Evolution of the human diet

October 14, 2006

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Filed under: Lectures, readings, test 2 — cambriaromance @ 7:10 am

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Printable View of:Lectures

 

File: Lecture 7 Coastal Migration and Aquatic Resources

The Evolution of the Human Diet

Lecture 7

Coastal Migration and Aquatic Resources

Anthro 4962/5962 Instructor Helen Alvarez

 

The mosaic habitats shown in this photograph of Rajegwesi in

Meru Betiri National Park on the south coast of Java illustrate the paleoecology hypothesized for H. erectus in Java by O. Frank Huffman and his colleagues. The photo is from

from www.eastjava.com

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 good condition of the skull and the large size of the ancient Mojokerto Delta favor the conclusion that the hominin died in the deltaic environment in which it was deposited. The Mojokerto child therefore provides evidence for a seacoast Homo erectus population in Southeast Asia, and raises interest in the role that maritime adaptation might have played in the dispersal and paleoecology of early hominins (Huffman et al. 2006 449).”

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.

 

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.

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 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.

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.

 

Assigned Reading
Erlandson, Jon M., 2001. The archaeology of aquatic adaptations: Paradigms for a new millennium. J Arch. Research. 9(4): 287- 350. For this lecture read pages 287-321.
Also Recommended

Barton, H., 2005. The case for rainforest foragers: The starch record from Niah Cave, Sarawak. Asian Perspectives 44(1): 56-72. (Asian Perspectives available as electronic edition Marriott Library)

Huffman, O. F., Y. Zaim, J. Kappelman, D. R. Ruez Jr., J. de Vos, Y. Rizal, F. Aziz, C. Hertler. 2006. Relocation of the 1936 Mojokerto skull discovery site near Perning, East Java. J. Hum. Evol. 50: 431-451.

Huffman, O. F., and Yahdi Zaim. 2003. Mojokerto Delta, East Jawa: Paleoenvironment of Homo modjokertensis First Results

Semah, Francois, Semah, Anne-Marie and Simanjuntak, Truman, 2003. More than a million years of human occupation in insular Southeast Asia. In Under the Canopy: The Archaeology of Tropical Rain Forests. pp 161-190. ed. J. Mercader New Brunswick, NJ: Rutgers University Press. (available from Marriott Library, Course Reserves)

 

 

File: Lecture 8 Optimal Foraging Theory

Anthro 4962 The Evolution of the Human Diet

Lecture 8

Optimal Foraging Theory

University of Utah Fall 2005 Helen Alvarez

 

Shores of Walker Lake (The North American Indian; v.15) from Northwestern University Library,  Edward S. Curtis’s ‘The North American Indian’: the Photographic Images, 2001. Curtis described the setting of this photo: Walker lake, one of numerous saline lakes remaining from a great inland sea that once covered western Nevada and northeastern California, is the seat of a major division of the Paviotso. In the western corner of Nevada it is fed by Walker river, the numerous branches of which head on the eastern slope of the Sierra Nevada in California. Although there is no outlet, the water is not too saline for the thriving of trout and suckers, which were taken on bone hooks, with double-pointed spears, and in gill-nets.

 

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.

 

The first efforts toward understanding resource capture were undertaken by a Canadian entomologist, C. S. Hollings who designed a simple experiment modeling predator and prey interactions. The prey were represented by 4 cm sandpaper discs tacked to a 9 sq. ft. table. The predator was his blindfolded research assistant who tapped her fingers over the table until she encountered a disc, which she removed and set aside before returning to the task of “capture”. The trials were iterated at various prey densities and the relationship between rate of capture and disc density was plotted giving the marginal value curve shown on the right. The rate of capture does not increase in a linear fashion with the increase in the density of the prey.

Hollings’s results became the basis for the distinction between search and handling in the subsequent development of OFT. His simple experiments showed that search time was a function of prey density on the landscape while handling time after encounter was a function peculiar to the prey type. These distinctions became important in all subsequent OFT models.

Optimal foraging theory drew attention to the empirical fact that every resource has a cost of acquisition and processing. These costs make comparisons of energy yields of food meaningless unless handling costs are considered in the comparisons. Here I have posted Table 1 from Bright, Ugan and Hunsaker (2002) showing the energy yield of certain resources taken by Australian and Great Basin foragers and the Kcal/hr return for those resources after processing. I have sorted their table from highest net gain to lowest and plotted the yield with and without processing to show you how the rank of resources changes when the handling costs are debited.

Notice on the figure, drawn from the table, that resources with high energy density (red line) are not necessarily the resources that yield the highest returns after processing (blue line). From this set, the highest ranked resources are all animals with tubers ranking just below small game. Compare the yield of rabbits (#3) and pine nuts (#11) on both curves to see how processing costs change the relative rank of resources. If meat is always the highest ranked resource, across many habitats, why should human foragers ever take any other resource? You should be able to answer this questions by the end of Section II.

 

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.

 

Risk, Uncertainty and the Forager’s Energy Balance

 

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).

 

Great Basin Archaeology and Optimal Foraging Models

 

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.

 

Assigned Reading

Sih, Andrew and Milton, Katherine. 1985. Optimal diet theory: Should the !Kung eat mongongos? Amer. Anthropol. 87(2):396-401.

Hawkes, Kristen and O’Connell, James F. 1985 Optimal foraging models and the case of the !Kung. Amer. Anthropol. 87(2):401-405.

Elston, R. G. and Zeanah, D. W. 2002. Thinking outside the box: A new perspective on diet breadth and sexual division of labor in the Prearchaic Great Basin. World Arch. 34(1):103-130.

Also Recommended
Winterhalder, B., Lu, F., and Tucker, B. 1999. Risk-sensitive adaptive tactics: Models and evidence from subsistence studies in biology and anthropology. J. Arch. Res. 7(4): 301-348.

 

 

 

 

 

File: Lecture 9 The Ache: Broadleaf Evergreen Foragers

Anthro 4962 The Evolution of the Human Diet

Lecture 9

The Ache: Broadleaf Evergreen Foragers

University of Utah Fall 2005 Helen Alvarez

 

Ache Man Carrying the Head of a Tapir

photo from anthrophoto.com

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.

Note the distribution of zones colored purple, pink, pale yellow and medium green. All of these zones are characterized by wet conditions during the season of maximum solar insolation.

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.

Summary: Five Years of Research

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.

Hill, Kaplan, Hawkes, and Hurtado (1987) Foraging decisions amount Ache hunter-gatherers: New data and implications for optimal foraging models. Ethology and Sociobiology 8: 1-36.

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.

Assigned Reading
Hawkes, K., Hill, K., and O’Connell, James F. 1982 Why hunters gather: optimal foraging and the Ache of eastern Paraguay. Am. Ethnol. 9(2): 379-397.
Also Recommended
Hill, K., Kaplan, H., Hawkes, K., and Hurtado, A.M. 1987 Foraging decisions among Ache hunter-gatherers: New data and implications for optimal foraging models. Ethology and Sociobiology 8:1-36.

 

 

 

 

 

File: Lecture 10 Semiarid African Strategies

Anthro 4962 The Evolution of the Human Diet

University of Utah Fall 2005 Helen Alvarez

Lecture 10

Semiarid African Strategies

 

!Kung Mother with Her Baby on Her Back Gathers Berries

photo by Marjorie Shostak from

www.anthrophoto.com

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.

Hadza Country

Hadza country is characterized by rocky hills covered with mixed Acacia scrub and grass leading down to narrow, grass covered valleys. Foragers walking through this landscape to hunt or gather have vistas and landmarks to guide them. In this area children are able to forage away from home and find their way safely back to camp. We will come back to the habitat differences when we discuss juvenile provisioning and ask why the !Kung children are not able to gather more of their own resources.

!Kung Bushland

The Kalahari is a vast plateau at 1100 m above sea level covered by grasses and woodland trees growing to about 30 m in height. It is a nearly featureless plain with few landmarks to guide the hunters and gatherers who make their living from its resources. Even though habitat variability seems low there is some topographic relief provided by lower lying, shallow, seasonal river beds, called molapo, and frying pan shaped dry lakes which hold water for a few days after a heavy rain but are otherwise hard and dry most of the time. After the rains, the Kalahari plains are grass covered. The higher land supports low shrubs that give way to taller woodland trees on the sand dunes and finally, on the outermost zone, open scrub plain. Each of these habitats provides a unique suite of resources for the !Kung. The woman introducing this lecture is gathering berries that grow on the scrub plain.

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.

The four resources pictured here are not quite to scale but give you an idea of the foods listed in Table 8. First is Citrullus lanatus, the wild version of our cultivated watermelon. Second in the top row is Acanthosicyos naudiniana, the Kan melon. Surprisingly these melons can be gathered and kept for many months to supply water and some nutrients. Notice the relative size of these two melons in Tanaka p 60. The tubers that are so important when melons are not available are illustrated in the next panel. This tuber is probably Coccinia rehmannii. The final panel shows a Moroccan truffle of the same genus as the Kalahari truffle, Terfezia. North African truffles are now marketed to the global gourmet food trade.

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.”

It is difficult to get a sense of the size of the eland in this photo but if you convert 300 kg to lbs these animals weigh about 660 lbs. Compare that to beef cattle weighing from 600 to 900 lbs. Lee estimates the edible fraction of game taken by the Dobe !Kung to be 50% of the live weight or 150 kg for an eland. In Table 5, reproduced below, he lists per capita consumption of meat at 230 grams. Calculating .230 kg per consumer day a large eland would provide 652 consumer days of meat. Dividing that by a camp size of 30 gives 21 days. However we know that the meat would spoil long before it was consumed at that consumption rate. Even though meat can be dried in the hot sun of the Kalahari it is most often consumed within a few days of capture.

 

In this photo of gemsbok you get a good sense of the red sand of the Kalahari and the formidable task of a hunter taking these impressive animals with only bow and arrow. Adult gemsbok males weigh from 167 to 209 kg with females smaller at 116 to 188 kg. Tanaka (p 77) reports that a gemsbok was brought into camp on Oct. 12 and nearly all consumed by the end of the next day. Again if we use Lee’s figures for edible yield and a mid-range figure of 188 for a male animal we can estimate 94 kg of meat in camp. On that same page, Tanaka reports 16 providers and 14 dependents so 30 people consumed, on average, 1.57 kg of meat each day for the two days the meat lasted. To convert these figures to caloric gain we can use Lee’s figures from Table 5 of 690 calories in 230 grams of meat or 4710 cal each of those days from meat.

The figures in the above panel give you some idea of why meat is so highly regarded. Remember they are not figures that could be used to rank meat in an optimal diet because they include no post encounter processing costs. But we might think of the costs in another way. The days with meat in camp are days that men do not go hunting nor women go out to gather. For those days no one pays the energetic costs of traveling to foraging patches, collecting and carrying in the hot sun. Tanaka reports a weight of 200 kg for the greater kudu, pictured here. In the last estimate of meat shared around the camp I neglected the portion eaten by the hunters in the bush. The amount of meat eaten by camp consumers should be reduced by the weight of internal organs and ribs the hunters sometimes eat before they transport the game (see Tanaka in the 3 pages on hunting). Hawkes et al. (1991) report slightly higher yield of edible fraction from live weight for Ache prey, 69 to 88% edible. An experiment at the University of Nevada Reno reported “dressing yield” of over 60% for grass fed beef. Keep in mind that hunters eating wild game would eat more parts of the animal than reported as yield by contemporary americans.

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.”

At the end of the dry season when resources around the water holes have been exhausted the !Kung women have to walk as much as 10 to 15 miles to the nut groves carrying babies as well as the nuts they gather. On these journeys the women have to make a second energetic choice, do they carry toddlers or suffer the time and energy constraint of going at the slow pace dictated by the capabilities of the youngsters. Older children are left in the camp with other caretakers but nursing babies are never left at home. In groups subsisting entirely on wild foods, interbirth intervals (IBIs) are nearly 4 years and babies are at least 3 years old before weaning. Even though !Kung babies are smaller than bottle fed American babies these women are carrying more than 30 lbs of baby and nuts.

Child care constraints always have to be factored into foraging decisions made by !Kung women. How far to travel, what to pursue, how much to carry, how long to gather are all decisions a women faces in the tradeoff between when to have another baby and how to feed the ones she already has. In the photo on the right a women is digging tubers. Blurton Jones and Sibly considered these factors in the backload model to predict the birth spacing that would optimize a !Kung women’s reproductive success. Considering the rigors of walking long distances in the hot sun and the number of nuts needed to feed a family, they predicted interbirth intervals of approximately 4 years, a prediction that matched the actual IBIs fairly closely.

 

Proteins and Calories from Vegetable and Animal Resources

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.

Hadza Hunter-Gatherers of Lake Eyasi Basin

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.

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.

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.

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).

Energy Conservation?

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.

Assigned Reading

Tanaka, J. 1980. Chapter 2 Subsistence Ecology In The San Hunter-Gathers of the Kalahari: A Study in Ecological Anthropology. pp 53-91. Tokyo: University of Tokyo Press. Go to Marriott Library Course Reserves to download the electronic file.

Tanaka, J. 1980. Hunting 3 pages from Chapter 1 subsection hunting. In Subsistence Ecology In The San Hunter-Gathers of the Kalahari: A Study in Ecological Anthropology. pp 30-35. Tokyo: University of Tokyo Press. Available from the Readings section webCt.

O’Connell, J. F., Hawkes, K. and Blurton Jones N. G. 1988. Hadza scavenging: implications for Plio/Pleistocene hominid subsistence. Current Anthropology 29(2): 356-363.

O’Connell, J. F., Hawkes, K. and Blurton Jones N. G., 1999. Grandmothering and the evolution of Homo erectus. J. Hum. Evol. 36:461-485. Posted under readings Lecture 5 Review p 465 from the Grandmother Hypothesis to p 468 Applying the argument to Homo erectus and p 470 beginning with Climate change and “children’s” resources to p 475 An evolutionary scenario grounded in the Plio-Pleistocene.

 

 

File: Lecture 11 Meriam Aquatic Foragers

Anthro 4962 The Evolution of the Human Diet

Lecture 11

Meriam Aquatic Foragers

University of Utah Fall 2005 Helen Alvarez

 

Mer Island

Home of the Meriam

photo © Great Barrier Reef Marine Park Authority 1996.

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.

Reef Flat Foraging

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.

 

Spinefoot to 38 cm
Sea Mullet to 78 cm
Sweet lips to 45 cm

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 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.

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.

 

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.

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.

Return rates

 

Costly Signaling

 

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.

 

Assigned Reading

Bliege Bird, R., Smith, E. A., and Bird, D. W. 2001. The hunting handicap: costly signaling in human foraging strategies. Behav Ecol Sociobiol 50:9-19.

Hawkes, K. and Bliege Bird, R. 2002 Showing off, handicap signaling, and the evolution of men’s work. Evol Anthro 11: 58-67.

Also Recommended

Bird, R. 1999. Cooperation and conflict: The behavioral ecology of the sexual division of labor. Evol Anthro 8(2): 65-75.

Smith, E. A., Bliege Bird, R. and Bird, D. W. 2003. The benefits of costly signaling: Meriam turtle hunters. Behav Ecol 14(1):116-126.

 

File: Lecture 12 Juvenile Foraging

ANTH 4962 The Evolution of the Human Diet

Lecture 12

Juvenile Foraging

University of Utah Fall 2005 Helen Alvarez

 

Overlapping Generations

photo by Marjorie Shostak @

http://www.anthrophoto.com/

 

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’s Foraging

 

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.

 

Table 2

 

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.

 

Table 6

 

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.

 

Juvenile Foraging Size or Experience?

 

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.

 

Figure 1 Age effects on large-hook beach fishing efficiency

 

Martu Children’s Hunting

 

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 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.

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.

 

Mikea Children

 

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.

 

Required Reading

Bird, D. W. and Bliege Bird, R. 2005. Martu children’s hunting strategies in the western desert, Australia. In Hunter-Gatherer Childhoods: Evolutionary, Developmental and Cultural Perspectives. eds. Hewlett, B. S. and Lamb, M. E. pp. 129-146. New Brunswick NJ: Transaction Publishers. Course Reserves Marriott Library

Tucker, B. and Young, A. G. 2005. Growing up Mikea: Children’s time allocation and tuber foraging in southwestern Madagascar. In Hunter-Gatherer Childhoods: Evolutionary, Developmental and Cultural Perspectives. eds. Hewlett, B. S. and Lamb, M. E. pp. 147-171. New Brunswick NJ: Transaction Publishers.

Also Recommended

Blurton Jones, N. G., Hawkes, K, and O’Connell, J. F. 1989. Modelling and measuring costs of children in two foraging societies. In Comparative Socioecology: The Behavioural Ecology of Humans and Other Mammals. eds. Standen, V. and Foley, R. A. pp. 367-390. Oxford: Blackwell Scientific Publications.

Hawkes, K. O’Connell, J. F. and Blurton Jones, N. G. 1995. Hadza children’ foraging: juvenile dependency, social arrangements, and mobility among hunter-gatherers. Curr Anthro 36(2): 688-700.

Bird D. W. and Bliege Bird, R. 2002a. Children on the reef: Slow learning or strategic foraging? Human Nature 13(2): 269-297.

Bliege Bird, R. and Bird, D. W. 2002b. Constraints of knowing or constraints of growing: Fishing and collecting by the children of Mer. Human Nature 13(2): 239-267.

 

 

 

Missing 8-1, 8-2, 9-1, 10-2







7-1





Journal of Archaeological Research,
Vol. 9, No. 4, December 2001 ( C 2001)


The Archaeology of Aquatic Adaptations:


Paradigms for a New Millennium


Jon M. Erlandson1


Although aquatic resources are often
seen as central to the development of post-


Pleistocene cultural complexity, most
models of human evolution have all but

ignored the role of aquatic or maritime
adaptations during the earlier stages of


human history. When did aquatic
resources, maritime adaptations, and seafaring


first play a significant role in
human evolution? I explore this fundamental question


by (1) reviewing various theories on
the subject; (2) discussing a variety of prob-


lems that prevent archaeologists from
providing a clear answer; and (3) examining


the archaeological record for evidence
of early aquatic resource use or seafaring. I


conclude that aquatic resources,
wherever they were both abundant and relatively


accessible, have probably always been
used opportunistically by our ancestors.


Evidence suggests, however, that
aquatic and maritime adaptations (including

seafaring) played a significantly
greater role in the demographic and geographic


expansion of anatomically modern humans
after about 150,000 years ago. Another


significant expansion occurred
somewhat later in time, with the development of


more sophisticated seafaring, fishing,
and marine hunting technologies.


KEY WORDS: aquatic resources; human
evolution; maritime societies; coastlines; boats.



INTRODUCTION


The average molluscan flesh is
certainly not very appealing in appearance and the earliest


humans apparently existed for
uncounted millennia before that anonymous hero ate the first

oyster. In any event, shell
middens of real antiquity are rare or absent in world archaeology


(Meighan, 1969, p. 417).


Central to the success of our
species—measured by our wide geographi-


cal range and astounding population
growth—is the combination of human


1 Department of Anthropology,
University of Oregon, Eugene, Oregon 97403-1218; e-mail: jerland@


oregon.uoregon.edu.



287


1059-0161/01/1200-0287/0 2001 Plenum Publishing
Corporation



C


288
Erlandson


intelligence, adaptive flexibility,
and technological sophistication. In a broad his-


torical or evolutionary framework,
humans are the ultimate in generalists and


opportunists, omnivores who thrived in
the widest range of earthly environments,


both natural and cultural. On a planet
whose surface is almost 75% water, where


life itself is dependent on water to
survive, and where our ancestors have success-

fully adapted for at least 2.5 million
years, it has always seemed strange to me that


modern anthropological theory has
maintained that aquatic resources and habitats


were not systematically used by humans
until relatively recently (e.g., Binford,


1968; Cohen, 1977; Osborn, 1977a,b;
Waselkov, 1987; Washburn and Lancaster,


1968; Yesner, 1987). As Bass (1972, p.
9) noted “even our land masses are crossed


and broken by rivers and streams or
dotted with lakes.” Yet among the 10 major


habitats listed by Gamble (1994, pp.
10–11, 1998) as significant to our early an-


cestors as they spread around the
earth, coastlines, lakeshores, and other aquatic


habitats are nowhere to be found.

As Washburn and Lancaster (1968,
p. 294) argued more than 30 years ago,


many archaeologists still seem to
believe that


During most of human history,
water must have been a major physical and psychological


barrier and the inability to cope
with water is shown in the archaeological record by the


absence of remains of fish,
shellfish, or any object that required going deeply into water or


using boats. There is no evidence
that resources of river and sea were utilized until this late

pre-agricultural period . . . for
early man, water was a barrier and a danger, not a resource.


More recently, Yesner (1987, p. 285)
stated categorically that the ”historical fact


that maritime resources were not
exploited until relatively late in the prehistoric


record has attracted a general
consensus. . . . A real commitment to maritime life-


ways did not precede late Upper
Paleolithic times.”


If such statements are accurate,
how did hominids spread around the globe,


colonizing much of Africa and Eurasia
by at least a million years ago, without the


aid of floats, boats, or the
capability to cross sizable bodies of water? How did

they survive in such a wide range of
landscapes when aquatic habitats were such


a physical and psychological
impediment? Why would our omnivorous hominid


ancestors—problem solvers and keen
observers of the world around them—ignore


aquatic resources when hundreds of
highly visible nonhuman predators and omni-


vores do not? Why is there so little
archaeological evidence for the use of marine


resources until postglacial times, long
after the well-documented maritime colo-


nization of island Southeast Asia and
greater Australia? Is it really possible that


aquatic resources were virtually
ignored for more than 99% of human history?


I believe the general perception
that humans only began to seriously adapt to

aquatic environments during the last
15,000 years or so has had a stultifying effect


on the evolutionary study of aquatic
adaptations and societies, maritime migrations,


and the development of boats and other
seafaring technologies. Such perceptions


peripheralize the significance of
aquatic habitats in human evolution, relegating


them to an essentially incidental role
in the broad-spectrum revolution leading to


The Archaeology of Aquatic Adaptations
289


agricultural societies and
civilizations. Thus maritime adaptations appear to play a


marginal role in a relatively brief
process during which the human developmental


trajectory departed from its natural
course as population growth forced humans

into increasingly artificial modes of
subsistence and production.


As Yesner (1987) noted, however,
the picture of aquatic resources as marginal


foods of “last resort” is out of
step with historical and archaeological data that


suggest that maritime or aquatic
hunter-gatherers were generally more sedentary,


populous, and culturally complex than
their terrestrially based interior neighbors


(Birdsell, 1953; McCartney, 1975;
Palsson, 1988; Townsend, 1980). Indeed, some


of the most complex and artistically
acomplished hunter-gatherers of all time de-


veloped in rich marine environments,
including many North Pacific peoples (the

Tlingit, Haida, Aleut, Koniag, etc.)
who lived adjacent to terrestrial environments


relatively unproductive for human
subsistence. Thus, while aquatic resources sup-


ported some of the most complex and
populous hunter-gatherer cultures on earth,


archaeological evidence for the
antiquity of aquatic resource use was extremely


limited. This results in a fundamental
paradox, where supposedly marginal aquatic


resources (although often both diverse
and abundant) appear to provide the eco-


nomic foundation for relatively complex
societies characterized by high popula-


tions and elaborated material cultures.
Despite some notable attempts to account


for problematic aspects of such models
(e.g., Osborn, 1977b; Yesner, 1987), this

aquatic paradox has yet to be
adequately explained or resolved.


In this paper, I discuss some of
these questions and problems by examining


the nature and antiquity of aquatic
adaptations. In the process, I address some of the


broader implications for our
understanding of human migrations, the evolution of


human subsistence and technology, and
current models of optimal foraging theory,


human economic intensification, and
the broad spectrum revolution. I begin with a


short summary of historical thought
about the archaeology of aquatic adaptations,


then discuss some epistemological,
methodological, and taphonomic problems

that currently prevent any real
consensus from being reached about the antiquity of


aquatic adaptations. I then review the
archaeological data available on early aquatic


resource use and maritime migrations
before discussing the broader implications


and some approaches I see as
potentially fruitful for the study of maritime and


aquatic adaptations as we embark on our
voyage into the twenty-first century.


A BRIEF HISTORY
OF THOUGHT


The study of coastal and other
aquatic societies has a long history in an-


thropology and archaeology, one that
closely reflects the general development of

the two fields. Despite this long
history, recent decades have seen a lively de-


bate about the nature of aquatic
environments, their economic productivity for


human societies, and the role they have
played in human evolution (e.g., Bailey,


1975, 1978; Binford, 1968; Claassen,
1991, 1998; Erlandson, 1988, 1994; Fischer,


290
Erlandson


1995a; Glassow and Wilcoxon, 1988;
Isaac, 1971; Jones, 1991; Moseley, 1975;


Osborn, 1977a; Parmalee and Klippel,
1974; Perlman, 1980; Price, 1995; Quilter


and Stocker, 1983; Raymond, 1981;
Sauer, 1962; Waselkov, 1987; Washburn and


Lancaster, 1968; Wilson, 1981; Yesner,
1980, 1987). Prior to the development of

the “New Archaeology” of the 1960s
and 1970s, however, there was little or no


coherent body of theory on the broader
evolution of aquatic adaptations. The opin-


ions expressed on such matters were
generally linked to regional discussions and


varied widely (see Clark, 1936, p. 140;
Morgan, 1877; Uhle, 1907). Nonetheless,


as Clark (1936) and many others
documented the close association of abundant


and widespread shell mounds with
postglacial shorelines, the development of shell


middens and relatively intensive
aquatic economies gradually came to symbolize


an important component of the
post-Pleistocene broad spectrum revolution (see


Bailey, 1978; Binford, 1968). As an
emphasis on theory, method, and broad syn-

thesis came into vogue in the 1960s and
1970s, moreover, considerable interest


focused on more global approaches to
the nature and antiquity of human adapta-


tions to aquatic environments.


In 1994, largely for heuristic
purposes, I characterized the more polarized


viewpoints in this debate as “Garden
of Eden” versus “Gates of Hell” models


(Erlandson, 1994, p. 273). Garden of
Eden theorists, I suggested, saw coastal or


aquatic habitats as veritable
cornucopia where a diverse array of foods—essentially


inexhaustible and easily harvested—was
available (e.g., Cutting, 1962; Fischer,

1995a; Hewes, 1968; Morgan, 1877, p.
21; Okladnikov, 1965, pp. 114–115; Sauer,


1962). On a global level, such
assertions may best be illustrated by Sauer’s de-


scription of the role of the sea in
human evolution.


. . . the path of our evolution
turned aside from the common primate course by going to the


sea. No other setting is as
attractive for the beginnings of humanity. The sea, in particular


the tidal shore, presented the
best opportunity to eat, settle, increase, and learn. It afforded


diversity and abundance of
provisions, continuous and inexhaustible. It gave the congenial

ecologic niche in which animal
ethology could become human culture (Sauer, 1962, p. 45).


Similar statements linked to specific
ethnographic accounts for some coastal groups


or to regional archaeological sequences
limited to the last 5,000 years were es-


poused by a number of authors. Such
glowing assessments often ignored the fact,


however, that archaeological records
for the same regions showed little evidence


for such aquatic largesse dating back
more than a few millennia. On a global


level, moreover, the accumulation of
archaeological data and the development of


chronometric dating techniques made
such statements increasingly problematic.

If aquatic resources were so
productive, why was there relatively little evidence in


the archaeological record for their
exploitation until very late in human prehistory?


After the 1960s, following the
lead of Uhle (1907) and others, a number of


scholars explicitly asserted that
aquatic habitats and resources, when compared


to the hunting of large terrestrial
game, were relatively unproductive for human


exploitation (e.g., Bailey, 1978;
Cohen, 1977; Gamble, 1986, pp. 35–36; Hogg


The Archaeology of Aquatic Adaptations
291


et al., 1971; Osborn, 1977a). These
Gates of Hell models articulated nicely with

the prevailing view of the time that,
prior to the development of agriculture, male-


dominated big-game hunting was the
driving force in human physical, cultural,


and technological evolution. Shellfish
and other aquatic foods, generally viewed


in such models as marginal or even
starvation foods, were portrayed as small and


costly to harvest or process, poor
sources of nutrition, relatively unpredictable or


unreliable, or requiring high
technological investments (boats, etc.) to access. The


fact that collecting shellfish and
other small aquatic foods was primarily women’s


work in most ethnographic societies
further marginalized their importance in hu-


man economies (Claassen, 1998, p. 175).
Gates of Hell models proposed, there-

fore, that the archaeological record
accurately reflected the low productivity of


aquatic resources and the relatively
low value placed on them by many forag-


ing peoples. They argue that humans did
not systematically or intensively harvest


aquatic resources until the
productivity of terrestrial hunting had been reduced by


the intensive harvest pressure of
growing human populations or by the postglacial


extinction of the Pleistocene
megafauna. Thus the use of aquatic resources was


(and is) often assumed to be evidence
for population pressure and environmental


degradation. Osborn, the most ardent
advocate of this position, argued that our an-


cestors “ignored” shellfish and
other aquatic resources for 99% of human history

(Osborn, 1977b, p. 301) and that the
low productivity of marine resources was


virtually universal.


. . . marine resources are
low-return subsistence resources due to a need for labor inten-


sification, in the case of
shellfish and small food package-sized organisms, and due to


their low protein content. A
number of factors combine to create an evolutionary threshold


that is too costly for human
populations to cross unless they are experiencing density-


dependent selection. This
subsistence-related threshold is so costly to cross, in fact, that,

given the option, we should expect
to see human groups shift away from the exploita-


tion of the sea, at least in
nonindustrial societies, whenever possible (Osborn, 1977a,


p. 177).


In practice, relatively few
published opinions can easily be categorized into


such polarized schemes, and most
scholars generally recognize that the situation


is considerably more complex.
Nonetheless, something closer to the Gates of Hell


model has heavily influenced the work
of some of the most influential scholars who

have worked with or discussed coastal
or other aquatic archaeological sequences


(e.g., Bailey, 1975; Binford, 1968;
Cohen, 1977; Fagan, 2001; Gamble, 1986;


Hayden, 1981; Isaac, 1971; Kelly, 1996;
Washburn and Lancaster, 1968).


To square such a dismal view of
the prospects of aquatic peoples with the


evidence that many coastal societies
were characterized by relatively high popula-


tion densities, sedentism, and cultural
complexity—the coastal paradox—required


further explanation. Osborn (1977b)
argued that the population density of aquatic


societies was exaggerated because their
offshore territories were not included

in density calculations, but he could
not resolve the more important issues of


sedentism and cultural complexity.
Cohen (1981) argued that the complexity of


292
Erlandson


Northwest Coast societies was a result
of their high population densities, but


never adequately explained how they
attained such high populations in supposedly


marginal environments. Yesner (1987)
developed the most explicit and sophisti-


cated explanation for the coastal
paradox, arguing that marine and other aquatic


environments were relatively
unproductive until the post-Pleistocene period, when


a combination of megafaunal
extinctions, climatic amelioration, sea level stabiliza-

tion, and the development of mature
coastal habitats allowed coastal populations


to bloom. Thus, he argued, humans did
not intensively utilize aquatic resources


until relatively late in human history,
but the growing productivity of postglacial


aquatic habitats ultimately fostered
the high populations, sedentism, and complex-


ity typical of many Middle or Late
Holocene coastal societies. Problems with this


model include significant variation in
the patterns and timing of megafaunal ex-


tinction or survival, the considerable
evidence for aquatic adaptations prior to such


widespread extinctions, and little
evidence that marine and other aquatic resources


were relatively unproductive prior to
sea level stabilization.

As we shall see, none of these
explanations adequately accounts for either the


basic paradox of supposedly low aquatic
productivity versus high human popula-


tions and cultural complexity, or for
the emerging archaeological data that suggest


that aquatic adaptations developed
earlier and were more widespread than pre-


viously believed. Nor do they explain
how archaeologists armed with essentially


identical data sets can come to such
radically different conclusions about the de-


velopment of such basic aspects of
human economies. To explore these problems,


however, we must first review some of
the different perspectives on the nature

of aquatic resources, then examine some
epistemological problems that inhibit a


comprehensive understanding of the
evolution of aquatic adaptations.


AQUATIC
RESOURCES


Much has been said about the
productivity of various classes of aquatic re-


sources: shellfish, fish, sea
mammals, waterfowl and seabirds, amphibians, plants,


and others. I do not review these
arguments in detail, for such a task could easily


be the subject of an entire paper. It
is important to my later arguments, however,


to examine some of the divergent
opinions expressed about the nature of various

classes of aquatic resources. Also
significant is the fact that aquatic resources are


often collectively lumped as “small”
resources, with the unwarranted assumptions


that they are therefore less productive
than terrestrial game animals for human


subsistence and their presence in
archaeological sites represents de facto evidence


for resource stress or economic
intensification. In this section, I examine some dis-


parate viewpoints about the major
classes of aquatic resources, recognizing that


classes of aquatic organisms not
discussed (amphibians, reptiles, insect larvae,


plants, etc.) may also be significant
resources in some areas.


The Archaeology of Aquatic Adaptations
293


Freshwater


Curiously, perhaps the single
most important aquatic resource for humans,


freshwater for drinking, is seldom
discussed. This may be because our dependence


on water is so fundamental and so
crucial to survival that it is taken for granted. It


is significant, however, because the
almost daily need for drinking water tethered


our ancestors to aquatic habitats for
most of human history. More than any other


resource, drinking water determined
where they settled and where they went,


especially in relatively arid regions.
Maintaining this crucial lifeline to aquatic

habitats, hominids would have spent a
great deal of time observing the behavior of


animals in such environments, including
many terrestrial and amphibious predators


or scavengers that fed on aquatic
animals (see Erlandson and Moss, in press).


Under these circumstances, it seems
unlikely that hominid hydrophobia would


have prevented similar opportunistic
harvesting of shallow water fauna by some


of our earliest ancestors living along
the shores of African lakes. With general


similarities between many of the
animals (fish, shellfish, birds, etc.) that live in


lakes, rivers, estuaries, and marine
habitats, it also seems unlikely that a significant


learning curve would have been required
to transfer such skills between aquatic

habitats. The intensity of such aquatic
harvesting probably varied tremendously,


of course, depending on the relative
productivity of such activities compared to


the other subsistence pursuits
available to a group at various times.


As noted above, the notion of a
long-standing inability of hominids to cope


with aquatic habitats is also difficult
to reconcile with the fact that our human


ancestors now appear to have spread
from Africa into southern Eurasia by about


1.7 million years ago. How did they
accomplish such extensive and early migrations


if they were afraid of the water and
incapable of either swimming or constructing

simple rafts, boats, or other flotation
devices?



Shellfish


No class of aquatic resources has
generated more debate among archaeol-


ogists than shellfish (e.g., Bailey,
1975, 1978; Buchanan, 1988; Claassen, 1991,


1998; Erlandson, 1988, 1991; Glassow
and Wilcoxon, 1988; Jones and Richman,


1995; Meehan, 1977, 1982; Meighan,
1969; Moss, 1993; Noli and Avery, 1988;


Osborn, 1977a; Parmalee and Klippel,
1974; Quilter and Stocker, 1983; Waselkov,


1987; Yesner, 1987). The generic term
shellfish is usually used to refer to a vari-

ety of aquatic invertebrates, dominated
by molluscs (bivalves and univalves), but


also including crabs, sea urchins,
barnacles, shrimp, and other relatively common


organisms. Although the size of
shellfish taxa utilized by humans varies consider-


ably, from large octopi or giant clams
to very small bivalves or gastropods, most


shellfish are relatively small
organisms. What they lack in size, however, many


294
Erlandson


shellfish make up for in quantity and
accessibility—many types are found in large


and sessile aggregations. While most
shellfish provide nutritious sources of com-


plete animal proteins and some vitamins
or minerals, most are relatively low in fat,

carbohydrates, and calories (see
Sidwell, 1981). Although shellfish beds have of-


ten been portrayed by anthropologists
as relatively unproductive, biological studies


indicate that mussel beds produce one
of the highest rates of biomass production


on earth (Jones and Richman, 1995).


Since at least the early 1900s,
many archaeologists have depicted these diverse


and seemingly innocuous creatures as
marginal, secondary, or even starvation foods


for humans.


. . . procuring the essentials of
life by collecting shells in itself indicates a low form of human

existence. In all parts of the
world, even today, people may be seen on the shore at low water


gathering for food the shells
uncovered by the retreating tide . . . these people always belong


to the lower classes of society
and lead in this manner a primitive as well as a simple life


(Uhle, 1907, p. 31).


Some archaeologists bolstered such
arguments with simple comparisons of the


nutritional content of shellfish
versus large land mammals. Bailey (1978, p. 39)


calculated, for example, that 156,800
cockles were required to provide the caloric

yield of one red deer. Some of these
comparisons were inaccurate, and others


ignored the fact that shellfish may
sometimes have been used primarily as a protein


source or that they were often a
relatively predictable and readily available meat


source that could be gathered by
virtually all members of society, including women,


children, and the elderly (see
Erlandson, 1988; Glassow and Wilcoxon, 1988;


Meehan, 1977). The fact that shellfish
gathering was done primarily by women in


most ethnographic societies (Claassen,
1998, p. 175; Moss, 1993, p. 632), in fact,


suggests that such comparisons of
shellfishing versus hunting yields may often be


inappropriate.

Some scholars have also argued
that the small size of shellfish, their relatively


low caloric content, and their
generally high ratio of shell to meat meant that they


were relatively laborious to process
(e.g., Osborn, 1977a; Waselkov, 1987). Others


countered that they required little
search time or technological investment and


could provide highly reliable and
relatively large meat yields that could buffer


the high failure rates of hunting
forays (e.g., Jones, 1991; Meehan, 1982). While


some researchers extolled shellfish as
an efficient protein source (Erlandson, 1988),


others noted that a heavy reliance on
lean shellfish meats could produce “protein

poisoning” (Noli and Avery, 1988),
and still others pointed out that a reliance


on many land mammals (bison, rabbits,
etc.) could produce the same problem


(Buchanan, 1988). While some criticized
shellfish as a resource highly susceptible


to periodic El Nino, storm, or red tide
events, others pointed out that such problems


could sometimes be predicted and
controlled for (Moss, 1993, pp. 640–641) and


that agricultural products and other
terrestrial resources were equally susceptible


to floods, droughts, disease, and
other problems (Quilter and Stocker, 1983).


The Archaeology of Aquatic Adaptations
295


Finally, though Osborn (1977a,b)
and others have used ethnographic or

historical accounts to support the
notion that shellfish were marginal or starvation


foods, Moss (1993) clearly exposed the
complexities and potential androcentrism


often inherent in such accounts. I was
present during her interview of an elderly


Tlingit friend, Richard Newton, who
insisted shellfish were not a major food for


his people. Responding to questions
about the incredible abundance of shellfish


remains in Tlingit village and camp
sites, however, Mr. Newton eventually char-


acterized their dietary role as similar
to bread and butter—long a staple in western


society (Moss, 1993, p. 643). When
asked about this contradiction, he patiently


explained that the ideal Tlingit man
needed to work hard to succeed, that shellfish

encouraged laziness because they were
too easy to collect, that they were gathered


primarily by women, but that they also
were regularly gathered and consumed


by Tlingit men (Moss, 1993). Certain
types of shellfish were especially prized by


Tlingit men, in fact, because they were
said to enhance the libido.


All this debate has had little
effect on the pervasive notion that shellfish


and other small resources are lower
ranked by human foragers (e.g., Broughton


and O’Connell, 1999; Renfrew and
Bahn, 1996, p. 282). Fagan (2001, p. 341)


concluded, for instance, that “no one
can believe that mollusks were the staple

diet” of any ancient society.
Following such assumptions, many archaeologists


continue to view the appearance of
shell middens in archaeological sequences


as evidence for human demographic
pressure, environmental degradation, and


economic intensification (e.g., Cohen,
1977; Hayden, 1981; Waselkov, 1987). The


postglacial florescence of shell
middens adjacent to aquatic habitats around the


world, therefore, has become
essentially synonymous with the anthropological


notion that human economies were
transformed by a global and relatively recent


broad-spectrum revolution.



Fish

Similar debates have taken place
over the nature and productivity of fishing


(e.g., Butler, 1996; Clark, 1948;
Garson, 1980; Kelly, 1996; Limp and Reidhead,


1979; Lindstrom, 1996; Morgan, 1877;
Osborn, 1977b; Rick and Erlandson, 2000).


Literally thousands of different
varieties of fish inhabit the wide range of aquatic


habitats, from the deep abyssal floors
of the oceans to high mountain lakes. Even as


adults, these fish range in size from
tiny gobies to the gigantic whale shark. Some are


largely solitary and relatively rare,
while others are incredibly abundant and swim


in concentrated schools numbering in
the millions. Some aquatic communities,

moreover, are characterized by a
diversity and abundance of fish; others contain


only one or two species and even these
are relatively rare. Still other communities


have low species diversity but a
relatively high piscine biomass.


The nutritional value of fish
also varies considerably, especially the fat and


calorie content of various species.
Although generally low in carbohydrates, fish


296
Erlandson


are a relatively nutritious source of
protein, vitamins, and minerals (Sidwell, 1981;


Watt and Merrill, 1975). Fish eggs,
which can sometimes be harvested in large

quantities, are also generally very
high in protein and calories. Fish flesh and protein


are also highly digestible and
metabolized more efficiently by the human body


than the meat of land mammals. High
rates of fish consumption in modern human


populations—especially certain fish
oils—also seems to be generally correlated


with lower rates of disease and greater
longevity.


Opinions expressed about the
economic productivity of fishing vary widely.


Some accounts have portrayed fishing
as an extremely productive activity, begin-


ning with Morgan’s idealized
statement that “Fish were universal in distribution,

unlimited in supply, and the only kind
of food at all times attainable” (Morgan,


1877, p. 21). In contrast, in comparing
fishing to more traditional hunting activities,


Kelly (1996, p. 209) stated that


. . . fish are different. Some
species, especially surface feeders, will give away their presence,


but not bottom feeders. And fish
cannot be tracked—this is a particular problem in exploiting


oceanic fish. The forager can
only go to a likely place to find fish, then begin searching


randomly. If there are no fish
there, the forager could waste quite a bit of time before

accepting this as likely.


Kelly’s characterization, however, is
at odds with many types of marine fishing,


including the extremely productive and
predictable fishing that can characterize


halibut or cod banks, kelp beds, and
some other nearshore habitats.


Others have argued that fishing
requires relatively sophisticated knowledge


and high technological investments.
Experimental work by Limp and Reidhead


(1979) suggested, however, that under
the right circumstances riverine fishing


could be extremely productive even
without complex technologies. In some aquatic

habitats, the seasonal drying of ponds
or pools can strand fish in shallow water or


on mud flats where they can be easily
collected. In some lakes, periodic hypersaline


or anoxic conditions can also lead to
massive fish kills in which large windrows


of dead fish are deposited on the
beach (e.g., Butler, 1996, p. 701). Quilter and


Stocker (1983, p. 549) described an
apparently regular Peruvian phenomenon


known as “anchovy beaching,” in
which hundreds of thousands of small fish strand


themselves on the beach roughly four
times a year. Spawning fish (salmon, herring,


lamprey eels, grunion, and many others)
also can be highly vulnerable to human


predation, and such spawning runs are
often highly predictable, facilitating the

logistical planning required for mass
harvesting and the processing of fish for


storage.


Even when more sophisticated
technologies are required to capture fish,


these need not be especially elaborate
or expensive to produce. Dip nets or small


tidal weirs, for instance, can greatly
facilitate the mass harvest of small fish in


truly impressive yields. Before
commercial overexploitation devastated many of


California’s marine fisheries,
enormous schools of sardines and anchovies were


available in nearshore and estuarine
habitats. With the aid of boats and dip nets,

The Archaeology of Aquatic Adaptations
297


huge quantities of these small fish
could be captured quickly and easily dried for


later consumption. Still, considering
the lack of evidence for weaving techniques


prior to the advent of the Upper
Paleolithic, even relatively simple fishing tech-


nologies involving cordage, baskets,
nets, or composite projectiles may have been


beyond the capabilities of hominids
prior to the appearance of anatomically mod-


ern humans. And some fishing
activities—especially those requiring large nets,


sophisticated boats, or elaborate weir
structures—would have required consider-


able investment in materials, labor,
and maintenance, as well as intellectual and

communication skills that may have been
beyond the capabilities of our archaic


ancestors.


Despite such technological
constraints, a number of cultural ecological stud-


ies have modeled the productivity of
various fishing activities relative to alterna-


tive terrestrial subsistence pursuits
(e.g., Osborn, 1977b; Perlman, 1980; Simms,


1987). Some of these estimates are
based on incomplete data or the use of inap-


propriate technologies in potentially
depleted modern environments, but they are


still informative, suggesting that the
productivity of fishing varies tremendously.

The most sophisticated analysis of
which I am aware is Lindstrom’s study of the


Truckee River fishery in the western
Great Basin (Lindstrom, 1996), which sug-


gests that fishing harvests using a
number of different aboriginal techniques were


higher than the return rates calculated
by Simms (1987) for terrestrial hunting.


Lindstrom’s projected return rates
varied considerably, however, and some meth-


ods of fishing produced yields that
were considerably less productive than many


terrestrial alternatives.



Aquatic Mammals

There has also been considerable
debate about the nature, antiquity, and eco-


nomic productivity of aquatic mammal
use, especially marine mammals such as


whales, seals, sea lions, sirenians
(sea cow, manatees, etc.), and sea otters (e.g.,


Clark, 1946, 1947; Colten and Arnold,
1998; Erlandson et al., 1998; Hildebrandt


and Jones, 1992; Jones and Hildebrandt,
1995; Lyman, 1995; Osborn, 1977b;


Workman and McCartney, 1998). Aquatic
habitats also are home to a variety of


freshwater animals (hippopotami,
beavers, otters, etc.) of various sizes, which


spend varying amounts of time in the
water and, like some marine mammals, may

sometimes be taken on land. Some
aquatic mammals are also not easily catego-


rized as clearly marine or freshwater:
some seals or dolphins swim considerable


distances up rivers; seals live
permanently in Lake Baikal, the Caspian Sea, and


other European lakes (Reeves et al.,
1992); some manatees are equally at home in


salt- or freshwater habitats; and river
otters and other typically freshwater mammals


may also spend time in brackish or
saltwater habitats.


Clearly most aquatic mammals are
not small resources. They include many


of the largest animals on earth, which
until devastated by commercial whaling or

298
Erlandson


hunting also were relatively abundant
along many of the world’s coastlines. Many


marine mammals weigh well over 500 kg,
with the largest whales weighing over


100,000 kg. Osborn (1977b) argued that
most aquatic mammals occupy positions


relatively high in the food chain,
which limits their numbers relative to the primary


productivity of the world’s oceans.
Such global modeling probably had little or


no relevance, however, to maritime
peoples such as the Koniag or Aleut, who


lived in proximity to biannual
migrations of hundreds of thousands of whales and


pinnipeds (see Haggarty et al., 1991).

Like virtually all mammals, the
meat and organs of aquatic mammals are


relatively rich sources of nutrients,
high in protein, vitamins, and minerals (see


Heller and Scott, 1967; Osborn, 1977b;
Sidwell, 1981). Many aquatic (especially


marine) mammals also have a thick layer
of subcutaneous blubber that provides


them with insulation and human hunters
with a rich source of fat and calories. These


fat deposits can also be rendered into
oil that may be stored for later consumption


or used in lamps as a source of heat
and light. The skins, bones, teeth, ivory, and


baleen of many aquatic animals also
provide valuable raw materials used in a

variety of technologies (houses, boats,
clothing, tools, ornaments, etc.). Among


many societies that actively pursue
large aquatic mammals, successful hunters may


also gain significant status,
prestige, and possibly even reproductive advantages.


Such potentially lucrative
economic payoffs must be measured against the


costs and risks of procuring aquatic
mammals. Sea mammal hunting can be a dan-


gerous and seasonal pursuit, especially
in offshore marine settings, and successful


hunting forays are often relatively
rare. Like fishing, some forms of aquatic hunting


may also require relatively complex and
expensive technology, including seawor-

thy boats and related hunting gear that
represent a significant investment of energy


to produce and maintain. This is
particularly true for many types of sea hunting


recorded among ethnographic marine
hunters. Many of these peoples had high


population densities and had hunted sea
mammals for millennia, however, with


negative effects on the distribution
and density of local prey populations (Jones and


Hildebrandt, 1995; Lyman, 1995). Prior
to such impacts, many pinnipeds may have


been taken relatively easily while
hauled out in breeding or birthing colonies on


islands or other isolated coastal
locales. Where abundant, scavenging of cetacean


and pinniped carcasses off the beach
could also provide large (and potentially

huge) subsistence dividends, with
minimal technological or search costs (Smith


and Kinahan, 1983). Finally, the costs
of manufacturing and maintaining boats


must be measured against the greater
overall efficiency achieved in a variety of


hunting, fishing, and transportation
activities.


As with virtually all classes of
aquatic and terrestrial resources, there is con-


siderable variability in the
characteristics of aquatic mammals and their economic


potential. This includes aspects of
their biology and behavior, their abundance and


availability to humans, the methods
used to procure them, and the relative produc-

tivity of various procurement
strategies versus subsistence alternatives. Given this


The Archaeology of Aquatic Adaptations
299


diversity, it should be no surprise
that different researchers have reached quite dif-


ferent conclusions about the general
role of aquatic mammals in human economies.


In recent discussions, for instance,
various researchers have viewed sea mam-


mals as either central or peripheral to
the development of maritime adaptations


along the Pacific Coast of North
America. Hildebrandt and Jones (1992; Jones


and Hildebrandt, 1995) proposed that
because of their large size and vulnerability


to predation in rookeries, some seals
and sea lions were the focus of early ma-

rine hunters, with later technological
developments (boats, etc.) representing labor


intensification as human impacts on
pinniped populations increased and hunting


strategies changed. Colten and Arnold
(1998) and Erlandson et al. (1998) noted


little evidence for an early focus on
pinniped hunting in the area, however, and


suggested that its general economic
importance may have been overemphasized


(see also Kent, 1989, p. 5; Workman and
McCartney, 1998, p. 362). Central to


resolving such debates are problems
related to recovering and interpreting repre-


sentative samples of sea mammal remains
and estimating their dietary contribution


within the larger economies of human
societies.

PROBLEMS IN
PARADIGMS


Underlying such debates, but
often pushed well into the background, is the


ambiguity of the archaeological record
itself. In some cases, diverging opinions


have been supported with data from
different regions. In others, nearly opposite


conclusions were drawn from virtually
the same archaeological record. How is


it possible for researchers to reach
such different conclusions based on the anal-


ysis of the same body of data? The
answer to that question lies in a variety of


taphonomic, methodological,
interpretive, and theoretical problems that make our

reconstructions of the history of
aquatic societies fraught with uncertainties. The


divergence of opinions about the
antiquity of aquatic adaptations can be attributed


to a variety of problems with the
archaeological record itself, to differences in


the way individual archaeologists
believe the record should be interpreted, and to


differences in the preconceptions of
various researchers.



Definitions


In part, different opinions can
be attributed to the general lack of defini-


tion for what constitutes a dietary
staple, systematic or intensive resource use,

or terms such as coastal, aquatic,
littoral, or maritime adaptations (Workman and


McCartney, 1998). Definitions for
coastal or maritime adaptations have varied, for


instance, from those groups who procure
some portion of their sustenance from


the sea to those who go to sea in boats
and rely on other specialized technologies.


Recognizing the complexity and
diversity inherent in northern cultures, Fitzhugh


300
Erlandson


(1975, p. 344) tried to bring some
order to the classification of maritime societies


by defining five broad adaptive
types: modified interior, interior-maritime, mod-


ified marine, maritime, and riverine.
In his work on the Oregon coast, Lyman

(1991) differentiated littoral from
maritime adaptations, the latter representing


groups who went to sea to obtain much
of their sustenance. Finally, in an attempt


to operationalize the definition of
maritime societies for anthropologists, Yesner


(1980, p. 728) defined “fully
maritime” peoples as those obtaining at least 50% of


their calories or protein from marine
sources. This definition is easily adapted to


riverine or lacustrine peoples, but in
practice it is difficult to accurately or precisely


quantify the dietary contribution of
aquatic versus terrestrial resources. Isotopic


and trace element studies of human bone
have improved our ability to quantify


general aspects of ancient diets, but a
variety of problems (diagenesis, varying

photosynthetic pathways, etc.) continue
to limit such studies.


At times, we must even confront
the issue of what constitutes an aquatic


versus terrestrial resource. How do we
classify a salmon or other anadromous fish


that may be caught in the ocean one
week, in a river or lake the next week, or


scavenged from the shoreline the next?
How do we classify the beaver, hippopota-


mus, crocodile, land otter, or many
other animals that spend a good deal of time


in aquatic habitats but may also be
captured on land? Are seabirds (or their eggs)


taken from terrestrial colonies aquatic
or terrestrial resources? What about seals

or sea lions taken from onshore
rookeries? Finally, how do we classify a deer or


elk captured—as they were sometimes
taken along the Northwest Coast of North


America—while swimming to or from
islands (Tveskov, 2000, p. 131) or nearly


paralyzed by the cold on the beach just
after such a swim? It might be argued that


such ambiguous cases are relatively
unusual, but I suspect they are more com-


mon than many of us recognize, and they
blur the arbitrary distinctions already


drawn between aquatic and terrestrial
resources or marine, estuarine, riverine, and


lacustrine habitats. If such
ambiguities can be recognized in modern habitats and


behaviors, moreover, how can we hope to
differentiate between such ambiguous

cases in the archaeological record?


Changing Sea Levels, Coastal
Erosion, and the Archaeological Record


Despite such ambiguities, the
single greatest problem in evaluating the his-


tory of aquatic adaptations lies in the
fact that sea and lake levels have varied


tremendously over the past 2 million
years, and erosion during high stands has


repeatedly obliterated the
archaeological record where evidence for early aquatic


resource use is most likely to be
found. Sea level today is among the highest of the


Quaternary, exceeded only by Last
Interglacial levels about 6 m higher than today.

Many scholars are rightfully hesitant
to assume that Pleistocene shell middens


were once widespread along submerged
shorelines. Geologically, however, there


The Archaeology of Aquatic Adaptations
301


is ample reason to believe the
archaeological record of coastal adaptations is se-


riously underrepresented (Kraft et al.,
1983). During the last glacial about 20,000


years ago, world sea levels stood
between about 100 and 125 m below present,


exposing broad coastal plains around
the world that have virtually all been inun-


dated as seas rose to their present
levels. Similar cycles have occurred numerous


times during the Plio-Pleistocene,
causing enormous and highly variable changes

in coastal geography around the world.


Worldwide, only Africa and
Eurasia were occupied by hominids when sea


levels were last comparable to today.
Along such Old World coastlines, the Last In-


terglacial sea stand of 125,000–130,000
years ago cut erosional platforms that may


have destroyed most evidence for
earlier coastal occupations. In fact, each time


global sea levels have risen
significantly the record of hominid occupation associ-


ated with lower shorelines has either
been inundated, destroyed by coastal erosion,


or both. Even today, with sea level
roughly 6 m below the Last Interglacial high,

many important coastal sites (e.g.,
Klasies River Mouth caves, Gorham’s Cave,


Grotta dei Moscerini, Daisy Cave)
occupied between about 125,000 and 10,000


years ago are being destroyed by marine
erosion. Uplifted shorelines associated


with earlier interglacials are present
in some areas, but the periods of sea level


maxima represent just a small fraction
of the Pleistocene. It should be no surprise


that associated occupation sites (e.g.,
Terra Amata) are rare. Much more common


are localities such as those in North
Africa and the Levant, where Lower Paleolithic


artifacts (hand axes, etc.) have been
found redeposited on raised marine terraces,


testifying to the destruction of
ancient sites located in coastal or pericoastal settings

(e.g., Bar-Yosef, 1994; Howe, 1967).


Equally important for
understanding the evolution of coastal and aquatic


adaptations are the effects of sea
level change on the paleogeography of coastal


localities. As sea levels rise or fall,
coastlines move laterally in response to such


changes; the environmental setting of
archaeological sites can change dramatically.


Reconstructions at coastal sites with
long occupational sequences have shown


that the exploitation territories of
many sites located on the modern coast were


entirely terrestrial during earlier
occupations (e.g., Parkington, 1981; Shackleton

and van Andel, 1980). The maximum
lateral movements of coastlines during the


last 20,000 years, for instance, have
varied from as much as 1000 km in some


areas (e.g., northern Australia) to
less than 1 km in others. Areas where shorelines


have moved less than about 10 km are
unusual and tend to be strongly correlated


with relatively early evidence for
coastal occupations (Erlandson, in press; see


ahead). Reconstructing the
paleogeography in the vicinity of coastal sites is crucial,


because a cave or open site located on
the coast today may have been 5, 10, 50 km,


or more from the coast at various times
during the last 25,000–125,000 years.


Study of modern coastal
hunter-gatherers suggests that they rarely travel more

than about 5 or 10 km from a home base
to gather foods (Bigalke, 1973, p. 161;


Meehan, 1982). When they do hunt or
forage further afield, the skeletal remains of


302
Erlandson


shellfish, fish, or sea mammals are
often not transported back to a residential base.


In most situations, therefore, sites
located more than about 5–10 km from an ancient


shoreline are unlikely to contain
substantial evidence for marine resource use.


Distances of even 1 or 2 km can
dramatically reduce the density of aquatic faunal


remains (Wing, 1977). During periods of
shoreline transgression or regression,


the intensity of aquatic resource use
at any given site should fluctuate depending

on its proximity to coastal habitats.
After the dramatic postglacial sea level rise


of the last 17,000 years, coastal sites
with long occupational sequences may show


evidence for an intensification of
marine resource use related primarily to changes


in local environments rather than a
regional diversification or intensification of


human subsistence (see Bailey, 1983a;
Parkington, 1981; Shackleton, 1988).


Some may argue that the loss of
early coastal sites can be mitigated by ex-


amining the antiquity of the human use
of lacustrine or riverine resources, but two


problems inhibit such comparisons.
First, it is not clear that the productivity and

diversity of most freshwater habitats
is comparable to marine or estuarine commu-


nities. Second, it is not clear if the
archaeological record of riverine or lacustrine


habitats is any more representative.
Such freshwater environments are also highly


dynamic, and climatic, glacial, and sea
level changes have had profound effects


on their structure and productivity.
Fluctuating lake levels also are common, and


shoreline erosion can produce
geological features essentially identical to marine


shorelines. In riverine systems,
moreover, erosive cycles can rapidly destroy sites


while depositional cycles can bury them
under large quantities of sediment. Thus


preservation and visibility problems
may be just as significant in some freshwater

systems as they are in marine
environments.


Differential
Preservation, Recovery, and Reporting


Another problem lies in the
differential preservation, recovery, and reporting


of organic remains. As we all know, the
shell and bone remains that constitute


the primary record of human use of
aquatic resources are not preserved in many


archaeological sites. Acidic soils, for
instance, or the gradual action of humic


acids in neutral soils, commonly lead
to the deterioration of shells and bones


in archaeological sites. In
comparatively recent sites, especially those occupied

by relatively sedentary peoples, the
accumulation of substantial shell middens can


mitigate the effects of soil acidity or
other factors that lead to the destruction of shell


or bone. For the Paleolithic or
Paleoindian periods, however, when most scholars


believe humans were relatively mobile,
the shell in many low-density middens may


have been insufficient to counteract
soil acidity. The same may be true of pericoastal


or other sites located some distance
from aquatic habitats, where the density of


aquatic food remains was limited by
transportation costs. My experiments with


shells and bones exposed to dilute acid
solutions also showed that shells generally


deteriorate faster than bones, probably
due to their higher calcium carbonate and

The Archaeology of Aquatic Adaptations
303


lower collagen or lipid content. At a
number of archaeological sites, including


Hidden Falls in southeast Alaska
(Erlandson, 1989, p. 139) and Die Kelders in


South Africa (Goldberg, 2000),
moreover, researchers found bone still recoverable,


while shells had either disintegrated
or were too deteriorated to recover or identify.


In the case of Die Kelders, despite the
fact that calcareous rock was abundant in the


site strata, decalcification
completely destroyed the shellfish remains in portions


of the site while bone fragments were
still relatively well preserved.


Among animal bones alone, the
denser and thicker bones of large land mam-

mals are more likely to be preserved in
most archaeological contexts (see Butler


and Chatters, 1994). There has been
relatively little experimentation on the compar-


ative survivability of skeletal remains
from terrestrial versus aquatic vertebrates,


but differential bone density is a
significant factor in preservation. The bones of


aquatic vertebrates generally have
lower densities and are probably more suscep-


tible to chemical dissolution and
mechanical breakdown. Except for the teeth of


some taxa (sharks, etc.), fish bones
are especially lightly built and often have very


high surface area to volume or mass
ratios, suggesting that they would be highly


vulnerable to chemical deterioration.
Some economically important fish (sharks,

rays, sturgeon, lamprey eels, etc.)
also have cartilaginous skeletons with very few


bony parts, and small bony fish (i.e.,
sardines, anchovies) are often eaten whole.


The bones of many aquatic mammals are
also relatively porous and may be prone


to differential deterioration from
mechanical and chemical processes.


Numerous studies clearly show
that the recovery techniques used by archae-


ologists dramatically affect the
interpretations drawn from the recovered assem-


blages. Studies of faunal recovery, for
instance, show that large proportions of the


fish bone and shellfish remains in
many sites are lost during screening of exca-

vated sediments through coarser (0.25
in. or larger) mesh sizes (e.g., Erlandson,


1994; Garson, 1980; Koloseike, 1968;
Moss, 1989). This is a crucial problem in


evaluating the evidence for aquatic
resource use in many early excavation reports,


where researchers had limited interest
in subsistence, faunal remains often were not


systematically recovered, or fine-screen
samples were not collected. Many inves-


tigators now routinely collect faunal
and floral samples through fine screening and


flotation, but others still rely on
cheaper and less systematic recovery techniques.


Because the importance of hunting
or scavenging large game animals has long

been emphasized, there have sometimes
been biases in the analysis or reporting


of other faunal remains from
archaeological sites. In many studies of Middle or


Upper Paleolithic subsistence, in fact,
the only subsistence remains reported on are


large land mammals (e.g., Barker, 1974;
Wolf, 1988), even in early coastal sites


that produced a variety of faunal
remains. Years ago, while visiting early sites in


Gibraltar, I was surprised to find a
number of bluefin tuna and mackeral vertebrae


in the Gibraltar Museum, materials
excavated from early Upper Paleolithic strata


at Gorham’s Cave. For some reason,
these fish bones were never mentioned in


any of the site publications, even
though reports on mammals, tortoises, birds, and

shellfish were all published (see
Baden-Powell, 1964; Eastham, 1968; Waechter,


304
Erlandson


1951, 1964; Zeuner and Sutcliffe,
1964). A similar problem is encountered for the


Middle and Upper Paleolithic levels at
Mugharet el‘Aliya, located near Tangier in


Morocco (Howe, 1967; Howe and Movius,
1947). One of the few Last Interglacial


sites from the south coast of the
Mediterranean, the Paleolithic cave deposits pro-


duced seal, fish, and “a series of”
mollusk remains (Howe and Movius, 1947, p.


21). Although vertebrate remains were
not quantified, they were at least identi-


fied (Arambourg, 1967). Description of
the shellfish remains was limited to the

statement that a “number of mollusks
were found in Layers 5, 6, and 9 of the ar-


chaeological deposits in the Mugharet
el‘Aliya, and were submitted to Dr. William


J. Clench of the Museum of Comparative
Zoology at Harvard. Nothing of value


for our purposes came of this however”
(Briggs, 1967, p. 187).


Such problems may have been due,
in part, to the dearth of specialists who


could identify and analyze the remains
of aquatic fauna. They are symptomatic,


however, of the lower priority
archaeologists traditionally assigned to resources


such as shellfish and fish that were
considered economically marginal or unimpor-

tant. In Howe’s synthesis of the
Mugharet el‘Aliya investigations (Howe, 1967),


for instance, the description of stone
tools is over 31 pages long, the vertebrate


remains are relegated to 5 pages in an
appendix, and the shellfish merit a single


short and obscure paragraph.


The
Hunting Hangover


Even if we can overcome such
analytical hurdles, another serious problem


still confronts us. This is the
persistent effect of the ”Man the Hunter” paradigm


on archaeology. The historical
overemphasis on hunting as central to early human

economies has been dealt with at length
elsewhere (e.g., Slocum, 1975; Zihlman,


1997). The remnants of this outdated
view are still with us, however, more than a


decade after most scholars recognized
that scavenging probably supplied much of


the meat early hominids consumed and
that gathering was much more important


than recognized in earlier
anthropological models. Comparative anatomy also tells


us that human dentition is
fundamentally adapted to omnivory and a relatively


eclectic diet (Scott and Turner, 1997,
p. 81). Evolutionary theory tells us that over


the long haul species are rarely well
served by excessive specialization. Modern


medicine and nutritional studies show
that dietary diversity is fundamental to

human health, growth, and reproductive
success. And common sense tells us that


as our hominid ancestors spread around
the globe, a fundamental part of their


success was their ability to adapt to a
variety of environments or situations and


their relatively eclectic and
opportunistic subsistence economies.


Nonetheless, many theoretical
discussions of subsistence inappropriately


compare the yields of shellfishing or
fishing to those of large-game hunting. If


many hominids relied heavily on
scavenging rather than hunting, for instance,


the relative productivity of gathering
shellfish should be compared to scavenging

The Archaeology of Aquatic Adaptations
305


yields in such cases and must have been
higher than previously estimated. Many


predictions based on optimal foraging
principles also inappropriately treat early


human societies as groups of generic
individuals, ignoring the gender or age-based


divisions of labor in hunting and
gathering activities typical of most recent foraging


cultures. Even for Holocene peoples,
therefore, comparisons of shellfishing and


hunting yields to predict dietary
breadth and subsistence choices may be inappro-


priate, since large-game hunting was
often a primarily male pursuit, and shellfish


and some other aquatic resources were
often collected mostly by women, children,

and older individuals. There is little
doubt, in fact, that the historical devaluation


of shellfish gathering in human
history is related to the fact that it was primarily


the work of women or commoners, to an
androcentric fascination with hunting,


and to biases in historical and
ethnographic accounts recorded primarily by men


(Claassen, 1991, pp. 278–279; Moss,
1993).


Until anthropology transcends some
pervasive misconceptions, the signifi-


cance of aquatic adaptations will
continue to be underemphasized in our recon-


structions of human evolution. These
misconceptions include (1) the notion that

large land mammals were virtually
always the most productive and highly ranked


resources for our hominid ancestors;
(2) that male-dominated hunting was always


the central force that shaped human
subsistence, settlement, and technological de-


velopments; (3) that the utilization of
aquatic resources is automatically evidence


for demographic pressure or resource
stress; and (4) that the archaeological record


preserves a representative picture of
our past.


ARCHAEOLOGICAL EVIDENCE FOR
THE ANTIQUITY


OF AQUATIC
RESOURCE USE

Given the nature and ubiquity of
such problems, is it any wonder that we


know so little about the history of
aquatic resource use? To understand the devel-


opment of aquatic adaptations we are
burdened with several fundamentally flawed


theoretical assumptions and blessed
with a relatively small number of assemblages


where faunal preservation is
exceptional and the full range of faunal remains were


systematically recovered and completely
reported. At the same time, any current


synthesis must rely on an
archaeological record that comes almost exclusively


from sites preserved above modern sea
level even though virtually all coastlines

dating between about 120,000 and 15,000
years ago now lie submerged and distant


from the modern coast.


Despite such problems, numerous
early sites with evidence for aquatic re-


source use have been listed over the
years by Osborn (1977a,b), Perlman (1980),


and Waselkov (1987) or mentioned by
others (e.g., Claassen, 1998; Klein and


Scott, 1986; Yesner, 1980). None of
these lists were exhaustive when they first


appeared, and additional data have
continued to accumulate in subsequent years.


In Tables I–III, I have compiled my
own lists of early “aquatic” sites—those that

306
Erlandson


Table I. Some Early Old World
Localities With Possible Evidence for Aquatic Resource Use


Description
of aquatic fauna


Locality/site and
associations Age (yr) Reference


Homo habilis


Senga 5, Possible use of
freshwater fish, 2.3–2.0M Harris et al., 1990;


Semliki molluscs, and
reptiles associated Meylan, 1990

River, Zaire with Oldowan
tools.


Olduvai Gorge, Possible use of
freshwater fish, 1.8–1.1M Leakey, 1971;


Tanzania crocodiles,
turtles, amphibians, Stewart, 1994


and molluscs.


Homo erectus


Olduvai Gorge, Possible use of
freshwater fish, 1.1–0.8M Leakey, 1971,

Tanzania crocodiles,
aquatic mammals 1994; Roe,


(hippo),
turtles, amphibians, 1994, p. 304;


molluscs, and
possibly salt. Stewart, 1994


Kao Pah Nam, Pile of
freshwater oyster shells 700K Fagan, 1990,


Thailand against cave
wall, associated with p. 120; Pope,


hearth and
land animal bones. 1989

Holon, Israel Freshwater turtle
(Trionyx sp.) shells 500–400 K Bar-Yosef, 1994,


and hippo
bones in Middle p. 246


Acheulian
assemblage of mostly


scavenged (?)
land mammals.


Mas des caves, Seal remains
found in cave site now ca. 400K Cleyet-Merle and


Lunel-Viel, located ca. 10
km from Madelaine,

France Mediterranean
coast. 1995, p. 306


Archaic Homo sapiens



∼350–300K


Hoxne, England Remains of fish,
otter, beaver, and Singer et al.,


waterfowl
associated with 1993; Stuart


Acheulian
deposits; distributions et al., 1993

similar to
artifacts, suggesting a


cultural
origin.



∼400–200K


Duinefontein 2, Sea bird
(penguin, cormorant) Klein et al., 1999a


South Africa remains in
Late Acheulian site


dominated by
land mammal bones.


∼300–230K


Terra Amata, Shellfish and
possibly fish remains de Lumley, 1969;


France associated
with multicomponent Villa, 1983


coastal
campsite.



∼186–127K


Lazaret, France Marine shellfish
in late Acheulian Cleyet-Merle and

context.
Madelaine,



1995



∼150 ± 50K?


Ramandils, Marine shellfish
(>300 fragments) in Cleyet-Merle and


France Middle
Paleolithic strata, probable Madelaine,

food remains.
1995



150 ± 50K


Kebibat, Rabat, Aterian shell
midden on Atlantic Souville, 1973,


Morocco coast,
associated with Neandertal pp. 73–81


remains.


∼130–40K


Presqu’ile du Aterian site on
coast near Berard, Roubet, 1969


Canal, contains
unspecified numbers of


Berard, limpets.


Algeria



∼130–50K

Haua Fteah, Marine shellfish
in Last Interglacial McBurney, 1967


Cyrenaica, strata.


Libya


The Archaeology of Aquatic Adaptations
307



Table I. (Continued)


Description
of aquatic fauna

Locality/site and
associations Age (yr) Reference



∼125–40K Arambourg, 1967;


Mugharet el ’Aliya, Marine
shellfish, fish, and monk seal


Morocco remains in
Mousterian/Aterian Howe, 1967


strata.



∼125–140K D´ b´ nath and

La Grotte Zouhrah, Aterian
assemblage with marine ee


Rabat, Morocco shellfish
(limpets, mussels, and Sbihi-Alaoui,


crab), Homo
sapiens remains. 1979



∼127–40K Roche and Texier,


Grotte des Aterian shell
midden on Atlantic


Contrebandiers, Coast
associated with Homo 1976; Souville,

Morocco sapiens
remains, abundant limpets. 1973, p. 112



∼125–50K Garrod et al., 1928


Devil’s Tower, “Thick
layers” of mussels over


Gibraltar Mousterian
hearths, and a “large


heap” of
marine shells.



∼125–50K Baden-Powell,

Gorham’s Cave, A variety of
marine shellfish remains


Gibraltar from several
Mousterian 1964; Waechter,


occupation
levels. 1951, 1964



∼115–65K Stiner, 1994


Grotta dei Diverse marine
shell remains (3100


Moscerini, fragments),
dominated by mussels

Latium, Italy and clams.
High rates of burning


suggest human
predation; chipped


shell tools.



>45K


Vanguard Cave, Mousterian strata
containing “clear Barton et al., 1999


Gibraltar evidence”
for marine shellfish use

by
Neandertals; includes mussels,


limpets,
cockles, etc., some burned.



>40K


Ras el-Kelb, Mousterian
occupation of coastal or Copeland and


Lebanon pericoastal
cave site, with small Moloney, 1998;


numbers of
marine shells recovered Reese, 1998

from various
occupation levels.



>40K


Salzgitter- Freshwater fish
and mollusk remains Butzer, 1971,


Lebenstedt, associated
with Mousterian p. 477; Cohen,


Germany assemblage.
1977



>37K

Grotta Breuil, Small numbers of
clam and limpet Stiner, 1994,


Latium, Italy shells from
Mousterian strata; p.189


probably not
an “economically


significant”
resource.


Gruta da Figueira Marine shells
(Patella sp.) in 31–30K Straus et al., 1993,


Brava, Portugal Mousterian
levels; density, origin, p. 15

and other
constituents unknown.


Anatomically Modern Humans (Homo
sapiens sapiens)


Middle Stone Age
use of shellfish, sea ∼130–55K Singer and


Klasies River


Mouth, South mammals, and
flightless birds. Wymer, 1982


Africa

Middle Stone Age
shell midden with ∼130–>40K Klein, 1999,


Boegoeberg II,


South Africa numerous
cormorant bones. p. 455; Klein



et al., 1999b


Abdur, Eritrea Middle Stone Age
shell midden? 125K Walter et al., 2000


Early Middle
Stone Age shell midden ∼120–80K Brink and Deacon,

Herolds Bay Cave,


South Africa with mussels
(Perna perna), other 1982


shellfish,
and otter remains


associated
with hearths.



(Continued)


308
Erlandson


Table I. (Continued)


Description
of aquatic fauna


Locality/site and
associations Age (yr) Reference



∼90–75K Brooks et al.,


Katanda 9 and 16, Thousands of fish
bones associated


Semliki River, with MSA barbed
bone harpoon 1995; Yellen

Zaire points in
riverine setting. et al., 1995



∼75–55K Marean et al.,


Die Kelders 1, Sea mammals,
birds, and shellfish


South Africa remains
abundant in MSA cave 2000; Tankard


deposits;
shellfish remains are and Schweitzer,


poorly
preserved. 1974


∼70–60K Volman, 1978


Hoodjies Punt, Open air MSA site
with evidence for


South Africa shellfish, sea
mammals, and fish.



∼70–60K Volman, 1978


Sea Harvest, South Open air MSA site
with evidence for


Africa the use of
shellfish, sea mammals,

and fish.



∼60–50K Henshilwood and


Blombos Cave, MSA shell midden
strata, with variable



or >100K


South Africa densities of
marine shell (mussels, Sealy, 1997;


limpets, etc.),
fish remains, and personal

formal bone
tools. communication,



2000



∼50–15K Johnston et al.,


Willandra Lakes, Abundant shellfish
and fish remains


Australia from numerous
lakeside camps, 1998


associated with
terrestrial fauna and

mixed economy.


Ksar ‘Akil, Numerous
freshwater and marine 43–22K Altena, 1962;


Lebanon shellfish
fragments in Early Upper Ewing, 1947;


Paleolithic
strata; pelican, swan, Kersten, 1991


goose(?), and
duck also found.



∼40–15K Arambourg, 1967;

Mugharet el ‘Aliya, Marine shellfish
and fish remains in


Morocco undated Upper
Paleolithic strata. Howe, 1967


New Britain, Several early
sites containing shell 36–15K Allen et al.,


Melanesia middens, fish
bones, etc.; several 1989a,b


substantial sea
voyages required for


colonization of
archipelago.

Riparo Mochi, Early Aurignacian
stratum produced 35–32K Kuhn and Stiner,


Liguria, Italy almost 5000
pieces of marine food 1998; Stiner,


shell (MNI ca.
500), plus 240 shell 1999


ornaments made
from 43 taxa.



∼35K


Castonet Shelter, Greenland seal
(Phoca hispida) bones Cleyet-Merle and

France in early
Aurignacian stratum. Madelaine, 1995


Mandu Mandu Low density
midden with shellfish, 34–20K Bowdler, 1990;


Rockshelter, crab, and fish
remains at pericoastal Morse, 1988


Western site ca. 5 km
from coast during early


Australia occupation.


Leang Burung, Abundant
freshwater shellfish remains 31–19K Glover, 1981

Sulawesi in cave site.


Gorham’s Cave, Numerous marine
shellfish remains in 30–25K Waechter, 1964;


Gibraltar Early Upper
Paleolithic levels; some Zeuner and


sea bird, seal,
and fish remains. Sutcliffe, 1964


Kilu Rockshelter, Shell midden with
fish bones and other 29–20K Wickler and


Solomon Islands, fauna;
colonization of island Spriggs, 1988

Melanesia required
several substantial voyages


by maritime
peoples.


The Archaeology of Aquatic Adaptations
309



Table I. (Continued)



Description of aquatic fauna


Locality/site
and associations Age (yr) Reference

Shuwikhat-1, Catfish and
large mammal remains at 25K Vermeersch and


Upper Egypt fishing and
hunting station. Van Peer, 1988


Ishango 11 and 14, Abundant fish
remains and some 25–16K Brooks et al.,


Semliki River, shellfish,
crab remains with barbed 1995; Yellen


Zaire bone points
in early LSA et al., 1995


assemblages
in riverine and

lacustrine
setting.


Site 1017 (Khor Khormusan
campsite produced 22.7K Greenwood, 1968,


Musa), Egyptian numerous
catfish bones as part of p. 100


Nubia mixed
economy.


Ohalo II, Jordan Thousands of
fish bones associated 21–18K Nadel and Werker,


Valley, Israel with house
floor on south shore of 1999

Sea of
Galilee.


La Riera, Asturias, Upper
Paleolithic cave strata with 21–14K Straus et al., 1981


Spain shellfish,
fish, and rare seal remains.


Ballana (Site Halfan
campsite with large quantities 19–18K Greenwood, 1968,


8859), Egyptian of burned
bone, mostly freshwater p. 108;


Nubia fish
(catfish, etc.). Wendorf, 1968,


p. 797



∼18–17K Straus, 1976–1977


Altamira Cave, Solutrean use
of shellfish and seal


Santander, Spain within a
predominantly terrestrial


site
economy.



∼17K

Balmori Cave, Upper
Paleolithic conchero containing Clark, 1974–1975


Asturias, Spain hundreds of
marine shells, mostly


limpets.


Coberizas Cave, Shellfish
remains and occasional fish 17–15K Clark and


Spain bones in
Upper Paleolithic strata. Cartledge, 1973


Small numbers
(n = 44) of salmon

Cueva Ambrosio,
16.5K L´ pez, 1988



o


Almeria, Spain vertebrae
and several hundred marine


shell
fragments—ornamental and



nonornamental—in Solutrean levels

of cave ca.
60 km from modern coast.


Note. M = million years; K = thousand
years.


have produced possible evidence for the
use of aquatic foods, other resources, or


maritime activities. These lists, too,
are illustrative rather than comprehensive—I


have compiled such data for years but
still frequently encounter sites with ap-


parent evidence for aquatic resource
use that I was unaware of. Because of the


proliferation of such sites, in fact, I
have limited myself to Old World localities


more than 15,000 years old and New
World sites more than 8,000 years old. There

is no question that aquatic resources
were systematically used in these areas af-


ter these times, and the different
thresholds for the Old and New Worlds also


help compensate for the fact that the
two areas were first colonized by humans at


very different times. Even so, early
aquatic sites are too numerous to discuss or


list individually. Instead, I first
discuss the evidence for the use of aquatic foods


310
Erlandson


Table II. Some Early New World
Localities With Evidence for Aquatic Resource Use



Description of aquatic fauna and


14 C


Locality/site
association age (Kyr) References


Monte Verde, Chile Pericoastal
site with evidence for 12.5? Dillehay, 1997


coastal
contact (seaweeds, etc.).


Broken Mammoth, Abundant
waterfowl remains, some 11.6–9.6 Yesner, 1996


Alaska fish,
otter, and beaver in mixed


economy.

Tule Lake, Fish and
waterfowl as a primary 11.4 Beaton, 1991


California resource in
basal layers of SIS-218



rockshelter.


Lewisville, Texas Several Clovis
hearths associated with 11 Storey et al., 1990


freshwater
shellfish, turtles, and fish


remains
within diversified economy.


Quebrada Jaguay, Faunal
assemblage dominated by fish, 11.1–9.9 Sandweiss et
al.,

Peru shellfish,
and seabird remains. 1998


Pedra Pintada Freshwater fish,
shellfish remains in 11.3–10 Roosevelt et al.,


Cave, Brazil several
Paleoindian occupation 1996


levels.


Marmes Use of
freshwater mussels and salmon 11–10 Caulk, 1988


Rockshelter, along with
terrestrial resources.


Washington

Healy Lake, Alaska Possible
freshwater fish use. 10.9 Borden, 1979


Quebrada Seabird, fish,
and shellfish use. 10.8–10.5 Keefer et al., 1998


Tacahuay, Peru


Kanaka Rapids site, Isotopic
signature of Buhl woman 10.7 Carlson, 1998;


Idaho skeleton
suggests marine (salmon?) Green et al.,


component
in Paleoindian diet. 1998


Ring site, Peru Basal levels of
multicomponent shell 10.5 Richardson, 1998;

dated to
terminal Pleistocene. Sandweiss et al.,



1989


Dalton Complex,
fish as a primary meat 10.5–9.9 Goodyear, 1982


Rodgers Shelter,


Missouri source.



10.4–7.8 Erlandson et al.,

Daisy Cave, San Abalones,
mussels, turbans, and other



1996; Rick et al.,


Miguel Island, shells in
island Paleoindian site;



in press


California Early
Holocene component rich in


shellfish,
fish, and pinniped remains,

with shell
beads, fish gorges, etc.


49-PET-408, Human skeletal
remains with strongly 9.2 Dixon, 1999,


southeast Alaska marine
dietary signature. pp. 180–181


Island
occupation and probable 9–8 Davis, 1989


Hidden Falls,


southeast Alaska maritime
economy—faunal remains


poorly
preserved.


9.6 Dunbar, 1997


Cutler Ridge, Shell midden
with tuna and shark


Florida remains,
located adjacent to narrow


continental
shelf.



9–8 Erlandson, 1994;


California coast Numerous Early
Holocene shell



Erlandson and

middens on
islands and mainland


with
diversity of maritime Moss, 1996



adaptations.


Sabine River site, Submerged Gulf
Coast shell midden 8.5 Dunbar, 1997


Texas with burned
and unburned fish bone.


Chuck Lake II, Island shell
midden with abundant fish 8.2 Ackerman et al.,


southeast Alaska remains
1985

Note. Kyr = thousand years.


The Archaeology of Aquatic Adaptations
311


Table III. Islands
Colonized or Explored by Pleistocene Seafarers


Locality
Description of evidence Date (Kyr) References


Flores Southeast Possible evidence
for Homo erectus 800? Morwood et al.,


Asia crossing of
initial water gap from 1998; Sondaar


Sunda to
Flores. et al., 1994

New Guinea and Oldest sites in
Sunda are the earliest 60–40 Clark, 1991; Groube


Australia evidence for
planned maritime et al., 1986;


voyaging,
involving several sea Roberts et al.,


crossings up
to 90 km long. 1990



∼50


Crete, Greece Homo sapiens
sapiens remains with Facchini and


poorly
documented context; Giusberti, 1992

calcareous
breccia in which bones


were cemented
dated by Pa/U to


51,000 ±
12,000 BP; colonization of


Crete
apparently required several short


sea crossings.

Bismarck Shell middens,
fishing, and seafaring at 35 Allen et al.,


Archipelago, several sites
dated from 15–35 Kyr, 1989a,b; Wickler


Melanesia with voyages
up to 140 km long. and Spriggs, 1988


Sicily, Italy Aurignacian
assemblage from 30 Chilardi et al., 1996


Mediterranean
Island involving short


voyage.


Ryukyu Islands, Human skeletal
remains found in 32–15 Matsu’ura, 1996

Japan Yamashita-cho
and other caves on


Okinawa and
other islands; involves


voyages of ca.
75–150 km.


Kozushima Island, Upper Paleolithic
peoples on Honshu 25–20 Oda, 1990, p. 64


Japan crossing 50 km
wide channel to obtain


obsidian.

Melos Island, Travel across ca.
24 km of open water to 13 Cherry, 1990


Greece obtain
obsidian for mainland trade.


Admiralty Islands, Settlement of
Manus Island required 12 Allen and Kershaw,


Melanesia 200 km voyage.
1996


Cyprus Occupation of
Aetokremnos site, 10.3 Cherry, 1990, p. 151


Akrotiri
Peninsula on southwest coast


of Cyprus.

Channel Islands, Boat and marine
resource use by coastal 11–10 Erlandson et al.,


California Paleoindian
groups, with sea crossings 1996; Johnson


of at least 10
km. et al., 2000; Orr,



1968


Southeast Alaska Presence on
islands indicates a maritime 10–9 Davis, 1989; Fedje


and British lifestyle and
seafaring capabilities. and Christensen,


Columbia
1999

Note. Kyr = thousand years.


during various stages of human
evolution, examining several key sites along the


way. After reviewing such “direct”
evidence for aquatic subsistence, I show that


questions often remain about the
cultural origin of the aquatic (and other) fau-


nal remains found in such sites.
Finally, I discuss some other lines of evidence


for early aquatic adaptations,
including early seafaring and maritime adaptations,


sites submerged on continental shelves
around the world, and the significance


312
Erlandson


of pericoastal sites that indicate some
use of coastal or other aquatic habitats or

resources.


Old
World Localities


For the Lower Paleolithic,
relatively little is known about hominid subsis-


tence. Preservation problems are
especially serious for sites of such antiquity, and


taphonomic issues related to the origin
of faunal remains and their association with


evidence for hominid activity are
paramount. The earliest evidence for the possible


use of aquatic resources by hominids
comes from East African Rift Valley localities


where the remains of a variety of
aquatic or amphibious fauna have been found with

stone tools between about 2.5 and 1.7
million years old (e.g., Auffenberg, 1981;


Greenwood and Todd, 1970; Harris et
al., 1990; Leakey, 1971, 1994; Meylan,


1990; Stewart, 1994). Probably left
primarily by Homo habilis, the contents of


these lacustrine sites record the
scavenging and foraging activities of early ho-


minids, as well as the background noise
of natural accumulation processes. Most


researchers today believe the remains
of large land mammals found at such sites


were accumulated primarily via
scavenging of animals killed by more efficient


predators or other natural causes.
Fernandez-Jalvo et al. (1999) have suggested,


however, that some of the small mammals
represented at such sites may have been

hunted by hominids. Several early Rift
Valley sites have also produced the bones of


aquatic or amphibious animals,
including hippos, crocodiles, fish, frogs, shellfish,


etc. (Leakey, 1971). Because many of
these sites formed in dynamic lakeshore


settings, however, any clear
association of aquatic (and terrestrial) fauna with ho-


minid activities is difficult to
demonstrate. At some sites, the remains of fish appear


to be closely associated with hominid
artifacts, but in others fish bones are rela-


tively abundant in both cultural and
natural strata. Greenwood and Todd (1970,


p. 240) and Stewart (1994) have argued,
however, that fish (especially the cat-


fish, Clarias sp.) would have been
relatively easy to procure in some Rift Valley

aquatic settings and are unlikely to
have been ignored by early hominids. This


seems logical, especially for hominids
living in lakeshore settings with economies


based on opportunistic scavenging and
foraging.


For Homo erectus, a series of
East African sites has produced similar asso-


ciations of artifacts and aquatic or
amphibious fauna. At Olduvai, Leakey (1994)


reported that the bones of catfish and
hippos are ubiquitous in artifact-bearing


sediments dated between about 1.1 and
0.4 million years ago, and the remains


of crocodiles, aquatic turtles, and
shellfish also are found in some sites. As was

the case with much of the Olduvai
fauna, Leakey (1994, p. 142) recognized the


difficulty in determining whether
these aquatic taxa were deposited by hominids,


but she argued that a cultural origin
for the catfish was most likely given their


fragmentary condition and close
association with artifacts (see also Auffenberg,


1981; Roe, 1994; Stewart, 1994). In Bed
III at Olduvai, dated between about 1.1


The Archaeology of Aquatic Adaptations
313


and 0.8 million years ago, Leakey
(1994; see also Roe, 1994) also found a series


of distinctive pits and furrows
possibly associated with the evaporative production


of salt by Homo erectus.

Along the coastlines of Africa
and the Middle East, there is also relatively


widespread evidence for Lower
Paleolithic occupation (e.g., Bar-Yosef, 1994,


p. 214; Howe, 1967; Wulsin, 1941). Most
of these localities are poorly dated,


however, and contain choppers, hand
axes, and other stone tools found in raised


interglacial beach deposits. Although
many of these clearly document the oc-


cupation of coastal plains, the precise
age and environmental context (coastal,


pericoastal, inland?) of such
occupations is not clear.


What appears to be relatively
unambiguous use of aquatic resources by

Homo erectus in Southeast Asia comes
from the site of Kao Pah Nam, a lime-


stone cave in northern Thailand
occupied about 700,000 years ago (Pope, 1989).


According to Fagan (1990, p. 120),
“considerable numbers” of freshwater oyster


shells were found piled against the
cave wall. In the same level, stone tools, a


cobble-ringed hearth, and the remains
of hippo, ox, deer, porcupine, and rat were


found.


Evidence for aquatic resource use
increases somewhat with the appearance


of archaic Homo sapiens after about
400,000 years ago. It is not clear, however,

whether this increase represents real
behavioral or environmental shifts or the


better preservation and greater
visibility of more recent occupations. At the Lower


Paleolithic site of Hoxne in England,
Clactonian artifacts and faunal remains have


been found in what have been
interpreted as lakeshore and alluvial deposits (Singer


et al., 1993). Although the dating of
the Hoxne occupations is still somewhat


tentative, much of the Clactonian
occupation appears to have occurred during an


interglacial period between about
350,000 and 300,000 years ago. The associated


fauna are dominated by large land
mammals (especially horse and deer), but include


numerous specimens of freshwater fish
(pike, roach, stickleback, etc.) and beaver,

and smaller numbers of otter and
waterfowl (Stuart et al., 1993). The cultural


origin of the aquatic and other faunal
remains, like those from the Olduvai sites,


has not been firmly established, but
Stuart et al. (1993, p. 198) noted


that the distributions of all of
the beaver Castor fiber and extinct beaver Trogontherium


cuvieri material . . . and most of
the fish material . . . follow the same broad distribution


pattern as the larger bones,
stones, and artifacts. This suggests that the remains of these taxa


also might be food remains
accumulated by man. . . .

Also in England, excavations at the
Lower Paleolithic site of Clacton-on-Sea pro-


duced fish and freshwater mussel
remains (Singer et al., 1973), although the dating


of the site (ca. 425,000 years (Singer
et al., 1993, p. 219) or ca. 250,000 (Gamble,


1986, p. 140)) and the cultural origin
of the aquatic fauna remain uncertain.


About 300,000 years ago, archaic
Homo sapiens also occupied Terra Amata


along the Mediterranean coast of France
(de Lumley, 1969; Villa, 1983). Mussels


314
Erlandson


and other marine shells were found at
Terra Amata, but their context and quan-

tity are poorly documented. Other early
Old World evidence for shellfish use


comes from several North African Middle
Paleolithic or Aterian sites like Haua


Fteah in Libya (Klein and Scott, 1986;
McBurney, 1967), Mugharet el’Aliya in


Morocco (Howe, 1967), and several sites
in Morocco and Algeria (D´ b´ neth and



ee


Sbihi-Alaoui, 1979; Roche and Texier,
1976; Roubet, 1969; Souville, 1973). In

southern Europe, Mousterian use of
shellfish is suggested by assemblages from


Monte Circeo (Stiner, 1994) and
Grimaldi caves (Stiner, 1999) in Italy, Ramandils


in France (Cleyet-Merle and Madelaine,
1995), and Devil’s Tower Rockshelter


(Garrod et al., 1928), Gorham’s Cave
(Waechter, 1964), and Vanguard Cave


(Barton et al., 1999) in Gibraltar. At
the Italian cave of Grotta Moscerini, ma-


rine shells with flaked edges suggest
that shell tools were used by Neandertals


between about 60,000 and 80,000 years
ago (Stiner, 1994, pp. 187–188). For


Neandertals and other archaic Homo
sapiens living in coastal areas, there is little


evidence for the exploitation of fish
(but see Cleyet-Merle, 1990; Cleyet-Merle and

Madelaine, 1995), and the exceptional
cases may represent scavenging from the


beach. Pinniped bones also are rare in
Middle Paleolithic sites and may represent


scavenging of stranded animals or
carcasses. Nonetheless, there is little doubt that


archaic Homo sapiens occupying the
Mediterranean littoral actively foraged for


shellfish and other intertidal
resources (Stiner, 1994, p. 216).


With the appearance of
anatomically modern humans (Homo sapiens sapiens


or AMH), beginning about 125,000 years
ago, Old World evidence for the use of


aquatic resources increases
dramatically. This disparity is even more pronounced

if the Aterian sites of northwest
Africa are considered to be associated with early


or nearly modern Homo sapiens sapiens
groups (see Klein, 1999). The earliest


evidence for such associations may come
from a recently reported locality near


Abdur in Eritrea along the Red Sea
coast, where what are described as Middle


Stone Age (MSA) stone tools were found
with the remains of marine shells and


other aquatic fauna in strata dated to
about 125,000 year ago (Stringer, 2000;


Walter et al., 2000). With the
information currently available, however, it is not


clear whether the stone artifacts were
left by anatomically modern humans or if


the faunal remains represent the food
refuse of hominids. More secure and better-

documented associations and evidence
come from a series of MSA coastal sites


in South Africa dating between about
120,000 and 50,000 years ago, including


Klasies River Mouth caves (Deacon and
Deacon, 1999, pp. 102–106; Singer and


Wymer, 1982), Die Kelders cave (Marean
et al., 2000; Tankard and Schweitzer,


1974), the Sea Harvest and Hoodjies
Punt sites near Saldanha Bay (Volman, 1978),


Herolds Bay Cave (Brink and Deacon,
1982), the Boegoeberg 2 rockshelter (Klein,


et al., 1999b), and Blombos Cave
(Henshilwood and Sealy, 1997). At these sites,


the earliest evidence for relatively
diversified coastal (or mixed) economies is


found, including the relatively
intensive use of shellfish, pinnipeds and cetaceans,

and flightless seabirds (i.e.,
penguins). Fish remains are virtually absent from these


coastal MSA localities (Klein and
Cruz-Uribe, 2000), except for Blombos Cave


The Archaeology of Aquatic Adaptations
315


where a significant number of large
fish bones have been found in MSA shell


midden strata associated with bone and
stone projectile points (Henshilwood and


Sealy, 1997). Initially estimated to be
between about 50,000 and 60,000 years


old, sediments capping the MSA levels
at Blombos Cave have now been dated


via thermoluminescence (TL) to
approximately 100,000 years ago (Vogel et al.,


1999). The dearth of fish in most
South African sites led Klein (1995, 1998) and

Klein and Cruz-Uribe (2000) to suggest
that fishing may have been beyond the


intellectual or technological
capabilities of early anatomically modern humans. It


is possible, however, that the higher
technological costs of marine fishing generally


discouraged such activities, just as
fishing seems to have been limited at most sites


along the California coast during the
early Holocene (Erlandson, 1994, but see


Rick et al., in press). Along with the
Blombos Cave fish remains, support for this


latter idea comes from the carefully
made barbed bone harpoon points found with


the remains of numerous large
freshwater fish at two MSA sites at Katanda on the


Semliki River in Zaire (Brooks et al.,
1995; Yellen, 1998; Yellen et al., 1995). Dated

to about 80,000 years ago, the Katanda
harpoons represent the earliest evidence for


complex composite fishing technologies
in the world and add to the evidence for a


significant expansion of aquatic
resource use among anatomically modern humans.


Similar barbed bone points also
have been found associated with numerous


fish bones at the Late Stone Age site
of Ishango 14 on Lake Rutingaze (Edward)


in Zaire, in strata dated to about
20,000 radiocarbon years before present (RYBP)


(Yellen, 1998). Fish bone is relatively
abundant at some other Late Pleistocene


African sites, including the White
Paintings rockshelter in Botswana (Stewart,

1994; Yellen, 1998) where the lower
levels are tentatively dated to ca. 20,000


years ago, and a series of Nile River
sites dated between about 40,000 and 15,000


RYBP. In coastal areas, little is known
about aquatic resource use during this time


period because sea levels were deeply
depressed during the Last Glacial and most


African coastlines were far removed
from sites now located along the modern


shore (see van Andel, 1989).


This same interval in southwest
Asia and Europe also is problematic due to


lowering sea levels and extensive
glaciation. Numerous Upper Paleolithic sites in

southern and southwest Europe have
produced evidence for shellfish collection


and consumption. Shellfish densities
increase in many of these sites near the end


of the Pleistocene (e.g., Straus, 1990;
Straus et al., 1980, 1981), but it is not


clear if this represents an
intensification of shellfishing in response to population


growth, increased sedentism, changes in
marine or estuarine environments, or


a combination of such processes (see
Bailey, 1983a,b; Clark and Straus, 1983;


Straus and Clark, 1983). Numerous
interior or pericoastal Upper Paleolithic sites


in Europe and southwest Asia also have
produced beads or other ornaments made


from marine shells or artistic
depictions of aquatic animals (Cleyet-Merle and

Madelaine, 1995; Clottes and Courtin,
1996; White, 1993). The presence of sizable


numbers of marine shell ornaments, in
some sites obtained from both Atlantic and


Mediterranean coastlines more than 100
km distant, suggests that interior people


316
Erlandson


traveled to the coast seasonally or
actively traded with peoples living along these


coasts.


In southern Asia, there is only
limited evidence for aquatic resource use


from this time period. In Indonesia, a
freshwater shell midden known as Leang

Burung attests to the systematic
exploitation of shellfish as much as 31,000 RYBP


(Glover, 1981). At Longrien, a long
Upper Paleolithic sequence contains very


limited evidence for aquatic resource
use, but produced a few bivalves from a


layer dated to about 30,000 RYBP.
Despite the current dearth of evidence, there


can be little doubt that maritime or
other aquatic peoples lived in Southeast Asia


since at least 50,000 years ago.


The peopling of Australia and New
Guinea testifies to this, since migrating


from Southeast Asia to Sahul would have
required several substantial sea crossings

even during periods of much lower sea
level (Clark, 1991). Not surprisingly, early


evidence for the use of freshwater fish
and shellfish comes from Australia, which


now appears to have been settled by
maritime peoples between about 50,000


(Roberts et al., 1990) and 60,000 years
ago (Thorne et al., 1999). Numerous


freshwater shell middens from the
Willandra Lakes area of southeast Australia have


been radiocarbon dated between 38,000
and 15,000 RYBP; Thorne et al. (1999)


recently argued that some of these
lacustrine occupations may date to as much


as 60,000 years ago. Although evidence
for intensive marine resource use in late


Pleistocene Australia is lacking,
several sites from western Australia have produced

limited amounts of marine shell from
strata dated between about 20,000 and 36,000


RYBP (e.g., Bowdler, 1990; Morse, 1988;
O’Connor, 1989; Veth, 1993). At Mandu


Mandu Creek rockshelter, located only
about 4–5 km from the coast just prior to


the Last Glacial, a low-density midden
deposit includes the remains of shellfish,


crab, fish, and terrestrial fauna
(Bowdler, 1990; Morse, 1988). These sites could


be interpreted as evidence for limited
Pleistocene use of marine resources, but sea


level and shoreline reconstructions
show a strong correlation between the presence


and density of marine resources and the
variable distance of each site from the sea.


The apparent abandonment of most of the
sites during the height of the last glacial,

and the fact that they were reoccupied
when sea levels again approached modern


levels, can be interpreted as evidence
that the lateral migration of coastal habitats


strongly influenced local settlement
and subsistence patterns. Several saltwater


shell middens located on the Melanesian
islands of New Ireland, New Britain,


and the Solomons—islands that
required additional sea voyages of 80–100 km to


reach—have been dated between about
35,000 and 15,000 RYBP (Allen et al.,


1989a,b; Wickler and Spriggs, 1988).
The aquatic focus of these early Melanesian


occupations is attested to not just by
the seafaring required to settle the islands,


but also by the abundance of marine
shellfish and fish remains found in the site

deposits. The presence of such sites in
western Melanesia, in contrast to Australia


and New Guinea, is due to the steep
local geography, where the bathymetry plunges


rapidly into deep water and changes in
sea level have had relatively limited effects


on the local shorelines and the coastal
archaeological record.


The Archaeology of Aquatic Adaptations
317


New
World Localities


In the New World, most early
evidence for human use of marine resources


comes from the Pacific Coast, where
relatively steep bathymetry also has limited

the lateral displacement of postglacial
shorelines (Erlandson, in press; Richardson,


1998). The earliest sites currently
come from South America. In Chile, the con-


troversial pericoastal site of Monte
Verde has been dated to ca. 12,500 RYBP and


reportedly contains evidence for
coastal foraging, including four types of seaweed


(Dillehay, 1997). At the coastal site
of Querero, which has produced a suite of


dates between about 11,600 and 10,900
RYBP, marine shellfish, sea lion, and


whale remains were all found associated
with those of mastodon, deer, and other


land mammals (Nu˜ ez et al., 1994). At
Quebrada de las Conchas on Chile’s north


n

coast, Llagostera (1979) also
documented the existence of a diversified maritime


economy including the use of a variety
of shellfish and fish between about 9700


and 9400 RYBP.


Along the south coast of Peru,
Sandweiss et al. (1998) reported an early


component from Quebrada Jaguay, where
shellfish, fish, and sea bird remains


have been found in strata dated between
about 11,100 and 9,900 RYBP. The


faunal remains at Quebrada Jaguay
suggest an almost exclusive reliance on marine


animals, but the presence of obsidian
from a distant interior source suggests that the

site may be just one aspect of a
seasonal round that included interior sites as well


(see Richardson, 1998). Also located on
the southern Peruvian coast, and nearly


as old (10,800–10,500 RYBP), is
Quebrada Tacahuay, where the faunal remains


from the earliest occupation are
dominated by sea bird (cormorant, booby, and


pelican) and fish (anchoveta, anchovy)
bones, with a few shellfish (clam, mussel)


remains (Keefer et al., 1998). Of the
3,775 faunal elements recovered from the


basal stratum at Quebrada Tacahuay,
only eight (0.2%) were from terrestrial taxa.


A third site on the south coast, the
Ring site, contains a shell midden that first may


have been occupied as early as 10,600
RYBP (Sandweiss et al., 1989). Along the

north coast of Peru, Richardson (1998)
has described several ephemeral camps


of the Amotape complex, where unifacial
tools have been found associated with


the remains of mangrove shellfish
(Anadara tuberculosa) dated to about 11,200,


10,000, 9200, and 9000 RYBP (see also
Llagostera, 1992). In Ecuador, coastal


shell middens of the Las Vegas complex
are now dated as early as 10,800–10,100


RYBP (Richardson, 1998; Stothert,
1985).


The meticulous work of Roosevelt
et al. (1996) on Paleoindian components


at Pedra Pintada cave in Brazil dated
to ca. 11,000 RYBP also shows that freshwa-

ter fish were an important component
of an early Amazonian economy that was


relatively eclectic and focused on
smaller plant and animal resources.


Along the Pacific Coast of North
America, the earliest and best-documented


maritime sites currently come primarily
from California. On San Miguel Island off


the California coast, Daisy Cave
contains a thin dark soil containing a few chipped


stone artifacts and a low-density shell
midden containing abalone, mussel, turban,


318
Erlandson


and other shellfish remains dated to
about 10,400 RYBP (Erlandson et al., 1996).

That humans were on California’s
Channel Islands by the end of the Pleistocene


has long been suggested by Orr’s 14 C
dating of the Arlington ”Man” (probably a


woman) skeleton to ca. 10,000 RYBP
(Orr, 1968). Recent redating of this skeleton


suggests that Arlington Woman actually
may have died closer to 11,000 RYBP


(Johnson et al., 2000), but a precise
date has yet to be established. Since the


Channel Islands have been separated
from the California mainland throughout the


Pleistocene, these two sites
demonstrate that Paleoindian peoples had seaworthy


boats during the terminal Pleistocene
and leave little doubt about their maritime


capabilities. Along the California
coast, there are also dozens of shell middens

dated between about 9,700 and 8,000
RYBP (Erlandson, 1994; Erlandson and


Moss, 1996; Jones, 1991). One of the
best examples comes from Daisy Cave,


where stratified shell midden deposits
dated between about 9,700 and 7,800 RYBP


contain abundant shellfish and fish
remains, smaller numbers of pinniped and sea


bird remains, numerous bone fishing
gorges and shell beads, and woven artifacts


made from sea grass (Connolly et al.,
1995; Erlandson et al., 1996).


Along the coastlines of northern
California, Oregon, and Washington, there


are only two shell middens reliably
dated to about 8,000 RYBP, Duncan’s Landing

Rockshelter on the northern California
coast and the Indian Sands site on the Ore-


gon coast (Erlandson, 1994; Erlandson
and Moss, 1996; Lightfoot, 1993; Moss


and Erlandson, 1995). The dearth of
early sites in this intermediate area of the


Pacific Coast now appears to be
related to a long history of occasional massive


subsidence earthquakes along the
Cascadia Subduction Zone, tectonic events com-


monly associated with tsunamis and
severe marine erosion (Erlandson et al., 1998;


Minor and Grant, 1996). In British
Columbia and southern Alaska, a number of


early coastal sites dated between about
8,000 and 10,000 RYBP have been docu-


mented (Carlson, 1998; Erlandson and
Moss, 1996; Fedje and Christensen, 1999;

Moss, 1998), including portions of a
human skeleton found in a cave known as 49-


PET-408 (On-Your-Knees Cave) on Prince
of Wales Island dated to approximately


9,200 RYBP (Dixon, 1999, p. 118). The
isotopic composition of this skeleton is


consistent with a diet comprised almost
entirely of marine foods. A bone tool


manufactured from a land mammal rib
found in another part of the same cave


has been dated to about 10,300 RYBP
(Dixon, 1999, p. 181), suggesting that the


site may have been occupied even
earlier. This terminal Pleistocene date is similar


to the estimated age of a basalt flake
recovered from the surface of a paleodelta


deposit located on the continental
shelf off the Queen Charlotte Islands of British

Columbia (Fedje and Christensen, 1999),
although these early dates should be


regarded as very preliminary.


Adjacent to the generally broader
and shallower continental shelves of the


Gulf of Mexico and Atlantic coasts,
early coastal archaeological sites are much less


common. On the Louisiana coast, where
shorelines of the Mississippi delta have


been prograding for millennia, Gagliano
(1970) reported estuarine shell associated


The Archaeology of Aquatic Adaptations
319


with an 11,000-year-old archaeological
site at Avery Island. Off the Gulf Coast,

near the intersection of a creek and
the submerged channel of the Sabine River by


the Louisiana and Texas border, Dunbar
(1997) noted the presence of a submerged


shell midden dated to about 8,500 RYBP
that contains both burned and unburned


fish bone.


Along the Atlantic Coast of North
America, shell middens dating earlier than


about 8,000 years are extremely rare.
Along most of the Florida coast, for in-


stance, the Clovis-age shoreline is
believed to have been between about 50 and


150 km offshore (see Dunbar et al.,
1992, p. 125). Consequently, postglacial shore-

line changes have been dramatic in most
areas, Florida coast shell middens more


than about 5,000 years old are highly
unusual, and a number of submerged shell


middens have been found. One exception
to this pattern is the Cutler Ridge site,


located adjacent to a narrow stretch of
continental shelf near Miami, where lateral


shoreline changes associated with
postglacial sea level rise have been minimal.


This important site, dated to as much
as 9,600 RYBP but largely unpublished,


reportedly has produced the remains of
a variety of marine fish (tuna, shark, etc.)


and shellfish (Dunbar, 1997).


Although interior Paleoindian
groups are often portrayed as relatively special-

ized big-game hunters, there is
evidence for the use of aquatic resources at a number


of early sites. These include the
Broken Mammoth site in south-central Alaska,


where two well-stratified terminal
Pleistocene components have been identified,


one dating between about 11,800 and
11,000 RYBP and another between about


10,300 and 9,600 RYBP (Yesner, 1996).
Faunal remains are well preserved in these


early components. Identifiable
elements from the older component are dominated


(>60%) by aquatic birds (swan,
geese, and ducks), but also include some large


and small land mammals (wapiti, bison,
etc.). The younger of these components

is dominated by the remains of large
ungulates, but also contains about 30% small


mammals, 10% waterfowl, and smaller
numbers of salmonid, beaver, and otter


remains. Another Paleoindian component
containing the remains of aquatic fauna


is the Lewisville Clovis site located
along the Trinity River in north-central Texas,


where archaeological deposits
associated with numerous hearths yielded a diverse


array of plant and animal remains. Of
the 16 hearths excavated, 9 contained the


remains of freshwater mussel and snail
shells—many of them burned. Also recov-


ered were the remains of box turtle,
fish, amphibians, prairie dog, rabbit, tortoise,


egg shells, raccoon, snake, etc. (Story
et al., 1990). At the Horn Shelter 2 site lo-

cated along the Brazos River,
Clovis-age deposits also yielded the remains of land


turtles and a few fish remains. A
younger component at the site, dated between


about 10,000 and 9,500 RYBP, also
produced a diverse faunal assemblage, includ-


ing the remains of many freshwater
mussels and fish such as drum, gar, and catfish,


along with a double human burial
associated with numerous beads made from the


marine shells Oliva sayana and Neritina
reclivita (Story et al., 1990, pp. 203–204).


Another early North American site is a
small rockshelter (CA-SIS-218) located


320
Erlandson


on the shore of Tule Lake in northern
California, where Beaton (1991) identified

a hearth dated to 11,450 ± 340
RYBP associated with charcoal, ash, and fish, wa-


terfowl, and mammal bones. At Marmes
Rockshelter in Washington, freshwater


mussels and salmon bones are reported
from deposits dated between about 10,000


and 11,000 RYBP.


Shellfish
Feeders and Carrion Eaters


Globally, the growing number of
early sites known to contain the remains of


shellfish, fish, sea birds, sea
mammals, and other aquatic fauna may indicate that

aquatic resources were used relatively
early in human history, by Homo habilis,


H. erectus, and H. sapiens. Due to a
variety of questions about the context, taphon-


omy, recovery, and interpretation of
many ancient faunal assemblages, however,


it is difficult to evaluate how
significant aquatic resources were in early hominid


economies. Moreover, lists like those
presented here suffer from another problem


that must be addressed before we can
conclude that even incidental use of aquatic


resources was both early and
widespread. This problem is the possible role various


animals and other noncultural processes
may have played in the accumulation of


aquatic animal remains in early sites
(see Butler, 1993; Erlandson and Moss, in

press). Although recent taphonomic
studies show that a wide range of scavengers


and predators transport bones into
caves and other sites, few have considered the


possibility that animals and not humans
may have transported marine shells, fish


bones, or sea mammal remains into early
coastal sites.


After visiting several
Paleolithic cave sites in Gibraltar in the mid-1980s, I


did not initially question whether the
remains of marine shellfish and fish found


in the site deposits (other than a
clearly defined Last Interglacial beach) could


have been deposited by anything other
than humans. In her detailed taphonomic

analysis of faunal remains from the
Monte Circeo caves in Italy, Stiner (1994)


considered a host of possible sources
for animal bones but considered only cul-


tural mechanisms for the accumulation
of shellfish remains. Recent research has


shown, however, that a wide range of
predators and scavengers—bears, hyenas,


coyotes, badgers, cats, and a variety
of birds—transport the remains of aquatic


vertebrates (seals, fish, birds, etc.)
and invertebrates (shellfish, etc.) to terrestrial


landforms (e.g., Erlandson and Moss, in
press; Jones and Allen, 1978; Klein et al.,


1999b). Caves and rockshelters, in
particular, provide shelter for a wide variety

of mammals and birds that hunt or
scavenge in aquatic habitats and may deposit


carcasses or skeletal remains at site
locations where they can be mixed with fau-


nal remains left by hominids.
Archaeologists, therefore, must carefully evaluate


the nature of terrestrial and aquatic
faunal remains found in both cave and open


sites to determine whether the activity
of nonhuman predators or scavengers has


contributed significantly to the
faunal remains present at a site.


Unfortunately, such careful
evaluations have rarely been done, and it is either


difficult or impossible to evaluate
the cultural origin of the aquatic fauna in many of

The Archaeology of Aquatic Adaptations
321


the sites listed in Tables I and II.
With a more critical eye toward the origin of aquatic


remains in early sites, the evidence
for early aquatic resource use at some key


localities may need to be reassessed.
Gorham’s Cave produced hundreds of marine


shells, for instance, but my
observations suggest that these were widely scattered


in the cave deposits. Gorham’s Cave
also produced the remains of a wide variety


of birds, including seagulls and others
known to feed on and transport shellfish


(Erlandson and Moss, in press). Without
further evidence to link these aquatic fauna


to cultural activities, we cannot be
certain how significant aquatic foods were to the

Neandertal and Upper Paleolithic cave
occupants (but see Barton et al., 1999). At


present, similar questions can be
raised about virtually all of the Lower Paleolithic


sites listed above, as well as the
Mugharet el’Aliya in Morocco where monk seal,


fish, and shellfish remains were
found in Paleolithic layers (Arambourg, 1967;


Howe, 1967). In the New World, similar
questions have been raised about some


Pleistocene or Early Holocene “shell
middens” located on California’s northern


Channel Islands (e.g., Erlandson, 1994,
pp. 183, 196; Erlandson and Morris, 1992;


Erlandson and Moss, in press).


For other sites, the Middle Stone
Age middens of South Africa promi-

nent among them (but see Klein et al.,
1999a,b), the evidence linking hominids


with aquatic resource use seems much
more secure. In the Mousterian levels at


Devil’s Tower, for instance, Garrod
et al. (1928, p. 42) described “thick layers” and


a “large heap” of shells associated
with hearths. At Grotta Moscerini in Italy, Stiner


(1994, pp. 181–184) found that a
significant percentage of the marine shells was


burned, suggesting that they too were
deliberately collected by Neandertals. Other


Old World examples include many of the
freshwater shell middens of Willandra


Lakes in Australia and the Pleistocene
middens of Melanesia (Allen et al., 1989a,b;


Wickler and Spriggs, 1988). In the New
World, there seems to be little question

about the predominantly cultural origin
of the aquatic fauna found at most of the


open air middens along the Pacific
Coast. Early components at Daisy Cave, Broken


Mammoth, and Lewisville also seem
relatively secure.


The Distribution
of Early Coastal Localities


Even allowing for such
uncertainties about the origin of aquatic faunal


remains—often even more serious for
the remains of terrestrial fauna found at


early sites—a significant number of
Paleolithic localities with secure evidence for


systematic early aquatic resource use
are now relatively well documented. The

spatial and temporal distribution of
these early sites, particularly the coastal ex-


amples, is of special interest. Yesner
(1987, 1998, p. 205) suggested, for instance,


that such sites are exceptional and are
located in areas of upwelling and unusu-


ally high marine productivity. Thus
such early coastal sites are often viewed as


rare examples of relatively intensive
aquatic resource use in a Pleistocene world


otherwise dominated by terrestrial
economies.


322
Erlandson


This association holds for some
early coastal localities, but it does not explain

the evidence for early marine resource
use at several early Mediterranean sites


in Italy, Lebanon, Libya, and Algeria
(see Klein and Scott, 1986; McBurney,


1967; Stiner, 1994), where marine
productivity is comparatively low by global


standards. My own comparison of the
distribution of early coastal sites leads to a


different conclusion. While a number of
early sites are found in areas of intense


upwelling (Peru, California, Gibraltar,
etc.), many others are not. Comparing the


distribution of coastal sites to
various physical and biological characteristics in


an atlas of the world’s oceans
(Couper, 1989), I found no clear correlation with


intensive marine upwelling, exceptional
primary (phytoplankton) or secondary

(zooplankton) productivity, sea
temperature, salinity, latitude, tidal range, tectonics


or vulcanism, marine habitat, or
terrestrial habitat. In fact, relatively early sites are


found in areas of coral reefs,
temperate seas, and even arctic or subarctic coasts


(by 8,000–10,000 years ago). They are
found adjacent to tundra environments,


boreal forests, savanna, chaparral, and
hyperarid landscapes, including some where


contemporary interior sites contain
relatively abundant remains of large terrestrial


game.


I found only one trait that seems
to link the early coastal localities: steep

bathymetry. From California to Florida
and from Melanesia to the Mediterranean,


all the early sites are located along
relatively steep shorelines where the offshore


topography drops off rapidly. The
opposite also holds true, with areas of broad and


shallow continental shelves generally
producing only relatively recent evidence


for marine resource use, regardless of
the intensity of marine upwelling. This


is due to the simple fact, clearly
demonstrated by several elegant studies (e.g.,


Parkington, 1981; Shackleton et al.,
1984), that most localities situated along


modern coastlines were far removed from
coastal habitats during most of the last


250,000 years and more. Studies of
historical foragers in coastal habitats have

shown that the skeletal remains of
edible aquatic animals are rarely transported to


residential sites more than about 10 km
from the coast (Bigalke, 1973; Meehan,


1982), except for those that have
ornamental or other utilitarian value. Where


shorelines are steep, however, sites
still preserved above sea level may sometimes


be found within the foraging radius of
ancient coastal habitats. The occupants of


sites located along shallow continental
shelves, on the other hand, may only have


had access to marine resources for the
last 5,000–8,000 years, as local sea levels


and shorelines approached the modern
condition.


This general bathymetric
correlation (which I call Richardson’s Rule)—in

which steep shorelines are associated
with relatively early evidence for marine


resource use, while shallow shelves
yield relatively recent evidence—is a much


stronger predictor of the location of
early coastal sites than upwelling or any of the


other aquatic or terrestrial traits I
examined. Furthermore, Richardson’s Rule helps


explain some puzzling anomalies. It
explains, for instance, why early coastal sites


are much more common along the
generally steep Pacific Coast versus the relatively


The Archaeology of Aquatic Adaptations
323


shallow Atlantic Coast of the United
States. It explains why along the Peruvian


coast, all of which is characterized by
upwelling and high marine productivity,

the earliest coastal sites are
differentially distributed in areas of relatively steep


bathymetry (Richardson, 1998). Finally,
it helps explain why along most of the


Florida coast, where beaches were as
much as 100 km offshore about 14,000 years


ago, the modern shoreline has produced
evidence for maritime adaptations no more


than about 5,000 years old, except for
the steeply plunging shorelines near Miami


where the Cutler Ridge site contains
evidence for marine fishing and other coastal


foraging dated to about 9,600 RYBP.


The correlation between steep
bathymetry and the location of early coastal

sites also seems to contradict two
tenets of traditional theories about maritime


adaptations, (1) that steep bathymetry,
which generally limits the extent of inter-


tidal and nearshore habitats, reduces
the productivity of such marine environments


and renders them relatively
unattractive to humans; and (2) widespread maritime


adaptations only developed in the
Holocene after sea level stabilization led to the


development of relatively broad,
shallow, and productive nearshore habitats. My


analysis of the distribution of early
coastal localities suggests that many coastal


habitats are more productive than
previously envisioned, that Pleistocene mar-


itime adaptations were more widespread
than previously thought, and that the

archaeological record for the antiquity
of coastal adaptations is fundamentally


biased in most parts of the world.


The
Antiquity of Seafaring


Further support for this
viewpoint comes from recent evidence for a relatively


early development of seafaring in
several parts of the world, including evidence


for Pleistocene maritime voyaging in
areas where the oldest coastal shell middens


date to the Holocene. For decades, the
idea that our Pleistocene ancestors may


have made substantial migrations by
boat suffered from the same theories that

marginalized maritime adaptations in
general and argued that our ancestors were


relatively unsophisticated
technologically. There is little question that hominids


must have crossed rivers and other
short water barriers in spreading out of Africa


and through Eurasia. Prior to 1980,
however, there was virtual unanimity that boats


were a very recent addition to human
technologies (e.g., Bass, 1972; Greenhill,


1976; Johnstone, 1980, p. xv). Due to
preservation problems, evidence for the


earliest use of boats—as opposed to
simple logs or floats that allowed hominids


to cross small water barriers while
partly submerged—remains lost in obscurity,


depending primarily on indirect
evidence for the colonization of island groups

(Table III). Except for the
long-distance voyaging evident among Austronesian


and other peoples in the last 5,000
years or so, such evidence requires the presence


of not-too-distant islands that have
been separated from continental land masses


in recent geological times, criteria
many regions of the world cannot meet.


324
Erlandson


Archaeologists have long argued
inconclusively both for and against the idea


that Homo erectus was capable of making
the relatively short crossing (<20 km)


of the Straits of Gibraltar from
Morocco to the Iberian Peninsula (see Cachel

and Harris, 1998; Rolland, 1998). As
evidence accumulates for a relatively long


isolation of Neandertals in western
Europe (e.g., Krings et al., 1997), however,


it seems increasingly unlikely that
archaic Homo sapiens had the capability to


routinely cross the potentially
hazardous Straits of Gibraltar. Elsewhere in the


Mediterranean, there is limited but
more convincing evidence for occasional island


exploration by Neandertals (Cherry,
1990). For Southeast Asia, recent evidence


may indicate that Homo erectus reached
the Indonesian island of Flores as much as


700,000–800,000 years ago (Morwood et
al., 1998, 1999; Sondaar et al., 1994), and


Bednarik (1998) and Bednarik et al.
(1999) proposed that relatively sophisticated

seafaring and maritime adaptations date
back a million years or more. So far,


however, there is little evidence for
any systematic use of seaworthy watercraft


by Homo erectus or archaic Homo
sapiens, and their voyaging capabilities appear


more likely to have been relatively
rudimentary.


Evidence for Pleistocene
seafaring by anatomically modern humans is much


more compelling and more widespread,
involving the dispersal of hominids across


a number of unequivocal and substantial
water barriers (Clark, 1991; Erlandson,


in press; Irwin, 1992). Evidence for
systematic and sophisticated Pleistocene voy-

aging comes primarily from eastern
Asia, Australia, and Melanesia, where voy-


ages in excess of 20–200 km have now
been widely documented between at least


50,000 and 15,000 years ago. The proof
that seafaring extended well back into


the Pleistocene requires a fundamental
paradigm shift, not yet fully realized, since


maritime voyaging was once thought to
be strictly a Holocene phenomenon. By


the 1970s, terminal Pleistocene
seafaring had been documented by the presence


of obsidian from the Mediterranean
island of Melos in strata at Franchthi Cave in


mainland Greece dated to as early as
13,000 RYBP (Cherry, 1990). The antiquity


of seafaring was extended with the
discovery that humans had reached Australia

by 20,000 years ago (Lampert, 1971), a
date rapidly pushed back to 33,000 years


ago (Bowler et al., 1970), 40,000 years
ago (Groube et al., 1986), and now as much


as 50,000–60,000 years ago (Roberts
et al., 1990; Thorne et al., 1999). Regardless


of the route chosen, colonization of
New Guinea and Australia required several


separate sea crossings, including
voyages of at least 80 km (Clark, 1991; Irwin,


1992). As a result, the colonization of
Australia is now widely viewed as the ear-


liest evidence for planned maritime
voyaging in human history and possibly some


of the earliest evidence for
anatomically modern human behavior anywhere in the


world (Davidson and Noble, 1992).

For a time, two puzzling facts
allowed some scholars to believe the Pleistocene


colonization of Australia may have been
accomplished by accident. First, in historic


times Australian Aborigines reportedly
had no sophisticated watercraft capable of


making substantial sea crossings
(Flood, 1990, p. 36), which raised questions


about their ability to travel through
island Southeast Asia by boat. Like much of


The Archaeology of Aquatic Adaptations
325


the rest of the world, Australia also
had no true coastal shell middens or other


direct evidence for maritime
adaptations dating to the Pleistocene. In fact, the vast

majority of such sites were less than
about 5,000–6,000 years old.


With the discovery in the late
1980s of several Pleistocene shell middens in the


Bismarck Archipelago and the Solomon
Islands in western Melanesia (see Allen


et al., 1989a,b; Wickler and Spriggs,
1988), any doubts about the role of deliberate


maritime voyaging in the peopling of
Australia essentially vanished. Settlement


of these islands, now dated to at least
35,000 RYBP (Allen and Kershaw, 1996,


p. 185), added several significant
maritime crossings to those already required to


reach Australia and New Guinea. More
importantly, these islands contain rela-

tively impoverished terrestrial flora
and fauna, and the sites themselves contained


the marine shellfish, fish, and other
remains expected of a maritime people. The


Melanesian evidence also suggests that
maritime voyaging capabilities improved


significantly between about 35,000 and
15,000 years ago. While the initial settle-


ment of New Guinea, New Britain, and
New Ireland required voyages of up to


100 km, colonization of Buka in the
Solomon Islands at least 28,000 years ago re-


quired a minimum sea voyage of 140 km
and possibly 175 km (Irwin, 1992, p. 20).


By 15,000 years ago, moreover,
Melanesian seafarers had reached Manus Island in


the Admiralty group, which required an
uninterrupted voyage of 200–220 km, 60–

90 km of which would have been
completely out of sight of land (Irwin, 1992, p. 21).


Further evidence for Pleistocene
seafaring comes from the islands of Japan.


Japan itself was connected to the Asian
mainland during periods of very low sea


level, so its settlement did not
necessarily require boats. Fagan (1990, p. 191) ar-


gued, however, that new blade and
edge-grinding technologies introduced about


30,000 years ago when Japan was
separated from the mainland probably involved


maritime contacts. This idea may be
supported by the discovery of human bones


beneath a charcoal-rich stratum in
Yamashita-cho Cave on Okinawa dated to

about 32,100 RYBP (Matsu’ura, 1996,
p. 186). Human remains dated between


about 15,000 and 26,000 RYBP also have
been found in several other limestone


caves on Okinawa and the smaller
islands of the Ryukyu chain (Matsu’ura, 1996),


which stretches southward from Japan
nearly to Taiwan. At Pinza-abu Cave on


Miyako Island, human remains found
below a calcareous flowstone were associ-


ated with charcoal dated to about
26,000 RYBP (Matsu’ura, 1996, p. 187). Given


the bathymetry of the Ryukyu Islands,
several sea voyages would have been re-


quired to reach Okinawa from Japan,
including a crossing about 75 km long.


Reaching Miyako Island, from either
Japan or Taiwan, would have required even

longer voyages of up to 150 km. In
Japan itself, archaeological evidence suggests


that by about 21,000 RYBP, maritime
peoples from Honshu were using boats to ob-


tain obsidian from Kozushima Island
approximately 50 km offshore (Oda, 1990).


Similar to Australia, despite
considerable evidence for Pleistocene seafaring, the


oldest shell middens in Japan date to
the Holocene (Aikens and Akazawa, 1996,


p. 224; see also Aikens and Higuchi,
1982). It seems likely, therefore, that earlier


coastal sites have been submerged by
rising sea levels.


326
Erlandson


The evidence for Pleistocene
seafaring in Japan is also significant because

it places competent mariners in the
cool waters and boreal climates of the North


Pacific at a date early enough to have
contributed to the initial colonization of the


Americas (Engelbrecht and Seyfert,
1995; Erlandson, 1994, in press). From Japan,


the Kurile Islands stretch to the
northeast like stepping stones to the Kamchatka


Peninsula and the southern shores of
Beringia. With the now well-documented


Pleistocene seafaring capabilities of
Homo sapiens sapiens, the presence of Pleis-


tocene seafarers in Japan, and the
geography of the North Pacific, maritime peoples


appear to have had the capabilities to
follow a coastal pathway to the Americas.


Whether they made such a journey is
still unknown, and the evidence that could

resolve the issue—like so many
questions related to the evolution of maritime


adaptations—lies largely unstudied
and submerged on the continental shelves of


the North Pacific.


Other Evidence for
Early Aquatic Adaptations


Two other sources of data need to
be considered in any comprehensive eval-


uation of the antiquity of aquatic
adaptations: the archaeological record from sub-


merged or “drowned” terrestrial
sites, and the nature of pericoastal sites that show


limited evidence for marine or
estuarine resource use. Although a detailed exam-

ination of either topic is beyond the
scope of this paper, it would be a mistake not


to consider such evidence at all.


Submerged
Terrestrial Sites


Scholars have long known that
human occupation sites lie submerged on con-


tinental shelves around the world (see
Negris, 1904; Smyth, 1854) and that these


may fundamentally bias our
understanding of the development of aquatic adapta-


tions (e.g., Emery and Edwards, 1966;
Flemming, 1998, p. 130; Kraft et al., 1983;


Richardson, 1981; Shepard, 1964). What
to do with such knowledge, however,

raises fundamental problems. We must be
governed by some rules of evidence,


after all, and simply assuming that
ancient shell middens lie submerged off coast-


lines around the world and that coastal
adaptations have always been a part of


the human story leaves many scholars
with a very uncomfortable feeling. On the


other hand, assuming that the
archaeological record is representative in the face of


clear evidence to the contrary is
equally problematic. The obvious solution is to


examine submerged coastal landscapes
for the presence or absence of submerged


shell middens or other evidence for
early aquatic resource use.


Unfortunately, this is not as
easy as it sounds, and such programs have been

limited so far. During marine
transgressions, the submergence of most terrestrial


sites would have been accompanied by
their essential destruction. Just as it is


destroying countless coastal sites
around the world today, wave erosion would


The Archaeology of Aquatic Adaptations
327


have redeposited most older sites into
the intertidal zone, leaving only lag deposits


on wave-cut platforms such as the Lower
Paleolithic localities documented on Old


World marine terraces. Along the
predominantly erosional outer coasts around


the world, intact submerged sites would
be preserved only in special situations


where local landforms are protected by
offshore islands, archaeological deposits

are cemented or sealed under
erosion-resistant strata, or earthquakes caused a rapid


subsidence of sites into the intertidal
or subtidal zone. In estuarine or lacustrine


settings, where wave energy and
shoreline erosion are generally less severe, the


potential for the preservation of
submerged sites is considerably better (Flemming,


1983, 1998). Even in these settings,
however, relatively little archaeological work


has been accomplished on submerged
terrestrial sites (but see Fischer, 1995b;


Masters and Flemming, 1983). Due to
technological and financial constraints,


moreover, the work that has been done
has been limited primarily to sites found


comparatively close to shore and in
relatively shallow water. Because sea levels

have risen up to 150 m in the past
17,000 years, these limitations have so far


prevented effective undersea
reconnaissance along shorelines more than about


10,000–12,000 years old.


Nonetheless, impressive numbers
of submerged coastal sites have been found,


and the number of sites is rapidly
growing (see Fischer, 1995b; Flemming, 1998).


In a recent summary, Flemming (1998, p.
129) noted that roughly 550 submerged


human occupation sites (SHOS) have been
located in coastal settings around the


world, about 100 of which are older
than 3,000 years. These include Acheulian

hand axes found in sediments underlying
a historical shipwreck 8 m below sea level


off the South African coast and
thousands of Mousterian artifacts eroding from a


creek bank submerged 18 m below sea
level near Cherbourg on the Atlantic coast


of France (Flemming, 1998, pp.
135–136). Elsewhere, numerous submerged sites


dating to the middle and early Holocene
are now known, and artifacts of Pleistocene


age also have been recovered from the
sea floor (e.g., Dunbar et al., 1992; Faught


et al., 1992; Fedje and Christenson,
1999; Flemming, 1983, 1998; Sanger, 1995;


Stright, 1990). In a number of
protected coastal areas, submerged sites that contain


evidence for aquatic adaptations have
been found. Some of these submerged sites

feature remarkable preservation, as
shown by the intact structural remains, human


burials, canoes, canoe paddles,
hearths, and other materials recovered from early


Holocene sites such as Tybrind Vig in
Denmark (Andersen, 1985, 1987) and Newe


Yam off the coast of Israel (Raban,
1983; Wreschner, 1983).


One of the earliest submerged
sites, and surely one of the most remarkable,


is the Upper Paleolithic Cosquer Cave
discovered in 1991 on the Mediterranean


coast of France (Clottes et al., 1992;
Clottes and Courtin, 1996). Cosquer Cave is a


partly submerged limestone cavern, the
small mouth of which lies 37 m below sea

level. A narrow and gradually rising
shaft extends for approximately 140 m before


opening into a large cavern, only parts
of which remain above sea level. Over 250


engraved or painted motifs have been
documented in the unflooded remnants of the


cavern, and radiocarbon dates indicate
that these were executed primarily during


328
Erlandson


two periods about 27,000 and 18,500
RYBP (Clottes and Courtin, 1996). Upper


Paleolithic artistic representations of
marine animals are rare in Europe, but at


Cosquer they make up about 12% of the
animal images and include depictions of


seals, the great auk (Pinguinus
impennis), and possibly fish and jellyfish. Although

the cave appears to have been located
more than 10 km from the sea during the


height of the Last Glacial, these
images testify to the significance of marine animals


to the artists. As Clottes and Courtin
(1996, pp. 44–45) noted, several Upper


Paleolithic skeletons excavated in the
late 1800s from the Grimaldi Caves about


150 km to the east were associated with
hundreds of marine shell ornaments, also


testifying to the symbolic importance
of the sea among Upper Paleolithic people


of the area.


Clearly, submerged terrestrial
sites do exist and may be preserved under the

right conditions. The questions that
remain are whether such submerged coastal


sites represent the proverbial tip of
the iceberg or isolated cases, whether even


earlier coastal sites lie offshore in
deeper water, and whether such sites can be


found and sampled to help unravel some
of the mysteries that remain about the


role of the sea in human history.



Pericoastal Sites


Like Cosquer Cave and the
Grimaldi Caves, there are scores of Pleistocene


sites around the world that are located
in pericoastal or even interior settings

that lack dense accumulations of
aquatic food remains, but nonetheless testify


to linkages to aquatic habitats. These
include numerous coastal sites listed in


Table I, which at various times in
their occupational histories appear to have been


located some distance from the coast.
In a number of well-dated sites with detailed


paleogeographic reconstructions, these
periods seem to correlate with reduced


densities of marine food remains, the
presence of strictly ornamental or utilitarian


objects (shell beads, baler shells,
etc.), a complete reliance on terrestrial foods, or


site abandonment. For a number of other
sites, less well documented or located


further from aquatic habitats, such
relationships are not as clear. Sites that contain

small amounts of aquatic food remains
can be viewed as evidence that aquatic


resources were relatively unimportant,
as part of a seasonal round that included


residential periods on the coast, of
exchange with more maritime people living


closer to the coast, or some
combination of such inferences. A similar range of


arguments has been made for interior
sites in Upper Paleolithic Europe where


marine shell ornaments are found far
from the coast. Such sites clearly indicate


some economic or ritualized use of
aquatic resources, but their significance can be


argued depending on the theoretical
stance of individual investigators.


In this regard, I find the
Australian case particularly compelling, where sev-

eral pericoastal sites (Mandu Mandu,
Shark Bay, etc.) more than 30,000 years


old contain small numbers of shells,
often ornamental or utilitarian types traded


The Archaeology of Aquatic Adaptations
329


between interior and coastal groups
ethnographically. It is currently impossible to


know for certain whether these shells
are evidence that early terrestrially adapted


Australians occasionally visited the
coast, that coastal people occasionally vis-


ited the interior, or that they
represent the exchange of goods between discrete


groups residing separately in coastal
and interior areas. How can a continent like


Australia, which we now know was
colonized by boat at least 50,000 years ago,

have so few early coastal sites? Did
maritime peoples reach Australia then aban-


don its coastlines in favor of more
productive terrestrial habitats and resources


for tens of thousands of years? If so,
how do we account for the evidence for the


systematic use of shellfish and fish
in the Willandra Lakes area at least 40,000


years ago? Why do we see further
evidence for maritime voyaging into western


Melanesia by 35,000–40,000 years ago?
To me, it seems most likely that the early


pericoastal sites in Australia are the
remnants of inland settlement and subsistence


by coastally adapted Pleistocene
peoples whose main settlements were submerged


or destroyed by rising postglacial
seas.


DISCUSSION


To some, none of the individual
lines of evidence I have examined may provide


a compelling argument for the early
development of aquatic adaptations. When


the theoretical and methodological
issues I have raised are combined with current


knowledge about world sea levels and
coastal paleogeography, as well as a variety


of lines of archaeological evidence, I
believe there are compelling reasons to doubt


the veracity of the current consensus
model. Comparatively speaking, it is still true


that there is only limited evidence for
the intensive use of aquatic resources prior

to the end of the Pleistocene. This is
not surprising in the New World, which


appears to have been colonized near the
end of the Pleistocene when sea levels


were much lower than they are today.
Even on a global scale, however, it is not clear


that this pattern accurately reflects
changes in human subsistence through time.


To understand the history of aquatic
adaptations, globally or in any particular


region, we must first determine if the
patterns evident in the archaeological record


result from actual changes in human
behavior, patterns imposed by geological


or taphonomic forces, or the recovery
and analytical methods of archaeologists


themselves. Unfortunately, except for
areas first occupied by humans relatively

recently (i.e., Polynesia), such
evaluations are fraught with difficulty and have


rarely been done in a manner that
inspires confidence in the results.


A New
World Test Case


It is possible, however, to
return to some of the fundamental tenets of recent


theory about aquatic adaptations and
examine them in light of current archaeolog-


ical data from the New World. If
shellfish and other aquatic foods are generally


330
Erlandson


smaller and less productive than
terrestrial alternatives—and their systematic use

reflects demographic pressure,
resource stress, or economic intensification—then


the antiquity of coastal adaptations in
the Americas should provide an excellent test


case. Traditional theory, including
many recent applications of diet breadth-prey


ranking models, suggests that until
population growth produced sufficient de-


mographic pressure to reduce the
productivity of “high-ranked” terrestrial game,


aquatic resources would not be
systematically used. Thus the relatively recent peo-


pling of two vast continents with
highly diverse and productive terrestrial land-


scapes, especially by small
hunter-gatherer groups generally thought to have mi-


grated through an interior route into
the heart of North America, should show little

evidence for marine resource use or
even coastal settlement until quite recently. If


aquatic resources are not necessarily
marginal, then we should find relatively early


evidence for their use, at least in
areas where the economics of aquatic resource


use compare favorably to those
available in adjacent terrestrial habitats.


To adequately evaluate the
evidence, of course, we need to know when humans


first settled the Americas and whether
the archaeological record is representative


of the full range of early adaptations,
especially in coastal zones. At present, we


cannot answer either question with any
certainty. In 1977, Osborn used a regional

analysis of the Peruvian archaeological
record to argue that marine resources were


inferior in the arid western slopes of
the Andes. When Osborn (1977a) evaluated


the Peruvian evidence, he believed
there was a lag between the earliest interior ver-


sus coastal occupation equal to more
than half of the total cultural sequence for the


region, well over 12,000 years. This
apparent gap was explained by proposing that


the earliest occupants of the region
did not systematically use marine resources until


their population densities had
effectively reached the carrying capacity of the terres-


trial environment. Thus the Andean
coast—one of the richest marine environments


on earth—was characterized as an
environment marginal for human occupation.

In retrospect, we now know
Osborn’s analysis had significant problems (see


Erlandson, 1988; Perlman, 1980; Quilter
and Stocker, 1983; Yesner, 1980). First,


he assumed the Andean archaeological
record was representative, based on the now


disproved claim that tectonic uplift of
the Peruvian landscape had outpaced sea


level rise since the last glacial.
Second, he assumed that the regional demographic


clock began with initial human
occupation of the Andean uplands at least 23,000


years ago, based on claims by MacNeish
(1971) for the presence of “artifacts”


(made from the same rock as the cave
walls) in the lower levels of Pikimachay

(Flea) Cave. Even today, with the
“Clovis barrier” seemingly broken, few scholars


accept the dubious evidence for
pre-Clovis occupation at Flea Cave (e.g., Dixon,


1999, pp. 100–101).


Today, the earliest widely
accepted (there are still doubters) Andean archaeo-


logical site is Monte Verde (Dillehay,
1997), which appears to date to about 12,500


RYBP. Monte Verde is located in a
valley roughly 30 km from the coast of Chile.


While there is evidence that big game
was hunted from the site, there is also ev-


idence for a relatively eclectic
economy in which plants and smaller resources

The Archaeology of Aquatic Adaptations
331


played a significant role. The
presence of seaweed also suggests that the site oc-


cupants had links to the coast. Recent
research also has pushed back the earliest


occupation of the Andean coast to
approximately 11,200–10,500 RYBP (Keefer


et al., 1998; Richardson, 1998;
Sandweiss et al., 1998), and the earliest site, Que-


brada Jaguay, shows interior links
(Sandweiss et al., 1998) that may represent


the opposite end of a seasonal
subsistence round represented at Monte Verde. If


we accept that Monte Verde is one of
the earliest sites in the Andes (which seems


likely) and assume that its occupants
had no coastal neighbors and used few aquatic

resources themselves (which seems less
likely), the age differential between the


earliest coastal versus interior
settlements has withered to less than 2,000 years.


This might be enough time for
terrestrial foragers to shift to a partly littoral or


maritime focus, but it seems highly
unlikely that a shift to marine resources about


11,000 years ago was due to demographic
pressure on the vast Andean landscape.


In North America, the situation
is very similar, with increasing evidence


for early coastal settlement and use of
marine and other aquatic resources. The


presence of people on California’s
Channel Islands 10,500–11,000 years ago

(Erlandson et al., 1996; Johnson et
al., 2000; Orr, 1968), for instance, is extremely


difficult to account for as a response
to human pressure on the highly produc-


tive and diverse terrestrial resources
of the adjacent mainland. The earliest people


of the Channel Islands, moreover, seem
to have subsisted primarily on shellfish,


small fish, plant foods, and
occasional seals or sea lions. There is currently no evi-


dence that they were big-game hunters
drawn to the islands by pygmy mammoths


or massive elephant seals. Rather, they
seem to have been eclectic foragers who


relied on a variety of resources,
including abalones, mussels, small turban snails,


and a variety of small fish. The very
presence of such people on the islands at

this early date suggests that shellfish,
fish, and other marine resources were highly


ranked, highly regarded, and highly
relied on. Along the Atlantic and Gulf coasts


of North America, the evidence for
early coastal adaptations is neither as early nor


as widespread, but in these areas the
presence of submerged sites and Paleoindian


artifacts on the continental shelves
indicates that we are missing early components


of the coastal archaeological record.


To me, the available data suggest
that marine and other aquatic resources


were an integral part of many early New
World economies, that their signifi-

cance has been underemphasized in
previous models, and that their presence in


archaeological sites does not
necessarily indicate the existence of environmental


deterioration, population pressure,
resource stress, or economic intensification. I


am not suggesting—as many have done
for large land mammals and other ter-


restrial resources—that aquatic
resources were universally productive. Nor am I


arguing that the use of aquatic
resources was not, at times, associated with resource


stress and economic intensification.
What I am suggesting is that, in the New World


and the Old World, the factors that
govern human decisions about what resources


will be used, when, and by whom are
highly complex and situational. Recognizing

this complexity, the diversity of
environments (terrestrial and aquatic) encountered


332
Erlandson


by our ancestors, and the flexibility
and opportunism of hunter-gatherers, I believe


global or universal statements about
the productivity of aquatic resources do not do


justice to the diversity and complexity
that we should expect of the archaeological


record.


SUMMARY AND
CONCLUSIONS


I began this paper by suggesting
that general conceptions of the history and

nature of aquatic adaptations have
marginalized the study of coastal, riverine, and


lacustrine societies, relegating them
to the last 1% of human history. The view that


aquatic resources are marginal and that
aquatic adaptations developed relatively


recently renders their study
essentially peripheral to many of the most compelling


issues addressed by archaeologists:
human evolution, early hominid migrations,


the appearance of anatomically modern
humans, peopling of the Americas, the


development of agriculture, the rise of
civilization, and others. I also argued that


a variety of taphonomic processes,
epistemological issues, methodological prob-


lems, and data gaps raise serious
questions about assertions that widespread and

systematic aquatic adaptations
developed only since the end of the last glacial.


Specifically, I suggested that


1. postglacial sea level rise has
submerged most of the shorelines older than


about 10,000 years along which
the evidence for earlier coastal occupa-


tions would logically be
found;


2. differential preservation,
recovery, or reporting of site constituents has


selectively underemphasized
the importance of aquatic resource use in

archaeological sites around
the world;


3. traditional models of
hunter-gatherer behavior have overemphasized the


role of hunting in general,
and large land mammal hunting in particular,


in many ancient societies;


4. normative cultural ecological
reconstructions have too often treated hu-


man societies as aggregations
of generic individuals rather than groups of

diverse people (men, women,
and children; young and old, rich and poor,


etc.) who often were engaged
in different activities;


5. prior to the development of
relatively effective hunting technologies, ho-


minids relied to a significant
extent on scavenging behavior that would have


increased the relative
productivity of and reliance on aquatic resources that


required no specialized
technologies to obtain or process;

6. our hominid ancestors have
always, with rare exceptions dictated by un-


usual environmental
conditions, been highly opportunistic and relatively


eclectic omnivores, an
economic orientation fundamental to our extraor-


dinary success in colonizing
virtually every habitable land and seascape


on earth.


The Archaeology of Aquatic Adaptations
333

With these issues in mind, I
reviewed the evidence for early aquatic resource


use in archaeological sites around the
world, focusing on Old World sites dating


earlier than about 15,000 years ago and
New World sites more than 8,000 years


old. I concluded that a variety of
unresolved problems continue to prevent us


from determining when aquatic
adaptations developed, how widespread they were,


and how important they were in the
broad scheme of human evolution. Some


of the earliest archaeological
localities associated with Homo habilis and Homo


erectus in Africa contain the remains
of aquatic or amphibious animals such as

fish, crocodiles, molluscs, and
hippos, as do some early European sites associated


with late Homo erectus or early archaic
Homo sapiens populations. Although the


distribution of fish and other aquatic
remains in some of these early sites coincides


closely with artifacts and other faunal
remains, the cultural origin of the faunal


remains (aquatic and terrestrial) and
their nature (scavenged or hunted) are difficult


to prove. Less equivocal evidence for
the use of shellfish by Homo erectus and


archaic Homo sapiens also is found at
several Old World sites. There is no doubt


that these hominids occupied coastal
and other aquatic habitats and little reason to


doubt that aquatic resources were used
by them at least occasionally. At present,

the intensity of such use remains
unknown, just as the overall nature of Lower


Paleolithic subsistence remains largely
obscure.


Hominids clearly crossed aquatic
hurdles in spreading out of Africa and


through much of Eurasia, indicating
that rivers and even some straits were not


necessarily the physical or
psychological barriers sometimes imagined. Prior to


the appearance of anatomically modern
humans, however, the use of aquatic re-


sources may have been limited largely
to shellfish and occasional “low-tech” uses


of fish, birds, mammals, and other
resources that could be collected without spe-

cialized technologies. Only with the
appearance of anatomically modern humans


do we find the evidence for a more
intensive use of shellfish and a wider range


of marine or aquatic resources. Not
surprisingly, this economic diversification co-


incides with the first evidence for
the development of a number of “modern” or


transitional technologies, including
the earliest relatively intensive use of chipped


stone blade and geometric or
microlithic industries, the first formal bone tools, the


earliest widespread evidence for the
use of red ochre, and probably the first use of


relatively sophisticated boats. In the
context of such transitional Middle Stone Age


technologies at Klasies River Mouth
caves, Die Kelders, and other Last Interglacial

localities in South Africa, for
instance, AMH appear to have regularly eaten a va-


riety of shellfish, marine mammals,
and flightless birds (Klein and Cruz-Uribe,


2000), although some or all of the
larger vertebrates may have been scavenged


rather than hunted (Binford, 1984).
From the Semliki River area in Zaire comes


the earliest evidence for complex
aquatic hunting gear, ∼80,000-year-old barbed


harpoons from Katanda associated with
numerous large fish remains (Yellen et al.,


1995). From Blombos Cave in South
Africa comes evidence for Middle Stone Age


marine fishing, probably dating to
over 60,000 years (Henshilwood and Sealy,


334
Erlandson

1997). And from the Boegeberg 2 shell
midden comes possible evidence for the


relatively intensive MSA use of
cormorants (Klein, 1999, p. 456).


Dated to about 90,000 years ago
are the earliest skeletal remains of Homo


sapiens sapiens found outside of
Africa—the Qafzeh and Skhul skeletons from


coastal Israel—suggesting that early
modern populations had begun to move out


of Africa by this time. Although
anatomically modern humans do not appear to


have moved into most of Europe for
another 50,000 years (Klein, 1998), current


evidence suggests that they spread into
southern Asia at least 60,000 years ago.

From there, within just a few
millennia, they probably made the multiple maritime


voyages through island Southeast Asia
required to settle Australia and New Guinea.


By about 30,000–35,000 years ago,
seafaring AMH peoples also had colonized


western Melanesia and the Ryukyu
Islands south of Japan.


Although the current state of our
knowledge remains somewhat fluid, the


earliest subsistence strategies that
included relatively eclectic and intensive use


of marine or other aquatic resources
may be associated with the appearance of


anatomically modern humans. When such
aquatic adaptations were combined with

the exploitation of a range of
terrestrial plants and animals, a more diversified and


stable resource base would have
resulted. Such economies may have contributed


significantly to the development of
greater sedentism (see Kelly, 1995, p. 125),


to the reproductive success of Homo
sapiens sapiens, and to our apparently dra-


matic demographic and geographic
expansion over the last 150,000 years. In this


regard, it is worth noting that current
(“Out of Africa”) models for the rapid spread


of anatomically modern humans allow
only about 10% of the time available in


multiregional models for this
demographic and geographic expansion (Erlandson,


in press). Wherever anatomically modern
humans first appear, they seem to have

carried with them a penchant for art
and symbolism, technological innovation, and


complex problem-solving and
communication skills (see Davidson and Noble,


1992; Klein, 1998; Mellars, 1998).
Aquatic adaptations and Pleistocene seafar-


ing played a more significant role
than previously supposed in the demographic


expansion, the geographic spread, and
the phenomenal success of our species.


The available archaeological
evidence contradicts aspects of both Gates of


Hell and Garden of Eden models, with
the most likely scenario falling—like


Aristotle’s “golden mean”—somewhere
in between. Despite a number of cate-

gorical statements to the contrary, we
simply do not know when aquatic resources


were first widely used by our hominid
ancestors or how important they were in hu-


man evolution. However, it makes no
sense that hominids would have completely


ignored aquatic resources for more than
2 million years. As long as scavenging


was a significant hominid pursuit, in
fact, it seems likely that aquatic resources


found in shallow water or on the shore
would have been utilized when reason-


ably abundant and available without
complex technologies. There undoubtedly


has been some intensification of
aquatic resource use during human history, but


it also seems likely that our ancestors
used such resources opportunistically and

situationally whenever and wherever it
made economic sense to do so. If aquatic


The Archaeology of Aquatic Adaptations
335


resources sometimes compare favorably
to terrestrial subsistence alternatives, it


raises significant questions about the
emblematic role shell middens have played


as anthropological indicators of the
broad spectrum revolution and postglacial


economies (see Bailey, 1978). If shell
middens are not diagnostic of postglacial


economies, is the broad spectrum
revolution still a revolution?


We cannot afford to ignore the
fact, however, that the efficient or intensive

exploitation of many types of marine
and other aquatic (and terrestrial) resources


requires relatively complex
technologies (e.g., sophisticated boats, nets, harpoons,


hook-and-line) that currently appear to
have been beyond the capabilities of ho-


minids other than anatomically modern
humans. We should also recognize that


aquatic habitats are extremely
variable, that they are juxtaposed with equally var-


ied terrestrial habitats, and that
together these offer such a diverse range of envi-


ronments that they defy broad
generalization (Erlandson, 1994, p. 278; Perlman,


1980). Given the nearly endless
diversity in the relative productivity and accessi-


bility of aquatic versus terrestrial
habitats around the world, it seems likely that

the antiquity and intensity of aquatic
adaptations varied widely through both space


and time. Today, rather than searching
for general rules of human behavior in


aquatic settings, we should be working
to overcome the taphonomic problems


that currently inhibit our
interpretations so we can more effectively document the


diversity of aquatic adaptations. More
interesting questions should take the place


of dichotomized debates about whether
the world’s aquatic habitats were Gardens


of Eden or Gates of Hell. Once we
recognize the diversity of aquatic habitats


through space and time, as well as the
almost limitless combinations of mosaic en-


vironments that result from juxtaposing
such aquatic habitats with equally diverse

terrestrial habitats, we can focus on
the complexity of human responses to aquatic


environments that took place as our
ancestors developed increasingly sophisticated


subsistence strategies on both land and
in the water. Given this diversity and the


innumerable adaptive responses possible
under various intellectual, technologi-


cal, demographic, and sociopolitical
circumstances, a search for a global model or


universal laws of aquatic adaptations
is almost certainly fruitless.


As we move into the twenty-first
century, I hope we can transcend the simple


models and polarized arguments that
have often characterized scholarly debates

about the evolution of aquatic
adaptations. Surely, as Claassen (1991, p. 275)


has suggested, “it is time to put to
rest the generic clam,” as well as the generic


fish, sea mammal, or coastal forager.
In the last century or so, archaeologists have


made great strides in understanding the
development of maritime and other aquatic


adaptations in human history. As our
studies continue in the next century, there are


numerous issues yet to be resolved and
numerous productive avenues of inquiry to


be studied. To truly understand the
role of the sea (and aquatic habitats) in human


history, however, a number of issues
need to be resolved.


Perhaps the most pressing are
questions related to the antiquity of seafaring

and the search for ancient sites
located along Pleistocene shorelines beneath the


sea. It is time to extend the search
for submerged terrestrial sites to a wider range


336
Erlandson


of shorelines around the world and into
deeper waters to look for coastal sites


dating to the crucial period between
about 60,000 and 10,000 years ago. Utilizing


new technologies, offshore
archaeological survey might be particularly productive


along steeply dipping shorelines of the
Mediterranean, where sites like Cosquer


Cave and submerged caves off Gibraltar
(Waechter and Flemming, 1962) have


been identified in limestone bedrock
where dripstone formations may have helped

to preserve evidence for early coastal
adaptations despite the problems of rising


sea levels and coastal erosion
(Flemming, 1998). Underwater reconnaissance and


excavation work might also be
particularly fruitful off some of the more protected


shorelines of the Japanese archipelago,
where evidence for the Upper Paleolithic


antecedents of the Jomon peoples may
lie submerged. We need to know not only


where such sites are located and when
they were occupied but also whether aquatic


resources played a significant role in
the lives of the occupants and how widespread


such adaptations were.


Critical to evaluating the idea
that anatomically modern humans may have

moved rapidly out of Africa along the
coastlines of southern Asia into Australia


and beyond is a search for early shell
middens or other sites on land, in ar-


eas associated with Last Interglacial
shorelines of East Africa, southern Asia,


and the islands of Southeast Asia.
Considering the spatial distribution of early


coastal sites elsewhere in the world,
such efforts are most likely to succeed if they


are focused along coastal stretches
characterized by relatively steep bathymetry,


where lateral shoreline movements
associated with sea level fluctuations have been


minimized.


Also needed are renewed
excavations at early sites that have already produced

the remains of aquatic resources, with
more sophisticated excavation and analytical


techniques, including fine screen
recovery, flotation, and more critical evaluation


of the origin of aquatic and other
faunal remains. We need more taphonomic and


actualistic studies to help distinguish
between aquatic animal remains of cultural


versus natural origin, work that will
complement the extensive studies that have


been done for terrestrial fauna in
interior areas.


Ultimately, we need more data
from “aquatic” sites of all ages and in all


regions to better document the
variability in aquatic adaptations through space

and time. In the twenty-first century,
the study of maritime peoples and aquatic


adaptations should focus on documenting
the remarkably diverse role that aquatic


resources played in human history as
hominids and humans spread around the


world, from sea to shining sea.



ACKNOWLEDGMENTS


This paper is dedicated to the
memory of J. G. D. Clark, who years ago


recognized the limits of what we could
reasonably say about the development


of aquatic adaptations. I would also
like to recognize the work of Carl Sauer,

The Archaeology of Aquatic Adaptations
337


Alan Osborn, Geoff Bailey, and David
Yesner, scholars whose provocative


views stimulated tremendous progress in
the study of coastal and aquatic adap-


tations. For freely sharing ideas and
information that contributed to my research,


I thank Virginia Butler, James Dunbar,
Anders Fischer, Leland Gilsen, Michael


Glassow, Nina Jablonski, Antoinette
Jerardino, Alan McCartney, Jerry Moore,


Greg Nelson, John Parkington, Geoffrey
Pope, Douglas Price, Mark Raab, Jim


Richardson, Torben Rick, Anna
Roosevelt, Dan Sandweiss, Judith Sealy, Ren´
e

Vellanoweth, Larry Wilcoxon, John
Yellen, and David Yesner. Jim Richardson,


Stephen Nash, and an anonymous reviewer
provided constructive criticisms that


helped me revise an earlier draft. I am
also grateful to Gary Feinman and Douglas


Price for inviting me to write this
paper, for their editorial comments and as-


sistance, and for their patience. Most
of all, however, I am deeply indebted to


Madonna Moss—my wife, colleague, and
best friend—for sharing so many of


my coastal journeys (physical and
intellectual) over the years and for her detailed


comments on an earlier version of this
paper. Only I am responsible, of course, for


the opinions expressed in this paper.


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8-1


Missing





8-2


Missing








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
Pleistocene/Holocene transition (PHT) demonstrates that the


technology and mobility of Prearchaic
hunter-gatherers differed dramatically from later Holocene

foragers, suggesting a hunting-oriented
subsistence. However, meager PHT faunal assemblages


imply a generalized, broad-spectrum
diet. Ethnographic analogy fails to provide a behavioral frame-


work for understanding this discrepancy
because the resource structure of the PHT differed utterly


from the ethnographic present.
Palaeoenvironmental data alone are incapable of retrodicting


ancient diets without an understanding
of foraging costs in extinct resource landscapes. This paper


reviews recent studies using behavioral
ecology as a theoretical framework for simulating foraging


behavior in a PHT resource landscape.
The simulation for Railroad Valley, Nevada, suggests the


explanation for the diversity of
subsistence remains in PHT records lies in different foraging


strategies for men and women, rather
than risk aversion alone. Furthermore, the simulation suggests

that Prearchaic hunter-gatherers
enjoyed a narrower diet breadth than later foragers, prompting


the mobility and technological pro les
evinced in the PHT archaeological record.


Keywords


Great Basin; Pleistocene/Holocene
transition; foraging behavior; simulation; sexual division of labor.


Introduction


The North American Great Basin (Fig. 1)
is renowned for its rich ethnographic record


documenting the ecological
relationships of hunter-gatherers and the arid setting in which


they lived. Informed by ethnographic
analogy, over fty years of archaeological research


has demonstrated the existence of
similar, although variable, ‘Archaic’ lifeways through

much of the Holocene (Jennings 1957,
1964; Willey and Phillips 1958). More problematic


is the pattern characterized by
dramatically different technological organization and site


World
Archaeology Vol. 34(1): 103–130 Archaeology & Evolutionary
Ecology


©
2002 Taylor & Francis Ltd ISSN 0043-8243 print/1470-1375 online



DOI: 10.1080/0043824022013428 7

104 Robert G. Elston and David W.
Zeanah


distribution in the Great Basin during
the Pleistocene–Holocene transition (PHT),


roughly between 11,200 and 8,000 BP.
Understanding the nature of PHT adaptive


strategies is an enduring problem in
Great Basin prehistory (Beck and Jones 1997;


Grayson 1993; Simms 1988; Willig and
Aikens 1988). This task is complicated by PHT


environments utterly unlike any of
historical times, leaving Great Basin archaeologists


without valid ethnographic analogs to
assist interpretation of PHT material culture.


Figure 1 Physiographic Great Basin
showing maximum extent of Pleistocene lakes, existing lakes


(after Benson et al. 1972) and research
areas mentioned in the text and in Fig. 4.


105



Thinking outside the box


The significance of differences
between PHT foragers and later hunter-gatherer s


divides archaeologists. Early
assemblages contain an array of large projectile points


and formal flaked tools resembling
those of Great Plains and Southwestern Palaeoin-


dians, but few ground stone tools,
while middle to late Holocene assemblages contain


smaller points, fewer formal flaked
tools and abundant ground stone. These contrasts

suggest PHT foragers emphasized hunting
more than subsequent hunter-gatherers .


However, lack of PHT megafauna hunting
or butchering sites, or evidence that such


animals survived in the Great Basin
after 11,300 BP (Grayson 1993), obviates special-


ized big-game hunting similar to that
in Palaeoindian models (Madsen 1982; Tuohy


1974). Moreover, direct subsistence
evidence retrieved from coprolites (Eisalt 1997;


Fry 1970, 1976; Napton 1997), and from
a growing number of faunal assemblages (Beck


and Jones 1997; Delacorte 1999; Pinson
1999), reveals a diet that included seeds, fish,


and small animals.


Thus, many archaeologists see in the
PHT the roots of later broad-spectrum hunting

and gathering, using the terms
Palaeoarchaic or Initial Archaic for PHT foragers (Beck


and Jones 1997; Jones and Beck 1999;
Pinson 1999), or eschewing categories altogether


(Madsen 1999; Simms 1988).
Nevertheless, while many aspects of technology and subsist-


ence appearing in the PHT persist
through the later Holocene, we, and others (Basgall


1988; Elston 1982, 1986a; Jennings
1986; Zeanah et al. 1995), emphasize the unique


features of PHT archaeology with the
term Prearchaic.


The problem concerns more than
cultural-historical classi cation, going to the root of


long-held notions about cultural
ecology and Great Basin adaptive change. If Prearchaic

adaptive strategies differ only in
degree from ethnohistoric broad-spectrum adaptations,


how can we explain the dramatically
different technologies and settlement patterns


manifested in the archaeological
record? On the other hand, if we let the archaeology


guide us to an inductive reconstruction
of an extinct, specialized subsistence adaptation,


how do we reconcile subsistence
evidence that seems out of sync with the material


culture?


The dilemma calls for a research
strategy that lets us think ‘outside the box’ imposed


by Great Basin cultural history and
culture ecology. Some aspects of this research strat-

egy concern purely methodological
issues of locating buried PHT deposits in the Great


Basin, developing chronologies,
extracting subsistence data from surface assemblages and


so forth (Beck and Jones 1997; Grayson
1993; Pinson 1999; Willig 1988). However, induc-


tion alone can succeed only in
identifying ancient archaeological patterns unexplainable


by reference to ethnography (O’Connell
and Elston 1999). A theoretically based


approach is required, one more
informative of behavior than induction, and independent


of ethnographic models. We are
presently developing such an approach to be used in


future archaeological tests.


Almost twenty years ago, behavioral
ecology was nominated as a suitable theoretical

approach to the Prearchaic (O’Connell
et al. 1982). In this paper, we describe salient


features of PHT environment and
Prearchaic technological adaptation, discuss develop-


ment of behavioral ecology (BE) as a
research strategy for investigating Prearchaic adap-


tations, examine three applications of
BE and discuss its implications for interpreting


likely hunter-gatherer ecology in PHT
resource landscapes. We concentrate on the inter-


val 10,500–8,000 BP to which most
extant archaeological data pertain.


106 Robert G. Elston and David W.
Zeanah


Environments of the
Pleistocene–Holocene transition


The Great Basin of North America is a
region of internal drainage, broken into many

mountain ranges and intervening valleys
(Fig. 1). Great Basin environments responded


to global climate changes (Fig. 2) of
the PHT, although climatic effects varied from place


to place (Benson 1999; Benson et al.
1995; Madsen 1999, 2000; Thompson et al. 1993).


Plant species continually shifted
range, location and abundance, forming communities


with no modern analogs. High
biodiversity and species turnover (Grayson 2000; Nowak


et al. 1994) suggest a dynamic
landscape in which climax was rarely approached. Never-


theless, some environmental
generalizations are warranted.


After the cold and relatively dry
last glacial maximum (LGM), a warmer and wetter

climate produced shrinking glaciers and
rising valley lakes (Madsen 2000; Thompson et


al. 1993). Between 14,000 BP and 13,000
BP, lakes achieved their maxima (Fig. 1), but


regressed rapidly after 12,000 BP,
perhaps to desiccation. The last records of Pleistocene


megafauna in the Great Basin occur
about 11,300 BP (Grayson 1993). The Younger Dryas


(YD) was an abrupt global return to
glacial climate between 11,200 BP and 10,100 BP,


brie y re lling lakes (Madsen 1999,
2000). The rst well-dated signs of human occupation


(Beck and Jones 1997; Grayson 1993)
appear with its onset.


The early Holocene (EH) began with
an abrupt return to a warming climate, 2–3°C

cooler and moister than present, with
greater seasonality. After the extreme variability of


the late Pleistocene, EH climate was
much more even (Fig. 2). Valley lakes regressed after


the YD, but many basins contained
shallow lakes and marshes until after 8,000 BP


(Grayson 1993; Madsen 1999). In the
northern Great Basin, a cool steppe comprised of


sagebrush, mesophilic shrubs, perennial
forbs and grasses extended from lake/marsh


margins into the surrounding mountain
ranges with stands of juniper, mountain


Figure 2 Late Pleistocene- Early
Holocene global temperatures based on GISP2 data (from Pinson


1999, used with permission).



107


Thinking outside the box


mahogany and riparian aspen (Madsen
1999; Nowak et al. 1994; Rhode 2000). Increasing


desert scrub species in pollen records
re ect the gradual desiccation of basins between


9,000 and 8,000 BP.


The relatively cool, even EH
climate, abundant surface water and complex steppe vege-


tation created productive habitats for
a rich biota of sh, waterfowl and mammals


(Broughton 2000; Grayson 1993, 2000,
Hockett 2000; Livingston 2000). In the northern


Great Basin, many small mammal species
(e.g. pygmy rabbits, woodrats, marmots, pikas)

occurred at lower elevations than
present (Grayson 2000). While early fossil and,


especially, archaeological records of
ungulates are rare, elk and bison infrequently occur


in northern Great Basin sites (Bedwell
1973; Dansie 1987; Jennings 1957). Somewhat


more numerous records of deer, mountain
sheep and antelope suggest these animals were


probably present in about the same
relative proportions throughout the Holocene


(Livingston 1999). To help assess their
abundance in the EH, consider that Great Basin


brushy steppe today supports large
numbers of domestic cattle, wild horses and ungulates


(Burkhardt 1995). Because there is a
direct positive relationship between annual precipi-


tation and plant biomass and animal
productivity (Coe et al. 1976; Zeanah et al. 1995),

grazing capacity in the cool, moist EH
should have been greater than later in the


Holocene. Bio-diversity, which also in
uences grazing capacity, remained high through


the EH (Grayson 2000). For example, a
fossil woodrat midden in the Lahontan Basin


(Nowak et al. 1994) contained many more
taxa of shrubs, forbes and grasses used as forage


by various animals at the PHT than are
now present at the site (Table 1).


Large magnitude, high-frequency
climatic uctuations of the PHT (Fig. 2) have been


implicated in possible reduction of
ungulate populations and food plants, with resulting


stress on human populations (Madsen
1999; Pinson 1999). However, little evidence

supports this contention. If numbers of
radiocarbon dates are a crude measure of human


abundance, Great Basin foragers appear
to have ourished between 10,500 BP and 8,500


BP (Fig. 3).


Table 1 Forage plant taxa (shrubs,
forbes and grasses) in a fossil woodrat nest in the Lahontan Basin


(data from Nowak et al. 1994).



Numbers of forage plant taxa



Present* Fossil** Percentage increase


Antelope 8
5 38.5

Mule deer 10
9 47.4


Mountain sheep 12
8 40.0


Rabbit/hare 13
6 31.6


Ground squirrel 7
6 46.2


Woodrat/marmot 3
2 40.0


Notes


* taxa present in both fossil and
modern ora at site.


** taxa present only in fossil record
or modern ora at higher elevation.


108 Robert G. Elston and David W.
Zeanah

Figure 3 Numbers of Great Basin
radiocarbon dates by 500-year interval (charcoal or organic items


only – no bone or bulk dates).


Prearchaic–Archaic contrasts in
technology, settlement and land use


Prearchaic foragers of the Great Basin
behaved differently from Archaic folks, even


though both employed many of the same
components of material culture, including


cordage, baskets, mats, bags, shnets,
atlatls and darts, woven fur robes, ber sandals, skin


moccasins, milling stones and aked
stone tools (Beck and Jones 1997). In part, we can


understand these behavioral contrasts
by reference to middle-range theory and the econ-


omics of resource procurement in PHT
and later Holocene environments.

People tend to locate themselves in
places most convenient to resources they pursue;


however, environmental change may
create or eliminate resources, or change their abun-


dance or distribution (Binford 1977,
1979; Pinson 1999; Raven and Elston 1989; Zeanah


et al. 1995). During the EH, roughly
similar foraging opportunities pertained from valley


to valley: highly productive shallow
lakes and marshes on the valley oors (Fig. 1), brushy


steppe from shoreline to the tops of
most mountains, riparian woodlands along streams


and juniper woodland with a brushy or
herbaceous understory on ridges. The marsh-


steppe ecotone was convenient to both
marshes and piedmont; hence, a good choice for

bases from which to forage in a variety
of low to mid-elevation habitats (Pinson 1999).


Extensive valley marshes offered
foragers seeds, shoots, pollen, birds and bird eggs, shell-


sh, sh and small mammals. Moreover,
large animals may have found marsh-side forage


and cover attractive. Lower to
mid-elevation steppe and riparian woodland provided


high-quality habitat for ungulates
(Grayson 1993; Jones and Beck 1999) comparable to



109



Thinking outside the box

modern upper montane habitats
(historically best for hunting large game). The most


extensive Prearchaic sites often lie in
what now are poor habitats where later Archaic sites


are rare, but which in the PHT were
prime for exploiting lake/marsh resources and


hunting in mid- to low-elevation
steppe. These include lowland settings (beach bars or


lunettes) associated with pluvial lakes
or marshes, elevated old surfaces on valley margins


or Pleistocene river and stream
terraces (Basgall 1988: 104; Beck and Jones 1994, 1997;


Elston and Bullock 1994; Elston et al.
1977; Jones et al. 1996; Pinson 1999; Rusco and


Davis 1979; Willig 1988; Zancanella
1988). Fewer Prearchaic sites occur in upland-


montane locations where most are small
lithic scatters and isolates.

Several lines of evidence indicate
low population density and high residential mobility


for Prearchaic foragers. Prearchaic
sites are relatively few and, while sometimes spatially


extensive (e.g. Graf 2001), artifacts
are usually sparse. Prearchaic lithic assemblages


exhibit little variability, suggesting
minimal functional differentiation between sites. Sites


lack midden accumulations or evidence
of stored food. Very few residential structures


date to the EH; none represents
long-term occupation (Connolly and Jenkins 1995;


Harrington 1957; Pinson 1999).
Toolstone distributions from known sources suggest either


more extensive social networks or
operation over larger territories than in the Archaic

(Basgall 1989; Beck and Jones 2001;
Graf 2001) (Fig. 4). With low population density, little


competition for resources, and similar
sets of resources valley to valley, people might have


employed a land-use pattern similar to
the megapatch strategy proposed by Beaton (1991)


where it would pay to pick off
high-ranked resources in short-term occupations and then


move on to the next patch offering
similar opportunities. Frequent residential moves


(especially on the predominant
north–south axes suggested by toolstone distributions)


may have helped avoid seasonal
shortages and the need for food storage (Basgall 1989;


Elston 1986b).


The few studies of EH coprolites and
other archaeological residues indicate that

Prearchaic people made use of plants,
small mammals, birds, sh and shell sh, as well as


large ungulates such as antelope,
mountain sheep, elk and bison (Beck and Jones 1997;


Dansie 1987; Delacorte 1999; Eisalt
1997; Fry 1970, 1976; Napton 1997, Pinson 1999). The


range of items in this diet is similar
to Archaic diets. However, Prearchaic and Archaic


lithic toolkits suggest greater
investments in opposite ends of the diet spectrum.


Prearchaic assemblages are rich in
formal, hafted aked stone tools (points, bifaces and


scrapers) good for hunting and
processing large animals (Fig. 5). Although light duty


milling stones are occasionally present
in Prearchaic assemblages, evidence of intensive


seed processing and storage
characteristic of the Archaic is rare. Seeds were as surely in

the Prearchaic diet as large game, but
seed harvesting received less investment and


hunting greater investment in
Prearchaic technology than in Archaic technology.


In the warmer, dryer climates of the
early Middle Holocene (MH) and later, overall


biological productivity was lower,
sources of surface water became more scarce and less


equably distributed as lakes and
marshes in most valleys disappeared (Grayson 1993).


Some valleys continued to offered
abundant resources, while others were deserts domi-


nated by bare playas. MH environmental
heterogeneity and patchiness increased further


by northward spread of pinyon pine
across the Great Basin. The degree to which people

operated logistically from xed bases or
frequently moved camp (Binford 1977, 1979) was


increasingly in uenced by local
resource structure and population density (Elston 1982).


110 Robert G. Elston and David W.
Zeanah


Figure 4 Obsidian sources for
Prearchaic lithic assemblages from the Sadmat and Coleman sites in


western Nevada (Graf 2001), and from
Butte Valley, eastern Nevada (Beck and Jones 2001).


From the MH, Great Basin people foraged
in many lowland and upland settings, result-


ing in a more complex archaeological
record where sites display higher assemblage vari-


ability. Some sites were short-term
camps or task sites, while others assumed a residential


character, with middens, substantial
structures and storage features (Elston 1986a). Along


111



Thinking outside the box


Figure 5 Prearchaic lithic tools: a)
uted point; b) crescent (after Fagan 1988); c–f) stemmed points;


g–i) unifacial scrapers (after Davis
and Rusco 1987).


112 Robert G. Elston and David W.
Zeanah


with storage, heavy-duty seed-milling
equipment became abundant. However, the


seasonal Archaic pattern (also observed
ethnographically) of living in winter camps on


stored food (especially seeds), and
becoming more mobile during the remainder of the

year, seems to have begun in the MH.
With the passage of time, Great Basin foragers


operated in smaller territories, became
more sedentary, invested more in storage, harvest-


ing and milling equipment and less in
formal aked stone tools (e.g. Bettinger 1999).


Theoretical perspectives of Prearchaic
subsistence based on behavioral ecology


Before 1982, archaeologists lacked a
guiding theoretical context capable of garnering


insights from extant data to guide
future research or re ne subsistence-settlement models.


A lack of suitable ethnographic
analogies for EH foraging adaptations hindered investi-


gations of the Prearchaic. Middle range
theory helped interpret functional and techno-


logical variability in sites and
artifacts, but did little to explain it. Big-game hunting

proponents eagerly awaited discoveries
of kill and butchery sites, but were unable to


reconcile mounting evidence for use of
a broader array of resources. Advocates of


generalized foraging were unable to
accommodate the hunting-oriented technology, or


evidence of extensive mobility, into
their notions of adaptive continuity.


In this context, O’Connell et al.
(1982: 234–35) advocated behavioral ecology (BE) as


a theoretically coherent approach for
predicting prehistoric subsistence patterns where


suitable ethnographic models do not
apply. Such attempts have drawn three modeling


tools from the theoretical arsenal of
BE: diet breadth, patch choice and risk. In this

section, we trace the development of a
behavioral ecological research strategy for investi-


gating Prearchaic adaptations, and
discuss the implications such models have for inter-


preting EH forager ecology.


Models of Prearchaic diet breadth


O’Connell and colleagues (1982) used
the diet breadth model (DBM) to explain the rarity


of EH milling stones. The DBM ranks
resources according to the ratio of calories to


handling time, assuming that foragers
take encountered prey only if it yields higher caloric


returns than those offered by searching
for higher-ranked resources. Two DBM predic-


tions are that the abundance of
high-ranked resources determines whether lower-ranked

resources fall into the optimal diet,
and that fewer encounters with higher-ranked prey


can expand diet breadth to include
lower-ranked resources (Schoener 1971). High-ranked


resources in EH environments were
presumed to have been suf ciently abundant for


Prearchaic foragers to bypass
low-ranked seeds. As climatic changes diminished the abun-


dance of higher-ranked prey, Archaic
foragers added seeds to their diets and milling


equipment to their technological
repertoire. This hypothesis relied on a reasonable


assumption that seeds ranked lower than
most other Great Basin foods (i.e. medium and


large-sized game, roots and sh).


Simms (1987) tested this assumption
by experimentally harvesting an array of Great

Basin plant resources, estimating
animal return rates from historical, ethnographic and


wildlife management sources. Seeds
ranked lowest, although their return rates varied



113



Thinking outside the box


widely by species, harvest timing,
procurement mode and abundance. Game proved


among the highest-ranked resources,
with rank correlating roughly with body size.1 Given


DBM predictions, it seemed reasonable
to infer that Prearchaic and Archaic foragers took


large game whenever available, but
harvested seeds only when higher-ranked resources

were scarce. However, Simms could not
demonstrate that diminished encounters with


large game and consequent diet breadth
expansion accounted for intensi ed seed usage


during the MH. Estimating
nineteenth-century encounter rates, he calculated the diet


breadth of ethnohistoric foragers,
predicting that women should have bypassed lower-


ranked seeds under all conditions,
including the absence of large game. Clearly, the DBM


contradicts the ethnographic record
documenting Great Basin women harvesting and


storing small seeds. Because the DBM
fails to predict ethnohistoric small-seed harvest-


ing, even in the absence of large game,
it also fails to predict seed usage by either


Prearchaic or Archaic foragers.
Consequently, the MH proliferation of milling stones

cannot re ect the inclusion of seeds
into MH diets because of declining large game


encounters. Noting this predictive
failure, Simms (1987: 82–3) speculated that intensi ed


MH seed use might have resulted from
seed storability, rather than short-term foraging


ef ciency; gatherers stored seeds
anticipating seasonal food shortages.


In contrast to the predicted
exclusion of seeds from all prehistoric diets, Simms (1987:


88) calculated that Prearchaic
encounters with large game would have had to have been


as much as twenty- ve times greater
than ethnohistoric encounter rates to bump small


game and higher-ranked plants out of
men’s optimal diets. Simms concluded that these

middle-ranked resources were rmly
ensconced in Prearchaic diet under any plausible


scenario of large game abundance. Based
on this assessment, Simms (1987: 98) conceded


that Prearchaic folk may have taken
large game more often than Archaic foragers did,


but argued that EH game could not have
been abundant enough to support a specialized


big-game hunting adaptation.
Fluctuations in resource abundance demanded that


Prearchaic foragers make frequent
adjustments to diet breadth, ranging from broad-


spectrum seed harvesting to big-game
hunting on a daily, seasonal or yearly basis. This


notion of Prearchaic adaptive exibility
accounted for occasional Prearchaic milling


stones, coprolite evidence that
Prearchaic foragers ate seeds and the persistence of some

Prearchaic point types into the MH
(Simms 1988).


Models of Prearchaic patch choice


The DBM merely ranks resources by
relative returns while assuming that resources are


randomly distributed. However, foragers
face a more complicated world. Observing that


Simms’ modeling exercise did not
consider the spatial and seasonal distribution of


resources in EH environments,
Intermountain Research (IMR) used the patch choice


model (PCM) to simulate Prearchaic
subsistence and settlement (Elston et al. 1995). The


PCM predicts foraging choices in patchy
environments by ranking resource patches


according to caloric return. Foraging
in a patch is economical if the returns for seeking

and handling resources within that
patch exceed the overall returns for traveling to and


foraging within higher-ranked patches.
The PCM predicts that foragers prefer the most


pro table patches, and that a change in
resource abundance may alter the array of patches


selected (MacArthur and Pianka 1966).


114 Robert G. Elston and David W.
Zeanah


Estimating the composition and
distribution of biotic habitats by soil type, IMR simu-


lated the nineteenth-century resource
landscape of the Carson Desert in western Nevada,


classifying the biota associated with
each soil as a patch and modeling patch choice

decisions by seasonality and sexual
division of labor (Raven and Elston 1989; Zeanah et


al. 1995; Zeanah 1996). This landscape
served as a baseline against which palaeoenviron-


mental data could be compared to
approximate EH resource distributions and Prearchaic


foraging decisions.


IMR assumed that higher EH
precipitation fostered abundant small and large game, as


well as plant foods around lowland
wetland communities in distributions dramatically


contrasting with discrete
nineteenth-century upland, desert and wetland patches. IMR


calculated the optimal diet of
Prearchaic men and women (Fig. 6) for an array of wetland

and terrestrial prey types reasonably
presumed available within foraging distance of an


EH lowland camp. IMR used Simms’
(1987) return rate estimates for the ethnohistoric


Great Basin, but increased his
encounters with game by 75 per cent, corresponding to the


greater densities of forage expected
for the EH Carson Desert.


Like Simms, IMR’s simulation
predicted that Prearchaic women would have bypassed


most seeds in favor of the
highest-ranked plants and small game. The optimal diet of men


always included medium and large game
but, unlike Simms’ estimates, excluded smaller


game. Unlike ethnohistoric foragers,
Prearchaic men should have achieved higher return

Figure 6 Simulated autumn foraging
returns for men and women foragers in early Holocene


wetlands.



115



Thinking outside the box


rates than women, with both men’s and
women’s foraging best in lowland patches. IMR


proposed that this encouraged a
residentially mobile adaptation in which men’s hunting


opportunities determined residential
movements and women’s foraging opportunities


determined site location. Although
Prearchaic foragers could not be stereotyped as ‘big-

game hunters’, the EH ecological
setting prompted a hunting-oriented adaptation signi -


cantly different from later
hunter-gatherers. IMR argued that this perspective best


accounted for specialized Prearchaic
lithic assemblages, point distributions and evidence


for high residential mobility.


A model of Prearchaic risk sensitivity


A third application of BE to model
Prearchaic foraging used the Z-score model (ZSM)


to consider foraging risk in EH
environments (Pinson 1999). The ZSM predicts forager


choices when return rates vary
stochastically by considering the mean and variance of


resource return rates. It assumes that
acquisition of a minimum quantity of food within a

limited period is critical. If average
foraging returns exceed a critical threshold (i.e.


starvation, reproductive success),
pursuit of low-variance resources minimizes the risk of


falling short. If foraging means fall
below the threshold, gambling on high-variance


resources maximizes chances that
foraging returns will exceed minimum requirements


(Stephens and Charnov 1982).


Pinson’s (1999) hypothesis is that
starvation risk imposed a minimum threshold for


Prearchaic foragers. Pinson assumed
variance in return rates correlates with prey size,2


and that Simms’ (1987) resource
ranking correlates with mean return rates. Prearchaic

hunter-gatherers should have been
either risk-averse or risk-prone.3 Risk-prone


foragers followed predictions of the
DBM and PCM, preferentially pursuing large


ungulates over smaller game and
foraging in habitats where encounters with large game


were most likely. In contrast,
risk-averse foragers preferred lower-ranked, but


predictable, small mammals, fish and
fowl, foraging in locations offering access to


multiple habitats. Pinson did not
consider the effects of sexual division of labor on


resource choice or risk sensitivity.


These predictions found support in
analyses of faunal assemblages and point distri-

butions in the Alkali Basin of
southeastern Oregon. EH faunal assemblages have rela-


tively low ratios of artiodactyl to
lagomorph bone compared with later Holocene


4


assemblages, and EH projectile points
cluster in locations offering access to multiple


biotic habitats. Pinson interprets this
as evidence that Prearchaic foragers avoided


starvation risk by emphasizing
small-game procurement at the expense of large-game


hunting.


Evaluation of Z score and patch choice
models of early Holocene adaptation

Given that three attempts to model
Prearchaic adaptations have been made in the twenty


years since BE was introduced to Great
Basin archaeology, it is worth considering


whether BE has contributed any
meaningful insight. Prospects seem bleak at rst glance


since the three applications make
contradictory inferences about Prearchaic foraging


116 Robert G. Elston and David W.
Zeanah


behavior, claiming to retrodict
broad-spectrum foragers, big-game hunters and small-


game specialists. However, all three
agree that specialized big-game hunting is ecologi-


cally untenable in EH Great Basin
environments, expecting that Prearchaic foragers


procured a more generalized array of
prey. Of contention is whether Prearchaic adapta-

tions were categorically different from
later Archaic patterns; if so, what roles did large-


game hunting and generalized foraging
play in determining the distinctive strategies? We


have seen that DBM alone is useful for
ranking resources, but that patch choice conditions


diet breadth, in turn conditioned by
environment. Therefore, in the following discussion


we evaluate the ZSM as applied to the
North Alkali Basin, Oregon (Pinson 1999), and


the PCM (in which the DBM is included)
in Railroad Valley, Nevada (Zeanah et al. 1999).


The ZSM in North Alkali Valley, Oregon:
risk-averse pursuit of low-ranked resources


Pinson (1999) interprets the prevalence
of small game in EH faunal assemblages of south-


eastern Oregon as evidence that
starvation promoted a distinctive, risk-averse, Prearchaic

foraging strategy. The Oregon faunal
pro le is consistent with coprolite evidence from


the Bonneville and Lahontan basins for
generalized diets (Eiselt 1997; Fry 1976), and is


duplicated by EH faunal assemblages
from south-eastern California (Delacorte 1999).


These ndings do not contradict DBM and
PCM simulations (Elston et al. 1995; Simms


1987) that predict such resources to
have been regular prey of Prearchaic women. The


real question is whether the scarce
remains of large ungulates in the faunal record demon-


strates that Prearchaic foragers
avoided risk by shunning large game.


Pinson (1999: 109–13) proposes
that EH foraging risk was induced by local variations

in water budget, precipitation,
watershed catchment and so on, causing lake basin produc-


tivity to vary unpredictably. When
moving into a new basin, Prearchaic foragers could not


anticipate the abundance of prey based
on previous encounter rates, and, thus, avoided


starvation risk by procuring the most
dependable resources. Yet Prearchaic foragers must


occasionally have entered basins where
foraging returns proved unlikely to surpass the


starvation threshold. The ZSM predicts
that such foragers should choose higher-variance


resources over alternatives that are
more predictable: risk-averse strategies should char-


acterize only situations where mean
foraging returns are securely above the starvation


limit. If risk was the primary
constraint of Prearchaic adaptations, there should be

evidence for both specialized
large-game hunting and small-resource procurement rather


than consistent evidence of generalized
foraging.


Pinson’s scenario does not address
why, if foragers are risk-averse, they should even


bother to move from one basin to
another. EH lowland environments supported abun-


dant and diverse populations of sh,
small mammals and waterfowl that should have been


resilient to over-exploitation by
Prearchaic foragers (cf. Winterhalder et al. 1988). Late


prehistoric and ethnographic foragers
exploiting such patches tended to stay put, moving


out only when extraordinary resource
opportunities became available elsewhere or in

response to catastrophic environmental
failures (Fowler 1990, 1992; Raven 1992; Raven


and Elston 1989). If EH environments
were really as patchy and unpredictable as


proposed by Pinson, Prearchaic foragers
should have employed land-use strategies similar


to those of the Archaic, generating
similar archaeological residues. We agree that EH


ungulate abundance was probably
sensitive to hunting pressure, and, as well, varied with



117



Thinking outside the box


seasonal migrations (cf. Winterhalder
et al. 1988). But, in fact, there seems little reason

for Prearchaic hunter-gatherers to move
from basin to basin except to maximize large-


game encounters in the more homogeneous
and productive EH environments we en-


vision. Such a strategy would account
for the hunting orientation and evidence of high


mobility in Prearchaic lithic
assemblages.


One constraint of the standard ZSM
is that risk-averse foragers forgo the opportunity


to seek higher-return prey by pursuing
low-variance resources. Making this assumption


of Prearchaic foragers fails to
consider the role of sexual division of labor in alleviating


scheduling con icts. The Hadza of
eastern Africa provide a useful analogy (Hawkes 1990;

Hawkes et al. 1991). Hadza men pursue
big game to the exclusion of smaller prey even


though many days of failure separate
hunting successes. Women and children avoid


starvation by procuring small game,
roots, berries and nuts. A similar division of labor


likely characterized Prearchaic
foraging strategies:5 men pursued high-return but high-


variance large game, while women
procured more secure, but lower-ranked resources.


Therefore, Prearchaic foragers would
not have had to abandon hunting in order to


provision themselves daily with smaller
resources.


We propose that small seed usage is
a more likely example of risk-averse foraging than

small-game procurement. Both Simms
(1987) and IMR (Zeanah et al. 1995) predicted the


exclusion of small seeds in optimal
diets of Prearchaic through ethnohistoric hunter-


gatherers. 6 Obviously, ethnographic
and archaeological data demonstrating that seeds


were a critical food resource after the
Prearchaic challenge this prediction.


Noting this predictive failure,
Simms (1987) rst suggested that Archaic women stored


seeds to avoid over-winter food
shortages. Although expensive to process for consump-


tion, seeds can be stored with
relatively little effort, allowing gatherers to defer high


processing costs for future periods of
scarcity (Bettinger 1999). If so, seed storage quali-

es as risk-averse foraging, because
Archaic women were, by de nition, forgoing oppor-


tunities to pursue higher-ranked prey
to create a predictable food patch by stockpiling


food resources too low ranked to enter
their optimal diet.7 If risk aversion rather than


energy maximization accounts for
Archaic seed usage, then there should have been less


of a need for food storage in the EH
than in later periods.


Patch choice and EH and MH foraging
ecology in Railroad Valley


To contrast EH and MH foraging
strategies, it is instructive to consider IMR’s (Zeanah


et al. 1999) simulation of foraging
ecology in Railroad Valley, eastern Nevada, using

similar techniques for estimating
biotic habitats by soil type that were employed in the


Carson Desert. However, more detailed
regional palaeoenvironmental data, better infor-


mation on the productive capacity of
modern soil types and improved GIS (Geographic


Information System) capabilities
permitted a more nely tuned reconstruction of prehis-


toric foraging landscapes than was
feasible in the Carson Desert.


Annual herbaceous biomass
productivity (Fig. 7) was estimated for EH and MH forag-


ing landscapes in Railroad Valley by
putting an EH lake on the modern playa and alter-


ing the modern productivity of soil
types to re ect expected EH and MH parameters of

precipitation, water table and erosion.
These resource mosaics formed the setting for


simulating the optimal diet breadth of
male and female foragers randomly encountering


118 Robert G. Elston and David W.
Zeanah


prey in each habitat by season.
Obviously, these palaeoenvironmental reconstructions and


foraging simulations must be cautiously
regarded as only tentative, rough approximations.


However, they allow us to gauge the
effects of palaeoenvironmental change on foraging


behavior in a theoretically consistent
manner.


Mean and standard deviation of
simulated men’s foraging returns by season for the

fteen most productive Railroad Valley
habitats are shown in Figure 8. The gure also


shows the thresholds under which
lower-ranked seeds and small game begin to enter


optimal diets. Compared to the EH,
hunting returns of MH men diminish as much 75 per


cent in all seasons, suggesting that
hunting was much less productive in the MH. Women’s


MH foraging returns (Fig. 9) lessen
much less dramatically, with autumn returns increas-


ing slightly (due to arrival of pinyon
as a new resource). During summer, autumn and


early winter, women’s foraging
returns were too high to allow small seeds into optimal


diets under both EH and MH climatic
scenarios. However, MH women experience a 60

per cent reduction in late
winter-spring foraging returns that should allow even the lowest-


ranked seeds into the diet, but in the
season when most small seeds are unavailable. This


diminution of women’s foraging
opportunities re ects the desiccation of lakeside


wetlands in Railroad Valley, removing
sh, waterfowl, eggs and wetland small mammals


from women’s springtime prey
opportunities.


Figure 7 Above-ground annual herbaceous
biomass estimated for early and middle Holocene Rail-


road Valley, Nevada.



119


Thinking outside the box


Figure 8 Men’s foraging returns (mean
and standard deviation) by season for the fteen highest-


ranked habitats in Railroad Valley,
showing thresholds at which different resource classes enter the


diet.


Figure 9 Women’s foraging returns
(mean and standard deviation) by season for the fteen highest-


ranked habitats in Railroad Valley,
showing thresholds at which different resource classes enter the


diet.


Conclusion

If broad-spectrum foraging and sexual
division of labor were always components of


Great Basin foraging strategies, what
accounts for the Prearchaic–Archaic contrast in


the archaeological record? We propose
that Prearchaic technology and settlement


120 Robert G. Elston and David W.
Zeanah


pattern are responses to low human
population amid a unique landscape and resource


structure. Our simulations point to key
ecological differences between Prearchaic and


Archaic subsistence strategies in the
Great Basin. Large animals should have been rela-


tively abundant in low to mid-elevation
brushy steppe from fall to spring where they


could have been hunted from lowland
bases. EH wetlands provided a relatively secure

late winter-early spring foraging patch
for women where they could take small game,


waterfowl and fish from the same sites.
The abundance of expansive wetlands, and lack


of competition throughout the Great
Basin, allowed Prearchaic foragers high mobility


that maximized men’s encounters with
large game without sacrificing women’s foraging


interests.


If lithic technology and tool
morphology are adaptively signi cant, the function of


Prearchaic lithic assemblages is clear.
In addition to ake tools and choppers, these assem-


blages include bifaces, projectile
points, steep-edged end- and side-scrapers, bifaces, ne

gravers and awls, and crescents, but
they rarely contain milling stones (Basgall 1988; Beck


and Jones 1997; Davis and Rusco 1987;
Elston 1986a; Willig 1988). Flaked stone tool


preforms are frequently blades or
blade-like akes; tools are notable for their large size.


Stemmed bifaces are often multipurpose,
serving variously as points, knives and scrapers


(Jones and Beck 1999). The large size
and abundance of formal tools in Prearchaic lithic


assemblages re ect both mobility and
the importance of capture and processing tasks for


which the tools were used (Bleed 1986;
Elston 1990, 1992; Goodyear 1979; Kelly 1988,


1992; Kelly and Todd 1988; Tomka 2001;
Torrence 1983, 1989; Ugan and Rogers 2000).


Bifaces provide a means for mobile
foragers to transport toolstone in a useful form far

from toolstone sources, and are
reliable, maintainable and exible (easily converted to


another form). These qualities, and the
ability to predict tool-use life, are enhanced in


bifaces and other formal tools by use
of high-quality toolstone, large tool size, standard-


ized symmetrical form and standardized
maintenance techniques. Such a formal, stan-


dardized lithic technology is more
expensive to master and maintain than expedient


approaches (Elston 1992). However, the
pay-offs are enhanced tool reliability and work


ef ciency (Bleed 1986; Tomka 2001).


Although one could use this tool kit
to capture and process fish, small mammals, and

plants, it seems more appropriate for
use on larger prey. Indeed, Tomka (2001) argues


that proportions of formal and
standardized tools (particularly knives and scrapers) in


lithic assemblages relate directly to
intensity of large-animal processing. He suggests


that the long-term trend in North
America toward greater reliance on expedient tools,


noted by Parry and Kelly (1987), is
better explained as a decreasing reliance on


strategies of high-volume animal
hunting and processing than as decreasing residential


mobility.


Extirpation of Great Basin wetlands
during the MH removed women’s critical forag-

ing patches from most basins, creating
signi cant risk of food shortages. Residential sites


were established at perennial springs
and associated wetlands (Fagan 1974; O’Connell


1975; Elston 1982). By accumulating
seed stores during periods of seasonal abundance,


MH women minimized resource variance,
decreasing the chances of falling short of food


during the winter. Dependence on
storage tethered Archaic foragers to their seed caches,


reducing their overall mobility and
spawning a profound reorganization of tool produc-


tion, toolstone procurement and hunting
strategies (e.g. Eckerle and Hobey 1999). Men



121


Thinking outside the box


continued to hunt, but more and more in
logistical forays from relatively xed bases. As


men’s and women’s diets
increasingly overlapped, men invested less in hunting equip-


ment, while women invested more in
plant-processing tools. Mass large-game procure-


ment techniques occurred too
infrequently for investment in specialized formal


processing tools to pay off. This
explanation accounts not only for the MH disappearance


of formal chipped stone tool industries
and the proliferation of milling equipment, but


also the appearance of storage features
and prolonged duration residential base camps

(Bedwell 1973; O’Connell 1975;
Hattori 1982; Schroedl and Coulam 1994).


We cannot base contrasts between
Prearchaic and Archaic foragers in the Great Basin


on either diet breadth or
risk-sensitivity alone. Broad-spectrum foraging spans at least the


Late Palaeolithic (Stiner et al. 2000),
and was always a feature of Great Basin foraging.


Moreover, Great Basin foragers
continued to hunt existing large game through the mid-


Holocene and into the ethnohistoric
period. Starvation was a risk faced by hunter-


gatherers throughout Great Basin
prehistory, moderated by a complex of tactics, includ-


ing variable diet breadth and mobility,
and sexual division of labor (Zeanah n.d.),

although the implications of the latter
for Great Basin foraging strategies have been little


appreciated. Great Basin men’s and
women’s diets overlapped somewhat, but like male


foragers elsewhere (Hawkes 1996), men
tended to pursue more high-ranked but highly


variable resources such as large game,
while women focused on lower-ranked, less vari-


able resources such as waterfowl, sh
and seeds that require little processing. This division


allowed Great Basin foragers to have it
both ways: women saw to it everyone had some-


thing to eat, while men brought home
the occasional high-return prey. Therefore, men


invested heavily in hunting technology,
while women had no need to invest in equipment


for intensive plant processing.

We have built a case that a BE
theoretical approach has yielded meaningful insight into


Prearchaic foraging adaptations, and
that DBM and PCM simulations better account for


the Prearchaic archaeological record
than the ZSM alone. However, our theoretical


expectations must be borne out by
further archaeological investigations. In large part, the


objectives of future research have
already been laid out, and involve continued search for


buried PHT deposits, and extraction of
subsistence and chronological data from surface


assemblages (Beck and Jones 1997). Our
BE approach adds testable expectations to this


research strategy. Our simulations
suggest that the known Prearchaic record is biased

toward sites positioned primarily to
access women’s resources, but bearing a technology


re ecting men’s subsistence and
mobility strategies. If we are correct, evidence of men’s


high-variance foraging behavior lies
buried in valley piedmonts and passes, away from the


wetland settings where the most
extensive Prearchaic sites are usually found. Effective


search for such evidence must combine
geo-archaeological identi cation of sediment


exposures of suitable age, with
theoretical simulations of EH large game habitat and


Prearchaic hunting behavior.


Acknowledgements


We thank Eric Ingbar, Ariane Pinson,
Steve Simms, our two anonymous reviewers for

their comments and suggestions and,
especially, Ariane Pinson for permission to use the


122 Robert G. Elston and David W.
Zeanah


graph in Fig. 2, and Gene Hattori and
Nevada State Museum for permission to use the


tool illustrations in Fig. 5.



Silver City, Nevada


Notes


1 A correlation between body size and
resource ranking seems well supported ethno-


graphically (O’Connell and Hawkes
1981; Winterhalder 1981; Hawkes et al. 1982), and

often serves to rank prey items
relatively (Broughton 1994). However, this assumption


is violated when smaller resources
occur in unusual abundance, returning exceptionally


high caloric yields. For example,
grasshoppers obtained from windrows along margins


of lakes provide higher caloric
return rates than large-game hunting (Madsen and


Kirkman 1988). Brine y larvae,
bulrush and pickleweed seed and sh are also known


to occur occasionally in windrows and
can be expected to return similarly fantastically

high caloric return rates (Raven and
Elston 1988; Bettinger 1993; Barlow and Metcalfe


1996; Madsen and Schmitt 1998).
Similarly, mass capture technologies and communal


hunting tactics can provide higher
caloric returns for hunting small and medium-sized


mammals than encounter hunting of
large mammals (Simms 1987). Therefore, the


ranking of any particular prey item
varies depending on mode of procurement.


2 Like the DBM assumption that prey
size correlated with resource ranking, this assump-

tion seems ethnographically justi ed
(Hawkes 1990, 1991) and is likely to be generally


true. However, mode of procurement
may allow smaller resources to be harvested,


unpredictably, at fantastically high
return rates. In addition, many small and medium-


sized animals are prone to periodic,
catastrophic population declines induced by


climatic change, disease and
overcrowding (e.g. Zeanah 1996; Kelly 2001). Therefore,


in many short-term circumstances, the
return garnered from smaller game may actually

be more variable than the returns
from large game.


3 Pinson (1999) characterizes possible
risk strategies as risk-sensitive or risk-indifferent.


However, risk-sensitive foragers can
be either risk-prone (variance-maximizing big-


game hunters) or risk-averse
(variance-minimizing small-game gatherers) depending


on circumstances. Because the outcome
of risk-indifferent foraging is the same


(variance-maximizing) as risk-prone
foraging, we refer to the dyad ‘risk-prone–risk-

averse’ for the sake of simplicity.


4 Pinson’s (1999) analysis does not
consider other factors that might contribute to low


artiodactyl index values, such as
processing tactics to minimize transport costs when kill


sites and residential bases were in
different locations (e.g. Metcalfe and Barlow 1992;


O’Connell et al. 1990).


5 This may appear to contradict our
initial assessment of ethnographic analogies as


incapable of providing insight into
Prearchaic adaptations. However, our conviction

that sexual division of labor was
likely a critical aspect of Prearchaic foraging strategies


is based on homologous rather than
analogous reasoning, and is theoretically justi ed


from a BE perspective. Sexual
division of subsistence labor is nearly universal among


ethnographic hunter-gatherers, and
most anthropologists perceive sexual division of



123



Thinking outside the box

labor as fundamental to
hunter-gatherer subsistence organization (Ember 1975; Hiatt


1978). In addition, current
evolutionary models see sexual division of labor as a signi -


cant trend in hominid evolution
selected by con icting responses of males and females


to the risks of successful
reproduction (Bird 1999; Hawkes 1996). Ethnographers


observed a division of subsistence
labor by gender among Great Basin foragers (Kelly


1932: 79; Steward 1938: 44, 1941:
312–13; Stewart 1941: 406). Moreover, a growing body

of bioarchaeological evidence builds
a compelling case for sexual division of labor


among Archaic hunter-gatherers in
the Great Basin (Hemphill 1999: 285; Larsen and


Hutchinson 1999; Rhode et al. 2000:
55–6). We are, therefore, con dent that sexual


division of labor was practiced by
Prearchaic foragers; models of Prearchaic adaptation


that ignore this factor cannot be
complete.


6 Critics point out that this nding
hinges on the use of foraging experiments to replicate

prehistoric resource return rates
(Bettinger 1994), and are skeptical that modern


scholars can reproduce the expertise
of prehistoric foragers or judge circumstances


under which they chose to take
resources. However, many estimates have been repli-


cated by independent experiments
(Larralde and Chandler 1981; Simms 1987; Jones


and Madsen 1991; Bullock 1994;
Barlow and Metcalfe 1996) and are comparable to


return rates procured by
ethnographic hunter-gatherers who take similar arrays of food

(O’Connell and Hawkes 1981; Cane
1987). More important, the experimental replica-


tions would have to be wrong by
several orders of magnitude to change DBM predic-


tions (Simms 1987: 49, 53).


7 Obviously, food storage also incurs
risk in that caches may be lost to spoilage and theft.


However, we agree with many previous
biological and anthropological models that


consider food storage to be a
strategy for minimizing variability in foraging returns

(Testart 1982; Vander Wall 1990;
Winterhalder 1990; Winterhalder et al. 1999).


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9-1 Missing


































































Missing















































































10-1





From Tanaka, Jiro (1980) The San
Hunter-Gatherers of the Kalahari: A Study in


Ecological Anthropology. Tokyo:
University of Tokyo Press


Bottom of page 30 through bottom of
page 35


Hunting


Representative hunting methods of the
≠Kade San include shooting large game with bow


and arrow, catching small antelopes in
a snare, and hunting the springhare with a hooked

pole. If he owns a dog, the San may
hunt with a spear with the help of the dog; in the case


of small birds or animals, he may beat
them to death with a club or bring them down with a


throw of a handy stick. Such beasts of
prey as lion, leopard, cheetah, and lycaon are "used"


in obtaining animal meat: when the
people observe a great number of vultures wheeling, they


dash to the point and usurp the prey.


Bow-and-arrow hunting is surprisingly
difficult and requires tremendous exertion. It is


used mainly on large antelopes, but
also on giraffes, ostriches, duikers, steenboks, and other


animals. Whenever a man goes out, he
slings over his shoulder a leather bag holding his

bow, arrows, and spear, and as long as
he is not out of poisoned arrows he is always ready


to shoot whenever a target appears.
Sometimes, after making the rounds of their snares, the


hunters will loiter around a spot where
gemsbok, eland, and other animals are known to


flock, meanwhile sticking their hooked
poles into holes in the ground in search of


springhare. People sometimes walk 20 km
a day over the practically shadeless bushveld.


And yet most days they return to camp
emptyhanded. If by chance the hunter runs across a


herd, or even just one animal grazing,
he approaches from downwind so as not to be


perceived by the animal.


The Kalahari has few trees, and grass
and shrubs a meter or so in height grow only in

scattered spots; this lack of cover
makes it extremely difficult to sneak up on game. Because


the San’s bow (Figure 13) is a small,
crude device, he must approach to within about 20 m


of his intended prey. If he manages to
get within range and hit the animal with a poisoned


arrow, and ascertains that the poisoned
point has firmly penetrated its body, he must then


observe the direction of the animal’s
flight and memorize its hoofprints. Shocked by its


wound and the effects of the poison,
the animal will flee for its life, and the San must track


it. Since game seldom approaches the
campsite, hunting is usually done some 10 or 15 km


away. The hunter who has scored a hit
returns temporarily to camp. Until he confirms the


death of the animal, he can take no
food or drink other than water, for the San believe that if

the archer eats food the wounded animal
will regain its health and escape. That night the


hunter sleeps in the camp, but the next
morning before dawn he departs, accompanied by


several hardy young men, and the
tracking begins. Although pained by the poison, the


animal gives all its strength and flees
practically in a straight line. Finally, weakening, it


begins to seek shade and rest under
some of the few trees to be found. As the distance


between their prey’s rest stops
shrinks, they know it is not far off. Soon they will find


imprints of its body, as it can no
longer rest on its feet but must lie down. Now, even if it


should muster all its dying strength
for a last run, it cannot go far. At last the hunting party


will come upon the animal lying under a
tree; if it is still breathing, they will approach with

their spears (Figure 14) and, striking
for the heart, administer the quietus.


Since dark is by now fast approaching,
the hunters hurriedly skin the animal and sever the


head and limbs. The body is also
chopped up into a load for each man. The horns are


usually thrown away, but may be brought
back to be used as an axe handle or the like. As


for the rest, almost nothing is left
behind except sometimes the contents of the stomach and


intestines. If the hide is thick and
untannable, like that of alarge male antelope or a giraffe, it


is sectioned and used as food; thinner
skin is stripped off carefully so as not to tear it, and


made into clothing or a carrying skin.
The contents of the stomach are full of stomach acid


and are sour and bitter, but in times
of drought they are a source of precious drinking water.

The blood is poured into the stomach
sac and hardened in the heat, making it easier to carry.


The fat sticking to the membrane
surrounding and supporting the intestines is mixed with


blood and then stuffed into the
duodenum and small intestine; the latter are then heated to


make sausage. The liver, heart, and
kidneys, which spoil easily, are cooked and eaten on the


spot; the meat on the ribs is also
roasted and eaten for dinner that night. Since the San


especially relish the liver and heart,
these are the first to be cooked and eaten. The heart in


particular is taboo for women; since
the San believe that if a woman should eat the heart the


men’s hunting would be unsuccessful, it
is almost never brought back to camp.


The men, having finally come upon some
meat after a long food drought, usually spend the

night gorging at the scene of the kill;
then the next morning each man shoulders his load of


meat and they return to camp. If the
animal weighed 200 kg, assuming they actually carry


back 150 kg of that, a party of five
men would each have to carry 30 kg; the camp might be


from 15 to 30 km away. Transporting
such a load across the burning Kalahari is no joke. In


the case of a giraffe weighing over a
ton, the tracking may take three or four days, making


the trip home that much longer, not to
mention the additional amount of meat to be carried.


In such cases the hunters will discard
the bones, then cut the meat into strips and dry it,


which reduces the carrying weight
considerably. Drying also serves to prevent spoilage.


It usually takes three to four days,
all told, to bring in a large antelope. Even with all this

effort, there are times when the poison
works poorly and the prey escapes, or when a lion,


leopard, or other carnivore gets to the
dying animal first. Bow and arrow hunting not only


requires much exertion, it is also
seldom rewarded. Nevertheless, the hunters put all their


heart into it, encouraged by the
prospect of getting so much meat at one time.


One night, !Kawamakue ate nothing when
he came back to camp. I knew at once that he had


shot some game and asked him if he had,
but his answer was, "I couldn’t find /o [gemsbok]


since there is no water around G/au/u
Pan." It was late at night when I was ready to go to

bed that I heard the news that he had
shot a gemsbok with a poisoned arrow.


Next morning at dawn, I joined the men
to follow the game. But it


was a trip of 50 km there and back, so
they would not take me with them on foot. I started


the engine of my truck, loading the
three young men of the camp, !Kawamakue, !Tebechu,


!Kana, and two twelve-year-old boys,
Daon//an and Daogu, on its bed, and made for the


spot where !Kawamakue shot the game. It
was a place a little way off from G/au/u Pan,


covered with endless bush. There was a
faint rut to G/au/u and I could drive straight there.


Beyond that, I would simply have to
drive in the direction !Kawamakue pointed. As there


were occasional trees that the truck
might run into, I drove slowly, at about 15 km/h.

Because the land is covered with
grasses, I could not see the contours of the land. If the


truck should break down here, I would
have no other recourse but to walk back to the


nearest village, 150 km away. It is, of
course, a possibility to reckon with in this sort of


fieldwork. The radiator grille was soon
covered with grass seeds and flowers, and we hadn’t


gone five minutes before the water
began to boil and the radiator started making sounds of


protest. Knowing I was overstraining
it, I still proceeded, but couldn’t go on more than 7 or


8 km. When I shut off the engine, half
the water had boiled over, and the remainder did not


stop boiling for five minutes. Removing
the grass from the front part of the radiator and


pouring in some more water, I again
proceeded, but not for more than 7 or 8 km. But it’s

still faster than walking. Indeed, the
Kalahari is no place for automobiles—it’s a place for


the San and the animals to walk about
freely.


When we arrived at the spot where
!Kawamakue shot the game yesterday, he told us all


about his activity and pointed out the
hoofprints of the poisoned animal. From there the


three men began to follow the prints on
foot. The game had run straight in the direction


opposite to the campsite. We also
started following the track. The animal must have been


suffering as the poison passed into its
system. We could find signs where it had rested here


and there under the trees. As we
proceeded, the distance between resting places became


shorter and we knew that the animal was
close at hand. At about 11 o’clock we finally found

the animal. Seeing us, the gemsbok
tried with all its strength to run away. But, being


poisoned, it could not go far. After
200 m it lay down under a tree, breathing hard. The


arrow which !Kawamakue had shot
yesterday was hanging broken at the joint of its foreleg.


The two beautiful horns leaned forward
heavily. It seemed just on the verge of dying. But a


careless approach was still dangerous,
since the poisoned animal might, out of pain, attack


with all its remaining strength.
!Kawamakue put an arrow to the bow about 10 m from the


animal to add more poison. !Kana also
shot another arrow. They threw sticks at the


gemsbok, but it no longer had the
strength to lift its head. Approaching from behind, a man


thrust a spear at its heart. The
gemsbok, though standing on its legs for a time, at last

collapsed to the ground, its body
convulsing, and took its last breath. The hunters at once


began to cut it to pieces. It was
marvelous to see them cut it open in a straight line from


chest to belly and skillfully disjoint
the body. The skin of the male gemsbok is too thick for


any use, so after chopping up the flesh
they roasted and ate it all. In an instant, they took out


the internal organs and cut off the
limbs. The mushy content in the stomach is an important


water supply when there is no water, so
they squeeze it and get water from it. Almost no part


is discarded. The bones, of course, are
smashed and the marrow eaten. They stuffed


themselves with the roasted liver, part
of the intestines, and the breast. Resting a while, we


started for home. The old people,
women, and children must be waiting for us.

Compared with the difficulty of hunting
with bow and arrow, hunting for springhare and


small antelope is relatively easy. The
springhare looks for food at night and spends the day


resting in its underground burrow, so a
barbed pole about 4 m long can be used to hook


him. He cannot simply be pulled out,
however; the hunter fastens the pole down at the


entrance to the burrow and, after
estimating the location of the springhare, digs straight


down a meter or so and grabs it.


Small antelopes like the duiker and
steenbok usually travel singly or in pairs, tending to


follow predictable routes at certain
times. They can be taken by a snare strung across their


route (Figure 15). The men have five or
six snares set at any one time and make an

inspection tour every day. If it rains,
the rope used in the snare becomes useless and must


be replaced.

























10-2


Missing


































11-1





58 Evolutionary Anthropology

ARTICLES


Showing Off, Handicap Signaling, and
the


Evolution of Men’s Work


KRISTEN HAWKES AND REBECCA BLIEGE BIRD



thropological puzzles,
including seem-


Zahavi’s1,2 handicap principle
makes “waste” a common outcome of signal



ingly maladaptive
cultural practices,

selection because the cost of a signal
guarantees its honesty. The capacity to



monumental
architecture, relatively in-


bear the cost reveals the show-off’s
hidden qualities. While displays take many



efficient foraging
behaviors, and gener-


forms, some also provide fitness-related
benefits to the audience in addition to



osity.8 –13 Bliege
Bird14 –16 has shown


information about the show-off. Zahavi3
has used the handicap principle to

explain both merely wasteful displays
and altruistic behavior. Here we focus on that attention to
the signaling content


the distinction between these two kinds
of display and the importance of of foraging strategies
can help explain


benefits other than information in
show-off explanations of a particular puzzle differences
between the efforts that


in human evolution: men’s work. Males
of other primate species do not con- men and women devote
to them.


tribute any significant fraction of
the food consumed by females and juveniles. Among the Meriam
Islanders of the


Our own species is different. When
people live on wild foods, hunting is usually Torres Strait,
turtle hunters supply


a specialty of men, and meat is
commonly a substantial component of every- meat that is
widely shared at feasts,


one’s diet. Here we explore the
hypothesis that this unique male subsistence while
spearfishers target prey that are


contribution may have evolved as
hunting large animals became a focus of too small to be
widely shared but are

competitive display.
especially difficult
to capture. Signal



content can help
explain why men



forego sardine fishing
or shell-fish col-



lecting which could
earn them a higher



production of the display and the


Displays are a form of communica-


rate of nutrient
acquisition than the ac-


tion, providing information about an
quality being advertised. Honesty is



tivities they do
choose.


individual, often in a widely observ-
ensured when individuals of higher



Costly signals are
enormously vari-


able forum. In order for a display to
quality can pay highest costs to pro-



able. One important
dimension of

be worth performing, there must be
duce a more elaborate display or



variation is between
signals that pro-


an audience. In order for observers to
when individuals gain higher benefits



vide little but
information and those


bother paying attention, it must ben-
for producing a display of given cost.7



that provide benefits
to the audience


efit them to do so. According to
costly The differential costs or benefits of

signaling theory,1,2,4 – 6 the
observer in addition
to information. For exam-



signal production make it highly un-


benefit for paying attention is the
in- ple, when a
display consists of provid-



likely that lower-quality individuals


formation about an otherwise hidden
ing feasts, others
gain from participat-



will be able to fake the signal: they


quality that is conveyed by the dis-
ing in the feast. By
signaling in this


cannot afford to. The honesty of the


play. The information is kept honest
way, the show-off
provides something



information provided in a display


through intrinsic links between the
besides information
about a hidden



gives an immediate benefit to observ-



quality to the
audience.



ers because they can use the informa-


Zahavi1,2,3,17,18
has applied his



tion to adjust their own behavior



handicap principle to
both kinds of



toward the show-off to benefit them-



display, proposing
that the “altruism”



selves. Show-offs benefit from the

Kristen Hawkes, Professor of
Anthropol-



of the second kind, in
which the fit-



treatment that follows. Bluffs about


ogy at the University of Utah, uses
evolu-


tionary ecology to investigate age
and sex



ness-related benefits
other than infor-


the show-off’s quality would not in-


differences in human foraging
strategies,



mation are supplied to
audiences, can



form the audience, so only signals that


with special emphasis on problems in
hu-


man evolution.
be explained by its
contribution to the


are too costly to fake are reliable. Za-



effectiveness of the
signal. For exam-



havi’s label, “the handicap principle,”


Rebecca Bliege Bird, Research
Assistant


Professor in the same department,
uses



ple, among the
cooperatively breeding


underlines the paradox that it is cost


behavioral ecology to investigate
age and



birds he has studied
for decades (Ara-



to the signaler that makes displays


sex differences in human foraging
strate-


gies, with special emphasis on
explaining bian babblers,
Turdoides squamiceps),


honest enough to be worthy of audi-


ethnographic variation.



dominants display
their quality by



ence attention.



standing sentinel
duty, distracting or



Costly signaling models are proving


fending off predators,
and presenting



useful for unraveling an array of an-


Evolutionary Anthropology 11:58 – 67
(2002)



Evolutionary
Anthropology 59


ARTICLES



men. When hunters target large prey,


food to subordinates. Displays that
ity of this sharing strategy
depends on

supply such benefits are readily noted
the exclusion of
nonreciprocators.



and when others can learn about and


by other babblers. Demonstration of
However, no quantitative
studies of



compare their successes, hunting rep-


the capacity to bear the cost of these
food sharing have found
consumers



utation becomes a prominent deter-


displays substitutes for overt threats,
repaying the hunter by
returning


minant of how desirable a neighbor


which potentially lead to fights that
shares of meat to him (see
Box 2).



and ally, and how dangerous a rival, a


could be even more costly for the sig-
Without strictly contingent
sharing, in



man might be.24,25


naler and at least some of the audi-
which providing meat at one
time is



One treatment of the show-off hy-

ence. Using the label “conspicuous
do- the necessary price for
claiming meat



pothesis for men’s work emphasizes


nations,” Zahavi underlines, as did
at another, those who
free-ride on the



the value the audience places on meat


Veblen19 in his classic 1899 analysis,
work of others net greater
nutritional



and the nutritional gains they realize


the similarity between public generos-
gains than do the workers.


from preferential association with


ity and “conspicuous consumption.”12
If meat distributions
do not exclude



show-offs, both of which give success-



free-riders, then the meat
of large an-



ful hunters latitude in pressing their



imals can be rather like a
public good.


own interests.23 It is proposed that a


HUNTING AS DISPLAY



If supplied by one,
consumption ben-



man’s reputation as a hunter affects


Recognition that signal content



efits will be claimed by
many. When


the way that others treat him because


may play a role in the evolution of



individual effort supplies
goods that



of the nutritional benefits they expect


socially productive behaviors has im-



also benefit others,
individual and col-


from living in the same group with him.


plications with respect to variability



lective interests in those
goods can be



This argument depends on an im-


in the relative subsistence contribu-



at odds, resulting in
collective-action



portant inference about the wide shar-

tions of males and females. Among



problems.28 Because
consumers can


foragers, men produce, on average,



benefit whether or not they
them-


from 30% of all calories to nearly



selves are providers, public
goods are



Recognition that signal

100%, if one does not count a wom-



often under-supplied.29 As
with com-


an’s processing and tool preparation
content may play a role munity defense or public
radio, non-


as contributing to production.20 These



in the evolution of providers can free-ride, so
that a pro-


long-term averages often include ex-



vider’s own expected
consumption

tremely high short-term variation, in-
socially productive gain from such goods is not
enough


cluding periods when men provide lit-



behaviors has incentive to supply them.
Why pay for


tle or nothing. Where men’s average



what you can get for free?30
This prob-



implications with respect


caloric production is disproportion-


lem arises with any good
that is not


ately greater than women’s, men al-



to variability in the the private property of the
producer.


most always spend most of their for-



The distinction between
private and



relative subsistence


aging time hunting large game


public goods turns on
whether or not


animals and then sharing them



contributions of males users can exclude other
claimants and


widely, producing more calories for



whether anyone’s
consumption of the



and females.


the group than for their own house-


good affects the consumption
benefits


holds. The traditional explanation for



available to others. A
perfectly public


this pattern relies on two often un-



good cannot be used
exclusively by


tested and somewhat paradoxical as-



anyone. Also, consumers can
use it

sumptions: that such hunting is part



ing of meat. Ethnographers have long concurrently; that is,
consumption by


of the most efficient energy or
protein



noted that people in small face-to-face one does not reduce what is
available


maximization strategy and that such



communities share food readily.26,27 to others. Ostrom and
Ostrom31 use-


hunting is inefficient when attempted


Anthropologists from diverse theoret- fully distinguished these
two indepen-


by women. Because these two as-



ical traditions have favored explana- dent dimensions, labeling
them “ex-


sumptions may not always hold,14



tions for this sharing that focus on clusion” and
“subtractability.” Because


some other explanation is warranted.



reducing the nutritional income vari- both excludability and
subtractability

Men’s contribution to subsistence



ability of unpredictable resources (see are more continuous than
discrete,


may have evolved and may persist be-



Box 1). This explanation has been es- few goods are perfectly
private or per-


cause men establish and maintain



pecially favored for meat sharing. fectly public. But some
goods allow


their relative social standing by show-


Game animals are large enough, and more concurrent consumption;
and


ing off their hunting prowess. Various



hunting is risky enough, that if hunt- sometimes exclusion is
impossible, or


versions of that show-off hypothesis



ers owned the meat of the prey they its costs are too high to
pay.32,33


propose that hunters attract the favor-



killed they would increase the average Because hunting successes
can be

able attention of many potential con-



nutritional utility they derive from it so unpredictable and the
meat then go


sumers by acquiring foods that are



by giving shares they value little when mostly to others, whether by
demand


widely consumed.12,15,16,21–23 The
in-



they have a lot in exchange for the sharing, tolerated theft, or
normative


terest all have in the meat acquired by


larger benefit of shares to be repaid rules of distribution, the
nutritional


hunters makes hunting a central


arena for social competition among
benefits a hunter can
expect for him-



later when they have little. The stabil-


60 Evolutionary Anthropology

ARTICLES


Box 1.
Reciprocity in the Sharing of Display Game



gauging the interest and appetite of
others would do better

Models of sharing used by
behavioral ecologists typi-



not to start fights they would be likely
to lose.


cally incorporate Trivers’86 model
of reciprocal altruism in



The exchange model, in contrast,
assumes that suppli-


which an individual pays a short-term
cost that benefits



ers own whatever they acquire. They thus
incur a cost

someone else but nets an overall
profit when the benefi-



when giving up shares, but net a
compensating benefit


ciary later returns the favor.
Trivers highlighted the sub-



from subsequent repayment. The repayment
is necessary


stantial benefits that reciprocal
altruists would accumulate



for reciprocal altruism to be stable.
Nonreciprocators must

as compared to nonsharers if they
could somehow over-



be excluded, something that becomes
increasingly difficult


come their vulnerability to
exploitation by recipients who



when strategies are more variable and
groups include


did not repay. His ideas were
supported by formal model-



more than a few individuals.90 –93
Repayment can be espe-

ing and computer simulations designed
to tease out the


precise conditions that allow
reciprocal altruism to be evo- cially difficult to enforce when
sharing is highly visible, as


lutionarily stable. That work showed
that reciprocators when very large game animals are acquired
and successes


could do well if they clustered
together and if the range of are unpredictable, with the hunter’s
daily risk of failure


alternative strategies was sharply
limited. In the case of generally increasing with prey
size.24,25,54 When a big-


humans, Trivers suggested that
features of our emotional game hunter is successful, there is
a great deal of meat

architecture such as guilt and
moralistic aggression indi- and many who are hungry for it, and
many of them are


cate an evolutionary design for
reciprocal altruism. Pair- armed with lethal weapons.33 Hunters
themselves often do


wise exchanges of private goods among
humans can be not control the distribution, so they
cannot direct shares to


highly reciprocal and subject not
only to stringent control, or away from particular individuals
based on either debts


but also conventions about valuation
and expectations or prospects.25,40,50,55–57
Quantitative records of meat dis-


regarding the timing and quantity of
returns (for example, tributions over time often find
claimants continuing to get

Hxaro gift exchange,87 Kula88).
shares whether or not they ever supply
them, and hunters



continuing to supply more meat even when
others are


But human sharing can also involve
multiple recipients



deeply in their debt.48,51,56,57,94 When
this is so, the ques-


and have few, if any, of these
features. In cases where food


flows to multiple recipients and few
are excluded, sharing tion is why hunters continue to expend
their effort supply-

may not be exchange at all. Instead
of trading shares with ing goods that go mostly to others.


each other, claimants may be
appropriating shares from It could be that in spite of
the evidence that they do not


the “public domain.” Blurton
Jones32,33 noted that food control distributions of their
prey, hunters are repaid by



recipients in some other currency.47 This
would mean that


sharing need not imply owners paying
a cost to give up


shares. Instead of ownership rights
falling automatically on the meat is not like a public good after
all, but instead that

the acquirer, there could be a cost
for defending shares consumers are, in some undetermined way,
paying every


from the claims of other users. The
costs of not sharing hunter for each share. While this would
not conform to the


could sometimes be too high to be
worth paying.89 The nutritional variability reduction
models of sharing,27,95,96 it


Blurton Jones “tolerated-theft”
model showed that sharing would conform to Trivers’86 model.
Although the search for


could result if resources came in
large but divisible lumps, the currency in which to find
repayments continues, both


but not to everyone at once, and if
consumers were pre- theoretical and empirical work has
increasingly stimulated

pared to press claims for a share
according to the nutri- researchers from many fields to
consider other explanatory


tional value of the resource to them.
As he noted, this need pathways to the evolution of cooperation,
sharing, and the


not mean incessant squabbling, since
potential claimants provisioning of public
goods.8,18,83,97–102



some paid special attention to a here.12,35,36 If
audience members can


self and his family are not great



hunter, others might be able to free-

enough to make hunting an effective
pay attention to
the hunter and thus eat



ride, consuming meat brought down


provisioning strategy. This is strong
more of his meat
without doing any-



by a hunter without paying special at-


provocation to look for other reasons
thing that benefits
him, the hunter’s se-



tention to him. In hypothesizing that


why men hunt. Olson28 noted that se-
lective incentive
will disappear.


preferential attention is the selective Smith and
Bliege Bird12,37 and


lective incentives, benefits that went



incentive motivating hunters, Hawkes23


only to suppliers, could motivate a
Bliege Bird,
Smith, and Bird15 ad-


supply of collective goods. Selective
dressed these
issues by using costly



defined that problem away. Like the


incentives could draw men into hunt-
signaling as a
model for the payoffs to


attention foragers pay to more pro-


ing. Men may be enticed to hunt be-
hunters and
observers. These models



ductive patches by monitoring them


cause doing so earns them differential
stress the
information that others gain



more closely, preferential attention to


social attention. Rather than because
as the key to
evolutionary stability.



better hunters was assumed to increase

of the nutritional value meat has for
More than its
value as a source of



the chances for consuming meat.23,34


them or their families. This could pose
nutrition, meat is
a medium of com-



However, some still see a second-


another collective-action problem. If
munication through
which the hunter



order collective-action problem lurking



Evolutionary
Anthropology 61

ARTICLES



greater social, political, and reproduc- women as lovers and as
secondary fa-


transmits information to potential



thers of more children.
(Secondary fa-


mates, allies, and competitors. Collec-
tive success than do poorer competi-



thers are men other than
a mother’s


tive-action problems do not arise in
tors. Ache foragers of eastern Para-


husband who were
sexually involved


handicap models because it is mutu-
guay are an especially well-studied



with her at the time of
her pregnancy).


ally beneficial to both show-off and
case.43 When living in the forest, Ache



Ache women did not
nominate hunt-


audience to have the information
men spend nearly fifty hours a week in



ing skill as a criterion
for choosing a

about the show-off’s qualities re-
food acquisition,44 supplying a very



mate, but men emphasized
its impor-


vealed. Show-offs obtain differential
large fraction of a diet that is uncom-



tance for success with
women.43


treatment only by paying the signal
monly ample among modern hunter-


cost; signal recipients obtain informa-
In other ethnographic
cases, hunt-



gatherers.45 The generous nutrient av-

tion about a signaler’s quality only
by ing success is also
associated with ad-



erages result not from high hourly


attending to the signal.
vantages in male
competition. Hadza



foraging return rates but from these


If men hunt to display their
relative men foraging in
northern Tanzania



long hours,46,47 with better hunters


quality, then the benefits they earn
for are big game
specialists52–54 (Fig. 1).


spending the most time hunting48 (see


that effort come not from exchanges
As among the Ache,
hunters do not



Box 3). Wide sharing is especially well


of meat for other goods and services,
control the distribution
of meat.55–57



documented in this case.49,27 While


but from the different ways that oth-
In this case, the wives
and children of


ers treat them in light of the quality
better hunters do have
more positive


Collective-action


they reveal. Others use the informa-
weight gains,58 and
those wives have


tion of hunting reputations to their
surviving children
faster.59 But these



problems do not arise in


own advantage in the numerous deci-
differences are directly
associated



handicap models


sions of social life.
with the foraging effort
of the women


themselves.60,61 As with
the Ache, the



because it is mutually wide sharing of meat
means that


THE HUNTER’S INCENTIVES



beneficial to both show- Hadza women and
children receive


If hunting is a signal of quality,
then little of their meat
from kills by their



off and audience to

the hunter’s benefit does not depend
husband and father.
Consistent with



have the information


on collecting repayments from each
this, a father’s death
or parental di-


recipient of meat. Rather, hunters in-
vorce has no effect on
child survival.62



about the show-off’s


crease their prestige by contributing
However, better Hadza
hunters tend



qualities revealed.

more than others do (See Box 3). Za-
to be married to
harder-working


havi has observed babblers demon-
wives.61 Older men who
are better



Show-offs obtain


strating superior stamina by compet-
hunters have younger
wives,59 sug-



differential treatment


ing to invest more in costly sentinel
gesting they are more
likely to leave


duty.3,17,18,38 While others benefit



only by paying the an older wife to raise a
second fam-

from this public good, the providers
ily—another way they
have increased



signal cost; signal


earn prestige for their quality.
“Social success in
competing for paternity.


prestige functions like a peacock’s
tail recipients obtain Meriam turtle
hunters also have


or the song of a songbird. It attracts
higher age-specific
reproductive suc-



information about a


collaborators and deters rivals.”3
cess than do
nonhunters and, as with



signaler’s quality only by

In the human case, a reputation for
the Hadza, this seems
due to assorta-


good hunting generally affects a man’s
tive mating: hunters
claim more fer-



attending to the signal.


social standing relative to other men
tile wives than do
nonhunters.63


in foraging communities.20,39,40 In the


northern Kalahari, traditional hunters


usually bagged no more than two or
COMPARISONS WITH


three large antelope in a year.41


the Ache are on forest treks, any hunt- CHIMPANZEE
HUNTING


Thomas42 reported the characteriza-



er’s prey is distributed to all. The



Studies of the
benefits chimpanzees


tion of a famous man with the hyper-



hunter himself has no hand in this



obtain by hunting red
colobus mon-

bole that underlines the value placed



distribution, and so does not direct keys suggest some
parallels with hu-


on hunting success:



shares preferentially to particular re- man hunting. In that
species, hunting


It was said of him that he never
cipients.50 The families of better hunt- is also a male
specialty, and meat is


returned from a hunt without
ers end up with no more meat than more widely shared than
are other


having killed at least a wilde-
other families.27 Hill and Hurtado’s43 foods. Accumulating
evidence sug-

beest, if not something larger.
demographic data show little differ- gests that chimpanzee
hunting is best


Hence the people connected
ence in survival risk for the children of explained as a male
strategy for gain-


with him ate a great deal of
better hunters. But men rated as bet- ing and maintaining
higher sta-


meat and his popularity grew.
ter hunters had much higher fertility. tus.64,65 Stanford and
coworkers66



In a smaller data set,51 better Ache


There is increasing evidence that
found that the three
variables most

good hunters in many societies enjoy
hunters were more often named by strongly associated with
the likeli-


62 Evolutionary Anthropology

ARTICLES


Box 2. Sharing Among the Hadza and
the Meriam: Do Those Who Supply More Meat Receive



More Meat From Others?



the Meriam, those
households who


When sharing is exchange, some-
patterns suggests that reciprocity-



supply meat in greater
quantities (A

one is giving up a share, and so in-
based explanations are more likely to



and B) or who supply it
more fre-


curring a cost to get a repayment in
apply to the sharing of nongame or



quently (C and D) do
not seem to be


return. What people say about their
nonforest resources. Game, espe-



rewarded for their
generosity with

“rights” as claimants is relevant
to cially large game, which often is



more meat supplied by
others or


this question, but it is also of
interest shared in public, seems not to follow



more frequent
receipts.57,94 The Me-


to see, whatever the stated rules of
the same sharing rules as do other


ownership, whether there is a “quid
resources. riam data come from a
systematic

pro quo” pattern in the actual flow
of Are sharers rewarded for the survey of the
distribution patterns of


shares. Tests of reciprocal altruism
in shares they supply with future shares all turtles acquired
by island house-


human food-sharing patterns some-
of the same high-variability resource? holds during one year.
The Hadza


times show that recipients do repay
This is the central assumption of the meat-sharing data come
from a sam-


suppliers.103 Sometimes this applies
“risk-reduction reciprocity” hypothe- ple of shares from
large game (Fig. 1)



sis that hunters exchange meat to re- carried to households
in a study

to certain kinds of food and not oth-



duce the variability in their nutritional camp that moved its
location and


ers. For example, Gurven and co-



income. Among both the Hadza and changed in membership
over time.


workers,104 analysis of Hiwi sharing


Evolutionary Anthropology 63

ARTICLES



Box 2. (Continued)



the Hadza, we can
immediately reject



just as much and just as frequently


Men could only be recorded as re-



this possibility
because the majority of


as those who supplied it most gen-


cipients or suppliers of shares
while



calories that
acquirers supply to other



erously. Sharing turtle among the Me-


co-resident with the observers.
Those



nuclear families are
from large game


riam and sharing large game animals


who were in the sampling window



animals, with only a
small proportion



among the Hadza seems not to be


longer were, other things the same,



coming from honey.
Could Meriam


conditional on getting portions from


more likely to be caught doing both,



households be trading
turtle calories



others. The benefits of supplying


which accounts for the slight but
still



for fish? This does
not seem likely: Me-


meat do not appear to come from


insignificant positive trend. After
ac-



riam households who
shared turtle dur-



getting more meat from others.


counting for the large variation



ing the nesting season
were not pref-


There is the possibility that shares of


among hunters in days resident, par-



erentially given fish.
Among eight



Hadza game and Meriam turtles are


tial correlations actually indicate
a



households sampled,
five never shared


traded for other resources. For the Me-


slight negative trend between both



turtle, yet received
an average of 50



riam, turtle makes up more than 80%


Kg shared and received (R
.243)



grams of fish per
capita per sample


of all calories of food transferred be-


and the frequency of sharing and re-



day, while the three
households that



tween households during the nesting


ceiving (R .194). Indeed,
among



shared turtle received
an average of 4


season, with fish and other marine re-


both the Hadza and Meriam, those



grams of fish.



sources making up the remainder. For


who never supplied meat received


hood of chimpanzee hunting over a
Chimpanzee males
could be drawn


sharing session that follows can


ten-year period at Gombe were the
into hunting because
control of meat



be enormously costly relative to


number of males in a party, total party
is an effective way
to display relative



the quantity of meat that is usu-


size, and the number of females with
quality. Unlike the
human case where,



ally available.

estrous swellings in the party. The
last as argued here, a
hunter’s own benefit



At Ngogo,64 males who shared meat


of these was the strongest predictor,
depends on credit for
the kill rather



with each other also shared coalition-


suggesting that hunting is more likely
than on control of
the meat, chimpan-



ary support. Mitani and Watts64 con-


when male mating competition is
zee males compete for
possession of

most immediate. Teleki67 found that
cluded that Ngogo males hunt to ob- the meat. Sometimes
they rip the car-


Gombe females were both more likely
tain meat they then use to develop and cass apart in the
process. Unlike other


to beg meat from males and more
maintain social relationships with forms of display
among male chim-


likely to get meat when in estrus.
other males. panzees, however,
hunting and meat


Stanford68 observed “meat for sex”
ex-


changes. But at other times and at


other sites this is less
common.65,69,70


During Mitani and Watts’65 observa-


tions at Ngogo, estrous females re-

ceived meat more often and anestrous


females less often than expected by


chance, but sharing did not affect the


probability or frequency of mating.


Overall, adult males consume most of


the meat, little going to females and


even less to juveniles.65,68 –71


At Ngogo, hunting was not a strat-

egy for meeting a nutritional
shortfall.


To the contrary, Mitani and Watts64


found that hunting increased when


there was more ripe fruit available.
This


is consistent with Stanford’s72
conclu-


sion based on observations at Gombe:


Most members of the hunting


party receive very little meat
for


Figure 1. Hadza men women and children returning home from a
death, butchery, and


their effort, and the number of
consumption site where all have been eating meat. They are
carrying household meat


chimp-hours expended on the
shares57 (see Box 2) back to the residential base. (Photograph
courtesy of J. F.


hunt plus the long begging and
O’Connell.)


64 Evolutionary Anthropology

ARTICLES



Box 3. Competition Intensifies Work Effort


to spend time as group sentinels.38 Ache
men hunt very


Kaplan and coworkers47 have
recently interpreted age-


related changes in foraging
productivity as evidence that long hours,44,47 and better Ache
hunters spend even more


foraging ability accumulates over
the life span as a result of time hunting.48 The same appears to be
true for the !Kung


increased practice and experience.
We emphasize an al- and Hadza as well, with better hunters
spending more time


ternative and additional influence
on productivity profiles: hunting.41,57 Better Meriam spear
fishers also spend more

changes in the benefits for working
longer hours or forag- time out on the reef.15 When some
individuals gain com-


ing with greater intensity.
Sometimes working longer hours petitive advantages over others
by engaging in activities


does not mean more food for one’s
own household but with display value, and when productive
activities are ef-


increased relative status among
group members. Arabian fective displays, increased
productivity can be the out-


babblers compete among themselves
for the opportunity come of status competition.



observation shows that this danger is

sharing result in more than informa-
to talk quietly of
the day. Then, with-


tion for the audience. At least some
out fanfare, someone
else, perhaps a



minimized because it is not a hunter


others get to eat meat as a conse-
boy, will step
outside the circle of fires



himself who touts his own exploits.


quence, a possible evolutionary foun-
and drag in any prey
left discretely at



Lee’s73 famous anecdote about the

dation for the much greater benefits
the margins of the
camp. This absence



properly self-effacing behavior of


that flow to others from some kinds of
of
self-aggrandizement among hunt-



!Kung hunters captures the common


human showing off.
ers seems initially
inconsistent with



pattern.74 But !Kung men talk end-


The male competition for status
the proposition that
hunting is dis-

seen in modern chimpanzee hunting
play, but if the
message is in the meat,


provides a hominoid foundation for
it is the reliable
links between a



The male competition


the evolution of human hunting. Mod-
hunter and the prey
that matters. The


ern human hunters often display in a
stories told by
others, accumulating



for status seen in


way that provides more nutritional
from the interwoven
observations of



modern chimpanzee

benefits to all. That difference can
be many, make that
link. As in other do-


related to other differences between
mains of male
contest, “trash talk”



hunting provides a


us and our sister species. One of those
may have its uses,
but reputations for



hominoid foundation for


differences is, of course, that human
delivering the goods
cannot be built



the evolution of human

hunters have the technology to cap-
upon it.


ture prey larger than themselves. An-



hunting. Modern human


other may be the inability of chimpan-



WHY IS MEAT THE
SIGNAL?



hunters often display in


zee hunting techniques to distinguish

effectively among the varying skill
lev- If it is
merely information that is



a way that provides


els of hunters. Another difference is
being transferred,
does this mean that



more nutritional benefits


especially relevant in the context of
signal form is
arbitrary (as in Fisher-


arguments here: We have language.
ian runaway sexual
selection) as long



to all. That difference

Among people, the story of a hunter’s
as information is
transmitted honestly



can be related to other


success spreads to a wide audience,
to the appropriate
observers? There


though few, if any, of its members
are good reasons to
think not. While it



differences between us


actually saw him capture the prey.
is just as costly
for signalers to give an



and our sister species.

Human reputations can be built
altruistic signal as
an equivalently


through storytelling, but chimpanzee
wasteful display,18
it may not be


reputations cannot: only those on the
equally beneficial.
Some observers


scene of hunting and meat sharing ep-
may be more
interested in “altruistic”


isodes can be signal recipients. Lan-
signals than
“wasteful” ones because



lessly about hunts and hunting, re-


guage broadens the audience to in-
such signals provide
more than sim-



hearsing the “minutest details.”41,75,76

clude all who hear the story and thus
ply information,25
because they pro-



All those who listen to the storytelling


may vastly increase the signaling ben-
vide different sorts
of information,37



soon know which man it was that


efits of displaying skill through game
or both. Peacock
tails,77 conspicuous



made every kill. The self-effacing style


acquisition.
leisure,19 or
spear-throwing accuracy15


also characterizes Ache hunters, who


Talk, however, is cheap. Tales of
give only
information to signal recip-



arrive at the evening’s forest camp


hunting might allow show-offs to
ients. Sentinel
duty, hunting, feasting,



without a word, whether they have


bluff about their successes, or at
least political pork,
and group defense send



taken any prey or not. Ten minutes or

claim near misses, without paying
information but also
benefit the audi-



more may pass before the men begin


real costs. Widespread ethnographic
ence in other ways.
Benefits other


Evolutionary Anthropology 65


ARTICLES



In the last few years, a combination of


than information can play a role in the
ioral ecology: an
evolutionary approach. Oxford:


Blackwell
Science. p 155–178.



empirical and theoretical work has


dynamics of selection for displays be-



7 Getty T. 1998.
Handicap signaling: when fecun-



found costly displays to be much


cause it is audience attention that de-
dity and
viability do not add up. Anim Behav


more widespread than was previously


termines gains to the show-off.
56:127–130.



recognized. Darwin focused on mate


Audience sensitivity (receiver bias)
8 Boone JL. 1998.
The evolution of magnanimity:



when is it better
to give than to receive? Hum Nat



choice and competition for mates, but


affects the nature of displays because


9:1–21.



if displays are signals, selection can


signals must be detected effectively by
9 Neiman FD.
1998. Conspicuous consumption



favor wasteful expenditure on them in


appropriate recipients.78 – 82
Because as
wasteful advertising: a Darwinian perspective



on spatial
patterns in the Classic Maya terminal


any kind of social interaction, includ-


public goods are consumed by many,



monument dates.
In: Barton CM, Clark GA, edi-



ing those between parents and off-


identification with those goods
tors.
Rediscovering Darwin: evolutionary theory



spring or predators and prey.


reaches a wide audience of consum-
and
archaeological explanation. Washington


D. C.:
Archaeological Papers of the American An-



Our arguments seek to explain the


ers. This “broadcast effectiveness”



thropological
Association, No. 7. p 267–290.



evolution of men’s subsistence work


might help explain the recurrence of



10 Miller GF.
1999. Sexual selection for cultural


as a strategy to compete effectively for


common goods provisioning among
displays. In:
Dunbar R, editors. The evolution of



social advantage in a world where


humans.12,83 Signals designed to ac-
culture.
Edinburgh: Edinburgh University Press.



p 71–91.



honesty is at a premium and political


quire or maintain higher social stand-


11 Miller GF.
2000. The mating mind: how sex-



alliances substitute for body size and


ing in a group should be directed to
ual choice shaped
the evolution of human na-



canine weaponry in gaining the ad-


the group at large; other more special-
ture. New York:
Doubleday.



vantages of status. We highlight the


ized signals may be directed to
12 Smith EA,
Bliege Bird R. 2000. Turtle hunting


and tombstone
opening: public generosity as



fact that, at least among humans, both


smaller subsets of the population. Sig-



costly signaling.
Evol Hum Behav 21:245–261.



signaler and audience preference for


nalers competing for popular prestige
13 Sosis R. 2000.
Costly signaling and torch fish-


more effective and competitive signals


should seek to gain a larger and larger
ing on Ifaluk
Atoll. Evol Hum Behav 21:223–244.



can drive the evolution of displays to-


share of the advertising market. They
14 Bliege Bird R.
1999. Cooperation and conflict:



the behavioral
ecology of the sexual division of



ward increasing social benefits. The


gain a larger share by providing more


labor. Evol
Anthropol 8:65–75.



hypothesis that men’s work evolved


of what the viewers want to see or
15 Bliege Bird R,
Smith EA, Bird D. 2001. The



and often continues to be shaped by


consume than the competition pro-
hunting handicap:
costly signaling in human for-



showing off does not imply that men


vides.25 The provisioning of collective
aging strategies.
Behav Ecol Sociobiol, in press.


16 Bliege Bird R,
Bird D. 2001. Gendered fishing



contribute little to subsistence. On the


goods may serve the purpose of reach-



among the Meriam:
implications for sexual divi-



contrary, the showoff hypothesis and


ing a wide audience better.37 Both
sion of foraging
labor, submitted.


costly signaling can help explain how


competition among signalers and au-
17 Zahavi A.
1990. Arabian babblers: the quest



individuals seeking competitive ad-


dience preference for particular sig-
for social status
in a cooperative breeder. In:



Stacey PB, Koenig
WD, editors. Cooperative



vantages can increase their own


nals can play a role in shaping the
breeding in
birds: long-term studies of ecology


standing and so earn preferential


display.
and behavior.
Cambridge: Cambridge University



treatment by acting in ways that sup- Press. p 103–130.



18 Zahavi A,
Zahavi A. 1997. The handicap prin-



ply highly valued benefits to others.


CONCLUSIONS
ciple: a missing
piece of Darwin’s puzzle. Oxford:


Oxford University
Press.


Darwin84 developed the theory of



19 Veblen T.
1992. The Theory of the Leisure



ACKNOWLEDGMENTS


sexual selection to explain the evolu-
Class. Mills CW,
editor. New Brunswick, NJ:


tion of armaments and ornaments,
Transaction
Publishers.



We thank Helen Alvarez, Douglas 20 Kelly RL.
1995. The foraging spectrum: diver-

which seemed so extravagantly costly



Bird, Nicholas Blurton Jones, Jack sity in
hunter-gatherer lifeways. Washington:


and remarkably wasteful, given an ex-
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Hirshleifer, James O’Connell, Mark


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Pagel, Lars Rodseth, Carel van Schaik,


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Pauline Wiessner for helpful com-


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ments and advice.


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Articles in Forthcoming Issues


• Crotchets & Quiddities: Is
the Medium the Message?


Kenneth M. Weiss


• What is Molecular Anthropology?
What Can It Be?


Jonathan Marks

• Primate Evolutionary
Developmental Biology


Chi-Hua Chiu and Mark W. Hamrick


• Eocene Primates From Myanmar


Russell Ciochon and Gregg Gunnell


• Forest People: The Role of
African Rainforests in Human Evolution


and Dispersal

Julio Mercador


• Missing Links: Eugene Dubois and
the Origins of Paleoanthropology


`


Pat Shipman and Paul Storm


• The Behavioral Ecology of the
Spectral Tarsier, Tarsius spectrum


Sharon Gursky

• Primate Origins and Adaptations:
A Multidisciplinary Perspective


Mary Silcox








11-2


Behav Ecol Sociobiol (2001) 50:9–19


DOI 10.1007/s002650100338


O R I G I N A L A RT I C L E

Rebecca Bliege Bird · Eric Alden
Smith


Douglas W. Bird


The hunting handicap: costly signaling
in human foraging strategies


Received: 31 March 2000 / Revised: 10
January 2001 / Accepted: 5 January 2001 / Published online: 31 March
2001


© Springer-Verlag 2001



rates, provide widely shared goods without
repayment,


Abstract Humans sometimes forage or
distribute the


or incur an increased risk of injury.
These decisions have


products of foraging in ways that do
not maximize indi-



often been explained by the benefits of a
sexual division


vidual energetic return rates. As an
alternative to hypoth-



of labor in parental investment (e.g.,
Hurtado et al.


eses that rely on reciprocal altruism
to counter the costs



1992), the optimization of macronutrients
(e.g., Hill

of inefficiency, we suggest that the
cost itself could be



1988), or reduction of consumption
variance (e.g., Smith


recouped through signal benefit. Costly
signaling theory



1988). In particular, the practice common
to many hunt-


predicts that signals can provide
fitness benefits when



er-gatherer societies of widespread or
group-wide shar-


costs are honestly linked to signaler
quality, and this in-


ing of large prey captured by any subset
of the group is


formation is broadcast to potential
mates and competi-



conventionally explained as a form of risk
reduction


tors. Here, we test some predictions of
costly signaling



where all ultimately benefit from a
reciprocal sharing


theory against empirical data on human
food acquisition



of unpredictable harvests (reviews in
Hawkes 1992;

and sharing patterns. We show that at
least two types of



Winterhalder 1997).


marine foraging, turtle hunting and
spearfishing, as prac-



While some types of food seem to be
distributed in


ticed among the Meriam (a Melanesian
people of Torres



ways that conform to delayed reciprocity
(Gurven et al.


Strait, Australia) meet key criteria
for costly signaling:


2000), recent studies (Hawkes 1993; Bliege
Bird and


signal traits are (1) differentially
costly or beneficial



Bird 1997) have challenged its ability to
explain the


in ways that are (2) honestly linked to
signaler quality,



wide variety of sharing patterns to which
it has been


and (3) designed to effectively
broadcast the signal. We



claimed to apply. The cases that seem
least congruent

conclude that relatively inefficient
hunting or sharing



with strategies of delayed reciprocity are
those associat-


choices may be maintained in a
population if they serve



ed with public distributions of food in
which the acquirer


as costly and reliable signals designed
to reveal the



does not control access to the harvest or
its distribution,


signaler’s qualities to observers.


and may not even reserve a portion for
him- or herself


Keywords Costly signaling ·
Human behavioral ecology (Wiessner 1996). Under these
conditions, acquirers seem


· Hunting · Handicap
models unlikely to be able to
direct shares to reciprocators and



withhold them from free-riders, as is
required to solve



the collective action problem (e.g.,
prisoner’s dilemma)

Introduction
associated with enforcing delayed
reciprocity. Similar



challenges to the primacy of reciprocal
altruism (RA) on


Human foragers often make decisions
that lead them both theoretical and empirical fronts
are also found in


to bypass alternative activities with
higher energy gain recent literature on non-human behavior
(reviews in



Dugatkin 1997; Pusey and Packer 1997).
Many theorists



have suggested the paradigm may require
extensive


Communicated by M. Borgerhoff Mulder


revisions (Clements and Stephens 1995;
Connor 1996;


R. Bliege Bird (✉) · D.W. Bird
Dugatkin 1997; Roberts 1998).


Department of Anthropology, 270 S. 1400
E., University of Utah, An alternative explanation for such
“economically ir-


Salt Lake City, UT 84112, USA



rational” decisions is that under some
circumstances they


e-mail: r.bird@mindspring.com



could serve as an honest signal of one or
more dimen-

Tel.: +1-801-5814494, Fax:
+1-801-5816252



sions of fitness-related quality (Neiman
1998; Boone


E.A. Smith



1998; Sosis 2000; Smith and Bliege Bird
2000). Costly


Department of Anthropology, University
of Washington,



signaling theory (CST) provides a powerful
framework


Seattle, WA 98195, USA

10


for explaining two paradoxical
observations: the persis- successful turtle hunters signal
strength, risk-taking, and


tence of wasteful phenotypes when
natural selection is (in the case of hunt leaders) a variety of
cognitive and


assumed to create greater efficiency,
and the evolution of leadership abilities to potential allies,
mates, and com-


honest communication despite the
pervasive conflicts of petitors.


interest underlying evolutionary
processes (Zahavi 1975,


1977; Grafen 1990; Johnstone 1995,
1997; Getty 1998).



Methods

CST proposes that communication between
individuals


with conflicting interests can be
evolutionarily stable if



Mer (a.k.a. Murray Island) is a small (1.6×2.2
km) island on the


the signal honestly advertises an
underlying quality of northern end of the Great Barrier Reef,
140 km from New Guinea


interest to observers. Advertising is
kept honest and thus in Australia’s Torres Strait. The island’s
current population is 430


mutually beneficial to both signaler
and observer as long individuals of Torres Strait Islander
descent, scattered in approxi-



mately 85 households. The Torres Strait as a
whole is adminis-

as the cost or benefit of advertisement
is so closely tied



tered by the State of Queensland and the
Commonwealth of


to the quality of the signaler that
faking it costs more



Australia.


than the signal is worth. If this
holds, the correlation be- Prior to about 1975, when Australian
welfare payments were


tween the quality of a signaler and the
quality or intensi- first made available to all indigenous
Australians, the Meriam


ty of the signal will be maintained by
differences in mar- were nearly full-time subsistence
horticulturalists and marine for-



agers planting tropical yams, bananas, sugar
cane, and introduced

ginal cost or benefit, allowing
recipients to reliably dis-



new world crops such as manioc, sweet potatoes,
and corn, and


criminate among competing signalers.
When these con- harvesting marine fish, shellfish, and sea
turtles. Today, fishing


ditions are met, honest signals and
reliable communica- and shellfish collecting remain a critical
component of Meriam


tion will be evolutionarily stable,
even when signaler and subsistence economy: mean per capita
after-sharing consumption



rates average 630 kcal of meat and 40 g of
protein. More than 80%


recipient are antagonists or
competitors.



of these calories are supplied by turtle when in
season. For addi-

The hypothesis that hunting might
serve as a form of tional ethnographic description and previous
work among the


status competition among men interested
in “showing Meriam, see Haddon (1906), Beckett (1988),
Sharp (1993), Bliege


off” to a public audience is not new
(Hawkes 1990, Bird et al. (1995), Bird and Bliege Bird
(1997, 2000), Bliege Bird



and Bird (1997), and Smith and Bliege Bird
(2000).


1991). Here, we recast the “show-off”
model using a



Observations of Meriam foraging choices,
time allocation, and


payoff structure compatible with CST
(Smith and Bliege


food-sharing strategies reported here were
conducted over several


Bird 2000). We propose that individuals
of high pheno- periods totaling 27 months between January 1993
and July 1999.


typic quality might reap higher
benefits or pay lower Much of this field research was
specifically designed to test hy-


costs to acquire skill-based resources
or to uncondition- potheses related to the origin and maintenance
of the human sexu-



al division of labor, and to determine the
nature of the tradeoffs af-


ally share their harvest. These
benefits (material, politi-



fecting men’s and women’s foraging decisions.

cal, and reproductive) flow from
observers who find it in Time allocation to intertidal
activities (spearfishing, shellfish


their interest to behave in ways that
improve the relative collecting) was measured through focal
individual follows occur-


advantage, status, or social dominance
of signalers, ulti- ring during randomly selected days during the
spring ebbing tide.



Locations along the foreshore were observed from
the midpoint of


mately enhancing signaler fitness.
Analyzed according to



the ebbing tide, before the reef was fully
exposed, for at least 2 h


CST, this is not delayed reciprocity
involving an ex- or until the last forager had left the reef.
During the intertidal sam-


change of substance (to the observer)
for social status (to pling period, we recorded the moment-by-moment
behavior of at

the signaler), but rather a form of
by-product mutualism. least one individual to arrive during the
sample, and if the individ-



ual remained after the interval ended, we
remained to completely


The payoff to the observer comes from
the usefulness of



record the episode. We observed the reef flat
habitat for a total of


the information inferred from the
signal: he or she



118.5 h over 41 spring ebb-tide days, recording
338.1 forager-


should be able to evaluate the
signaler’s suitability as a hours of subsistence activity in 210
partial and full foraging fol-

competitor, mate, or ally by attending
to the signal rather lows of 94 men, women, and children entering
the reef flat. We


than through more costly or unreliable
means of assess- then analyzed the percentage of total foraging
time each forager



devoted to all potential intertidal activities
during the follow.


ing the signaler’s abilities or hidden
qualities. The high



We analyzed the reef as a bounded habitat
within which there


cost or low benefit of faking the
signal guarantees that are “hunt types” sensu Smith (1991).
All hunt types are simulta-


signalers will not engage in false
advertising, and that neously available options within the
ecological habitat “reef flat at


observers will pay attention. Costly
signaling can thus low tide”: reef flat collecting, rocky
shore harvesting, and spear-


fishing (Bird and Bliege Bird 1997, 2000). Reef
flat collecting in-


spread (by natural selection or
imitation) because of its



volves mobile search of shellfish, with in situ
processing to in-


mutual benefits to both signalers and
observers. crease the utility of the load (Bird and
Bliege Bird 1997). Spear-


Here, we test the costly signaling
hypothesis among fishing involves search and travel across the
reef looking for en-


the Meriam (a Melanesian people of
Torres Strait, Aus- counters with mobile prey (small fish and
squid 250 g+) to the ex-



clusion of all other prey; when prey are
detected, the hunter stalks

tralia). We evaluate two candidate
foraging activities



the prey and launches his spear from a distance.
Both hunt types


(spearfishing on the reef at low tide,
and hunting turtle



occur in the same patch during the same period
of time, while the


for public feasts), to determine
whether these hunt types reef is exposed at low tide. The hunt
types vary in how spatially


meet key criteria for costly signaling
in being (1) differ- exclusive they are within the habitat:
spearfishing and shellfish


entially costly or beneficial in ways
that are (2) honestly collecting are the only two major hunt types
in which the forager



engaged in exclusive search for prey frequently
encounters and ig-

linked to signaler quality, and (3)
designed to effectively



nores prey in the other hunt type. These are not
the complete com-


broadcast the signal to the intended
audience. We pro- plement of hunt types simultaneously
available, but they are the


pose that accomplished spearfishers
signal such qualities three in which adults spend more than 90% of
their foraging time


as hand-eye coordination, stealth, and
patience, while while on the reef at low tide. Other minor hunt
types are handline


11



turtle shares are distributed in large,
uncooked portions (10–11 kg)

fishing from the reef edge and in
deeper lagoons, netting sardines,



of meat, fat, organs, and eggs among
nearby households, with the


hunting octopus, and diving from the
reef edge to take underwater



size of portions kept by the butchering
household determined pri-


prey by hand. Gross foraging return
rates for each hunt type (not



marily by the number of demanders
(hereafter termed “household


including energetic expenditure –
assumed to be roughly equal for


consumption” sensu Bliege Bird and
Bird 1997). While hunting a


all reef hunt types) were calculated
from the moment an individual



turtle is a costly activity in which
the benefit is acquired through


stepped onto the exposed reef and began
to engage in targeted



the social value of the hunt,
collecting a turtle is an activity which


search for particular prey types. The
macronutrient content of fin-



has little signaling potential and in
which the benefit is primarily

fish and shellfish was obtained through
published sources and



nutritional. However, there may be some
signal value of displays


through individual analysis of samples
collected on Mer (Hirth



of generosity among neighbors as turtle
portions are shared ac-


1971; Sidwell 1981; Brand Miller et al.
1993).



cording to the Meriam ethic of debe
tonar (“the good way,” which


For a detailed description of
methods used to collect turtle


involves sharing without expectation of
return).


hunting and sharing data see Bliege
Bird and Bird (1997); the re-



All statistical analyses were
performed using Statview (SAS).


sults presented here on turtle hunting
and sharing are a reanalysis



Large-sample means tests were performed
after testing for normal-


of those data. The acquisition of over
120 turtles was noted



ity and equality of variance using
either one- or two-tailed t-tests

through daily sampling of the entire
village during the nesting sea-



depending upon the prediction tested,
while tests on small samples


son (October–April) in 1994–1995,
and ad lib sampling of the vil-



and those violating the assumptions of
parametric tests used two-


lage during the 1994 and 1998 hunting
seasons (May–September).



tailed Mann-Whitney U-tests. All means
are reported with associ-


Nearly every successful hunt or
collection of turtle was recorded,


ated SEs.


with information obtained on
acquirer(s), place acquired, method


of acquisition, and subsequent
distribution though informant inter-


view.


There are two primary types of
marine turtle acquisition on



Results


Mer: turtle hunting (nam deraimer) and
turtle collection (nam ter-

pei). Turtle hunting occurs throughout
the year, but is the only


way to acquire turtles between May and
September (Kob Kerker), Spearfishing as costly signal


when green turtles (Chelonia mydas)
feed and mate on shallow


reefs about 16–20 km from Mer. Field
observations indicate that



There are significant sex differences
in time allocation to


turtles captured on hunts range from
100 to 150 kg live weight,



hunt types in the intertidal: on
average, men spend 63%


with an average edible yield of 50.1 kg
(Bliege Bird and Bird

1997). Hunters head out to the hunting
grounds in open boats of their reef foraging time spearing,
while for women


powered with outboard motors, often in
cooperation with at least this value is only 9% (t=6.00, df=66,
P=0.001), and


one other hunting boat. Among Meriam
turtle hunters, there are



women spend 76% and men 31% of their
time shellfish


three distinct roles: hunt leader
(ariemer-le), jumper (arpeir-le),



collecting (t=4.50, df=66, P=0.001).
The majority of


and driver, or tiller-man (korizer-le).
Hunt leaders organize and di-



men never collect shellfish at all, nor
do they combine

rect the hunt; there is always only one
leader per hunt, regardless


of crew size. Hunt leaders bear the
cost of organizing the hunt and spearing with shellfish collecting
in a single visit to the


ensuring boats and fuel to spare. They
direct the crew to particular



reef at low tide: 78.6% of 21 men in
the time allocation


locations, decide whether prey
encountered is worth pursuing,



sample spent all of their foraging time
spearfishing.


orchestrate the chase, and direct
jumpers when to jump from the



Is spearfishing a signal? The
decision men make to

boat to secure the turtle. Hunt leaders
are invested with public


recognition and receive full credit for
the kill regardless of spear fish nearly exclusively (rather
than collect shell-


whether or not they directly
participate in capture. fish) violates simple
energy-maximizing prey choice


The hunt proceeds as follows. While
one man drives, the rest



models, because (1) on average,
continuing to search for


of the crew stands toward the bow
scanning the reef for signs of



fish to spear (292±135 kcal/h,
n=26) offers lower overall

turtle. The hunt leader directs the
driver and coordinates with


crews in other boats if present. When a
turtle is spotted, the hunt energy returns than shellfishing
(1,492±173 kcal/h,


leader makes a decision whether to
pursue it based on its size n=47) while in the reef flat at
low tide (t=4.672, df=71,


(large turtles have more meat) and sex
(female turtles have more



P=0.0001) and (2) on-encounter returns
for the most


fat). The boat(s) then give chase,
keeping the turtle away from the



skilled spearfisher (2,505±778
kcal/h) are equivalent to

reef edge. When the turtle tires, the
hunt leader usually directs his



the on-encounter returns for the
lowest-ranked shellfish


jumper to launch himself from the bow
of the boat with a rope at-


tached to his upper arm. The jumper
then attempts to secure the prey still in the optimal diet
breadth (2,214±414 kcal/h)


turtle by locking his arms around the
flippers and, if successful,



(see Table 1).


the crew then pulls him and the turtle
on board.


One explanation for this pattern
might be that spear-


Turtle hunting occurs primarily in
the context of public feast-



fishers are not maximizing calories,
but other macronu-


ing events: hunters choose to hunt in
response to a request from


feast organizers to provide turtles for
consumption at a previously trients. But this appears not to be the
case: protein return


announced feast. The biggest and most
elaborate Meriam feasts rates from shellfish collecting at
284±31 g/h are higher


occur in the context of coming-of-age
celebrations and funeral


than from spearfishing at 6.6±2.9
g/h (t=6.78, df=68,


ceremonies (see Smith and Bliege Bird
2000 for further details).



P=0.0001). Fat return rates are also
higher for shellfish


In contrast to turtle hunting, turtles
are also collected (n=88


events), primarily in the context of
household provisioning, but collecting at 22±3 g/h than
for spearfishing at 1.6±


also for feasts, by men of all ages,
women, and children. This oc- 0.7 g/h (t=4.808, df=68, P=0.0001).


curs only when they can be harvested on
beaches during the nest-


Another possibility is that
spearfishers prefer other


ing season (Nam Kerker: October–April),
although during these



benefits supplied by reef fish: they
may be more valu-


months some turtles are also hunted on
nearby reefs. In the nesting



able as trade goods than shellfish
prey. But this seems


season, turtles are collected at night
or during the early morning


hours as they crawl onto sandy beaches
above the mean high wa- unlikely: shellfish collecting
produces larger harvests

ter mark to lay their eggs. Turtles are
acquired by flipping them



(1,962±247 g, n=44) than
spearing (356±100 g, n=26)


onto their backs, trussing their
flippers with ropes, and hauling



(t=24.857, df=68, P=0.0001). Shellfish
prey are also


them by boat back to the acquirer’s
household where they are kept



more likely to be shared: following a
harvest of shellfish,

alive until butchered. When butchered
for “private” consumption,


12



Encounter return rates for all prey
types include time spent han-


Table 1 Mean±SE return rates and
mean time allocated to inter-



dling (pursuit and processing) the item
while in the foraging habi-


tidal hunt types by Meriam men and
women. On-encounter returns



tat: no cooking times are included
since cooking methods for each

for each prey type are calculated from
the moment a forager com-



item can vary from minutes to hours
depending upon the dish pre-


mitted to pursuing a particular
individual prey item. For shellfish,



pared. Mean time allocation per person
shows the mean percent-


pursuits begin when a forager bends
down to pick up the item; for



age of total “reef habitat at low
tide” time devoted to each hunt


spearing, pursuits begin when a forager
spots the signs of prey and


type averaged for each of 19 women and
21 men observed more


drops into a stalk stance to pursue it;
for rocky shore harvesting,



than once on the reef during the random
time allocation scans


encounters are defined as the moment a
productive patch is found.



Reef collecting Reef spearing
Rocky shore


On-encounter returns (kcal/h) by prey
type Tridacna gigas: 13,064±4750 All fish:
2,505±778 Nerita: 1,106±465


Hippopus: 6,859±464
Asaphis: 455±52



Tridacna maxima: 4,418±708



Trochus: 3,904±467



Lambis: 3,412±205


Cypraea: 2,214±414


Hunt type returns by macronutrient


kcal/h
1,492±173 292±135
575±56


Protein (g/h)
284±31 6.6±2.9
88±9


Fat (g/h)
22±3 1.6±.7
9±.95

Mean time allocation


19 Women
0.76±0.07 0.09±0.05
0.14±0.05


21 Men
0.31±0.07 0.63±.08
0.04±0.03


on average 22% of the take is shared to
another house- through cooperative gender
specialization in macronutri-


hold, while only 7.5% of a spearing
harvest is shared. ent harvesting, nor through
reciprocal sharing. We pro-


Finally, could choosing to spear
fish instead of col- pose that the benefits of
spearfishing are gained through

lecting shellfish as women do provide
greater consump- the honest signal value of acquiring
the prey rather than


tion benefits in the long-term? The
short-term costs are through consumption, and that
honesty is maintained


inescapable: while women maximize their
patch returns through differential benefits: men of
higher phenotypic


and the size of their meat harvests by
stopping to take quality benefit more than
lower-quality individuals


shellfish when encountered on the reef,
male spearfishers because they can signal more intensely
each time they


take a large cut in protein, fat, and
energy income by signal.


ignoring shellfish and continuing to
search for fish to


spear. There could be long-term
benefits of such special-


ization if by dividing labor and
pooling harvests, male Prediction 1: men who signal
more frequently

spearfishers and female shellfish
collectors maintain obtain greater benefits


long-term shellfishing productivity on
the reef. This


could happen if the cooperative pooling
unit (the house- If spearfishing is a competitive
display, signalers should


hold) defended a reef territory,
excluding other pooling reap social benefits associated
with more frequent spear-


units from foraging so that the future
benefits of conser- fishing, such as the benefits from
gaining status through


vation could be realized. But they do
not: while reef ter- building a reputation as a skilled
spearfisher. In inter-


ritories are owned, the group sharing
use-rights to sec- views with 33 Meriam men and women,
none would


tions of reef is not equivalent to the
pooling unit. Sec- nominate a slate of “the best
shellfish collectors,” claim-


tions of reef are considered extensions
of residential ing that “being better than others”
(i.e., getting larger

plots: use-rights to residential plots
are shared by all harvests) depends solely on working
long hours, not on


members of a patriline. Only bounded
clam gardens qualities intrinsic to the forager,
and most nominated the


within reef territories are excludable
and defendable by most frequently observed woman on the
reef as “hardest-


single households, and these gardens
are approximately working shellfish collector.”
Across individuals, shell-


2–4 m in diameter, not large enough
to permit spearfish- fish-collecting harvest sizes are
strongly contingent upon


ing. Any gains in long-term shellfish
productivity would foraging time, whereas spearfishing
harvest sizes are not


have to be shared by the entire
patriline, and all men (Fig. 1). All interviewees were
willing to nominate a


within the patriline would have to
forgo shellfish collect- slate of “the best
spearfishers.” Because there was little


ing, which is unlikely since men do
vary in the extent to variance in signal frequency among
the majority of men

which they collect shellfish. We cannot
definitively ex- observed on the reef, we divided the
signalers into the


clude this possibility, only note that
it seems unlikely. most frequent (1 man with 10 observed
bouts) and the


Spearfishers interested in
maximizing return rates least frequent (14 men with 19
bouts total). The most


should not ignore shellfish while
foraging on the reef, frequent signaler obtained 23 out
of 61 (37.7%) total


but most of them do. The costs of
choosing to spear fish nominations by the 32 respondents
(who nominated as


do not appear to be balanced by
benefits received many as three individuals).
More than 75% of respon-


13


Fig. 1 Relationship between harvest
size and bout length for nal elements contained in
spearfishing are subtle and dif-

spearfishing (A) and shellfish
collecting (B). Foragers can predict- ficult to observe. Yet
reputations are still built in the ab-


ably expect larger harvests with longer
foraging time in shellfish



sence of any overt boasting or
tale-telling on the part of


collecting but not spearfishing. Each
point represents one random-



the forager himself, who when
successful, displays the


ly selected, fully observed bout, with
the open circle representing



typical behavior of a male solitary
hunter: he returns

the most active individual. Episodes
are distributed throughout the


tidal season of 1994. A Linear
regression with 95% confidence in- home carrying his catch
quietly, albeit very visibly. The


tervals for mean harvest size in kcal
(y) on reef patch foraging



signal elements incorporated in
spearfishing are con-


time in minutes (x). Harvest
size=155+2x; r2=0.074, P=0.1990.



veyed to observers in three distinct
ways. (1) Reef activi-


B The same regression for mean reef
shellfishing (female foragers



ties are highly visible to a large
section of the village

only) harvest size in kcal (y) on reef
flat collecting time in minutes


(x). Harvest size=–6682+21.86x;
r2=0.479, P=0.001 surrounding one’s residence,
which contributes to the



ease of observer perception of bout
frequency. (2) Spear-



fishermen make perception of harvest
size easier by car-


dents named the most frequent signaler
as the best spear-



rying their harvests openly in hand,
rather than carrying


fisherman. None of the less frequent
signalers in our


them in a bucket, as line fishers and
shellfish collectors


sample obtained any nominations at all.
The remainder



do. (3) Although bout frequency is the
signal element


of the nominations went to deceased
individuals or those



most easily observed, harvest size is
usually made


currently not active as spearfishers.



known to a large number of individuals
through verbal


communication. Hunts with large
harvests are recounted



many times over, and stories about
such success become


Prediction 2: signal intensity is
linked



part of local legend.


to phenotypic quality


For the qualities signaled by
spearfishing to be honestly



Turtle hunting as costly signal

advertised, high-quality spearfishers
should signal more


intensely, because they gain greater
marginal benefits or



Turtle hunting could serve as a
competitive display if


endure lower marginal costs per unit
signal. In addition



foragers can distinguish themselves
from others based on


to signal frequency (see above), good
measures of sig-



their skill. The signal(s) thus
produced by turtle hunting

naling intensity are harvest size and
success rate. The



would depend upon the kinds of costs
incurred. If turtle


mean harvest size per bout (in kcal) of
the most frequent



hunting is associated with lower
foraging returns than


spearfisher (680±197, n=9) was
larger than that obtained



other options, signals of skill sent
through hunting could


by other spearfishers (209±101,
n=15; U=32, P=0.05).


honestly indicate the ability of the
forager to expend


His hunt type return rates were also
higher at



time and energy in a wasteful pursuit.
If hunting has a


616±356 kg/h versus 137±75
kg/h (U=32, P=0.49),



high return but is associated with
wide, unconditional,


although his bouts were of equal mean
length as those of


and costly distributions of food,
signals of skill could


the less frequent signalers (106±17.2
vs 102±15.6 min).



honestly indicate one’s quality
(prosocial tendencies, or


The most frequent signaler’s return
rates were still lower



ability to gain resource reserves
through skill in other ac-


than he could achieve shellfish
collecting (U=72,


tivities) by expending time and energy
in providing


P=0.002). The “frequent signaler”
also had a much lower



goods for public consumption.


bout failure rate than other men
(failure rate per


bout=22.2% vs 66.7%; χ2=4.444,
P=0.03). Hunting turtles is indeed
a competitive pursuit, with a



very different complement of
participants than collect-



ing. As the Meriam put it, anyone can
collect turtle in the


nesting season, but only certain men
have the ability to


Prediction 3: the signal is designed to
be perceived



succeed at turtle hunting. Older and
younger men, chil-


by the intended audience



dren, and women of all ages
participate in turtle collect-


Since spearfishing is a solitary
hunting activity that pro- ing during the nesting season:
21% of turtle collections


duces no large, widely consumed common
good, the sig- involve adult females. In contrast,
for turtle hunting, the

14


only participants are males between the
ages of 16 and per capita net return of 4,653 kcal/h if they
divided the


47 (mean age=31.6, n=38). Though a
substantial propor- turtle among themselves; however, all
hunts during this


tion of Meriam men participate in
turtle hunting at some season provision feasts, and hunters keep
no share of the


point in their lives, relatively few do
so with regularity. turtle they provide. Hunters deliver the turtle
whole to


In our sample of 37 turtle hunts
extending over two peri- the feasting location. Hunters thus
obtain negative per


ods (1994–1995, and 1998), there were
a total of 102 capita consumption returns of –1,086
kcal/h, including


man-days expended, or 2.76 hunters per
hunt (all hunts the costs of traveling to the patch (see
Bliege Bird and


last 1 day or less). For the 87
man-days where hunters Bird 1997 for details of the method of
calculating net

are individually identified, there were
a total of 40 returns from turtle hunting).


unique individuals, or 2.18 hunts per
hunter. But the 3


most active men participated a total of
16 times, or


18.4% of the hunter-days of known
individuals. Thus, Nesting season hunts


44.4% of the 90 Meriam males aged 16–47
hunted at


least once in the study period, but the
3 most active During the remainder of the year, hunts are
undertaken


participants (3.3% of males) were over
five times more for two reasons: to provision previously
arranged feasts


likely to engage in a turtle hunt than
the average Meriam (n=13), and to provision households (n=9). In
this sea-


male in this age range.
son, hunting is much easier and hunters take on
fewer

Turtle hunting entails a variety of
costs that could costs: turtles are found on nearby reefs
waiting to crawl


ultimately be linked to signaler
quality in a way that onto the beaches to lay eggs at night,
the tradewinds


ensures signals sent by hunting will
remain honest have largely ceased, and in between monsoon
storms, the


(Smith and Bliege Bird 2000). Hunting
may involve (1) water is clear, calm, and visibility is
excellent, allowing


high opportunity costs in the form of
passing over oppor- hunters to dog turtles more closely, to lose
fewer, and to


tunities to acquire other resources
with a higher rate of more finely discern size and sex.
Considering only suc-


return, (2) low consumption return
rates (Bliege Bird and cessful hunts, hunters in the nesting
season could poten-


Bird 1997) as hunters distribute meat
to non-hunters fol- tially obtain per capita returns of 8,061
kcal/h hunting

lowing the hunt, or (3) high energetic,
monetary, or time for household consumption if they did not share
the meat


investment costs of preparing for and
conducting the with others. But hunters share more widely
than collec-


hunt that reduce energetic return rates
below that of oth- tors during the same season, leaving their
personal return


er more easily acquired resources.
There may be other rate from hunting for a feast at –1,633,
and those from


costs or risks of a social nature that
provide a link be- hunting for household consumption at 814
(Table 2).


tween turtle hunting and hunter
quality, particularly the Hunts during the nesting season should
not happen at all


loss of social status should a hunter
fail to deliver turtles if hunters were interested solely in
maximizing energetic


to a feast (see below).
return rates: during the nesting season,
collected turtle



offers higher after-sharing energetic returns
than hunted


turtle. In addition, other available resources
are also


Hunting season hunts
more energetically productive: sardine netting
provides



11,008±1,705 (n=28) kcal/h before
sharing – sardine


During the hunting season, between
April and October, harvests are shared only 5% of the time,
leaving on aver-


the goal of all hunts is to capture
turtle for a previously age 10,864 kcal/h. If hunting turtle were
solely about en-


arranged feast. During major feasts,
all those who come suring feast-goers can eat plenty of meat
and fat, men


must be fed by the feast-giving family
and their allies, should net 50 kg of sardines with no chance of
failure

and the family can never predict
precisely how many rather than spending the entire day
chasing turtle to risk


people will come to the feast itself.
Theoretically, the coming back to the feast empty-handed.


entire population of the island (over
400 men, women, Turtle hunting in both seasons thus
satisfies two of


and children) plus many off-island
visitors could attend, our criteria for consideration as an
honest and costly sig-


though attendance by 200 or fewer from
both on and off nal. Unlike turtle collecting, observers can
distinguish


the island is typical (mean=174.9,
range=49–343, n=54). skilled from unskilled individuals and
participation is re-


Failure to provide turtle is a
substantial cost to the stricted to a subset of the community;
second, hunting


hunter’s status since all feast-goers
will note the absence has the potential to generate high return
rates among


of turtle meat at the feast. Since
hunts are only undertak- simultaneously available foraging
alternatives (second

en a day or two prior to a feast, there
are only as many only to sardine netting), but does not result
in much con-


hunts as there are feasts during this
season: in 1994 there sumption benefit to the hunters since they
share widely


were ten hunts for public feasts.
During the hunting sea- and take no portion for themselves. As
with spearfishing,


son, the turtle patch is more distant,
encounters with tur- we hypothesize that the benefits of hunting
are gained


tle are less common, and turtles are
more difficult to de- through the honest-signal value of acquiring
the prey,


tect and follow under water that is
often murky and bro- rather than through consumption, and that
honesty is


ken by swells and whitecaps created by
the nearly con- maintained through differential costs and
benefits: men


stant 20-knot southeast tradewinds
during this season. of higher phenotypic quality benefit more
or pay less


After a successful hunt, hunters could
expect to obtain a than lower-quality individuals per unit
signal.

15



and handling. Since travel costs are
primarily the cost of fuel ex-


Table 2 Turtle hunting per capita
mean±SE net returns (kcal/h),



pended in outboard motors, the cost of
fuel used (l) is converted to


number of hunts/collections (n), and
total number of participating



calories of meat that could have been
purchased in the local shop

acquirers by consumption type (house or
feast), season (hunting or



with the money used to buy fuel for the
hunting trip. Further de-


nesting), and acquisition method
(hunting or collecting). Net re-



tails in Bliege Bird and Bird (1997)


turn rates are calculated as gross
edible energy captured minus en-


ergy expended in travel divided by time
spent in travel, search,



Before sharing
After Sharing


Hunting Nesting
Hunting Nesting



season season
season season


House
Collecting (108 acquirers) 21,875±2279
(n=44) 5,068±922


Hunting
(27 acquirers) 8,061±970 (n=9)
814±526


Feast
Collecting (56 acquirers) 16913±1842
(n=44) 1329±1127

Hunting
(40 acquirers) 4,922±345 (n=10) 10,607±3566
(n=13) –468±660 –1,633±325


Total number of hunts
10 22


Total number of collections
0 88


Number of turtles
13 116


Prediction 1: turtle hunting is
differentially costly agility, leadership and
organizational abilities, and proso-


or beneficial
cial tendencies (e.g., expending time
and energy in pro-


viding goods for public consumption at
feasts). Such in-


The relative costs and benefits of
hunting turtle vary formation is presumably of interest
to potential female


with the season: hunts are less costly
in the nesting sea- mates, their families, potential
political alliance partners,


son because the prey are found closer
to home, but they or competitors for social status.


are more ambiguous as a signal, since
turtles may also be As detailed above, the most active
hunters are a sub-


collected at this time. Since there is
the potential for dis- set of the adult male population. The
best turtle hunt


honest signaling (acquiring a turtle
through collecting leaders are well known to community
members: in a


and passing it off as a hunted turtle),
we predict that the series of 32 interviews we conducted
with men and

benefits of hunting during the nesting
season are lower women in which informants were asked to
nominate any


as well. We would thus expect that more
skilled hunters three individuals of their choice for
“best turtle hunters,”


would be over-represented among hunting
season hunts, 3 men (all ones we had observed as the
most frequent


which have the highest cost and the
lowest signal ambi- hunt leaders) garnered 38 (39.6%) of
the 96 nominations


guity, while the least skilled hunters
would be over-rep- (a total of 30 men were nominated for
“best hunter” out


resented among nesting season hunts,
when hunts are of a pool of 90 or more males aged 16
and older). While


less costly as well as harder to
discriminate from collect- several deceased individuals or
older men no longer


ing. Among turtle hunt participants,
age is usually a active as hunt leaders were named
among the “best hunt-


good proxy for skill, since hunt
participants move up in ers,” no men currently active
solely as jumpers were so

the hierarchy of roles as they gain in
skill over time, and named. Jumpers are rarely individually
credited by


since the skills necessary to be a good
hunt leader are not others for acquiring a turtle, and
drivers are rarely pub-


dependent upon size and strength. We
thus expect jump- licly credited with participation in
the hunt. Feast-goers


ers to make the move to become hunt
leaders only during can readily name hunt leaders as
providers of the turtle,


the nesting season. Hunt leaders during
the hunting sea- even several years after the feast, but
when pressed for


son (n=8) are in fact an average of 7.9
years older than the names of other hunt participants,
lump all others to-


hunt leaders (n=22) during the nesting
season, a differ- gether as “the boys.” Jumpers may
begin in this role as


ence that is statistically significant
(U=41.5, P=0.05). In young as 15–17 years old; anecdotal
evidence indicates


contrast, there is no difference in age
of jumper accord- that those who excel in this role and
gain increasing

ing to season (df=37, t=–0.253,
P=0.80). Of the 17 knowledge and peer respect
eventually become hunt


known current hunt leaders, the 4
youngest (aged 18–22) leaders, while others remain
jumpers or cease participat-


were observed as hunt leaders only
during the nesting ing in turtle hunting.


season, and served only as jumpers
during the hunting


season.



Prediction 3: the signal is designed to
be perceived



by the intended audience


Prediction 2: signal intensity is
linked

to phenotypic quality
Signals do little good if they are not
perceived and the in-



formation contained is not deciphered.
We propose that


Turtle hunting provides evidence of
skills that could when signals are designed to acquire
general social status


honestly signal several relevant
qualities: environmental or political dominance, broadcast
efficiency will be in-


and ethological knowledge, risk-taking,
strength and creased by directing them at larger
audiences, rather than


16


small subsets of interested parties (as
might be predicted sult, the most active spearfisher gains the
benefits of an


if the signal elements of hunting were
directed solely at enhanced spearing reputation that more
infrequent sig-

potential mating partners). Thus, we
predict that hunted nalers do not seem to enjoy. Spearing thus
has the poten-


turtles should be shared more widely
than collected tial to serve as an honest signal of a
spearfisherman’s


turtles; in other words, that
individuals should choose to phenotypic quality.


hunt for larger audiences, and to
collect when audiences Although we have demonstrated the
potential for sig-


(the number of households attracted to
the turtle and con- naling, and the bias in men’s foraging toward
resources


suming portions) are predicted to be
smaller. Because which have high competitive signaling
potential, precise-


turtles cannot be collected during the
hunting season, ly how both signaler and observer ultimately
benefit


we control for the effects of season.
We thus predict that from the signal is not clear. We do know that
good spear-


audience size should have a significant
effect on acquisi- ing men are widely recognized, and that this
adds to their

tion method within the nesting season
only: when larger social status within the community. Potential
competitors


audiences are available, men should
prefer to hunt rather might also gain from knowing who is a
better man with a


than collect. During the 1994–1995
nesting season, hunts spear, although the knowledge certainly
would have


for which the number of consumers were
known (n=22) been more useful in the past when spears were
the major


averaged 26.7±23.7 households,
while collections (n=80) means of ambushing competitors.
Spearing signals may


averaged 18.7±19.0 households
consuming, a significant not be designed to appeal to
potential mates interested


difference (t=–1.649, df=100,
P=0.05). only in marrying provisioners. A man
who demonstrates


Individuals who acquire turtles for
feasts have a large his intention to provision a household by
collecting

built-in audience; however, turtles
acquired for house- shellfish may be more attractive to such
women than a


hold provisioning will be shared among
a smaller num- man who demonstrates his intention to engage
in a com-


ber of households. If hunting serves as
an honest signal petitive, status-enhancing pursuit.


and hunters wish to broadcast the
signal widely, turtles Turtle hunters seem to have very
different goals than


acquired for household provisioning
should also be turtle collectors. Compared to collecting,
hunting is more


shared more widely when hunted than
when collected costly (in time, energy, and risk),
provides meat less effi-


(controlling for season). During the
1994–1995 nesting ciently, and is associated with wider
distributions of


season, there were 9 hunts and 44
collections to supply meat and larger audiences to witness the
hunters’ prow-


turtles for household consumption
(Table 2). As predict- ess. Hunters keep no meat for themselves
unless (quite

ed, hunted consumption turtles were
shared to more rarely) hunting for household consumption,
in which


households (7.5±3.0) than
collected consumption turtles case they still keep less and
share more than turtle col-


(4.6±2.5), and the statistical
difference is strongly signif- lectors. Hunters take on a variety of
costs for which they


icant (U=64, P=0.015). Because the
number of house- are not materially compensated: they
expend more time


holds receiving portions of a hunted
turtle is larger than and energy in hunting than they do
collecting, they spend


the number receiving portions of a
collected turtle, por- more money for fuel, they spend time
organizing and


tions shared to each household should
be smaller, but preparing the hunting team and its equipment
prior to the


they do not seem to be so, because
hunters kept less for hunt, and they deliver the meat to be
consumed by large

themselves and their own households
than did collectors audiences at feasts. The ability to bear
such costs ap-


(mean portion kept (kcal) for
hunters=7,780±2,194 kcal, pears to be linked to hunter
quality. Because a hunter is


n=38; for collectors=14,387±2,631
kcal, n=123) al- an organizer and decision-maker, his
abilities peak as he


though the difference is not
significant (t-test on log- gains skill and experience: Those
named as the best


transformed kcal kept, t=0.682, df=159,
P=0.50). hunters are older than other hunt participants,
such as



jumpers. As we predicted, when hunting was less
costly



and more difficult to discriminate from
collecting (i.e.,

Discussion
during the nesting season), hunters were
younger, in fact



composed almost entirely of ambitious jumpers
prepar-


Although our sample of spearfishers is
too small to de- ing to be hunt leaders themselves.
Furthermore, the sig-


fine a continuum in level of skill,
spearfishing does seem nals sent by hunting are efficiently
broadcast: hunts were


to exhibit strong potential for
competitive signaling. The associated with larger numbers of
consumers overall


most frequent signaler obtains much
higher gains per than collections during the nesting season
and during


unit patch residence time than other
spearfishers. Har- household consumption events. Most
feast-goers (audi-


vest size (the most easily observed
indicator of return ence members), when quizzed, know the
identity of

rates) is likely to be linked to
phenotypic quality in the hunters, but not the identity of
jumpers.


form of hand-eye coordination, stealth,
and patience, and Since the low take-home returns of hunters
providing


provides an immediate signal of forager
quality. Success turtle for household consumption and the
negative take-


rate is a signal element contributing
to the long-term rep- home returns of hunters providing turtle for
a feast are


utation of the spearfisher. High
success rates in conjunc- due to widespread sharing, the argument
could be made


tion with large harvest size ensure
that large harvests are that the signal cost is eventually
recovered in the form of


more likely due to skill
(forager-dependent variability) meat or other goods or services
returned as payback for


than to luck (forager-independent
variability). As a re- the hunter’s gift. In other words,
perhaps RA by the re-


17

cipients of a hunter’s largesse erases
the cost of his signal The results of additional analysis in
progress show


(Sosis and Hill 1997). The stable
maintenance of RA re- that the display of such qualities is
correlated with higher


quires that the provisioning of a
turtle be contingent up- social status and higher age-specific
reproductive suc-


on the eventual receipt of
counterbalancing benefits cess of hunters and their mates
than other Meriam (E.A.


which compensate the hunter for the
marginal cost of Smith, R. Bliege Bird, D. Bird, unpublished
data). As


giving up turtle that could have been
used in other fit- explained to us by Meriam hunters and their
mates, the


ness-enhancing ways. Previous tests of
predictions based signals sent by hunters are likely to be part
of both politi-


on RA in the form of risk reduction
reciprocity for shar- cal strategies, in which hunters
demonstrate to other men


ing of collected and hunted turtles
combined received their honest intention to work for the
public good, and

little support (Bliege Bird and Bird
1997). Further tests reproductive strategies, in which hunters
demonstrate


of alternative forms of reciprocity are
being conducted their “willingness to work hard” in order
to gain access


to evaluate this line of explanation
(R. Bliege Bird, G. to the “best girl” (Kaddy, personal
communication;


Kushnik, E.A. Smith, D. Bird, C.
Hadley, unpublished Passi, personal communication). Hunters
know that


data), as will be detailed in future
publications. spreading one’s influence widely via the
provisioning of


While we have measured the cost of
hunting using di- collective goods at feasts increase social
status over the


rect material currencies such as time,
energy, and money, long-term among the community as a whole,
while less


there are additional social costs that,
though difficult to public-minded status-enhancing activities (like
stealing

measure, may be of equal or even
greater importance. land or selfishly hoarding resources)
provides narrow,


Turtles provided for feasts are needed
by community el- short-term status in only a small pool of
competitors.


ders (feast hosts) to enhance their own
status by ensuring


there is plenty of meat for guests. If
successful hunting is


a reliable signal of the hunter’s
underlying quality (as de- Implications for the sexual division of
labor


fined above), lower-quality individuals
would be expect-


ed to fail on turtle hunts more often
and thus pay a higher One intriguing result of our analysis are the
gender


marginal cost per turtle delivered to a
feast (i.e., per unit differences in participation in those
activities with high


signal). Failing to deliver a turtle to
a feast also entails signaling potential. With spearfishing
particularly, the

significant social costs, since the
feast-holders are expect- differences were not consistent with the
common notion


ed to serve turtle to their guests, and
everyone will know that a sexual division of labor in humans
functions to


when a hunter has failed. High-quality
individuals should maximize the productivity of a cooperating
male-female


have a lower probability of failure and
thus a high ratio of pair (see Bliege Bird 1999). We propose that
the signal-


social benefits (from successful hunts)
to costs (from fail- ing benefits attached to certain foraging
strategies may


ures). Low-quality individuals may face
a large enough often change the valuation of certain prey
items for some


risk of failing to produce a net social
deficit from their individuals. While foraging on the reef, males
make


signaling attempts, and hence may avoid
signaling at all, choices that fail to maximize their
macronutrient return


making it difficult to test this
hypothesis. We do have an- rates. These choices appear to be due to
the tradeoffs

ecdotal evidence that some men have
attempted turtle men face over foraging for highly
productive resources


hunting in the past, and given up when
they found they which have little signaling value (shellfish)
against for-


were “no good at it.” In addition,
the facts (detailed aging for less productive resources with
high signaling


above) that only certain men engage in
turtle hunting, that value (speared fish). Likewise, choosing to
hunt turtle


only an older and more experienced
subset of these be- for feasts provides much lower consumption
benefits


come hunt leaders, and that hunt leader
status is widely than collecting turtle for the household, but
greater sig-


recognized in the community, all
indicate that success in naling benefits. Adult women only
participate in cooper-


this endeavor is not equally available.
ative turtle collection, comprising 21% of all
partici-


To qualify as costly signaling,
hunting must not only pants, and do not hunt.

be honest, but must also reveal
signaler quality. Given These data suggest that foraging sex
differences may


the three distinct roles played by hunt
participants, we not simply be a result of women preferring
plants and


propose (but cannot currently test)
that at least four dis- men meat, or women preferring small
harvests while men


tinct dimensions of underlying quality
could be signaled prefer large ones, or even women preferring
prey which


through turtle hunting: (1) physical
quality (such as can be more easily harvested while children
are present


strength, stamina, agility, and
risk-taking); (2) cognitive (shellfish are just as difficult as
fish to collect with small


skills (involving the ecological and
ethological knowl- children). Where there is a choice in
foraging method,


edge needed to successfully locate and
capture turtles); men seem to prefer to acquire meat through
more risky


(3) leadership skills (charisma and
organizational abili- methods that more easily differentiate the
skill of indi-

ties), and (4) generosity (ability and
willingness to bear vidual foragers, while women seem to prefer
less risky


the high immediate cost in time, money,
and energy of methods carrying little potential for
discrimination of for-


providing collective goods without
direct compensation). ager quality. We propose that sex-biased
foraging prefer-


We expect the first dimension to apply
primarily to the ences may arise due to the differing benefits
each sex re-


younger men who serve as “jumpers,”
whereas the other ceives from investing in competition for
status through


three dimensions refer primarily to
hunt leaders. signaling certain genotypic or phenotypic
qualities.


18



Acknowledgements We thank the Meriam community
foremost,

Foraging as communication



especially Chairman Ron Day for his foresight and
tolerance, our



Meriam families, particularly the Passis, and the
turtle hunters who


As Hawkes (1990, 1991, 1992, 1993)
first hypothe- volunteered information or invited us on
hunts. We also wish to ac-


sized, some kinds of hunting may
persist in human knowledge the invaluable field assistance
of Andrew Passi, Ron


populations because some foragers gain
benefits from “Sonny” Passi, Edna Kabere, Del Passi, and
Craig Hadley. For many



helpful discussions and comments on the
manuscript, we thank

widely disseminating knowledge about
their prowess



Monique Borgerhoff Mulder, Thomas Getty, Craig
Hadley, Polly


relative to other individuals through
the hunting of Wiessner, Kristen Hawkes, Kim Hill, Frank
Marlowe, Rich Sosis,


large and risky prey items. While
Hawkes (1993) Steven Siller, and Bruce Winterhalder.
Research in 1998–1999 was


stressed the benefits that hunters
obtain from providing supported by NSF grant SBR-9616096 to R.B.B.
and E.A.S., NSF



grant SBR-9616887 to D.W.B. and E.A.S., and
research in 1994–


collective goods (as with turtle
hunting and uncondi-



1995 was supported by grants to R.B.B. and D.W.B.
from AIATSIS,

tional sharing), our results show that
benefits may also



the L.S.B. Leakey Foundation, the Wenner Gren
Foundation for An-


come in the absence of any material
good provided to thropological Research, and an NSF
predoctoral fellowship to


observers (as with spearfishing) as a
result of honestly R.B.B. and an NSF Dissertation Improvement
Grant to D.W.B.


revealing hidden information to
interested parties. Dif-


ferential costs and benefits for
signalers of higher and



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7



Growing Up Mikea


Children’s Time
Allocation and Tuber


Foraging in
Southwestern Madagascar


Bram
Tucker and Alyson G. Young


I. INTRODUCTION


Human childhood poses
interrelated evolutionary, economic, and ecolog-


ical puzzles. The
evolutionary puzzle is that the human juvenile period is


considerably longer than
the premature developmental periods in other


great apes, making long
childhood a derived trait of potential significance


(Bock and Sellen 2002b).
Humans invest in growth and delay the start of

reproduction for 15 to 20
years (Bogin 1999; Hill and Kaplan 1999). This


seems contrary to
Darwinian logic because a long childhood period has


the potential to increase
risk of mortality prior to reproductive maturation.


Natural selection could
favor prolonged investment in growth and de-


layed reproduction for
several reasons. First, the costs of reduced fertility


could be overridden by the
mortality-reducing benefits of a long training

period, such as a longer
period to learn difficult foraging tasks (Kaplan et


al. 2000). Second, long
childhood could be a side effect of selection for a


long lifespan more
generally. Among mammals, body size, lifespan, and


the length of the juvenile
period are highly correlated (Charnov 1993, 2001;


Charnov and Berrigan
1993).


The economic dimension
of the puzzle has to do with the labor costs of

supporting children.
Children cannot meet all their resource needs from


their own labor, so
parents and alloparents must cooperate to provision


children and take care of
their needs, qualifying humans as “cooperative


breeders” (Hrdy 1999).
Children like all household members are simulta-


neously resource producers
and consumers. They are both labor that ben-


efits the household and
hungry mouths that must be fed. Before the age of

“positive net
production,” children consume more food than they produce



147


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148
Bram Tucker and Alyson G. Young


from their own labor; they
are a net cost to the household. The rest of the

household must work harder to
support them (Chayanov [1925] 1986; Ka-


plan 1997; Lee and Kramer
2002). After the age of positive net production,


young people can take on
alloparental responsibilities and care for their


younger siblings, easing the
childcare responsibilities of adults (Kramer


2002).


The ecological dimension
of this puzzle is that the costs of raising chil-

dren vary with environmental
conditions. Some environments favor chil-


dren’s active participation
in the household economy more than others


(Hawkes et al. 1995a). In
some environments, children can dig their own


tubers and pick their own
fruit. In other environments, children foragers


are endangered by the
elements or frustrated by difficult or costly forag-


ing tasks. When children can
produce their own food, they reduce the

physical and economic burdens
on adults, potentially facilitating adults’


future reproduction (Nag et
al. 1978; Kramer and Boone 2002). Ecological


variation in children’s
work roles may be one of the reasons that forager


fertility rates are so
variable (Kelly 1995:206-208; but see Early 1985).


Kaplan and colleagues
(Kaplan 1997; Kaplan et al. 2000) argue that de-


layed reproduction and a
lengthened juvenile period were favored by nat-

ural selection because of the
mortality-reducing benefits of a long learning


time, to learn how to forage
for high-quality foods. They note that while


other apes forage primarily
for “collected” resources such as fruits and fo-


liage that are easy to
procure and low in food value, humans specialize on


“extracted” resources
such as tubers, roots, nuts, honey, and hunted game


that are difficult to
procure and high in food value. They argue that human

foraging is so
skill-intensive that it takes most of the lifetime to learn, so


that children’s foraging is
more about learning than food procurement.


Their model leads to two
predictions. First, children and adolescents are


dependent on adults’
foraging success, and particularly men’s large game


hunting, to meet their food
needs. Kaplan (1997) found that in three


neotropical foraging
societies, Machiguenga, Piro, and Ache, young peo-

ple do not reach the age of
positive net production until around the age of


20. Second, because
age-related increases in foraging success are primarily


the result of increased
learned skill, foragers ought to continually improve


even once they have stopped
growing, achieving mastery at middle age.


Kaplan et al. (2000) claim
that Hiwi women of Venezuela master root dig-


ging at around ages 35 to 45,
and Hiwi males master honey foraging at age

25. Hambukushu of the
Kalahari master mongongo nut cracking at age 35.


Ache women of Paraguay reach
peak palm extraction rates during their


early twenties. A more
thorough study of Ache men (Walker et al. 2002)


combining longitudinal
hunting data with an archery contest showed that


men almost always increase in
foraging efficiency with age, and that


archery skill peaks around
age 40.

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149



Growing up Mikea


Other researchers have
disagreed with parts of the theory advanced by


Kaplan and his colleagues.
Hawkes and her colleagues disagree that house-

holds depend on men for
food and that male hunting is an efficient strat-


egy for provisioning a
household (Hawkes 1990, 1991, 1993, 2001; Hawkes


et al. 1991, 2001a,
2001b). They argue that hunted meat is a poor resource


choice for feeding
children because daily hunting success is highly vari-


able, and children need to
eat every day. When hunters are successful, they


are obliged to give most
of their meat away and cannot control the distri-

bution. Hawkes argues that
men hunt and generously distribute meat be-


cause of fitness-enhancing
benefits that may conflict with the cooperative


goals of the household.
Men may hunt as a form of social competition to


win allies, mates, and
deferential treatment for their families. Or they may


hunt as a way to
communicate to others the qualities they possess. Such


communication is “honest”
because men without the advertised qualities

find hunting success too
costly to mimic (Smith and Bliege Bird 2000; Bliege


Bird et al. 2001; Hawkes
and Bliege Bird 2002; Smith et al. 2003).


While Kaplan (1997)
finds that the age of positive net production for


Machiguenga, Piro, and
Ache is around 20, there is good reason to suspect


that this varies among
foraging groups according to environmental cir-


cumstances. Ju/’hoansi
San children near the Botswana/Namibia border

rarely accompany adult
mongongo nut foragers, while Hadza children of


the Eastern Rift Valley of
Tanzania are active foragers of baobab fruit and


wild tubers and berries
(Blurton Jones et al. 1989, 1994a, 1994b; Hawkes et


al. 1995a) Even if it does
take a lifetime to learn how to forage, one would


expect Hadza to achieve
positive net production at an earlier age than San.


While Kaplan et al.
(2000) argue that juveniles forage less productively

than adults because they
have not yet learned to forage efficiently, others


argue that children make
optimal foraging decisions for foragers of their


smaller size and lesser
strength. Bird and Bliege Bird (2002) note that


among intertidal
collectors in the Torres Straits, Australia, children are less


selective reef flat
collectors than are adults. Children gather almost all ed-


ible species of shellfish,
while adults target specific high-yielding Hippopus

species. This difference
is consistent with the predictions of the prey choice


model from optimal
foraging theory (MacArthur and Pianka 1966;


Schoener 1971; Stephens
and Krebs 1986). Because children are slower,


their encounter rate for
highly ranked resources such as Hippopus is effec-


tively lower than that of
adults, so they include more species in their diet


breadth.

In another study of
this population, Bliege Bird and Bird (2002b) con-


trast age-related trends
in foraging efficiency for activities that are cogni-


tively challenging but
physically easy, such as fishing and spearfishing,


versus activities that are
cognitively easy but physically difficult, such as


shellfish collecting.
Their data reveal significantly stronger age-related


07 5/5/04 13:27 Page 150

Kitty’s TS • Aldine •
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150 Bram
Tucker and Alyson G. Young


trends in the latter,
suggesting that strength is the more important deter-


minant than learning for
these marine exploitation tasks.


Our goal with this chapter
is to examine children’s foraging behavior in


what is probably one of the
best environments for young foragers. The

Mikea Forest of southwestern
Madagascar is a safe place for children.


The dense dry tropical forest
contains almost no predators and few poi-


sonous plants and animals.
Within this forest, Mikea forage for wild tubers,


small game, and honey in
addition to cultivating maize and manioc, herd-


ing livestock, and
participating in market-oriented activities. Mikea chil-


dren as young as four or five
successfully unearth wild Dioscorea tubers for

household consumption. Mikea
children tuber foragers have exceptionally


high net acquisition rates,
averaging between 536 net kilocalories/hour for


girls and 504 net
kilocalories/hour for boys. As such, the Mikea example


offers a potentially
interesting test case for theories of juvenile dependency


and foraging strategies.


We present scan-sampling
time allocation data and foraging return rate

data from the Mikea community
of Behisatse. We find that while Mikea


children experience over
twice as much leisure time as do adolescents and


adults, they allocate similar
time to foraging, especially for the wild tuber


called ovy (Dioscorea
acuminata). Mikea children make a significant contri-


bution to the household food
procurement effort. Although they do not


appear to achieve positive
net production, it is possible that they are capa-

ble of doing so, at least
during some seasons. Mikea children harvest ovy


patches with strategies that
are optimal for their lower strength and skill,


as predicted by Charnov’s
marginal value theorem (Charnov 1976). We


conclude by questioning
whether “efficiency” is commensurate with chil-


dren’s foraging goals.
Rather than children’s foraging excursions being


rigorous training sessions,
children appear to forage as a leisure activity,

an extension of play.


II. THE
MIKEA OF MADAGASCAR


The southwest coast of
Madagascar is a heterogeneous dry environment


conducive to foraging (Figure
7.1). The landscape is a mosaic of dense, dry,


deciduous forest covered in
vines; anthropogenic clearings residual from


agricultural and herding
activities; thorny forests of octopus trees (Didiera

madagascariensis) on sandy
dunes; open savanna and savanna woodland;


grassy lakebed pans; and
dense coastal mangrove swamps. For the past


four or more centuries, the
Mikea Forest has provided refuge and suste-


nance to Malagasy escaping
political, personal, or economic crises. During


the 17th through 19th
centuries, southern and western Madagascar was


dominated by several
cattle-herding kingdoms. Petty elites raided each

07 5/5/04 13:27 Page 151


Kitty’s TS • Aldine •
Hewlett&Lamb • 173043



Growing up Mikea



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~-


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a


s in


-B


(Other forest camps


throughout)

rest


Behisatse


Vorehe


(Market)


onal


N a mon t e

Nati


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-


-


-


Mike


a Fo


Androka River


r


ive


oR

mb



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no


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Ma


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Kilometers


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0 5 10


Savanna



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~


~


Dunes and thorny scrub


Dry forest


(mixed deciduous /


euphorb / thorn)


~~~~

~~~~


Permanent standing water



er



Fiherena Riv


~~~~


~~~~

~~~~


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——


—–


Lakebed


——

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——



:



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Nation



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TOLIARA

Figure 7.1. Map of the
Mikea Forest, with placenames mentioned in text. Ap-


proximate forest
extent based on 1994 Landsat imagery processed by


James Yount.


07 5/5/04 13:27 Page 152


Kitty’s TS • Aldine •
Hewlett&Lamb • 173043


152
Bram Tucker and Alyson G. Young

other for booty of cattle and
slaves within each polity, uniting periodically


in warfare against
neighboring polities (de Flacourt [1660] 1908; Drury


[1729] 1826; Fagereng 1950).
Cattle and slaves were sold to passing Euro-


pean ships in exchange for
glass beads, silver coins, and firearms (Parker


Pearson 1997). Today’s
Mikea are the descendants of neighboring Masi-


koro herders and Vezo
fishermen who fled into the forest to resist tribute

demands and threats of
slavery and livestock loss, or in other cases, to


avoid interpersonal disputes
and accusations of witchcraft. Other Mala-


gasy became Mikea during the
twentieth century, when they adopted for-


est residence and foraging as
an alternative to French colonial policies of


forced relocation, taxation,
and mandatory labor (Yount et al. 2001; Tucker


2003). Mikea are foragers of
recent vintage.

For most Malagasy, the
word “Mikea” connotes a forest-dwelling


hunter-gatherer (Poyer and
Kelly 2000; Yount et al. 2001). Mikea forage for


a wide variety of tubers
(Dioscorea bemandry, Tacca pinnatifida, others), but


the ovy tuber Dioscorea
acuminata is the most important foraged food


source.1 Other gathered foods
include honey, wild cucurbits, baobab fruit


(Adasonia digitata), and a
few other wild fruits (Flacourtia indica, Zizyphus

vulgaris). There are no large
game animals in the Mikea Forest except for


the exceedingly rare wild
boar (Potamocorus larvatus). Small game include


a variety of birds (Numida
meleagris, Lophotibis cristata, Coua coquereli, oth-


ers) and small mammals,
including tenrecs (Tenrec ecaudatus, Echinops


telfairi, Setifer setosus),
feral cats (Felis sylvestris), and lemurs (Chierogaleus


medius, Microcebus murinus,
Lepilemur ruficaudatus). Mikea also forage in

the intertidal zone for fish,
crabs, octopus, and sea cucumbers (for more


details, see Tucker 2001).


Despite their reputation
as forest-dwelling foragers, all Mikea house-


holds rely to some extent on
nonforaging activities. Their most significant


agricultural ventures include
slash-and-burn maize agriculture in the for-


est, and semi-intensive
manioc cultivation in the savanna and lakebeds.

Many households own cattle,
goats, chickens, and guinea fowl, and more


rarely, turkeys, ducks, and
swine. Mikea are regular participants in the


local market economy. They
sell their production at weekly village mar-


kets such as the Wednesday
market in Vorehe, where they purchase to-


bacco, soap, clothing, and
other necessities. Many practice mobile


retailing, purchasing goods
in one place to resell at a markup elsewhere.

Mikea also participate in the
labor market. Mikea girls replant rice shoots,


boys guard cattle, and men do
agricultural or herding labor for neighbor-


ing villagers.


The data discussed in
this chapter were collected in the Mikea hamlet of


Behisatse,2 a group of
bark-thatched huts in the north-central part of the


Mikea Forest. Mikea
households are mobile, moving flexibly throughout

the year to tend economic
activities at multiple sites. During the study pe-


riods in 1996–99, eleven
different households resided at Behisatse for a


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153



Growing up Mikea

month or more. They used
Behisatse as a home base when foraging for ovy


and attending
slash-and-burn maize fields and goat herds. Other locations


in their yearly round
included the villages of the Iovy Floodplain, where


they tended manioc fields
and performed wage labor; and the villages and


hamlets of the Namonte
Basin, where they attended family ceremonies,


foraged in the lakes and
dunes, tended livestock, and occasionally planted

small gardens in the
lakebeds. Many households also spent time in other


forest hamlets and camps
functionally similar to Behisatse, hosted by in-


laws. Some practiced
complete nomadic foraging for a week or two each


year.


Our data sample across
the four seasons recognized by the Mikea.


Eighty-five to 95 percent
of rainfall falls within the wet season, litsake

(December–February).
Rainfall varies stochastically from 100 to 1500 mm


per year (Tucker 2001).
The rest of the year is divided into an early dry sea-


son, fararano (March-May);
a middle dry season, asotre (May-August); and


a late dry season, faosa
(August-December). The dry seasons differ signifi-


cantly by temperature.
Temperatures in fararano are similar to those of lit-


sake (mean 33, min 21,
maximum 41° C). Asotre is the southern winter and

is the coolest season of
the year (mean 26, minimum 7, maximum 36° C).


Faosa is the hottest
season (mean 33, minimum 18, maximum 42° C).3


III.
METHODS, ANALYSES, AND RESULTS


A. Age Classification


Exact age of the study
subjects is unknown. Instead, we use age rank

order to explore
age-related trends, and an age classification scheme in


tests of significant
differences. Our categories correspond roughly to those


used by Mikea when
describing people’s age. These categories can be fur-


ther distinguished by
weaning status, degree of personal mobility, and


marital status:


• Infant (aja mena):
Babies that have not yet been weaned, and have no

mobility beyond that
supplied by caretakers. Infants do not forage or


engage in any other
work.


• Children (olo
kely): Children who are weaned and prepubescent, are


mobile enough to
leave camp and travel/work with others in the en-


virons of the camp,
but are not old enough to travel alone.


• Adolescents (olo
be-be; kidabo lahy): Young people nearing, experienc-

ing, or just past
puberty, unmarried, with complete independent per-


sonal mobility, who
can travel to the well or to the market alone. They


frequently talk
about marriage and sex. Some are sexually active.


• Adults (olo be):
Married people and parents.


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154
Bram Tucker and Alyson G. Young


B. Time Allocation


Scan-sampling time
allocation data were collected in the Mikea hamlet


of Behisatse during eight
noncontiguous months between 1996 and 1999


(see Table 7.1). The
procedure worked as follows. “Sampling months” con-


sisted of 19- to 24-day
blocks during which the research team resided at

Behisatse. “Sampling
events” were half-hour increments spanning day-


light time, which was from
6:00 to 18:30 during summer months and 6:30


to 18:00 during winter
months. A randomized schedule of sampling events


was constructed at the
beginning of each sampling month. All sampling


events were scheduled twice
during each month so that we sampled twice


or three times per day,
either 52 times (summer) or 48 times (winter) per

month.


During each sampling
event, we recorded the location and behavior of


Behisatse residents and
visitors of all ages. Our direct observation was lim-


ited to in-camp activities,
which we categorize as leisure and housework.


If people were absent from
the camp at the time of the observation, their


location was recorded as “not
in camp” and the purpose of their absence

was ascertained through
subsequent observation and interview. People


Table 7.1 Summary of Time
Allocation Sampling Strategy



No. of No. of No. of



days observation data Average


Month & year Dates
times points population Season

Jul 1996 Jul 7–15,
24 48 672 23.19 Asotre,


19–Aug 3
Middle dry


Nov 1997 1–10, 14–18,
24 52 1071 21.38 Faosa,


20–28
Late dry


Jan–Feb 1998 Jan 8–15,
19 52 1136 23.54 Litsake,


22–29; Feb 7,
Wet season

8, 14


Mar 1998 5–10, 12–16,
20 52 1139 36.30 Fararano,


21–29
Early dry


July 1998 Jun 30–Jul 2,
22 48 720 17.15 Asotre


8–14, 16–20,
Middle dry


24–28,

30–Aug 1


Oct 1998 6–26
20 48 547 12.88 Faosa,



Late dry


May 1999 9–17, 20–30
20 48 771 16.67 Asotre,



Middle dry


July 1999 Jun 27–Jul 17
21 48 581 12.19 Asotre,


Middle dry


TOTAL
6637


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were either absent for a
few hours, usually on a food production chore


near camp (foraging,
farming, or herding). Or they were away for a few

days, often with a specific
objective in mind (tending fields in other loca-


tions, attending the
market, attending a ceremony, etc). Visitors are ex-


cluded from the current
analyses, for they likely behave differently than


residents. Infants are
also excluded, since by our definition they do no


work. The resulting
dataset includes 6637 observations of 46 individuals.4


Figure 7.2 summarizes
these data by age and sex. Data are expressed as

percents of the total
number of observations within each age/sex category.


Figure 7.3 examines time
allocation to food production activities in further


detail. We make four
observations from these data:


1. Children spend their
time differently than people in older age cate-


gories, while there
is little difference between adolescent and adult


Food Production

Housework


17.0 %


Housework 8.8 %


155


Males


Other/Unknown

Away


19.1 %


Housework


26.9 %


Away


Leisure Leisure

Adults


Other/Unknown


22.1 %
36.6 %


27.6 %
30.2 %


Food Production


23.4 %

Food Production
Food Production


20.1 %
24.6 %


Other/Unknown Other/Unknown


Away Leisure
Away


Leisure


Adolescents 19.9 %

36.8 %
34.3 %


31.8 %


Children


100



Other/Unknown



Away
9.4 %

Food
Production


8.4 %



Housework 7.9 %



Growing up Mikea



Females


60

Leisure


71.6 %


40


20


Leisure


61.8 %

80


0


20


Housework 5.5 %


40


60

80


100


Percent daylight time allocated



Housework 5.8 %



Food
Production


15.4 %


Away
13.1 %



Other/Unknown


Figure 7.2. Percent
daylight time allocation by age/sex group. “Leisure” in-


cludes resting, chatting, eating, sleeping, dancing,
smoking, listening to


cassette, playing games, grooming, etc. “Housework”
includes meal


preparation, food processing, direct childcare,
manufacturing and repair-

ing tools and structures, fetching firewood, fetching
water, etc. “Food pro-


duction” includes foraging, herding, agriculture (see
Figure 3). People


were “Away from camp” to attend ceremonies, markets,
etc.


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156


25


Herding 1.1 %


Bram Tucker
and Alyson G. Young


Females
Males


5


0

Maize Other
Other


Tuber
Foraging Maize


3.9 % Foraging
Foraging


9.5 %
6.1 %


3.0 % 7.5 %


Herding

3.5 %


Adults


Herding 0.8 %


4.8 %


Other


Tuber

Other
Tuber Foraging


Maize


Adolescents Foraging


3.7 %


6.3 %


Herding 0.2 %

Foraging Maize


12.4 %


Maize 1.4 %


Other Foraging


Foraging


6.0 %

8.1 %


2.0 %


Maize 1.9 %


Herding


3.4 % 7.1 %


Tuber

Tuber


Children



20



15



10


5


0


Foraging Foraging


5.6 %


4.9 %



5



Other
Foraging

4.3 %



10


Herding


3.6 %



15



20


25



Percent daylight time allocated


Figure 7.3. Percent of
daylight time allocated to food production activities, by


age and sex.


time allocation. This
implies that Mikea adopt adult roles around the


age of puberty, many
years before marriage.

2. The most important
activity for people in all age/sex groups is


leisure, as is commonly
found in time allocation studies (Johnson


1975; Hill et al. 1985;
Hurtado et al. 1985; Hurtado and Hill 1987:179).


Leisure includes
individual and social non-work activities such as


resting, eating,
playing, chatting, listening to the cassette player, and


playing dominoes.5
Percent time spent in leisure is the main differ-

ence between children’s
time allocation and that of older people.


Children enjoy roughly
twice as much leisure time as adolescents or


adults. Figure 7.4
displays the percent of leisure observations for


each of the 46
individuals in the dataset by age rank. There is a clear,


negative relationship
between age and leisure time for both sexes.


The trend is more
linear for females (R2 = 0.82) than males (R2 =

0.23), for males appear
to increase leisure time again as they reach


middle age.


3. People in all age/sex
categories make important contributions to


household food
production. Adults and adolescents of both sexes


spend similar amounts
of time on food production tasks, while chil-


dren, especially girls,
spend somewhat less time. The most signifi-

cant food production
task for all age/sex groups except adult males


is foraging, and more
specifically, tuber foraging. Figure 7.5 plots


the percent of tuber
foraging time for each of the 46 individuals in the


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Growing up Mikea


Age rank


157



Female Male


Adults



40



30

Adolescents
20


Children



10



R-square = 0.83 R-square = 0.23


100 75 50
25 0 25 50 75 100



Percent daylight time spent in leisure

Figure 7.4. Percent of
daylight time that each of the 46 individuals in the time


allocation dataset
were observed to be engaging in leisure activities, by


age rank and sex.


dataset by age
rank. While the five youngest children spend less time


digging tubers than
others, beyond this point there is little age-


related tendency in
time spent tuber foraging (females, R2 = 0.03;

males, R2 = 0.06).


4. Males and females
do many of the same activities. Differences in sex-


ual division of
labor are less important for children than for adoles-


cents and adults.
The main differences between female and male


labor are time
devoted to housework and herding. Housework, es-


pecially food
processing and meal preparation, is primarily prac-

ticed by females,
while males do most of the livestock care. Adult


males spend less
time foraging for tubers and more time foraging for


honey and small
mammals, although it should be noted that women


do all the same
foraging activities but less frequently.


C. Foraging Return Rates


Foraging returns were
monitored during eight time allocation sampling

months in 1997–99, and
again during a nine-day period in October 2003.


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158
Bram Tucker and Alyson G. Young



Female



Age rank


Male


Adults



40



30


Adolescents
20


Juveniles


10


R-square = 0.03
R-square = 0.06


30 20
10 0 10 20 30


Percent
daylight time spent tuber foraging


Figure 7.5. Percent of
daylight time that each of the 46 individuals in the time


allocation dataset were
observed to be foraging for wild tubers, by age

rank and sex.


Data were collected in a
foraging log that recorded the time people left


camp to forage, the time they
returned to camp, and the number and


weight of resources captured.
Given the significance of wild tuber forag-


ing in the time allocation
data discussed above, we limit discussion to ovy


foraging.

People foraged for ovy in
teams of 1 to 12 individuals, with an average


team size of 2.25 people. In
some cases people returned to camp with bun-


dles of tubers representing
their individual effort, while in other cases they


pooled the tubers into team
bundles before returning to camp. During the


1997–99 field seasons we
observed 114 team foraging events; of the indi-


viduals within these teams,
128 were clearly carrying tubers they them-

selves had procured. In 2003
we sought to enlarge the individual return


rate data. Because our time
was limited to only nine days, we offered a


small cash incentive (500
Malagasy francs, enough to purchase a cup of


coffee and a rice cake in the
market) to foragers each time they allowed us


to weigh their tubers. We
asked that individuals present only the tubers


they dug themselves. We
recorded 124 individual return rates. Parti-

cipants received the same
cash gift regardless of how long they spent for-


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159



Growing up Mikea


aging or how many tubers
they brought back, so these data ought to accu-

rately reflect
individuals’ return rates. However, the cash incentive en-


couraged people to forage
more frequently, and in teams of different


composition than they
would have normally.


“Foraging success”
is quantified as net acquisition rate (NAR). Net ac-


quisition rate is the
number of kilocalories harvested minus the caloric costs


of foraging, divided by
the time spent foraging. We converted the weight

of tubers harvested into
gross caloric gain using the rate 119 kilocalo-


ries/100 grams, based on a
nutritional analysis of an ovy sample conducted


by Kelly and Poyer (1997).
We estimate the costs of foraging at 4.3 kilocalo-


ries/minute (258
kilocalories/hour). We base this estimate on focal follow


observations in which a
minute-by-minute record was kept of tuber for-


agers’ specific
actions-walking, digging, etc. The number of minutes spent

on each action was
multiplied by the caloric expenditure per minute for


that action using data
from Passmore and Durnin (1955) and Durnin and


Passmore (1967). This
estimate is within the range that Durnin and Pass-


more (1967:47) classify as
“moderate work.” Tuber foraging is more costly


than walking (2.1 to 3.1
kilocalories/minute) and less costly than digging


(5.0 to 10.5
kilocalories/minute), the two main actions involved.6

First we explored the
relationship between age and NAR, using the in-


dividual return rate data
from both 1997-99 and 2003. The plots in Figures


7.6a and 7.6b explore age
trends cross-sectionally in each dataset and Fig-


ure 7.7 examines
longitudinal trends for eight individuals who shifted age


categories between
datasets. We offer the following observations:


1. Both cross-sectional
and longitudinal analyses reveal that foraging

efficiency
increases with age, for both sexes. Figures 6a and 6b dis-


play the
cross-sectional data from 1997–99 and 2003 respectively, by


sex. Age rank
accounted for 18 percent of the variance in NAR for fe-


males and 39 percent
for males, in 1997–99; it accounted for 48 per-


cent for females,
and 33 percent for males in the 2003 dataset.


Combining the two
datasets (N = 254), we examine statistical dif-

ferences in the
means of the age categories with a series of inde-


pendent t-tests on
log-transformed data (an analysis of variance


comparing all
categories simultaneously was not possible due to sig-


nificant
differences in variances even when the data are logarithmi-


cally transformed).
The data reveal significant differences in all


pairwise
comparisons. Children experienced significantly lower

mean NAR than
adolescents (t = 5.923, df = 137, p = .000), and than


adults (t = 11.372,
df = 111, p = .000). Adolescents averaged signifi-


cantly lower NAR
than adults (t = 11.372, df = 111, p = .000).


Longitudinal
trends, differences in eight individuals’ mean NAR


in 1997–99 versus
2003, are displayed in Figure 7.7. To control for


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160
Bram Tucker and Alyson G. Young


A


Female


Age rank


Male

Adults
40



30


Adolescents
20


Children



10


R-square = 0.17
R-square = 0.39

5000 4000 3000
2000 1000 0 0 1000 2000 3000 4000
5000



Net Acquisition Rate (net kcal/hr)


B



Female



Age rank



Male

Adults



20



15


Adolescents



10


Children


5


R-square = 0.48
R-square = 0.33


5000 4000 3000
2000 1000 0 0 1000 2000 3000 4000 5000



Net Acquisition Rate (net kcal/hr)


Figure 7.6. A: Individual
foraging returns from 1997–1999, plotted by age


rank and divided by sex
(N=128). Each row of points represents observa-

tions of the individual
of the corresponding age rank on the y-axis. Some


rows have one point,
indicating that the individual was only observed


once. Others have
multiple points representing to multiple observations


of the same individual.
B: Individual foraging returns from 2003, plotted


by age rank and divided
by sex (N=124).


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Net Acquisition Rate



controlling for seasonal / interannual variation


(unstandardized residuals)


161


Growing up Mikea

adolescent to adult


1800 child to adolescent


1300


800


300


-200

-700



-1200



-1700


-2220


2003



1997-

1999


Figure 7.7. Longitudinal
trends in NAR for the eight individuals present in


both 1997–1999 and
2003. The circles and squares indicate the forager’s


mean NAR in both
datasets. The y-axis variable controls for seasonal and


interannual variation
in NAR.


seasonal and
interannual variation, NAR was first regressed by sea-

son and the
unstandardized residuals used as the dependent vari-


able. For seven of
the eight individuals mean NAR increased as


children became
adolescents, and as adolescents became adults.


2. Within most age
categories, males and females harvest tubers at sta-


tistically similar
rates. Mean NAR for female children versus male


children does not
differ significantly (t = .509; df = 77; p = .612), nor

does female versus
male adolescents (t = .268; df = 58; p = .789).


Adult males average
significantly higher NAR than adult females


(t = 2.114; df =
103; p = .037).


Next we explored
foraging team composition, and the success rates of


individuals when foraging
in teams with differing numbers of children.


These analyses are only
performed using the 1997–99 data, since the teams

observed in 2003 may not
have been “natural” due to the cash incentive.


We offer the following
observations:


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162
Bram Tucker and Alyson G. Young


1. Children rarely forage
alone. Children were involved in 41 of the 114

observed team events
in 1997–99. Children foraged alone in two


cases (5 percent);
they foraged with other children in four cases (10


percent); with
adolescents in 12 cases (29 percent); and with adoles-


cents and adults in
23 cases (56 percent). Adult males rarely foraged


with children.


2. Children experience
the same success rates when they forage in the

company of other
children as when they forage with adolescents and


adults (t = 1.537,
df = 6, p = .175). Adolescents and adults forage at


the same rates when
accompanied by children as when not accom-


panied by children
(t = .369, df = 47, p = .714).


D. Economic Dependency and
Self-Sufficiency


Finally, we combine data
from the analyses above to estimate the degree

to which each age/sex group
meets its own food needs through tuber for-


aging. The first column in
Table 7.2 lists the mean tuber foraging net ac-


quisition rate for each
age/sex group. These values are multiplied by the


average time spent tuber
foraging from the time allocation data (converted


to minutes) to arrive at an
estimate of daily caloric production.


We do not know age- and
sex-specific daily energy requirements for

Mikea. Assuming a 1500–2000
kilocalories/day requirement for children


and adolescents and a
2000–2500 kilocalories/day requirement for adults,


then children do not meet
their own caloric needs, while adolescents and


adult females probably do.
Mikea may achieve positive net production


during adolescence,
especially when the caloric contributions of their


other productive activities
are considered. Children only acquire a fraction

of the energy they need to
survive, but in doing so, they reduce adults’


Table 7.2 Average Daily
Caloric Production from Tuber Foraging



Average time Average



Average NAR spent foraging daily production



(net kcal/hr) (minutes/day)a (kcal)


Female children
537 35 313

Male children
505 41 345


Female adolescents
1196 89 1774


Male adolescents
1372 59 1349


Female adults
1851 68 2097


Male adults
2419 22 887


a Duration of activities was
estimated as follows. The time allocation values for tuber forag-

ing reported in Figure 7.3
are percent daylight time. Assuming 12 hours (720 minutes) of day-


light time, 4.9 %
observations = 35/720 minutes.


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Growing up Mikea

provisioning
responsibilities. Children would be capable of positive net


production if they
increased their time allocation to three to four hours


daily.



IV. DISCUSSION


In the remainder of this
chapter we consider how Mikea children achieve


their high return rates,
and also why they forage at lower rates than ado-

lescents and adults. First
we argue that the Mikea Forest is a safe environ-


ment for Mikea children.
Then we discuss why ovy foraging is a feasible


task for young foragers,
in reference to the plant’s ecology and age-specific


foraging strategies.
Finally, we question the degree to which children


strive to forage
“efficiently.”


A. The Mikea Forest as a
Foraging Environment for Children

In a series of papers,
Blurton Jones et al. (1994a, 1994b) and Hawkes et


al. (1995a) explore why
San children of the Kalahari spend little time for-


aging while Hadza children
of Tanzania forage avidly. Although the Kala-


hari Desert of Botswana
and the dry savanna around Lake Eyasi in


northern Tanzania are
similar environments in many ways (and not unlike


the Mikea Forest), these
studies conclude that important ecological differ-

ences affect how young
people best contribute to the household economy.


For Ju/’hoansi San
living along the Botswana/Namibia border, dry season


residential sites are
located along dry watercourses, while the most pro-


ductive foraging
microenvironments are mongongo nut groves on sand


dunes located 13 to 18
kilometers away. While children are efficient nut


gatherers within the
patch, the high travel costs are prohibitive. Children

cannot carry heavy loads
of nuts. There is little shade or water along the


way, so children are
susceptible to dehydration and heat stress. Children


who find it difficult to
keep pace with adults could become lost, and could


die of exposure or be
preyed upon by wild animals. The extra load of nuts


children could contribute
is too small to offset these hazards. Once back in


camp, the nuts must be
boiled and cracked before they yield food value.

Team rates are maximized
if children stay at camp and process nuts while


their mothers forage. By
contrast, Hadza foragers have lower travel costs


and fewer hazards. Hadza
children dig //ekwa (Vigna frutescens) and maka-


lita (Eminia atenullifera)
tubers in the immediate environs of camp without


adult supervision. They
also accompany adults to Cordia and Grewia berry


groves five to seven
kilometers away. Loads of berries are considerably

lighter than loads of
mongongo nuts. The Hadza savanna woodland pro-


vides more shade and
sources of water. Berry foragers consume berries as


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164
Bram Tucker and Alyson G. Young


they pick, providing them
with energy and water for the walk home.

Hadza women could achieve
higher return rates if they left their children


at home and foraged for
tubers, but the costs of foraging with children are


offset by the children’s
own foraging successes. Team rates are maximized


by foraging together.


In comparison to the
Kalahari or Lake Eyasi, the Mikea Forest is both a


safe and productive
environment for children foragers. Children can for-

age alongside adolescents and
adults without depressing their foraging


rates, at least during the
early dry season. Ovy grow within several kilo-


meters of camp. Heat stress
and dehydration are less of a threat than in the


Kalahari, for temperatures
are generally lower, the forest provides shade,


and Mikea children know
several sources of water including rainwater in


tree bowls, wild watermelons,
and water-engorged tubers such as babo

(Dioscorea bemandry) and ba
(scientific name unknown). Parties of children


can forage in the forest and
anthropogenic clearings with little fear of


physical danger, for there
are very few predators, poisonous snakes, sting-


ing insects, or irritating
plants. The only dangerous wildlife are scorpions


and fossa (Cryptoprocta
ferox), a rare predatory mongoose. Zoologists claim


that there are no poisonous
snakes in Madagascar (Preston-Mafham

1991:104) although Mikea
believe a few species to be deadly.


Nevertheless, children
sometimes expressed fear about wandering too


far from their parents and
caretakers. They feared encounters with olo raty,


“bad people,” including
cattle thieves, evil sorcerers, brain stealers, and


vazaha, a term referring
collectively to foreigners, white people, policemen,


military, gendarmes, and
other representatives of authority. They also

feared encounters with
monsters, such as the evil one-horned ungulate


tsongaombe, the undead wraith
tsiboko, and the sinister biby maseake (“cruel


animal”).


Ovy is a more profitable
tuber than Hadza’s //ekwa and makalita tubers.


Ovy grows deeper than
makalita but perhaps at similar depths as //ekwa,


both requiring on average a
hole 75 to 125 cm deep. But ovy is easier to dig

because it grows in sandy
soil devoid of rock. Hadza foragers digging


//ekwa face “a complex
underground jigsaw puzzle and a large excavation,


circumnavigating or removing
large rocks and boulders” (Blurton Jones


and Marlowe 2002:209). Mikea
children can successfully dig ovy with just


their bare hands, although a
metal spade and wooden scoop are preferred


tools. Ovy also has better
food value. Ovy contains 119 kilocalories/100

grams (Kelly and Poyer 1997)
whereas makalita has 73 kilocalories/100


grams and //ekwa has 85
kilocalories/100 grams (Hawkes et al. 1995a:


691). The entire ovy tuber is
edible, whereas the flesh of //ekwa and makalita


are chewed and a quid
expectorated. While some tubers require drying,


grinding, and leaching to
remove toxins, ovy requires only roasting or


boiling.

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Growing up Mikea


Mikea children are
significantly better tuber foragers than Hadza chil-


dren. The child and
adolescent return rates for the Hadza are actually

quite low. Rates reported
in Hawkes et al. (1995a) are expressed in grams/


hour. When converted into
net kilocalories/hour, many of these rates be-


come negative or near
zero.7 Hadza youths averaged 85 net kilocalories/


hour, while Mikea children
averaged 656 net kilocalories/hour. A t-test


finds significant
differences in the means of these two datasets (t = –4.040,


df = 44, p = .000).

B. Children’s Foraging
Strategies


We suggest that young
people have qualitatively different foraging


strategies than older
people. The costs and benefits of these strategies are


explained in reference to
the lifecycle and patch dynamics of the ovy tuber,


and the logic of Charnov’s
(1976) marginal value theorem.


Mikea informants
explained that the ovy plant begins life when a seed

falls from a nearby vine
and germinates during the wet season. A vine


grows up from the seed and
climbs the surrounding vegetation, while a


small, carrot-sized tuber
grows below the ground. When the dry season


begins, the leaves dry up
and fall, and the vine dies and breaks into small


fragments. When the next
wet season rolls around, the small tuber dies


and turns to mush
(fatin’ovy). The upper tip of the ovy, called the “ovy

head” (lohan’ovy),
regenerates a new, larger tuber in its place, and a new


vine grows above the
surface. This continues for many years. Each year,


the ovy tuber is larger in
size and deeper beneath the ground. After two


years the ovy is the size
of a zucchini and the head is 20 centimeters or


more beneath the ground;
after four years the ovy is the size of a person’s


arm and the head is 30
centimeters or more deep. So long as the ovy head

remains in the ground, a
new tuber will regenerate each year, even if the


previous year’s tuber
has been harvested; thus ovy foraging is potentially


sustainable.


Ovy grows wild
throughout much of the forest and anthropogenic


clearings. However,
foragers rarely search for virgin patches. Rather, they


identify a patch by finding
a spot, often around the base of some bushes,

where many holes have been
dug in the ground. Ovy patches have com-


plex, multiyear histories
of partial exploitation and reexploitation that can


be read in the ground by
counting the number and size of holes dug by


previous foragers. A
single ovy patch contains plants of many different


ages. Once the forager
arrives at a patch, he or she begins a process called


mifaokovy, which involves
scanning through the leaf litter and sand for the

white, underground portion
of the vine called the firambony. The forager


can tell by the thickness
of the firambony how old the plant is, and thus the


depth and size of the
tubers. It appears that adults target mature ovy


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166 Bram
Tucker and Alyson G. Young

plants, which provide large
tubers but with considerable digging effort.


Children target young ovy
plants, which provide small tubers with less


digging (adolescents may have
an intermediate strategy).


Charnov’s (1976)
marginal value theorem posits that the length of time


a forager spends in a patch
is determined by the rate of gain within the


patch and the travel time to
the next patch. Foragers remain in a patch

until the gain rate, net of
the cost of travel to the next patch, is maximized,


which normally occurs long
before the patch is depleted. Foragers who ei-


ther travel or harvest at a
more efficient pace ought to spend less time in


one patch before moving to
the next. Because adults are capable of exca-


vating larger tubers, they
maximize their in-patch return rates more rap-


idly than do children, and so
exploit patches faster but less thoroughly.

Children forage in patches
that adults have already exhausted of large


tubers.


Two analyses support this
story. First, there are interesting seasonal dif-


ferences in foraging return
rates and team compositions. Adolescents and


adult females often foraged
alongside children during the early dry season


fararano, and rarely during
the other seasons (Figure 7.8). In fararano, the

recent rains have led to the
regeneration of the tuber patches. Tuber forag-


ing is significantly more
profitable during this time of year (mean fararano


= 2116 kilocalories/hour;
mean other seasons = 1299; t = –4.167, df = 126,


p = .000). On 22 March 1998
over 120 kilograms of ovy were brought into


camp! Children forage with
older people during fararano because both are


interested in the same
patches, even though within the patches they

choose different plants to
harvest. By the middle and late dry seasons,


adults have already exhausted
the patches near camp of the large tubers


they prefer, and are
traveling increasing distances each day. Children con-


tinue to forage near camp for
the smaller (younger, shallower) tubers that


the elders do not consider
worth their while.


In 2003 we counted the
number of tubers harvested in addition to the

weight of the tuber bundle.
This allowed us to calculate the average


weight of each tuber. The
average tuber dug by children weighed 203


grams (carrot sized); by
adolescents, 337 grams; and by an adults, 499


grams. An analysis of
variance reveals that these means are significantly


different (F = 37.433; df =
2; p = .000). There are no significant differences


by sex (t = .172; df = 112; p
= .863).

C. Children’s Foraging
Goals


The main characteristic of
children’s time allocation is prolonged


leisure time. Most of this
time is spent in play. Play activities include


games of tag (especially at
night during full moon), tug of war, gymnas-


tics, dominoes, and
kiombiomby. Kiombiomby is pretend oxcart; the team of


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Growing up Mikea



Females Males


Adults


Adolescents

Children


12 10 8 6
4 2 0 2 4 6 8 10 12



Frequency



fararano



(Early dry season: Mar-Apr)



asotre, faosa


(Mid & late dry season: May-Sep)


Figure 7.8. The frequency
that individuals of different ages and sex foraged


for tubers in teams
that included juveniles, by season.


“oxen” are either two
children (holding forked sticks to simulate the


oxen’s horns), two
blocks of wood, two wild watermelons, or two rats, tied

together. These pairs are
tethered to another stick, melon, or block of wood


to simulate the cart.
Another popular play activity for children is digging


holes in the sand, perhaps
practice for ovy excavation. For girls, grooming


and coiffing are popular
preoccupations. Everyone enjoys listening and


dancing to music played on
to the cassette player or by wandering bands


of minstrels.

Despite enjoying twice
as much leisure time as adolescents and adults,


children spent almost as
much time as older people foraging for tubers.


Table 7.2 suggests that
children could achieve positive net production if


they made modest increases
in their time allocation. So, why don’t they?


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168 Bram
Tucker and Alyson G. Young


Few studies have tried to
predict time allocation to foraging. Hawkes et


al. (1985) argue that
foragers should increase their time allocation to for-


aging on days when foraging
returns are high. Smith (1987) criticized this


study for not fully
appreciating the role of opportunity costs. Hawkes et


al. state that because
foraging has multiple fitness-enhancing benefits, in-

cluding provisioning the
household, generous transfer, and competitive


display, more food
acquisition per unit time (rate maximization) is always


better. Smith counters that
foragers may also be time-minimizers, when ei-


ther the marginal utility of
foraged products diminishes rapidly, or when


activities alternative to
foraging have high fitness-enhancing benefits. A


mother with a young child may
allocate less time to foraging on a high-

returns day, so that she can
return home earlier to nurse. A young man


may return earlier from
hunting when hunting is particularly profitable,


so as to have more social
interaction with potential mates.


We argue that Mikea
children are not trying to be “efficient” at all. There


is little reason for them to
be either rate-maximizers or time-minimizers.


Because parents provision
children from their surplus, children are not

energy-limited. Because they
have few alternative uses for their time, and


their alternatives are not
likely to bestow fitness advantages, they are also


not time-limited. The life of
a child in a small foraging camp is often quite


dull. Children forage for the
physical and mental challenge, and because


it is an enjoyable social
activity. During one focal follow, the senior author


witnessed a “food fight”
between the boys and the girls. Several kilograms

of edible tubers were
destroyed in the ensuing volley. For children, forag-


ing is an extension of play
that occurs outside camp.



V. CONCLUSION


We have examined children’s
time allocation and foraging return rates in


the Mikea Forest of
southwestern Madagascar in order to contribute to


current debates about the
evolution, economy, and ecology of childhood

in foraging societies.


To Kaplan and colleagues
(Kaplan 1997; Kaplan et al. 2000) long child-


hood is the evolutionary
result of a dietary shift to difficult-to-acquire but


high-quality resources such
as wild tubers and hunted game, tasks that re-


quire high intelligence and a
long training period to master. In their the-


ory, lesser learned skill
accounts for why children target different

resources than adults, and
why children forage at lower rates. Consistent


with their explanation, Mikea
increase in foraging efficiency with increas-


ing age, achieving their
highest rates during adulthood. However, several


aspects of age-specific
foraging behavior among Mikea deviate from Ka-


plan et al.’s (2000)
predictions.


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Growing up Mikea


First, Mikea children
do not specialize on collected resources such as


fruits and foliage, but
rather, dedicate similar amounts of time to foraging


for wild tubers, a
high-quality “extracted” resource, as do older people.

Second, Mikea children
do not appear to be actively trained by older


people. Mikea children
experience the same return rates when foraging


with potential trainers
(adolescents and adults) as when foraging with


other children. Women only
forage with children during the early dry sea-


son, when both children
and adults are interested in the same patches.


Third, children may
make rational, “educated” decisions for foragers of

their smaller size and
lesser strength. Children preferentially dig young


ovy plants, whose tubers
are small but shallow, while adults target deeper,


larger tubers. Children
exploit patches more thoroughly than adults.


Fourth, Mikea children
are neither pressured to bring home a full load


of tubers, for they know
adults will provision them, nor are they pressed


for time when foraging,
for they have little else to do. So while they prob-

ably do learn while
foraging, they learn at their own leisurely pace. Their


objectives when foraging
may be primarily social and recreational.


Fifth, despite the
fact that Mikea children are probably not striving for


efficiency, they approach
the age of positive net production during ado-


lescence, considerably
earlier than Piro, Ache, Machiguenga, Ju/’hoansi


San, and Hadza.
Achievement of positive net production is more-or-less

coincidental with the
adoption of adult sexual division of labor and work-


load. Young people
increase their foraging efficiency when opportunity


costs increase.


If children’s
observed foraging return rates are not an accurate repre-


sentation of their maximum
foraging potential—of how efficiently they


could forage if efficiency
were their goal—then age of positive net produc-

tion is a somewhat
ambiguous concept, and so is juvenile dependency. The


question may not be when
children can feed themselves, but rather, When


do children want to work
hard enough to feed themselves? This makes it


difficult to argue that
juvenile dependency early in life permits one to be a


provisioner later in life.
The contrary is also possible: provisioning by


older people may permit
juvenile dependence.

Children’s limited
motivation to self-sufficiency may be part of an in-


herent parent-offspring
conflict. To the degree that children can meet their


own food needs, they free
up parental resources for investment in future


offspring. It may not be
in children’s best individual interest to do this, for


new babies would increase
children’s competition for resources. Previous


studies of the
relationship between children’s production and household

reproduction implicitly
assume that children’s goals are necessarily coop-


erative (Chayanov [1925]
1986; Kaplan 1997; Lee and Kramer 2002).


The possibility that
Mikea children foragers are not trying to be efficient


illustrates a problem with
using age- and sex-specific economic data to test


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170
Bram Tucker and Alyson G. Young


theories about the
profitability of children’s foraging. Contests in which


foragers compete for the best
success rate may produce better data for


evaluating absolute
age-specific ability (see Blurton Jones and Marlowe


2002; Bock 2002a; Walker et
al. 2002).


Mikea children foragers
are more productive than children in other

hunter-gatherer and
forager-horticulturalist populations in large part be-


cause of the ecology of the
Mikea Forest and the ovy plant. Ecological vari-


ation may be just as
important as ethnographic generalization when


constructing and testing
theories about the evolution of childhood. We


hope that the Mikea case will
expand anthropological imagination as to


the range of possibilities in
children’s foraging production.


ACKNOWLEDGEMENTS


We dedicate this chapter to
Dadebo and Tekely, with hopes that they one day be-


come great tuber foragers. We
wish to thank Doug Bird, Frank Marlowe, Nick Blur-


ton Jones, Pat Draper, and
Barry Hewlett for stimulating discussions with the


senior author at CHAGS9 in
Edinburgh. Useful feedback was provided by Paul


Sciulli, Nick Blurton Jones,
Doug Crews, Jim Yount, Bob Kelly, Clark Larsen, Ivy

Pike, Tsiazonera, Jaovola
Tombo, Veve Tantely, Scott Young, and the editors of this


volume. Tucker’s research
among the Mikea was funded by a Fulbright IIE grant,


a National Science Foundation
Dissertation Improvement Grant, a Sigma-Xi grant-


in-aid of research, and
travel grants from the University of North Carolina at


Chapel Hill (1996) and the
Ohio State University (2003).



NOTES

1. In the rest of
Madagascar, the word ovy refers to any tuber, including the do-


mestic potato Solanum
tuberosum. Most Malagasy people refer to wild Dioscorea


yams as ovi’ala, “forest
potato.” We conform to Mikea word choice. Mikea reserve


the term ovy specifically
for Dioscorea acuminata.


2. “Behisatse” is a
pseudonym.


3. Temperatures were
recorded at Behisatse as part of the scan sampling time al-

location project, using a
manual thermometer hung in the shade of the ethnogra-


pher’s hut.


4. We also recorded
absences of entire households during the sampling months.


The inclusion of these
observations allows a tracking of nine households through-


out the entire study period.
These observations will be useful for future studies of


household mobility, but are
not useful here because they reveal no differences in

activity age or sex. The
entire dataset includes 20,241 observations.


5. Actually, people
enjoyed somewhat more leisure time than is measured here,


for only leisure within camp
was visible to the data collectors. Undoubtedly in


some of the cases recorded as
“food production” or “away,” at the observation time


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Growing up Mikea


the subject was actually
taking a break from foraging or farming to enjoy a rest,


chat, cigarette, game of
tag, romantic interlude, etc.


6. While this estimate
is inexact, it is probably workable. If tuber foraging is


more or less costly than
we estimate, say 3 kilocalories/minute or 5 kilocalo-

ries/minute, then
expenditure per hour is 180 or 300 kilocalories/hour rather than


258. This would make minor
changes to our net acquisition values, and no changes


in statistical results and
interpretations.


7. We calculated Hadza
net acquisition rates the same way as for Mikea, as-


suming that foraging costs
4.3 kilocalories/hour. Makalita may be slightly less dif-


ficult, and //ekwa
slightly more difficult, due to the comparative depths and

difficulties caused by
rocks and roots.


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Lectures 7-12

Filed under: Lectures, test 2 — cambriaromance @ 12:09 am

File: Lecture 7 Coastal Migration and Aquatic Resources

The Evolution of the Human Diet

Lecture 7

Coastal Migration and Aquatic Resources

Anthro 4962/5962 Instructor Helen Alvarez

 

The mosaic habitats shown in this photograph of Rajegwesi in Meru Betiri National Park on the south coast of Java illustrate the paleoecology hypothesized for H. erectus in Java by O. Frank Huffman and his colleagues. The photo is from from www.eastjava.com

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 good condition of the skull and the large size of the ancient Mojokerto Delta favor the conclusion that the hominin died in the deltaic environment in which it was deposited. The Mojokerto child therefore provides evidence for a seacoast Homo erectus population in Southeast Asia, and raises interest in the role that maritime adaptation might have played in the dispersal and paleoecology of early hominins (Huffman et al. 2006 449).”

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.

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.

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 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.

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.

Assigned Reading

Erlandson, Jon M., 2001. The archaeology of aquatic adaptations: Paradigms for a new millennium. J Arch. Research. 9(4): 287- 350. For this lecture read pages 287-321.

Also Recommended

Barton, H., 2005. The case for rainforest foragers: The starch record from Niah Cave, Sarawak. Asian Perspectives 44(1): 56-72. (Asian Perspectives available as electronic edition Marriott Library)

Huffman, O. F., Y. Zaim, J. Kappelman, D. R. Ruez Jr., J. de Vos, Y. Rizal, F. Aziz, C. Hertler. 2006. Relocation of the 1936 Mojokerto skull discovery site near Perning, East Java. J. Hum. Evol. 50: 431-451.

Huffman, O. F., and Yahdi Zaim. 2003. Mojokerto Delta, East Jawa: Paleoenvironment of Homo modjokertensis First Results

Semah, Francois, Semah, Anne-Marie and Simanjuntak, Truman, 2003. More than a million years of human occupation in insular Southeast Asia. In Under the Canopy: The Archaeology of Tropical Rain Forests. pp 161-190. ed. J. Mercader New Brunswick, NJ: Rutgers University Press. (available from Marriott Library, Course Reserves)

File: Lecture 8 Optimal Foraging Theory

Anthro 4962 The Evolution of the Human Diet

Lecture 8

Optimal Foraging Theory

University of Utah Fall 2005 Helen Alvarez

 

Shores of Walker Lake (The North American Indian; v.15) from Northwestern University Library,  Edward S. Curtis’s ‘The North American Indian’: the Photographic Images, 2001. Curtis described the setting of this photo: Walker lake, one of numerous saline lakes remaining from a great inland sea that once covered western Nevada and northeastern California, is the seat of a major division of the Paviotso. In the western corner of Nevada it is fed by Walker river, the numerous branches of which head on the eastern slope of the Sierra Nevada in California. Although there is no outlet, the water is not too saline for the thriving of trout and suckers, which were taken on bone hooks, with double-pointed spears, and in gill-nets.

 

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.

 

The first efforts toward understanding resource capture were undertaken by a Canadian entomologist, C. S. Hollings who designed a simple experiment modeling predator and prey interactions. The prey were represented by 4 cm sandpaper discs tacked to a 9 sq. ft. table. The predator was his blindfolded research assistant who tapped her fingers over the table until she encountered a disc, which she removed and set aside before returning to the task of “capture”. The trials were iterated at various prey densities and the relationship between rate of capture and disc density was plotted giving the marginal value curve shown on the right. The rate of capture does not increase in a linear fashion with the increase in the density of the prey.

Hollings’s results became the basis for the distinction between search and handling in the subsequent development of OFT. His simple experiments showed that search time was a function of prey density on the landscape while handling time after encounter was a function peculiar to the prey type. These distinctions became important in all subsequent OFT models.

Optimal foraging theory drew attention to the empirical fact that every resource has a cost of acquisition and processing. These costs make comparisons of energy yields of food meaningless unless handling costs are considered in the comparisons. Here I have posted Table 1 from Bright, Ugan and Hunsaker (2002) showing the energy yield of certain resources taken by Australian and Great Basin foragers and the Kcal/hr return for those resources after processing. I have sorted their table from highest net gain to lowest and plotted the yield with and without processing to show you how the rank of resources changes when the handling costs are debited.

Notice on the figure, drawn from the table, that resources with high energy density (red line) are not necessarily the resources that yield the highest returns after processing (blue line). From this set, the highest ranked resources are all animals with tubers ranking just below small game. Compare the yield of rabbits (#3) and pine nuts (#11) on both curves to see how processing costs change the relative rank of resources. If meat is always the highest ranked resource, across many habitats, why should human foragers ever take any other resource? You should be able to answer this questions by the end of Section II.

 

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.

 

Risk, Uncertainty and the Forager’s Energy Balance

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).

Great Basin Archaeology and Optimal Foraging Models

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.

 

Assigned Reading

Sih, Andrew and Milton, Katherine. 1985. Optimal diet theory: Should the !Kung eat mongongos? Amer. Anthropol. 87(2):396-401.

Hawkes, Kristen and O’Connell, James F. 1985 Optimal foraging models and the case of the !Kung. Amer. Anthropol. 87(2):401-405.

Elston, R. G. and Zeanah, D. W. 2002. Thinking outside the box: A new perspective on diet breadth and sexual division of labor in the Prearchaic Great Basin. World Arch. 34(1):103-130.

Also Recommended

Winterhalder, B., Lu, F., and Tucker, B. 1999. Risk-sensitive adaptive tactics: Models and evidence from subsistence studies in biology and anthropology. J. Arch. Res. 7(4): 301-348.

 

 

 

 

File: Lecture 9 The Ache: Broadleaf Evergreen Foragers

Anthro 4962 The Evolution of the Human Diet

Lecture 9

The Ache: Broadleaf Evergreen Foragers

University of Utah Fall 2005 Helen Alvarez

 

Ache Man Carrying the Head of a Tapir

photo from anthrophoto.com

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.

Note the distribution of zones colored purple, pink, pale yellow and medium green. All of these zones are characterized by wet conditions during the season of maximum solar insolation.

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.

Summary: Five Years of Research

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.

Hill, Kaplan, Hawkes, and Hurtado (1987) Foraging decisions amount Ache hunter-gatherers: New data and implications for optimal foraging models. Ethology and Sociobiology 8: 1-36.

 

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.

Assigned Reading

Hawkes, K., Hill, K., and O’Connell, James F. 1982 Why hunters gather: optimal foraging and the Ache of eastern Paraguay. Am. Ethnol. 9(2): 379-397.

Also Recommended

Hill, K., Kaplan, H., Hawkes, K., and Hurtado, A.M. 1987 Foraging decisions among Ache hunter-gatherers: New data and implications for optimal foraging models. Ethology and Sociobiology 8:1-36.

 

File: Lecture 10 Semiarid African Strategies

Anthro 4962 The Evolution of the Human Diet

University of Utah Fall 2005 Helen Alvarez

Lecture 10

Semiarid African Strategies

 

!Kung Mother with Her Baby on Her Back Gathers Berries

photo by Marjorie Shostak from www.anthrophoto.com

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.

Hadza Country

Hadza country is characterized by rocky hills covered with mixed Acacia scrub and grass leading down to narrow, grass covered valleys. Foragers walking through this landscape to hunt or gather have vistas and landmarks to guide them. In this area children are able to forage away from home and find their way safely back to camp. We will come back to the habitat differences when we discuss juvenile provisioning and ask why the !Kung children are not able to gather more of their own resources.

!Kung Bushland

The Kalahari is a vast plateau at 1100 m above sea level covered by grasses and woodland trees growing to about 30 m in height. It is a nearly featureless plain with few landmarks to guide the hunters and gatherers who make their living from its resources. Even though habitat variability seems low there is some topographic relief provided by lower lying, shallow, seasonal river beds, called molapo, and frying pan shaped dry lakes which hold water for a few days after a heavy rain but are otherwise hard and dry most of the time. After the rains, the Kalahari plains are grass covered. The higher land supports low shrubs that give way to taller woodland trees on the sand dunes and finally, on the outermost zone, open scrub plain. Each of these habitats provides a unique suite of resources for the !Kung. The woman introducing this lecture is gathering berries that grow on the scrub plain.

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.

The four resources pictured here are not quite to scale but give you an idea of the foods listed in Table 8. First is Citrullus lanatus, the wild version of our cultivated watermelon. Second in the top row is Acanthosicyos naudiniana, the Kan melon. Surprisingly these melons can be gathered and kept for many months to supply water and some nutrients. Notice the relative size of these two melons in Tanaka p 60. The tubers that are so important when melons are not available are illustrated in the next panel. This tuber is probably Coccinia rehmannii. The final panel shows a Moroccan truffle of the same genus as the Kalahari truffle, Terfezia. North African truffles are now marketed to the global gourmet food trade.

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.”

It is difficult to get a sense of the size of the eland in this photo but if you convert 300 kg to lbs these animals weigh about 660 lbs. Compare that to beef cattle weighing from 600 to 900 lbs. Lee estimates the edible fraction of game taken by the Dobe !Kung to be 50% of the live weight or 150 kg for an eland. In Table 5, reproduced below, he lists per capita consumption of meat at 230 grams. Calculating .230 kg per consumer day a large eland would provide 652 consumer days of meat. Dividing that by a camp size of 30 gives 21 days. However we know that the meat would spoil long before it was consumed at that consumption rate. Even though meat can be dried in the hot sun of the Kalahari it is most often consumed within a few days of capture.
In this photo of gemsbok you get a good sense of the red sand of the Kalahari and the formidable task of a hunter taking these impressive animals with only bow and arrow. Adult gemsbok males weigh from 167 to 209 kg with females smaller at 116 to 188 kg. Tanaka (p 77) reports that a gemsbok was brought into camp on Oct. 12 and nearly all consumed by the end of the next day. Again if we use Lee’s figures for edible yield and a mid-range figure of 188 for a male animal we can estimate 94 kg of meat in camp. On that same page, Tanaka reports 16 providers and 14 dependents so 30 people consumed, on average, 1.57 kg of meat each day for the two days the meat lasted. To convert these figures to caloric gain we can use Lee’s figures from Table 5 of 690 calories in 230 grams of meat or 4710 cal each of those days from meat.
The figures in the above panel give you some idea of why meat is so highly regarded. Remember they are not figures that could be used to rank meat in an optimal diet because they include no post encounter processing costs. But we might think of the costs in another way. The days with meat in camp are days that men do not go hunting nor women go out to gather. For those days no one pays the energetic costs of traveling to foraging patches, collecting and carrying in the hot sun. Tanaka reports a weight of 200 kg for the greater kudu, pictured here. In the last estimate of meat shared around the camp I neglected the portion eaten by the hunters in the bush. The amount of meat eaten by camp consumers should be reduced by the weight of internal organs and ribs the hunters sometimes eat before they transport the game (see Tanaka in the 3 pages on hunting). Hawkes et al. (1991) report slightly higher yield of edible fraction from live weight for Ache prey, 69 to 88% edible. An experiment at the University of Nevada Reno reported “dressing yield” of over 60% for grass fed beef. Keep in mind that hunters eating wild game would eat more parts of the animal than reported as yield by contemporary americans.

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.”

At the end of the dry season when resources around the water holes have been exhausted the !Kung women have to walk as much as 10 to 15 miles to the nut groves carrying babies as well as the nuts they gather. On these journeys the women have to make a second energetic choice, do they carry toddlers or suffer the time and energy constraint of going at the slow pace dictated by the capabilities of the youngsters. Older children are left in the camp with other caretakers but nursing babies are never left at home. In groups subsisting entirely on wild foods, interbirth intervals (IBIs) are nearly 4 years and babies are at least 3 years old before weaning. Even though !Kung babies are smaller than bottle fed American babies these women are carrying more than 30 lbs of baby and nuts.

Child care constraints always have to be factored into foraging decisions made by !Kung women. How far to travel, what to pursue, how much to carry, how long to gather are all decisions a women faces in the tradeoff between when to have another baby and how to feed the ones she already has. In the photo on the right a women is digging tubers. Blurton Jones and Sibly considered these factors in the backload model to predict the birth spacing that would optimize a !Kung women’s reproductive success. Considering the rigors of walking long distances in the hot sun and the number of nuts needed to feed a family, they predicted interbirth intervals of approximately 4 years, a prediction that matched the actual IBIs fairly closely.

Proteins and Calories from Vegetable and Animal Resources

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.

Hadza Hunter-Gatherers of Lake Eyasi Basin

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.

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.

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.

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).

Energy Conservation?

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.

Assigned Reading

Tanaka, J. 1980. Chapter 2 Subsistence Ecology In The San Hunter-Gathers of the Kalahari: A Study in Ecological Anthropology. pp 53-91. Tokyo: University of Tokyo Press. Go to Marriott Library Course Reserves to download the electronic file.

Tanaka, J. 1980. Hunting 3 pages from Chapter 1 subsection hunting. In Subsistence Ecology In The San Hunter-Gathers of the Kalahari: A Study in Ecological Anthropology. pp 30-35. Tokyo: University of Tokyo Press. Available from the Readings section webCt.

O’Connell, J. F., Hawkes, K. and Blurton Jones N. G. 1988. Hadza scavenging: implications for Plio/Pleistocene hominid subsistence. Current Anthropology 29(2): 356-363.

O’Connell, J. F., Hawkes, K. and Blurton Jones N. G., 1999. Grandmothering and the evolution of Homo erectus. J. Hum. Evol. 36:461-485. Posted under readings Lecture 5 Review p 465 from the Grandmother Hypothesis to p 468 Applying the argument to Homo erectus and p 470 beginning with Climate change and “children’s” resources to p 475 An evolutionary scenario grounded in the Plio-Pleistocene.

File: Lecture 11 Meriam Aquatic Foragers

Anthro 4962 The Evolution of the Human Diet

Lecture 11

Meriam Aquatic Foragers

University of Utah Fall 2005 Helen Alvarez

 

Mer Island

Home of the Meriam

photo © Great Barrier Reef Marine Park Authority 1996.

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.

Reef Flat Foraging

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.

 

Spinefoot to 38 cm

Sea Mullet to 78 cm

Sweet lips to 45 cm

 

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 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.

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.

 

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.

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.

Return rates

Costly Signaling

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.

Assigned Reading

Bliege Bird, R., Smith, E. A., and Bird, D. W. 2001. The hunting handicap: costly signaling in human foraging strategies. Behav Ecol Sociobiol 50:9-19.

Hawkes, K. and Bliege Bird, R. 2002 Showing off, handicap signaling, and the evolution of men’s work. Evol Anthro 11: 58-67.

Also Recommended

Bird, R. 1999. Cooperation and conflict: The behavioral ecology of the sexual division of labor. Evol Anthro 8(2): 65-75.

Smith, E. A., Bliege Bird, R. and Bird, D. W. 2003. The benefits of costly signaling: Meriam turtle hunters. Behav Ecol 14(1):116-126.

File: Lecture 12 Juvenile Foraging

ANTH 4962 The Evolution of the Human Diet

Lecture 12

Juvenile Foraging

University of Utah Fall 2005 Helen Alvarez

 

Overlapping Generations

photo by Marjorie Shostak @ http://www.anthrophoto.com/

 

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’s Foraging

 

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.

 

Table 2

 

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.

 

Table 6

 

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.

 

Juvenile Foraging Size or Experience?

 

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.

 

Figure 1 Age effects on large-hook beach fishing efficiency

 

Martu Children’s Hunting

 

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 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.

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.

 

Mikea Children

 

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.

 

Required Reading

Bird, D. W. and Bliege Bird, R. 2005. Martu children’s hunting strategies in the western desert, Australia. In Hunter-Gatherer Childhoods: Evolutionary, Developmental and Cultural Perspectives. eds. Hewlett, B. S. and Lamb, M. E. pp. 129-146. New Brunswick NJ: Transaction Publishers. Course Reserves Marriott Library

Tucker, B. and Young, A. G. 2005. Growing up Mikea: Children’s time allocation and tuber foraging in southwestern Madagascar. In Hunter-Gatherer Childhoods: Evolutionary, Developmental and Cultural Perspectives. eds. Hewlett, B. S. and Lamb, M. E. pp. 147-171. New Brunswick NJ: Transaction Publishers.

Also Recommended

Blurton Jones, N. G., Hawkes, K, and O’Connell, J. F. 1989. Modelling and measuring costs of children in two foraging societies. In Comparative Socioecology: The Behavioural Ecology of Humans and Other Mammals. eds. Standen, V. and Foley, R. A. pp. 367-390. Oxford: Blackwell Scientific Publications.

Hawkes, K. O’Connell, J. F. and Blurton Jones, N. G. 1995. Hadza children’ foraging: juvenile dependency, social arrangements, and mobility among hunter-gatherers. Curr Anthro 36(2): 688-700.

Bird D. W. and Bliege Bird, R. 2002a. Children on the reef: Slow learning or strategic foraging? Human Nature 13(2): 269-297.

Bliege Bird, R. and Bird, D. W. 2002b. Constraints of knowing or constraints of growing: Fishing and collecting by the children of Mer. Human Nature 13(2): 239-267.

September 24, 2006

Lecture 8

Filed under: Lectures, test 2 — cambriaromance @ 4:10 am

Anthro 4962 The Evolution of the Human Diet

Lecture 8

Optimal Foraging Theory

University of Utah Fall 2005 Helen Alvarez

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Lecture 7

Filed under: Lectures, test 2 — cambriaromance @ 4:00 am

WebCT

 

The Evolution of the Human Diet

Lecture 7

Coastal Migration and Aquatic Resources

Anthro 4962/5962 Instructor Helen Alvarez (more…)

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