure on any day. Remember from earlier discussions that one of the problems with the simple OFT models is they ignore variance in acquisition. The only measure in the OFT models is the rate of return, in calories per hour, for that resource after the time cost of handling and processing is debited. Remember also that the high variance in individual hunter acquisition doesn’t necessarily translate into high variance in individual consumption because a large animal taken by any hunter is shared over all the individuals in the camp as well as others who come from longer distances when the “bush radio” indicates there is meat in the camp. The chance of consuming meat, in a camp with 6 hunters, is about once a week. For a large animal, such as a zebra, the meat might last 3 days increasing the probability of eating meat. In the picture below a Hadza hunter carries a portion of meat that was butchered at the kill or scavenging site. 
I have assigned, O’Connell, Hawkes and Blurton Jones (1988) for an argument using the ethnographic example of Hadza scavenging to assess the probability that ancestral Homo might have regularly eaten meat even though the weapons they had were much less effective than the weapons of the modern Hazda. In the final exam you should be able to integrate the ideas and information from O’Connell et al. (1999), Lecture 5, with the information in this lecture’s reading assignment. In the 1988 paper the researchers report a subset of the observations I have summarized above. In the period from 1985 through 1986 the Hadza took 54 medium/large animals of which 14% by carcass weight were acquired by scavenging. Table 2 shows the high variability of success in both hunting and scavenging. The Late dry season, Sept.- Oct. 1985 was the most successful of the four seasons tallied. During that period Hadza hunters brought 28 carcases into camp compared to 13 in the late dry, Aug.-Oct. 1986. Pay attention to “day-to-day availability” reported in column one, mid-page 359. Note the variation from one season to the next. Average income from scavenging was very high in in the early dry season 1986, 245 g per camp resident per day, but all the gain came from one animal, consumed in 3-4 days, leaving consumers many days without meat in spite of high average income from scavenging. 
As reviewed in O’Connell et al. (1999) berries and tubers provide the dependable daily nutritional requirements of the Hadza. Hadza children are effective foragers on berries, baobab fruit and some shallow rooted tubers but they do not have the strength to gather the deeply rooted tubers that provide the highest returns for adult women. This photo of a senior Hadza women illustrates the strength needed to gain access to these resources. Hadza women, with unweaned babies and children over the age of 6, leave camp in early morning to gather. They are usually accompanied by armed teenage boys and perhaps an adult male as protection against pastoralists who range through the same area. Men and boys who accompany women on these trips sometimes take honey from wild bees. Honey is a valued resource with high caloric return. Hadza men can achieve a mean daily return rate of .78 kg per man-day by spending an average of 41 mins a day in honey collecting. 
Both men and women forage in the shallow water of the reef flat in the 2 to 4 hour low tide periods from March to end of September. Men walk to the reef edge and fish with spears catching a variety of small and large fish. The largest fish they take is giant trevally, pictured on the left, which can grow to 
1.7 m, but they also take a variety of very small fish such as spine foot, sweet lip, and mullet about 38 cm, 45 cm and 78 cm, respectively. The catch for the day is displayed openly as the men leave the reef at the end of the fishing bout. If you can imagine these small fish, pictured in order below, flashing through the coral and the distorted optical perspective from the surface of the water to the shallow depths where fish school and hide, flash and dart you can get a sense of the skill needed to take these fish with wooden spears. In the methods section of the assigned reading, Bliege Bird, Smith and Bird (2001), the authors note that the spearfishing forager ignores other prey types as he travels across the reef. During the low tide men spend 63% of their time spearing and 31% of their time shell fish collecting. This time allocation bears no relationship to the potential returns from the prey, 292 plus or minus 135 kcal/h for spearing and 1492 plus or minus 173 kcal for shell fish collecting.
October the Meriam forage on reef flats at ebb-tide. In October through April they switch to collecting nesting turtles and eggs. Everyone participates in turtle collection during this season as the turtles come onto the sandy beaches to dig nests and lay their eggs. The foragers go out at night or early morning and wait on the beach for turtle to come ashore. When a turtle is spotted, it is flipped over, the flippers tied back and the turtle hauled home by boat. Nesting season collections are undertaken mostly for household provisioning but raw meat from the butchered turtle is always shared out with neighbors. In some cases nesting turtles will be taken for pre-arranged feasts. 
Hunting Returns from turtle collecting and hunting are reported in Table 2. The after-sharing returns during the nesting season are comparable to the returns from reef shellfish collecting. Notice the large variation between return rates from collecting in the nesting season and hunting in the hunting season, 21,875 kcal/h vs. 4922 kcal/hr. The very large difference is caused by the inclusion of search time and energy expended in travel. The energetic costs of travel in the hunt are calculated from the conversion of the cost of fuel for the boats into calories of meat that could have been purchased at the store. When per capita returns are reported after-sharing, open water hunting results in a net loss to the hunters because of the time and fuel costs of the activity. Notice that hunting in the hunting season is more costly than hunting in the nesting season. “In this season, hunting is much easier and hunters take on fewer costs: turtles are found on nearby reefs waiting to crawl onto the beaches to lay eggs at night, the tradewinds have largely ceased, and in between monsoon storms, the water is clear, calm and visibility is excellent, allowing hunters to dog turtles more closely, to lose fewer and to more finely discern size and sex (Bliege Bird et al. 2001 p 14).”
Men often hunt on open water for other resources such as tuna, mackerel and large marine sea mammals. This picture of men displaying their spanish mackerel catch comes from Rebecca Bliege Bird’s home page at Stanford. This looks like an impressive catch but you can see, in Fig. 7 below, how large fish rank in return compared to other resources. The returns graphed in Fig. 7 represent after-sharing returns. The bar represents mean returns with the long line to the right of the bar representing one standard deviation. The longer the line the higher the variance in return. In the case of large pelagic fish, the standard deviation indicates that men may achieve a positive or negative 5500 Net E/hr return per hour. Remember a negative return results from an unsuccessful hunt as the cost of fuel, calculated in purchasing power of store-bought meat, is debited from the return to quantify the costs of search. Only returns from netting sardine, 11,008 kcal/h before sharing, are reported in the assigned reading (p 14) but the figure from Bliege Bird and Bird (1997) illustrates returns from the most important resources. Both men and women use casting nets to intercept schools of sardine along the perimeter of the island and hand-line fishing using hook, line and bait to catch reef carnivores, herbivores and surf fish from the tidal margin. On this chart the returns from sardine netting are lower than those reported above but I assume acquisition costs are not debited from the figure of 11,008 kcal/hr.
rate at which foragers encounter prey in the two patches. Adults are able to move at faster speeds across the sand flats encountering goanna at about .90 items/hour searching whereas children would encounter prey in this same patch at .68 items/hour because of their slower walking speeds. Larger children gain nearly the same caloric gain foraging on the sand plain as in rocky outcrops but smaller children do much better on the rocky outcrops, 448 kcal/hour compared to 385 kcal/hour on the sand plain. The authors suggest children forage in patches that minimize time and effort expended. “Although the gross foraging return rates are the same, foraging in sandhills as compared to rocky outcrops requires 2.3 times greater time investment and 2.2 times more walking as does rocky outcrop foraging for only 1.3 times as many calories per hour (Bird and Bliege Bird 2005 p 142).” The young girls in this picture, copyright Rebecca Bird from her home page at Stanford, are holding a small lizard they have dug from a rocky outcrop burrow.