Little, M. - Human Biology of African Pastoralists

YEARBOOK OF PHYSICAL ANTHROPOLOGY 32:215-247 (1989)
Human Biology of African Pastoralists
MICHAEL A. LITTLE
Department ofdnthropology, State University of New York,
Binghamton, Binghamton, New York 13901
KEY WORDS Pastoral nomads, Environmental adaptation, Biobehavior,

Nutrition, Growth, Reproduction, Health
ABSTRACT Pastoralism in Africa is a subsistence tradition that has a

long and complex history. During the course of several thousand years of
nomadic and semi-nomadic herding practices, pastoral populations have
successfully exploited African savannas and rangelands by maintaining
highly mobile and low-density human populations and by maintaining sym-
biotic relationships between the people and their livestock. This review will
cover the pastoralists’ adaptations to their environments and the measures
of health and adaptability of pastoralists within a biobehavioral framework.
The measures that are considered are: diet and nutrition; infant, child, and
adolescent growth; adult size and body composition; activity and physical
fitness; reproduction; and disease. Several integrated issues (seasonal hun-
ger, drought, sedentarization) are discussed as important areas for current
and continued research.
Pastoralism is a pattern of subsistence that has been practiced by human pop-
ulations for a t least 10,000 years. Today, a form of traditional pastoralism involv-
ing nomadic or semi-nomadic movement and non-Western technology exists in
many parts of the world and supports millions of pastoral people (Dyson-Hudson
and Dyson-Hudson, 1980; Galaty et al., 1981; 1’Equipe Ecologie et Anthropologie
des Societes Pastorales, 1979; Monod, 1975; Weissleder, 1978). These population
numbers, however, are declining because pastoralists’ vast territories are dwin-
dling as the result of expanding non-pastoral populations, ranching schemes, and
attempts by national governments to sedentarize or settle the nomad. As pastoral
populations decline and disappear, we will lose a significant amount of the cultural
and biological diversity that enriches our species. The purpose of this review is to
document some of the known biobehavioral diversity of existing pastoral popula-
tions while these populations still retain some of their traditional culture and
subsistence. At the same time this will be done within the context of several major
problems that pastoralists face in maintaining this pattern of subsistence.
AFRICAN PASTORALISTS AND THEIR ENVIRONMENT

Tropical savanna ecosystems and grazing lands
Throughout the world, pastoralists are distributed across lands in which a large
proportion of the plant community is constituted of grasses, forbs, and shrubs. In
the tropics, these grazing lands are identified as savanna. Situated between the
moist, tropical forest and the arid and semi-arid desert zone, savanna lands cover
as much as one quarter of the world‘s surface. Compared with other biomes, sa-
vanna lands have a pattern of plant productivity in which most of the biomass is
available to fauna a t higher trophic levels. Hence, savanna lands are often occu-
pied by large populations of grazing and browsing mammals.
0 1989 Alan R. Liss, Inc
216 YEARBOOK OF PHYSICAL ANTHROPOLOGY [Vol. 32, 1989
The principal climatic factors that structure savanna ecosystems are tempera-
ture and a highly seasonal, monsoon pattern of precipitation. Precipitation can
range from less than 200 mmlyear to more than 1,000 mmlyear, but it is the highly
seasonal nature of the rainfall that limits forest growth and maintains the grass-
lands. Harris (1980) noted that the most crucial variable in defining savannas is
the duration of the dry season. For example, a tropical climate with adequate
rainfall and a dry season of less than 2.5 months will result in a closed-canopy,
moist forest. He defined three savanna zones: 1) humid, with 1,000-2,000 mm
annual rainfall and a dry season of 2.5-5.0 months; 2) subhumid, with 500-1,000
mm rainfall and a dry season of 5.0-7.5 months; and 3) semiarid, with 250-500
mm rainfall and dry season of 7.5-10.0 months. At the mesic or wet end of the
continuum is the woodland savanna, but with an open-canopy distribution of trees
to allow grasses to grow. Toward the xeric or arid end of the continuum is a full
grassland that grades into a scrub savanna. Seasonal precipitation much below
150 mmlyear leads to desert conditions and failure to support either wild grazing1
browsing animals or pastoralism.
Primary plant productivity (annual growth) of the savanna varies according to
dry-season duration and annual precipitation. Bourlihre and Hadley (1970) and,
later, Jewell (1980) reviewed published values that ranged from a net primary
plant production during drought of 32.6 gJm’Iyear (dry biomass) to nearly 2,000
glm’iyear in a highly productive humid savanna. Grasses tend to grade from a high
proportion of perennials in humid savannas to more annuals in dry savannas. In
the dry savanna, the resulting flush of annual vegetation with the onset of the first
rains after many months of seasonal drought is extraordinarily rapid (several
days) and quite spectacular. In woodland and shrub savanna and along riverine
woodland tracts, trees provide browse vegetation which is important to browsing
wild mammals as well as some domestic species (camels and goats). Walker (1980)
contends that woody vegetation in arid savannas helps to maintain ecosystem
stability. Stability is maintained by trees contributing to soil preservation, aiding
in water infiltration during heavy monsoon rains, and storing considerable bio-
mass that can be consumed by herbivores throughout the year.
African savanna lands cover about a half of the continent. They can be parti-
tioned into three main zones: 1)a sub-Saharan belt zone, and two others in 2) East
Africa and 3) South-Central Africa (Harris, 1980). The sub-Saharan belt can be
divided into Sahelian (dry) and Sudanic (wetter) sub-zones (Bernus, 1988). Most
pastoralism on the continent is practiced within these three zones. Figure 1 illus-
trates the distribution of savanna lands in Africa as modified from satellite imag-
ery synthesized by Tucker and his colleagues (1985). What is clear from this work
is that an “average” map such as this does not represent very well the substantial
seasonal variation and year-to-year variation in rainfall and drought that charac-
terize most of the continent, even the tropical forests.
Up until the beginning of the present century large wild herbivores were widely
distributed throughout African savannas (Jewell, 1980). Widespread declines in
large mammalian wildlife have resulted from expansion of pastoralism, human
population increase, and successful hunting practices by colonial Europeans.
Poaching for ivory, rhinoceros horn, and meat has contributed to recent declines in
wildlife, and further reduced competition between wild and domestic herbivores.
Carrying capacities of savannas for large herbivores depend on variation in rain-
fall and vegetation properties. Although wild herbivores are thought to be more
efficient at forage utilization than domestic herbivores, there do not appear to be
major differences in the carrying capacities of the land for the two classes of
animals (Jewell, 1980). Grazing strategies of pastoralists, such as maintaining
species diversity (grazing succession) and livestock mobility, probably enhance the
efficiency of foraging and increase the domestic livestock carrying capacity (Dyson-
Hudson, 1980). Another major variable contributing to the present distribution of
wild and domestic herbivores is the presence of the tsetse fly (Glossina spp.) and
animal trypanosomiasis (Desowitz, 1980, 1981). Wild herbivores are more resis-
Little] AFRICAN PASTORALISTS 217
20 30 40 ;o
SEMIARID U
20
3a
Fig. 1. Vegetation and climatic zones of Africa. Based on LANDSAT (Tucker et al., 1985) and other
sources of data (Harris, 1980).
tant to trypanosomiasis than are domestic species, a circumstance that limits

movement of domestic herds and their herders over vast areas of East and West
Africa.
There are several pronounced stresses which arise from savanna environments
that have an impact on humans. These include: limited vegetation and water
resources; high ambient temperatures, which impose a heat load on livestock and
humans, alike; seasonal and longer-period aridity; and endemic tropical diseases,
some of which affect both livestock and humans (Little, 1980).Human responses to
these environmental stresses are structured and constrained by the requirements
of a pastoral existence, internal operations of cultural systems, and biobehavioral
adaptability of the people.
History and the spread of African pastoralism
The prehistory of pastoralism in Africa is a dynamic montage of cultural inno-
vation, population movement of both humans and domestic species, and diffusion
of language and livestock practices. A variety of sources of information can be used

to document the presence and development of pastoralism. Rock paintings, al-
though difficult to date, depict cattle a t about 3400 B.C. in the central Sahara and
later times in Eritrea (Phillipson, 1985). Archaeological evidence confirms that
animal domestication was present in Africa as early as 4000 to 5000 B.C. (Phillip-
son, 1985; Smith, 1980; Williams, 1984).Considerable research has been conducted
on linguistic reconstruction (Ehret, 19841, but archaeologists caution against un-
critical acceptance of glottochronological dates (Phillipson, 1985). There is some
agreement that goats, sheep, and some varieties of domestic cattle were introduced
into the continent from Southwest Asia, but there is tentative evidence from
Capelletti cave in northern Algeria that suggests local domestication of cattle as
well (A.B. Smith, 1980, 1984).
Animal domestication spread westward along the Mediterranean coast of North
Africa during the fourth millennium B.C., when, by about 2000 to 3000 B.C., it
entered the highlands of Ethiopia and crossed the Sahara into the Sudanic zone
(Smith, 1984). These movements may have been stimulated by a drying trend in
the Sahara which moved the tsetse fly belt (and trypanosomiasis) further south
and allowed people to occupy previously infested areas (Smith, 1984). Penetration
into the Sudanic zone from the north with cattle and small stock continued until
about 1500 B.C. and was associated with both cultivation and the increasing aridity
of the Sahara (Clark, 1980). Linguistic evidence suggests that around the same
time or earlier, Eastern Cushites from southwest Ethiopia moved with their cattle
along the Azanian coast and inland into much of East Africa (Clark, 1980). With
linguistic evidence, however, it is difficult to determine whether populations ac-
tually moved into an area or whether the language was introduced through diffu-
sion. Rightmire (1984), for example, observed that there is little archaeological or
skeletal evidence to support the linguistic model of large-scale population move-
ment. Also, early dates of 2500 to 3000 B.C. have been cited for the presence of
livestock in the Lake Turkana Basin (Barthelme, 1984), so the Cushitic movement
or diffusion might have simply reintroduced domestication practices. A detailed
description of the archaeological remains and adaptations of these Savanna Pas-
toral Neolithic peoples is provided by Ambrose (1984).
Murdock (19591, from Egyptian historic sources, identified the Cushitic Beja of
southern Egypt and the Sudan as among the earliest of identifiable pastoral no-
mads in Africa (2700 B.C.). Since they still maintain this way of life, the Beja may
very well be the oldest extant pastoral nomadic population in the world.
By A.D. 300, a Bantu-speaking expansion, associated with iron use, carried pop-
ulations eastward into the area around Lake Victoria, including Kenya, Tanzania,
and Uganda (Phillipson, 1985). Many of these East African Bantu peoples, who
appeared to displace the resident Cushites, acquired livestock as well as herding,
which led, ultimately, to the mixed subsistence that characterizes these people
today. Another population intrusion involved the movement of the Nilotes (also
referred to as Para-Nilotes) from the upper Nile River Valley southward into East
Africa. Based on linguistic evidence, this expansion may have begun any time
between 0 and A.D. 1000 after the Nilotes had developed the “milking complex,”
and led to displacement of both Cushites and Bantu (Ambrose, 1984). Nilotic ex-
pansion continued well into the 19th century, when later movement was then
limited by European colonial rule (Were and Wilson, 1972). For example, the
Nilotic Turkana, an offshoot of the Jie, entered their present area west of Lake
Turkana, Kenya, in the early 18th century and had expanded to their present
distribution by 1900 (Lamphear, 1976a, 1988). At about the same time as the
beginning of the Nilotic expansion in East Africa (A.D. 1100), West Africa was
experiencing the Ful or Fulani pastoral expansion from the western Sudanic zone
in Senegal to as far east as Cameroon and the Central African Republic (Hiernaux,
1975). The Fulani were expanding into southeast Niger as late as the early 1900’s
(Horowitz, 1975).
Southward movement of pastoralism to Botswana and the Cape area probably
took place around 0 A.D. with the diffusion of ovicaprid (small stock or sheep and
goats) livestock practices to the existing hunting/gathering populations (Klein,
1984). The western and southern Cape Khoikhoi or Hottentot pastoralists (now
extinct) were closely related linguistically and physically to the Bushman hunter/
gatherers (Ehret, 1984; Hausman, 1984).
These population movements and diffusion spread domestic animal species
throughout North and sub-Saharan Africa. The question of cattle domestication in
Africa is unresolved at present (A.B. Smith, 1980). If domestication took place in
North Africa, then one of the longhorn species of Bos primigenius or Bos ibericus
was the wild ancestor. If not, then some cattle are derived from the Mediterranean
Bos taurus. Bos indzcus, the Indian humped zebu species of cattle, which is more
resistant to dry conditions, entered the continent at a later time. The introduction
of sheep, which are represented by a distinct non-wool-bearing and fat-tailed va-
riety (Ouis aries), and goats (Capra hircus) may even antedate that of cattle (A.B.
Smith, 1980,1984). The date of entry of dromedary camels (Camelus dromedarius)
into the continent is not clear at this time, but could have been about 3000 B.C. or
earlier (Brandt, 1984; Phillipson, 1985). The only animal that was certainly do-
mesticated in Africa is the donkey (Equus spp.), an essential transport animal for
nomads and semi-nomads (Clark and Brandt, 1984).
Distribution of African pastoral populations
Today, pastoral populations in Africa are distributed widely over grassland and
woodland savannas, from the most arid, near-desert zones to the moist woodlands
that border on the tropical forest. It is important, a t this point, to distinguish
between agropastoralists and pastoralists. Agropastoralists can be identified as
sedentary cultivators who also keep livestock for milk or meat. The proportional
mix of food in the diet is less important as a criterion of subsistence pattern than
is the agropastoralist’s settled existence and farming practice. Pastoralism, on the
other hand, can be defined as a subsistence strategy that involves a dependence on
spatial mobility where livestock are moved to seasonal or ephemeral pastures
(Dyson-Hudson and Dyson-Hudson, 1980). It is generally the case that more food
is derived from livestock and trade of livestock products in pastoralists than in
agropastoralists. Of course, the distinction between pastoralists and agropastoral-
ists blurs when some nomadic pastoralists cultivate during very wet periods and
some agropastoralists migrate during drought. The geographic distribution of pas-
toralists is more a reflection of the need of agropastoralists for adequate rainfall
for cultivation. As a result, pastoralists can be found more commonly in the drier
areas of savanna lands where marked seasonality and variability in rainfall are
the norm, and where cultivation is impractical.
Geographic distribution of some African pastoral populations is presented in
Figure 2. Many of the populations in Figure 2 are reasonably well known from the
ethnographic literature. Principal ethnographic references by pastoral population
are the following: Baggara (Cunnison, 19661, Barabaig (Klima, 1970), Bedouin
(Asad, 19701, Borana (Dahl, 19791, Dassanetch (Almagor, 19781, Fulani (Dupire,
1962; Stenning, 19591, Galla (Ensminger, 19841, Jie (Gulliver, 1955; Lamphear,
1976b), Karimojong (Dyson-Hudson, 19661, Maasai (Jacobs, 1975), Nuer (Evans-
Pritchard, 1940), Rendille (Spencer, 19731, Samburu (Spencer, 1965), Sebei (Gold-
Schmidt, 19761, Somali (Lewis, 19611,Tuareg (Bernus, 1981; Swift, 19751,Turkana
(Gulliver, 1951, 1955; McCabe, 1985), and Zaghawa (Tubiana and Tubiana, 1977).
Only a handful of these populations have been studied from perspectives of health
and human biology.
BIOBEHAVIORAL ATTRIBUTES OF PASTORALISTS
Although there is a remarkable degree of biobehavioral variation among African
pastoral populations, some generalizations still can be outlined (Dyson-Hudson,
1972; Dyson-Hudson and Dyson-Hudson, 1980; Goldschmidt, 1979, 1981). Inspec-
tion of these generalizations should guide readers toward a more complete under-
Fig. 2. Distribution of some pastoral populations in Africa.
standing of how conditions of behavior, within environmental contexts, influence

pastoralists’ health and biological adaptability,
Grazing territory
Most pastoral populations identify grazing lands as a public resource (Gold-
Schmidt, 1981), although such agreements usually apply only to members of a
given cultural unit. Since pastoralists are often in competition with agricultural
peoples for land resources, they are usually confined to what is identified as the
least desirable land for cultivation, or “marginal lands.” These are often semi-arid
or cool savanna, steppe, or prairie grazing lands that are subject to periodic
drought and strongly pulsed annual climatic variation. In addition, pastoralists
require large land areas to exploit because of the low and unpredictable biotic
productivity of the environment. Seasonal and longer-term fluctuations in rainfall
produce marked variations in the availability of forage resources, which, in turn
effect the availability of livestock food resources and lead to periodic human hun-
ger and, occasionally, starvation. Although pastoralists’ livestock do buffer major
environmental cycles of resource availability, such buffering is always imperfect,

and the people experience considerable periodic nutritional stress. These condi-
tions lead to pastoralists’ need to be highly mobile by moving herds from place to
place in search of palatable forage for their livestock.
Mobility
Many anthropologists have emphasized the essential requirement of spatial mo-
bility for pastoralists (Burnham, 1979; Dyson-Hudson and Dyson-Hudson, 1980;
Horowitz, 1981). Pastoralists must have adequate grazing lands within which to
move in search of forage with adequate protein and energy to meet the needs of
their livestock. Mobility is crucial when forage resources are limited and herds
must be moved quickly to t a p ephemeral grass flushes. Constraints on pastoral
movement that are imposed from the outside are often based on rainfall data from
average or good years, whereas years of poor rainfall are the times when mobility
is most needed (Horowitz, 1981). It is well known by rangelands ecologists that the
carrying capacity of pastoral grazing lands, particularly the semi-arid savannas, is
a function of the driest time of the year or a drought year rather than a n “average”
year (Coughenour et al., 1985; Ellis and Swift, 1988).
Livestock numbers
African pastoralism usually involves one or more of five species of livestock:
cattle, goats, sheep, donkeys, and camels. Camels and goats, as mixed feeders
(browse and graze), are better adapted to dry conditions, whereas cattle and sheep
require grass vegetation and are better adapted to wetter savannas (Wilson et al.,
1985). Ellis and Swift (1988) demonstrated that there are advantages to maintain-
ing large numbers of livestock by one pastoral group, the Turkana, since periodic
drought is the principal livestock population control. Droughts, and corresponding
declines in herd numbers, are sufficiently frequent in this relatively arid savanna
to allow ecosystem persistence despite maximization of livestock population
growth. Whether the primary goal of pastoralists is maximization of livestock
rather than of human populations has been asked, since periodic drought and
raiding tend to serve as livestock population controls, but have a limited influence
on humans. Rada Dyson-Hudson (1981) has suggested that it is the pastoralists’
objective to support a maximum number of people, not livestock. Maximization of
human numbers by pastoralists, as Neville Dyson-Hudson (1980) noted, may be a
response to the rapid rate of livestock recovery in contrast to the slow rate of
human population growth. Many of these questions are unresolved at present.
Human and livestock demography
Human population structure and dynamics are linked closely with those of the
livestock. Numbers and growth rates of both livestock and people must be balanced
for the system of subsistence to operate successfully. Pastoralism is a labor-inten-
sive activity that requires a close association between numbers of the food-pro-
ducing livestock and the numbers of the livestock-managing human labor force.
Moreover, many pastoralist societies have complex systems of exchange and reci-
procity where livestock serve as a kind of currency (Goldschmidt, 1981) and where
bridewealth and other socially sanctioned practices serve to redistribute animals
throughout the population. I n many pastoral societies, livestock are loaned to
relatives or friends, and sons who wish to marry and set up their own households
must be provided with enough animals to begin their own herding operation.
Another characteristic of pastoralists is a distinct pattern of division of labor by sex
and age that produces differentials in physical activity, food requirements, access
to food, and exposure to disease. Among the Turkana, adult male herd owners,
aside from walking, are relatively sedentary, whereas adult females are very ac-
tive and have responsibilities for carrying firewood and water and milking ani-
mals.
TABLE 1.Measures of health and adaptability

Cateeories Measures
Diet and nutrition Food availability; adequacy of intake of protein and energy;
regularity of intake; adequacy of intake by sex, age, and
activity groups; unusual foods and food habits
Infant, child, and adolescent growth Size at any given age; weight for height; rates of growth;
proportions; body composition (fat and muscle); environmental
and biobehavioral influences on growth processes
Adult size and body composition Height; weight for height; fat and body muscle content;
annual and seasonal variations
Activity and physical fitness Day-to-day activities; cardiovascular status; aerobic and
submaximal work capacity; age changes; environmental
influences; division of labor
Reproduction Fertility and reproductive health; completed fertility; limiting
factors to reproductive health
Disease Presence or absence and classes of morbidity; rates of
morbidity by age and sex; history of morbidity by individual
and bv ~ O U D mortalitv
: and links with morbiditv
HEALTH AND ADAPTABILITY ASSESSMENT

There are a number of approaches that can be taken to explore the biology of
pastoralists. One approach is to assess the health, well-being, and general status of
pastoral peoples according to a number of broad categories of biobehavioral adap-
tation. Some of these indicators of health and well-being were identified by Baker
(1977)as “measures of biological fitness” and characterized by Mazess (1975,1978)
as “adaptive domains.” In most cases, a “transdisciplinary approach must be
taken to solve problems within each of these categories (Baker, 1982, 1988). Also,
it should be understood that each category is an artificial construct to facilitate
analysis, and, hence, a comprehensive understanding of the biobehavior of any
population will depend on cross linkages among the categories (Little and Haas,
1989).
Measures of health and adaptability are identified in Table 1with brief lists of
items appropriate for data collection and analysis. This approach to an understand-
ing of pastoralists was taken in an earlier review of East African populations
(Little, 1980). The present review will serve both as an update incorporating a
decade of new research and as an expansion of the base of information to areas
outside of East Africa.
MEASURES OF HEALTH AND ADAPTABILITY

Diet and nutrition
About a decade ago, it was observed that no systematic surveys of the nutritional
status of pastoralists had been conducted (Little, 1980; Wheeler, 1980). Since that
time, we have begun to accumulate considerable dietary data on at least five
pastoral populations (Baggara, Fulani, Maasai, Tuareg, and Turkana). It is evi-
dent from these studies that pastoral diets show considerable among-population
variation and that this variation is dependent both on degree of cultivation vs.
pastoralism and patterns of trade and exchange.
Agropastoralists such as the Awlad Hamid, who are Arabic-speaking Baggara
nomads of the northern Darfar in the Sudan, have a diverse diet including millet,
meat, milk, okra, onions, garlic, red papper, dried tomatoes, and sesame and
groundnut oil (Holter, 1988a). Camel milk and milk products only constitute about
20-25% of the food energy of the diet (Holter, 1988b). Malian Tuareg (Kel
Tamasheq) pastoralist foodstuffs include millet and rice, milk and meat, with milk
and cereals consumption varying by season (Wagenaar-Brouwer, 1985). At the
other extreme are the Maasai and Turkana, Tanzania and Kenya pastoralists,
whose diets of milk and other animal products from livestock may amount to
nearly all of their caloric intake a t certain times of the year (Galvin, 1988; Nestel,
1985). Despite a heavy reliance on milk in diets of Maasai and Turkana, even these
two groups differ somewhat because many Kenya Maasai are now confined to
ranching schemes where commercial meat production is emphasized and traded
foods are important in the diet (Grandin, 1988). Also, the East African practice of
bleeding livestock to supplement the diet with blood during times of low milk
production is seldom practiced any longer by pastoral Maasai (Nestel, 1985), al-
though it is quite common in the Turkana (Galvin, 1985). There is substantial
variation in diet even among subsections of a tribal unit. For example, in nomadic
Ngisonyoka Turkana, milk intake is more than 90% of calories in the rainy season,
while neighboring Ngikamatak Turkana almost never consume more than 50% of
their calories in the form of milk a t any time of the year (Galvin, 1988). Fulani,
who are distributed widely throughout West Africa, also show marked variation in
food practice. Some Fulani populations subsist largely on milk, millet, and sor-
ghum (Benefice et al., 1984; Hilderbrand, 1985; Loutan, 1985), while others con-
sume staples of rice and fish in addition to milk and grains (Wagenaar-Brouwer,
1985).
Despite the variation expressed by pastoralists in dietary intakes, there are a
number of commonalities. Milk, meat, and in certain groups, blood, provide sub-
stantial animal protein, and relatively high fat, intakes. For the Maasai (Nestel,
1985) and the Senegalese Fulani (Benefice et al., 1984) protein intakes were re-
corded that were more than 200% the recommended daily intake (RDI), while
Galvin (1985) found that protein intakes in some age groups of the Turkana were
as high as 400% of RDIs. Serum protein levels were also high in small samples of
Maasai, Turkana, Boran, Rendille, and Samburu pastoralists from Kenya
(Gatenby Davies and Newson, 1975). Fat intakes varied from 73% of dietary cal-
ories in the Fulani (Murray et al., 1978a) to 30 or 40% of calories in the Maasai
(Nestel, 1985) and Turkana (Galvin, 1985). Much of the variation in fat intake is
a function of how much milk is consumed at different times of the year (Galvin and
Waweru, 1987; Nestel, 1986). Despite the high intakes of fat, pastoralists have not
manifested the signs of cardiovascular disease in such high prevalence in Western
populations with comparable fat intakes (Ho et al., 1971; Mann et al., 1972; Shaper
et al., 1961, 1963).
Energy or caloric intake, on the other hand, appears to be low for African pas-
toralists. Energy intake for Maasai and Turkana ranged between 55 and 75% of
RDIs for men and women (Galvin, 1985; Little et al., 1988a,b; Nestel, 19851, sug-
gesting that some of the excess protein intake is utilized for energy. These values
may not be quite as low as they seem, since FAO/WHO (1973)energy requirements
may be set too high (Leslie et al., 1984). Ascorbic acid was marginally low in a
number of pastoral populations (Benefice et al., 1984; Gatenby Davies and New-
son, 1974a,b, 1975; Nestel and Geissler, 1986), although clinical signs of vitamin
C deficiency were absent. Compared with many vegetables, milk is relatively low
in ascorbic acid (1.5-1.7 g/lOO ml), but since many pastoralists rely on milk as a
primary source of this vitamin, intake must be marginally adequate (Gomez,
1982).Hilderbrand (1985)found night blindness in Malian Fulani a t the end of the
dry season which she interpreted as symptomatic of vitamin A deficiency. Some
confirmation was provided by Shelley (1985), who suggested that vitamin A defi-
ciency was a problem in the Turkana based on relatively high frequencies of Bitot’s
spots and xerophthalmia of the eyes. Iron deficiencies were found also in the Tur-
kana (Murray et al., 1980) and in Somali nomads (Murray et al., 1978b). About
25%of the Maasai women and children studied by Nestel and Geissler (1986) were
anemic, which is suggestive of iron deficiency. The iron content of milk is low;
hence, high milk and low blood and meat intakes could be associated with iron
deficiency anemia. This problem of limited iron intake in women would be wors-
ened by blood losses and increased iron needs during menstruation. Many or all of
these nutritional conditions vary on a seasonal basis, and are not present in all
agelsex groups.
Fig. 3. Seasonal variation in milk consumption as a percentage of total energy intake for Ngisonyoka
Turkana of Kenya. (Data from Galvin, 1985.)
Seasonal food availability exerts a considerable influence on the diets of most

pastoral populations. Intakes during and immediately following the rainy season
tend to be well above RDIs for energy, protein, and other nutrients, while intakes
at the end of a dry season or prolonged drought fall below RDIs. Seasonal changes
in energy intakes and activity requirements are reflected by cyclic fluctuations in
body weights of adults and children that have been documented for several popu-
lations (Benefice and Chevassus-Agnes, 1985; Galvin, 1985; Hilderbrand, 1985;
Little et al., 1988b;Loutan, 1985; Wagenaar-Brouwer, 1985). Body weight changes
will be discussed in a later section.
In many nomadic pastoral populations, milk is the staple food and any fluctu-
ation in this supply will require incorporating other foods such as meat, grains, or
blood. Figure 3 illustrates seasonal variation in human milk consumption for the
Turkana. The marked seasonal variation in milk production of Turkana livestock
is a function of the health and productivity of several species, each of which has a
characteristic reproductive cycle and pattern of lactation (Wilson et al., 1985).
Turkana nomads as well as neighboring Dassanetch bleed animals only when milk
production declines or milk is unavailable (Galvin, 1985;Almagor, 1978:59). Other
populations almost certainly follow this practice. Livestock blood, then, serves as
a supplementary food that has seasonal use rather than as a staple food. The
consumption of grains such as millet, sorghum, and maize is also seasonal in many
populations and is also a function of milk production. Some populations, such as
the Turkana, trade for grain products (Galvin, 19851, while closely related popu-
lations, such as the Karimojong, cultivate some of their own grains (Dyson-Hudson
and Dyson-Hudson, 1970). Another important seasonal dietary pattern concerns
meat consumption. When animals are in poor condition at the end of a dry season
or prolonged drought, animals that are about to die will be slaughtered and meat
consumption will be artificially inflated. This practice has been documented for
the Fulani (Loutan, 1985) and the Turkana (McCabe, 1985,1988) and is probably
typical of all pastoralists who reside in arid or semi-arid savannas.
There is considerable variation in dietary intake and nutritional status by sex
and age. Those who have access to certain foods are likely to consume more than
others, and, in many cases, women do have greater opportunity to consume milk
than men. This has been recognized in ethnographic studies of pastoralists (Du-
pire, 1963; Wienpahl, 1984). Wagenaar-Brouwer (1985) found that Tuareg
(Tamasheq) women consumed more milk than men, with up to 58% of their calories
Fig. 4. Milk consumption in Ngisonyoka Turkana by age and sex groups. (Data from Galvin, 1985.)
in the form of milk. These women consumed about one and a half times as much
milk as the men. The same pattern held for the Turkana, where women consumed
about 56% of their calories and men consumed about 50% of their calories as milk
(Galvin, 1985). For Turkana children, milk was consumed in greater amounts than
in adults. Age and sex variation in milk intake for the Turkana is presented in
Figure 4 to illustrate milk consumption patterns for a heavy milk-using popula-
tion.
A final note on diet and nutrition concerns lactose intolerance and the practice
of dairying, the most common pastoral subsistence pattern in Africa. This question
of intolerance or malabsorption of lactose or milk sugar in milk-consuming pasto-
ral populations is an interesting one. Most peoples around the world lose the
ability to digest lactose several years after infancy. It appears that tolerance of
milk and milk products containing lactose beyond the period of infancy can occur
in two ways: 1) by a hereditary ability to produce the lactose-digesting enzyme
P-galactosidase, and 2) by “adaptation,” that is, “an increase in the amount of
lactose that can be consumed without significant gastrointestinal symptoms by a
lactose maldigester with continuing lactose intake” (Scrimshaw and Murray,
1988). Many pastoral populations would be in serious trouble if their members
became ill by consuming milk. Despite this problem, frequencies of lactose mal-
absorption show considerable variation across pastoral populations. For example,
Sudanese Beja had 17% (Bayoumi et al., 1982) and Zaire Tutsi 10% malabsorbers
(Cox and Elliott, 19741, while Maasai had 62% (Jackson and Latham, 1979) and
Dinka had 75% malabsorbers (Bayoumi et al., 1982). Most non-pastoralist, Bantu-
derived agricultural populations have high prevalences of lactose malabsorption at
levels greater than 80% or 90% (Cook and Kajubi, 1966; Jackson and Latham,
1978; Kretchmer et al., 1971), despite the fact that some Bantu populations prac-
tice dairy pastoralism (Segal et al., 1983). Table 2 provides a list of populations and
their prevalences of lactose malabsorption.
It is clear that non-Bantu pastoralists show considerable variation for lactose
intolerance, but most “intolerant” pastoralists appear to be able to tolerate milk in
a variety of forms. It is likely, among non-Bantu pastoralists, that there is a
combination of hereditary ability to digest lactose and an “adaptation” to a life-
long ingestion of lactose (also, a reliance on fermented milk products). Among
Bantu pastoralists, the ability to digest lactose may be largely a function of “ad-
aptation” to milk ingestion. Rosado et al. (1987) have identified a lactose-intol-
erant subpopulation of individuals who absorb lactose efficiently but show the
symptoms of maldigesters. Among Bantu cultivators, the hereditary ability and an
TABLE 2. Prevalences of lactose malabsorption in some pastoral and non-pastoral populations’

Population N % malabsorption Method Reference
Pastoral non-Bantu populations
Beja (Sudan) 303 17 BH,T (1)
Dinka (Sudan) 208 75
.. BH~T iij
Fulani (Nigeria) 33 58 LTT (2)
Maasai (Tanzania) 21 62 LTT (3)
Nuer (Sudan) 23 78 BH,T (1)
Tutsi (Zaire) 27 10 LTT (4)
Pastoral Bantu populations
Xhosa (South Africa) 17 82 BH,T (5)
Zulu (South Africa) 32 81 BH,T (5)
Agricultural Bantu populations
Bantu (Tanzania) 127 92 LTT (6)
Yoruba (Nigeria) 41 98 LTT (2)
Bantu (Ueanda) 12 100 LTT (7)
‘BH,T is the “breath-hydrogen test,” in which 10 g fluid milkkg body wt is given and expired air collected before and
after ingesting milk. Hydrogen gas is released in the large intestine associated with fermentation of undigested lactose.
This is measured by gas chromatography. LTT is the “lactosetolerance test,” where 50 g of lactoseim’ of body surface area
or 2 g k g body wt is administered in water and blood samples are taken before and after ingesting the lactose solution.
Glucose curves are determinedfrom the capillary blood samples (Scrimshaw and Murray, 1988).References: (1) Bayoumi
et al. (19821, (2) Kretchmer et al. (19711, (3) Jackson and Latham (19791, (4) Cox and Elliott (1974),( 5 )Segal et al. (19831,
(6) Jackson and Latham (1978), (7)Cook and Dahlqvist (1968).
“adaptation” to digest lactose seem t o be virtually absent. Further definition of

some of these relationships is needed because of fascinating evolutionary implica-
tions. For example, what is the prevalence of lactose intolerance among the heavily
milk-using Turkana? A curious point concerning the Turkana is that these pasto-
ralists generally do not like donkey’s milk, which has about 50% more lactose than
camel’s milk and 33% more lactose than goat‘s, sheep’s, or cow’s milk (Galvin,
1985; Scrimshaw and Murray, 1988).
Infant, child, and adolescent growth

As has been noted on many occasions “growth is a good, but not very specific,
indicator of health.” Hence, growth processes and health conditions (hygiene, dis-
ease, nutrition) in pastoralists can be described, but correlations are limited in the
literature. Also, many of these pastoralist relationships between health and
growth must be inferred from data on non-pastoralist populations.
As Eveleth and Tanner (1976) point out, “In looking at the growth of African
children, the two most obvious factors that confront us are malnutrition and dis-
ease.” Compared with Europeans, African infants show weight decrements after 6
months, and children show height decrements after 12 months of age (Eveleth and
Tanner, 1976). There is, however, considerable population variation among Afri-
can patterns of child and adolescent growth. For example, growth curves for height
and weight of several Nilotic pastoral populations are given in Figure 5 (Little et
al., 1983). Growth in height represents a pattern of slow but prolonged growth with
only slight adolescent velocity increases (Little and Johnson, 1987). Adult height
is achieved in the early 20s, yet males and females reach values that are equal to
or greater than European, American, or African-American norms. Roberts (1960)
first observed this for Sudanese Nilotic children (Shilluk and Dinka). Roberts and
Bainbridge (1963) also found that Dinka men were among the tallest people in the
world. Hence, although a typical pattern of African growth development occurs,
compared with European patterns, adult Nilotic pastoralists do achieve European
standards. Growth in weight is quite different. With body weight, values lag well
behind Western norms and children are leaner a t all ages up t o and including
adulthood. Children’s and adults’ weights seldom exceed the fifth or tenth percen-
tile of Western norms. Hence, many African pastoralists are tall and very lean as
adults, being among the world’s truly tall populations (Eveleth and Tanner, 1976).
The tall stature of many pastoral peoples is not simply a function of an enriched
HEIGHT
cm
r
160 -
140 -
120-
100 -
80 -
-
1 1 1 1 1 1 1 1 1 1 1 1 I 1 I I I I I I I I 1 1
B 2 4 6 8 10 12 14 16 18 20 22 B 2 4 6 8 10 12 14 16 18 20 22
AGE, y r AGE, y r
Fig. 5. Growth in height and weight of African Americans and several African pastoral populations.
(Data from Hiernaux, 1964; Little et al., 1983; Orr and Gilks, 1931; Heintz, 1963; Verghese et al., 1969.)
diet or favorable environmental conditions. This can be demonstrated by compar-

isons of the growth of well-off Baganda schoolgirls from Uganda with tribal Hutu
and Tutsi girls from Rwanda (see Fig. 6). Baganda schoolgirls are taller and
heavier than both Rwanda groups up to age 14 years, after which the Tutsi con-
tinue to grow in height despite limited dietary intake (Heintz, 1963). Baganda
girls surpassed both Rwanda groups in weight and were close to United States
standards (Hamill et al., 1979). Hiernaux (1964) found a similar pattern for Tutsi
boys.
Growth status has been based on a number of measures including height for age,
weight for age, and weight for height. Weight for height, which is considered a
measure of present nutritional status and degree of wasting, has been assessed for
a number of pastoral populations. Weight for height is taken as a percentage of
standards (50th percentile or median values) established for well-off Western pop-
ulations. Children who fall below 80% of the Western standard (roughly 2 SD
below average) are identified as experiencing moderate to severe wasting, and
hence, malnutrition. Wasting reflects more immediate conditions of poor nutri-
tional status, whereas stunting (short stature for age) is an indicator of long-range
problems with nutritional status or genetic shortness. A note of caution may be
added concerning interpretation of these percentile standards, in that z-scores
(based on standard deviations) give a more accurate representation of growth
status than percentiles since percentile variations and standard deviations change
by age.
Wagenaar-Brouwer (1985) compared Malian (Niger Delta) Tamasheq (Tuareg)
and Fulani children in weight for height. There were no Tamasheq children aged
0-5 years below 80% of the standard, while between 3 and 7% of children and
STATURE
cm
160 -
-
140 -
- c-1------
120 -
-
100 -
-
80 -
- 20
FEMALES
'1 0
1 1 1 1 1 1 1 1 1 1 1
2 4 6 8 10 12 14 16 18 20 22
AGE, yr
Fig. 6. Growth of height and weight in Rwanda Tutsi and Hutu girls and well-off Baganda schoolgirls.
(Data from Burgess and Burgess, 1964; Heintz, 1963.)
adolescents aged 5-17 years were below 80% of the standard. For the Fulani, many
more children fell below 80% of the standard for weightlheight than for the
Tamasheq. In Fulani children aged 0-5 years, up to 16% fell below 80% of the
standard, while in those aged 6-15 years, up to 22% fell below this standard. In
both Tamasheq and Fulani, there were marked seasonal variations where the
proportion of children and adolescents below 80% of standard increased during the
hungry season. Other differences that were found in the Fulani included: 1) a
markedly higher proportion of girls aged 6-15 years than boys who were below the
80% standard, and 2 ) social class variation in weightlheight where the Fulbe
nobles showed poorer weightheight by standard than the lower-class Rimaibe.
Hilderbrand (1985) found that Fulani in a more eastern part of Mali (Seno-Mango
Fulani) had a higher proportion of children aged 6 months to 5 years with poor
nutritional status compared with the Delta Fulani studied by Wagenaar-Brouwer
(1985). Between 16 and 20% of these Seno-Mango Fulani children were below the
80% standard and seasonal variation was present as well (20% below standard
during the period with less food and 16% below standard during the period with
more food). The seasonality of growth was quite striking among the Seno-Mango
Fulani in that children aged 1 to 5 years actually lost body weight during each of
the months of January through March (Hilderbrand, 1985). The author noted that
there are also seasonal variations in disease patterns. Hilderbrand's (1985) study
supports the observations of Wagenaar-Brouwer (1985) to the effect that although
Rimaibe may have lower social status than Fulbe, their nutritional status is prob-
ably better. This is an important observation, since high social status is commonly
linked with superior health conditions.
Benefice et al. (1984) worked with Senegalese Fulani (Ferlo) and found that the
weightheight as a percentage of the Western standard declined with increasing
Fig. 7. Comparison between Senegalese Fulani and Kenya Maasai for percentage of children and adult
women below the 80% standard for weightlheight. (Data from Benefice et al., 1984; Nestel, 1985.)
age of the Fulani children. These results are consistent with the observations on
East African pastoralist children and adolescents that indicate a progressive lin-
earization of physique from infancy to adulthood. Nestel (19851,for example, found
Maasai children t o be disproportionately represented in the group below 80% of
standard for weightheight compared with West African Fulani, although few
Maasai women were in this “moderate wasting” category. It can be assumed that
late adolescent girls begin accumulating body fat and then move out of this “mod-
erate wasting” category. These relationships are illustrated in Figure 7.
Growth in body composition of pastoral children and adolescents is represented
in Figure 8 by derived cross-sectional areas of the arm and leg of Kenyan Turkana.
Compared with United States norms, the Turkana have less adipose tissue (fat)
and less lean tissue (muscle) in both upper and lower limbs. Lower limb fat of
Turkana is considerably less than in the U.S. from early childhood on to adulthood.
Further, muscle from arms and legs and upper limb fat show less growth a t all ages
to produce generally lower values. These characteristics of body composition con-
tribute to the linear physique of the Turkana, which is so typical of Nilotic pasto-
ralists of East Africa. The slow pattern of growth to produce small muscle mass
should not be limited by dietary protein intake by Turkana children and adoles-
cents, since their dietary protein has been shown to be up to four times the FAO/
WHO (1973) requirements (Galvin, 1985).
Adult size and body composition

As noted above, adult pastoralists tend to be tall and lean, where this physique
is characteristic of both West African (Hilderbrand, 1985; Loutan, 1985) and East
African (Little and Johnson, 1986; Orr and Gilks, 1931; Roberts and Bainbridge,
1963) populations. Although there is marked sexual dimorphism in lean tissue in
U S . adults, Turkana lean tissue is almost equivalent in men and women (see Fig.
9). Fat thickness is uniformly smaller in the pastoralist adults compared with the
U S . adults. These values represent limited reserves of adipose tissue and metab-
olizable lean tissue in adult pastoralists, and particularly male pastoralists.
Limited energy reserves increase health risks associated with a t least two
stresses to which pastoralists are exposed. The first is the physiological stress of
pregnancy and lactation in women. The second is the seasonal stress of limited food
resources a t the end of the dry season when livestock productivity is low.
cm
- 80
us. -
MALES - 70
AREA Turkana
cmz
60- - 60
- 50
40 - - 40
30 - - 30
- 20
- 10
0- - 0
-
-
Emerson and others (1972) estimated caloric costs of a full-term pregnancy to be

on the order of 27,000 kcal or an average of 100 kcallday over a 38-week pregnancy.
Others have placed the energy requirement value at three times this amount, or
85,000 kcal (King et al., 1987). This latter value, however, is based on Western
norms, where nearly 50% of energy goes into fat and lean tissue stores. Daily
energy costs for lactation are based on an 80% efficiency of milk production by the
mother, requiring a 600-kcal energy intake to produce 500 kcal of breast milk for
a newborn infant (Thomson et al., 1970). Hence, energy needs by women within the
reproductive years are likely t o be considerably more than what only basal and
activity requirements might suggest. Such constant pressure of energy require-
ments for reproduction combined with limited energy intake in pastoralists prob-
ably contributes t o tissue depletion in pastoral women. This is illustrated in Figure
U
u.s.
5 4 3 2 1 0 1 2 3 4 5 ( . ,
Fig. 9. Cross-sectional areas of muscle and bone (stippled area) and subcutaneous fat tissue (open area)
for the upper arm of Turkana men and women. (Figure from Little and Johnson, 1986.)
10, which shows Turkana age changes in fat composition as estimated by the sum
of six skinfolds. Women peak in fat deposition during the late teens and then lose
these stores throughout their adult years. The effects of these losses of tissue
reserves on women’s health and reproductive capacity are not known.
The seasonal stress of limited food calories in the Third World has become of
increasing interest in recent years (Chambers et al., 1981). Pastoral populations
are particularly susceptible to seasonal food deficits because they exploit environ-
ments with marked seasonality of rainfall and availability of resources. Grazing
lands are characterized by a flush of vegetation during the wet season that must
be rapidly converted into milk and meat by livestock in order for the pastoralists
to exploit this food resource. The pattern is one of seasonal rainfall contributing to
plant productivity which leads to livestock productivity which leads to human
productivity. We speak of “loss of condition” of livestock during the long annual dry
season of the savannas. Although humans are, to some degree, buffered from
seasonal fluctuations, they also suffer from a mild form of “loss of condition” in
weight loss (and gains) during the annual cycle. Figure 11 illustrates the changes
in body weight in two groups of Fulani and in the Turkana. The human weight
changes are slight compared with seasonal livestock changes in body weight which
may be as high as 20% (Wilson et al., 1985). Individual human weight losses in
normal years may be as high as 5 or 6 kg (10 to 12% of body weight), although
average losses (see Fig. 11)for adult members of the population seldom exceed 5 to
7% of body weight. Nevertheless, a loss of 5 to 7% of adult body weight can be quite
significant to health status when tissue reserves are severely limited, as already
described.
Activity and physical fitness
Information on day-to-day physical activity and physical work capacity of Afri-
can pastoralists is limited. Data on these parameters is important for an assess-
ment of the health and adaptability of any population because of the relationship
between activity, nutrition, subsistence, and work productivity. A diagram show-
ing some of these relationships is presented in Figure 12. Aerobic power is a
measure of the ability of working muscles to use oxygen, and is represented by a
maximal value of oxygen consumption, VOz max. Weiner (1980) argued that phys-
I 9 FEMALES
v)
20-, I I I I I I , ,
I I 1
20
B 2 4 6 8 10 12 14 16 18 20 22" 26-30 31-40 41-50 51+
AGE,yr
Fig. 10. Age changes in the sum of six skinfolds (triceps, midaxillary, subscapula, suprailiac, perium-
hilical, medial calf) in Turkana females and males. (Figure from Little et al., 1983.)
65-
NGISONYOKA
TURKANA
m 60 -
y. +-/
I-
I
a
iii 55-
45 -
1 1 1
wm M
1 1 1 1 1 1 1
WH
1 1 1
Litt1eI AFRICAN PASTORALISTS 233
General Nutrition
Protein Nutrition I nfection/Parasites
OPERATION AND FITNESS

Environmental
I AEROBIC POWER
Other
PRODUCTIVITY Factors
Fig. 12. Flow diagram showing relationships associated with physical work capacity (aerobic power)
and productivity. (Figure from Little, 1985.)
iological work capacity, as determined by VO, max, is one of the most useful
measures of overall health. This single measure provides simultaneous assessment
of the health of the cardiovascular, nervous, and musculo-skeletal systems. A state
of physical fitness, which requires training or regular work performance, is one in
which aerobic power or VO, max is high. There is a close positive correlation
between physical fitness and work productivity when the work involves strenuous
labor (Davies et al., 1976; Spurr, 1984). It is also the case that the training or
regular work performance leading to a state of physical fitness has a high cost in
the form of food energy to fuel the work.
Several important questions concerning activity and physical work performance
in African pastoralists have yet to be fully answered. Activity patterns of pastor-
alists are quite different from those of cultivators. Although there is considerable
variation within each of these subsistence modes, pastoralists’ physical activities
tend to be uniform and moderate throughout the year, whereas cultivators are
more likely to engage in intensified periods of activity during planting, harvesting,
and crop processing. There are, however, exceptions to the uniform pastoral model
of activity. Pastoralists move great distances over uneven terrain in search of
green forage for their animals (McCabe, 1983, 1988; McCabe et al., 1988), partic-
ularly during the dry season. Also, activities, such as well-digging and water-
hauling intensify during the dry season (Dyson-Hudson, 1973). What, then, are
some of the relationships between food energy intake and activity that contribute
to body composition, size, and growth processes and other measures of health in
pastoralists?
Inspection of division of labor by sex and age is one way to begin to assess
activity and energy expenditure. Using the Turkana as a n example, boys and
young men tend the livestock, which involves walking long distances and chasing
stray animals, while girls and young women fetch water and firewood, assist in
activities at the settlement, and milk livestock. Young women’s activities may be
quite strenuous when they must carry heavy burdens (e.g., water, firewood, in-
fants) over long distances. As older men become herd owners, marry, and have
children to tend the herds, their physical activity levels decrease. Women, on the
other hand, maintain relatively high levels of activity throughout their adult
years. Based on timed activity schedules, it was found that Turkana men younger
than 30 years of age expended more energy in physical activity than women of
comparable age. Above age 30 years, women, despite their smaller average size,
expended more energy in activity than men (Little et al., 1988a). Conant (1982), in
an excellent review, discussed sexual division of labor in East Africa and noted
that the practice of women doing substantial work well into their middle years is
quite common in agricultural as well as agropastoral societies. Conant (1982)
noted, also, that the subsistence tasks required of most African cultivators or
pastoralists are well within the physical capacity of healthy women. Again, draw-
ing on the Turkana work, the lifting activities of young women probably contrib-
uted to a muscular hypertrophy of the upper limb, such that upper arm muscle
areas differed little by sex and forearm strength was only slightly greater in men
than women (Little and Johnson, 1986; Little et al., 1988b).
There are a number of questions that bear on seasonality in food intake and
activity requirements in pastoralists. Since it is clear that pastoralists undergo
some seasonal hunger and sustain weight losses during these periods, does phys-
ical activity also decrease during these times of hypocaloric stress? Ferro-Luzzi
(1984:95) stated the question most clearly when she indicated that “The major
issue in free-living conditions is not whether energy equilibrium is achieved or not
at a given energy intake level, but the determination of which activities are cur-
tailed in order to re-equilibrate a negative energy balance.” There is good evidence
from Keys et al.’s (1950) classic study during World War I1 that activity dramat-
ically declines during semi-starvation. Careful attempts to test this in malnour-
ished girls from Cali, Colombia, were not successful (Reina and Spurr, 1984);
however, subtle differences in activity are likely t o be difficult to detect. Based on
the use of timed activity schedules, Turkana pastoralists were found to be more
active in the wet season than the dry season (Little et al., 1988a). During the wet
season, when milk production from livestock was relatively high, the increased
physical activity of the people was attributed largely to games, dancing, and other
social forms of energy expenditure. This observation is similar to that made by
R.G. Whitehead, who reported that when lactating Gambian women were given
food supplements, the only difference observed was that “they sang while they
worked” (Beaton, 1984).
Finally, if there is seasonal variation in food availability and in body composi-
tion of pastoralists, does work capacity change seasonally? This is not known, but
the detraining effect of reduced physical labor during the dry season might be
exacerbated by loss of muscle and fat tissue due to the seasonal energy deficit,
which is likely to reduce physiological work capacity. The effect of repeated years
of this annual pattern on body composition and fitness is also unknown a t present.
Reproduction
Pastoralist patterns of reproduction are important to understand because they
are linked closely to environmental fluctuations and to livestock as an essential
resource. From the Turkana herd-owner’s view, daughters are important in that
they assist their mothers in camp activities and as young women bring livestock
into the household unit through brideprice, while sons are valuable in that they
serve as herders, an essential part of the labor force of pastoral subsistence (Dyson-
Hudson, 1973; Gulliver, 1951).This pattern is probably quite common among other
nomadic, as well as semi-nomadic, pastoralists and even more settled pastoral
populations. Hence, there are strong values for both boys and girls, who contribute
equally to herd management and the increase of herd numbers.
Some of the central questions concerning pastoralist reproduction are: 1)What
is the range of variation in fertility in pastoral popluations? 2) What limits fertility
B
NOMADS E TUAREG FULANI TURKANA
1 2 D 1 2 3 1 2 3 1 2
0
'1
N
Fig. 13. Completed fertility in a number of pastoral populations. Populations are: Sudan nomads,
including Baggara (1)and Kawahla (2) (Henin, 1968,1969); Bedouin (Muhsam, 1950); Niger Tuareg (1)
(Ganon, 1975); Mali Tuareg of Niger delta (2) and Gourma area (3) (Hill, 1985b): Niger Fulani (1)
(Ganon, 1975); Mali Fulani of Niger delta (2) and Seno-Mango area (3) (Hill, 1985b); Kenya Ngikitak
Turkana (1)(Brainard, 1981) and Ngisonyoka Turkana (2) (Leslie et al., 1988).
in pastoralists? 3) What are the causes of seasonal variation in fertility? 4) What

are the health effects of arid lands residence on reproduction? 5) Among nomadic
pastoralists, what are the effects of settlement or a sedentary life style on repro-
duction? Current investigations have only begun to address some of these ques-
tions.
Variation in completed fertility for some pastoral and non-pastoral African pop-
ulations is presented in Figure 13. Completed fertility refers to the number of live
births that women have had upon completion of their childbearing years (in post-
menopausal women). This single measure of fertility refers to events (births) that
took place up to three decades ago; hence, it does not reflect current fertility rates.
Total fertility rate takes into account the fertility of women at all ages and is a more
accurate measure of current reproductive performance. The values in Figure 13
indicate the wide range of variation in completed fertility, indicating, as Leslie et
al. (1988) suggest, that fertility of pastoral populations is certainly not uniform. In
general, pastoral populations have shown lower fertility than permanent agricul-
tural populations (Hill, 1985b).The exceptions are Malian Fulani (Hill, 1985b) and
Kenyan Turkana (Brainard, 1981; Leslie et al., 1988; see Fig. 13).
Hill (1985b) attributed much of the variation that he observed among Bambara
cultivators and Tuareg (Tamasheq) and Fulani pastoralists to birth interval dif-
ferences due to breastfeeding and consequent lactational amenorrhea. Brainard
(1981, 1986) suggested that Turkana nomads had high fertility because they had
relatively high infant mortality. These two explanations have a common basis in
their effects from breastfeeding practices. For example, an infant who dies con-
tributes to the cessation of breastfeeding by the mother, which, in turn, soon
returns the woman to a normal ovulatory pattern. This, then, increases the prob-
ability of conception, shortening the birth interval, and increasing the overall
fertility of the woman (Leslie et al., 1988). It may be the case among pastoralists,
also, that early termination of breastfeeding occurs because livestock milk is avail-
able as an infant food supplement (P. Leslie, personal communication). Based on
the Fulani (Hill, 1985b) and Turkana (Leslie et al., 1988) fertility data, it is ap-
parent that some pastoral populations are capable of maintaining relatively high
fertility. These values are in marked contrast to other completed fertility data on
huntedgatherer populations where fertility is quite low. For example, Aka Pyg-
mies of the Central African Republic had completed fertility of 5.0 live births per
woman beyond reproductive years (Cavalli-Sforza, 1986), while Dobe !Kung Bush-
men of Botswana were slightly lower at 4.7 completed fertility (Howell, 1979:123),
and Efe Pygmies of Zaire were extremely low with a value of 2.6 for completed
fertility (Bailey and Peacock, 1988). Sterility produced by gonorrhea was sug-
gested to explain some of these low fertility values (Bailey and Peacock, 1988;
Howell, 1979). The Ovaherero, who are Bantu-speaking cattle and goat pastoral-
ists of eastern Namibia and western Botswana, also have very low completed
fertility (about 3 live births/woman), but it is seen in an older cohort of women
(Harpending and Pennington, 1989). A higher completed fertility (about 6 live
birthdwoman) was found in a current cohort of Ovaherero women, indicating that
rapid generational changes can occur in non-Western peoples who maintain tra-
ditional life-ways.
In addition to the effects of savanna seasonal rainfall and resource availability
on nutrition and body weight, seasonality also has an influence on births, and,
hence, on human reproduction. Two pastoral populations have been identified
with marked seasonality of births to date: Malian Fulani (van den Eerenbeemt,
1985) and Ngisonyoka Turkana of Kenya (Leslie and Fry, 1989). The magnitude
of seasonality in the Turkana is greater than that recorded for any other
population, and is several times greater than in the Fulani. Seasonality in births
is not unusual and has been recorded for other populations such as the Kalahari
San Bushmen (Wilmsen, 1978). Explanations have ranged from social mecha-
nisms for Fulani (van den Eerenbeemt, 1985), to abstinence for Herero (Wilmsen,
1982), to dietary intake for San Bushmen (Wilmsen, 1978), to seasonal labor
migration in rural Lesotho (Huss-Ashmore, 1988), and to frequency of coitus in
rural Bangladesh (John et al., 1987). Leslie and Fry (1989) caution against
obvious explanations based on nutritional and body composition variation,
although they acknowledge that there is considerable indirect evidence for such a
relationship.
In a pastoral environment with limited food energy and high physical activity
demands on women, there is often a negative energy balance in women (Galvin,
1985; Nestel, 1985). Since weight loss and physical activity have been implicated
in menstrual and ovarian function changes in girls and women (Beaumont et al.,
1976; Ellison and Lager, 1986; Frisch, 1978; Frisch et al., 1981), there may be
seasonal or longer-range effects on reproduction in pastoralists from this complex
set of relationships. Another process that can further obscure our understanding of
these mechanisms is that of metabolic adaptation. One form of metabolic adapta-
tion involves the lowering of resting metabolic rate (RMR) after weight loss, but
with a lowering of RMR that is beyond what would be expected from lean tissue
loss alone. This has been demonstrated for pregnant and lactating women in Gam-
bia (Prentice, 19841, New Guinea (Durnin, 1980), and Taiwan (Adair, 1984). Mar-
ginally malnourished women from these three populations maintained energy bal-
ance over time despite the increased needs of pregnancy and lactation. Another
form of metabolic adaptation is not well documented, but is intriguing in its pos-
sibilities for pastoral peoples. The pattern is one of periodic weight loss and gain
that leads to increasing “energetic efficiency” and reduction in RMR. It has been
demonstrated in laboratory animals (Bjorntorp and Yang, 1982), wrestlers (Steen
et al., 19871, and a single woman dieter (Ghali and Durnin, 1977), but not in
Western women who are frequent (yo-yo) dieters (van Dale and Saris, 1989). The
implications for pastoral women are a function of their seasonal weight losses; the
potential combined effects of seasonal changes in dietary intake, activity patterns
and metabolic adaptation; and the net effect on reproductive function.
Disease
A comprehensive discussion of disease and morbidity in African pastoral popu-
lations is beyond the scope of this paper. Rather, what will be focused on is mor-
bidity and mortality that is more closely associated with pastoral environments
and modes of life.
The most comprehensive work on morbidity and mortality of any pastoral pop-
ulations was compiled by Hill (1985a) and concerned studies of Malian Fulani and
Tuareg (Tamasheq). The research design of this integrated study involved a num-
ber of populations at several sites in Mali (Niger River delta, Seno-Mango,
Gourma), and also separated Fulani (Fulbe and Rimaibe) and Tamasheq (Nobles
and Bella) into their constituent ethnic subdivisions. What is most apparent from
these studies is the remarkable degree of variation among the populations and
ethnic subdivisions.
One of the most common diseases is malaria, in which up to 35% of all groups
showed symptoms at specific times of the year. Chabasse and colleagues (1985)
found that malaria blood titres of Gourma Tamasheq were nearly twice as great as
those of Gourma Fulani. Hilderbrand (1985)compared malaria prevalence in Fulbe
and Rimaibe Fulani from the Seno-Mango area and hypothesized that the lower
prevalence of the disease in Rimaibe was a function of their ethnicity and probable
higher frequency of hemoglobin S (sickle-cell).Among Fulani infants and children,
the most frequent disorders (in addition to malaria) were diarrhea and conjunc-
tivitis. In any single month, about 20% of children and infants had symptoms of
diarrhea, whereas conjunctivitis was present in 40% of the same age group (Hilder-
brand, 1985). Chabasse et al. (1985) confirm these values for conjunctivitis but also
indicate that trachoma is very rare. This suggests that the presence of conjunctivitis
is a function of aridity, wind-blown dust and sand, and poor personal hygiene rather
than the outcome of other eye disorders. They also found 15% cataracts in Gourma
pastoralists of all ages, possibly resulting from trauma or chronic conjunctivitis.
Other diseases of adults included Guinea worm (local prevalence) and trepanomal
infections (about 50% prevalence in Gourma pastoralists), and Brucellosis (25%
prevalence in Gourma populations), which is a livestock disease and a common
hazard among pastoralists (Chabasse et al., 1985; Hilderbrand, 1985).
Infant (birth to 1 year) and child (birth to 5 years) mortality was very high in all
pastoral groups, but with some variation. Hill (198513) found infant rates at about
25% and child rates at 33%, with Delta Fulani the highest. In this Fulani popu-
lation, there was 51% mortality in children born alive of mothers aged 45-49
years. This was more than 10% greater mortality than that of Seno-Mango Fulani
or Tamasheq. On the other hand, Gourma Fulani had much lower infant and child
mortality than did. Gourma Tamasheq (Chabasse et al., 1985).
Limited information on other pastoral populations is consistent with the picture
presented for Malian pastoralists. Jelliffe and co-workers (1964) surveyed a large
sample of Karimojong children from Uganda and found similar prevalence of ma-
laria and eye disorders but little or no cases of protein-energy malnutrition or
wasting. This was also the case in the neighboring Kenyan Turkana in which
children were characterized as relatively healthy (Little et al., 198813; Shelley,
1985). About a third of the Turkana children had hepatomegaly, which is indica-
tive of malaria or other parasites, and this prevalence was confirmed by interviews
(Shelley, 1985). Stool analysis in Ngisonyoka Turkana gave very low parasite
counts, which is, again, consistent with data from the Malian project, and also
suggests that the hepatomegaly resulted from malaria infection (Little et al.,
1988b).
Measures of cardiovascular health in adults and the aged have repeatedly dem-
onstrated the absence of atherosclerosis, hypertension, and other indicators of
cardiovascular disease despite the relatively high fat diets of most pastoralists
238 YEARBOOK O F PHYSICAL ANTHROPOLOGY [Vol. 32, 1989
such as the Maasai, Samburu, and Turkana (Biss et al., 1971; Mann et al., 1964,
1972; Mugambi and Little, 1983; Shaper et al., 1961, 1963, 1969).
It would be inappropriate to close any discussion on disease in pastoralists, albeit
brief, without some discussion of trypanosomiasis. Desowitz (1981) stated that 15
million km2 of African savannas is unusable by pastoralists because of the tsetse
fly (Glossina ssp.) and livestock trypanosomiasis (Trypanosoma ssp.) infestation.
Although the effects of sleeping sickness on human populations can be disastrous,
the effects on livestock, and indirectly on human food supply, are even greater.
Foulkes (1981) estimated that the cattle population in Africa could double if the
tsetse fly were eradicated, but noted that several years ago, it would cost between
$300 and $500 to eradicate the tsetse fly from only 1km2 of savanna land!
The West African or Gambian trypanosomiasis is transmitted by Glossina pal-
palis, which inhabits the humid woodland savanna, while the East African or
Rhodesian variety is transmitted by Glossina morsitans, which inhabits dry sa-
vanna woodland (Desowitz, 1980). Pastoralists must balance the need for pasture
vs. the probable deaths to livestock when making decisions on whether or not to
invade the tsetse fly environment. Since the tsetse fly has a preference for live-
stock, the risk to humans of being bitten and contracting trypanosomiasis is re-
duced (Ford, 1971). However, the treatment for humans who have sleeping sick-
ness is still arsenic, which is highly toxic to humans (Foulkes, 1981). Fortunately,
the number of human cases of sleeping sickness is very small compared to other
endemic diseases of the African savanna (Foulkes, 19811, so the principal concern
of the pastoralist is to avoid starvation in his animals, while at the same time,
avoiding contacts with Glossina.
CURRENT ISSUES IN RESEARCH
There are many research issues concerning the human biology and biobehavior
of pastoralists that should be pursued. Brief discussions of three major topics fol-
low.
Seasonal hunger and its health effects
We have only become aware of the dimensions of seasonal hunger within the
past few years (Huss-Ashmore et al., 1988). Indications are that it is widespread
and regular among pastoralists. Future research designs must take into account
not only human variation, in general, but seasonal variation in nutrition, disease,
growth, reproduction, and overall health status, as well. In addition to the prac-
tical need for information in this area, there are also some very fundamental
scientific problems in human biology that can be explored within the framework of
seasonal stress and variation.
Seasonal stress usually has rainfall as its “driving variable,” where rainfall
regulates the agricultural cycle or the cycle of green forage availability for livestock
(Little et al., 1984). There are several effects of these seasonal rainfall patterns: 1)
food becomes more available sometime after the rains, 2) physical activity or work
levels usually increase during the wet season, and 3) infection rates also increase
at the time of the rains (e.g., malaria, gastrointestinal infections). Among pasto-
ralists, two broad questions might be asked: What happens during the period of
seasonal stress when nutritional deficiency, high work loads, and infection simul-
taneously strike members of the population? What happens with repeated cases of
seasonal stress during a lifetime of exposure to these conditions? The first question
bears on health and maintenance of the individual, family, and population; the
second question deals with longer-range problems and adaptation to periodic stress.
Studies of diet and body composition have shown that food intake is reduced
during late dry and very early rainy seasons in pastoralists and that this results in
body weight losses that probably reduce working capacity (Galvin, 1985; Hilder-
brand, 1985; Loutan and Lamotte, 1984; Nestel, 1985). Evidence is beginning to
accumulate, also, that children’s growth patterns are disrupted by seasonal hunger
(Galvin and Little, nd; Pagezy and Hauspie, 1985; Thomson, 1970), although per-
Little1 AFRICAN PASTORALISTS 239
manent effects are not completely known, particularly in pastoralists. One of the
most intriguing questions concerns the forms of adaptation to limited food supply
(Grande, 1964). We know that individuals reduce their activity (when possible) to
compensate for limited food intake, and that a reduction in basal energy require-
ments occurs with loss of lean body mass (James and Shetty, 1982). However, other
patterns of metabolic adaptation are less clear, such as during pregnancy and
lactation (Prentice, 19841, and during repeated exposures to stress, as in season-
ality. This is truly an exciting area for investigation.
Pastoralist responses to drought

Some pastoral populations seem to be able to combat drought conditions very
well, while others do not. Pastoralist behavioral adaptations to drought are mo-
bility and the maintenance of sizable livestock holdings. The principal require-
ment for these behavioral adaptations is a vast territory for movements of their
substantial numbers of animals. When drought strikes, herdsmen can move their
livestock to rangelands with remaining forage, and then sustain considerable an-
imal losses but retain enough livestock, to rebuild the herds. Ellis and colleagues
(1988) conducted studies of Turkana pastoralists and agropastoralists from four
regions in Turkana District, Kenya. They focused on the drought periods of 1979-
80 and 1984. The Ngisonyoka and neighboring Ngikamatak Turkana (subtribal
sections) fared quite well during these drought periods, whereas Ngiboceros Tur-
kana and both pastoralists and agropastoralists from the Tarach River area in the
north were hit very hard by the droughts. Computer simulations were carried out
on these Turkana populations and their subregional ecosystems, and predictions
were made on survivability of each population under a range of drought conditions.
The factors that were related to poor responses during drought and famine in this
relatively arid ecosystem were: sedentarization, limited access to dry-season graz-
ing areas, high population density, and a dependence on agriculture (Ellis et al.,
1988).
Drought may be viewed as an extension of seasonal aridity, such that rains will
fail from time to time and the customary wet period will be skipped or be inade-
quate. When rains fail, the failure may not be apparent for several months, and
full drought conditions will be identified only when rains fail completely. In agri-
cultural societies, drought effects are quite distinct when crops fail. However, in
pastoral societies, recovery may come a t any time if rains occur and vegetation
begins to flush. Therefore, to pastoralists, drought might be considered an indis-
tinct phenomenon. What, then, are the early warning indicators of pastoral pop-
ulations beginning to experience the stress of food deprivation from impending
drought? Galvin (1988) discussed drought in Turkana pastoralists within the con-
text of early warning prediction of dietary stress before drought becomes full-
blown and starvation is endemic. She identified two major ways to anticipate and
monitor nutritional stress due to drought: anthropometric assessment of body com-
position, and monitoring diet composition. Both have shortcomings. Human body
composition responds very slowly to food shortages, and diet composition monitor-
ing is a labor-intensive and expensive activity. Galvin (1988) suggested that both
be used, along with other measures of declining health and resources, including
personal interviews.
African pastoral populations experience seasonal drought on an annual basis,
and longer-period drought in a pattern that may range from once in 5 years to once
in 10 years (Little and Johnson, 1985). Hence, drought of differing periodicities is
a part of the way of life of pastoralists. Investigation of both behavioral and bio-
logical patterns of response to these periodic events should be encouraged.
Sedentarization in pastoralists
Nomadic pastoralists are being settled in large numbers on agricultural and
ranching schemes, or they are leaving the pastoral sector t o migrate to towns and
cities. What influences do these new conditions have on the health and biology of
the pastoralist?
At this time, we really have a very incomplete picture of the effects of sedenta-
rization on the health of pastoralists. The transformation of pastoral nomads or
semi-nomads to settled farmers or agropastoralists carries with it a number of
changes in diet, work patterns, cultural traditions, disease patterns, and the like.
These changes are by no means uniform, and depend on the context of the change.
Certain conditions under the new sedentary existence may be an improvement
over the nomadic way of life (e.g., hygiene, health care, schooling).Other sedentary
conditions may not be improvements and contribute to deterioration of health
conditions (e.g., less balanced diet, less physical activity, loss of social integration).
Any changes that are likely to take place in general health and biological status of
the people are: 1) related to how rapid the transformation was; 2) dependent on the
amount of time in the new sedentary life; and 3) a function of the ages of the people,
since the elderly are less adaptable to change and the very young may adapt quite
well (Little and Baker, 1988). An assumption that might be used in the case of all
changes in subsistence and lifestyle is that the new style is likely to impose
stresses on the population that lead to health problems. This is probably not al-
ways the case, but the assumption is based on the premise that populations (cul-
tures) which have maintained a subsistence pattern for a reasonably long period of
time have established biobehavioral and cultural adaptations within that environ-
mental context (Baker, 1977). Any changes, then, are likely to be disruptive.
No studies of health and adaptability of a population have been conducted be-
fore, during, and after a permanent settlement. This research design would involve
substantial commitment of time and resources. There are a number of studies that
have compared closely related populations, one of which is settled and the other
nomadic. Sandra Gray (1988) analyzed growth of Turkana nursery and primary
schoolchildren (aged 4-9 years) settled in the town of Katilu, Kenya, and com-
pared them with nomadic Turkana children. Nomadic children had diets much
higher in protein than settled schoolchildren, but the settled children were taller
and heavier than the nomads. These schoolchildren were provided breakfast and
lunch a t school, so their caloric intake may have been greater than the nomadic
children?or their levels of activity may have been less. We have been no informa-
tion on adolescents, but Kathleen Galvin (Ellis et al., 1988) compared settled
Katilu Turkana with nomadic Turkana adults. She found that body weight and
both cross-sectional areas of arm muscle and fat were less in the settled Katilu
adults than in the nomadic Turkana adults, suggesting poorer health and nutri-
tional status in the unsettled Turkana farmers. The settled Turkana were drawn
from the category of “failed” pastoralists who may have undergone more nutri-
tional stress during their pastoral lives than the “successful” pastoralists who
continued the pastoral pattern.
The picture is even less clear when mortality is considered. Brainard (1981)
studied fertility and mortality a t the Nakwamoru irrigation scheme close to Katilu
and found that Turkana who had previously been nomads had higher infant mor-
tality than a population of Ngikitak Turkana who had been settled for several
generations. This suggests that settled infants suffered less morbidity and mor-
tality than nomadic infants, perhaps because the settled farmers had greater ac-
cess to a nearby clinic. At this time the effects of settlement on Turkana growth,
adult size, morbidity, and mortality are equivocal, and more research is required.
A final note considers the influence of settlement on population fertility. Earlier
work, which compared settled farmers who had been pastoralists with currently
nomadic pastoralists, found that settled farmers had higher fertility (Henin, 1968,
1969; Muhsam, 1950). Brainard‘s (1981) results of fertility studies of settled and
nomadic Turkana were counter to earlier research in that she demonstrated that
nomadic women had higher fertility. Leslie and colleagues (1988), who confirmed
Brainard’s work in studies of nomads, argued that there is much less homogeneity
in fertility in pastoralists than is apparent from the literature.
This heterogeneity of fertility in pastoralists is only part of the remarkable

amount of variation found in diet, disease, growth, activity, and other realms of
health. Most of the research conducted to date has only covered the surface details
of what will prove to be a rich field of investigation.
CONCLUSIONS: THE NEED FOR INTEGRATED RESEARCH
African pastoralism is a mode of subsistence that, when compared with agricul-
ture, is of low productivity and expensive in terms of land resources needed. At the
same time, pastoralism is a very effective way to exploit the arid and semi-arid
lands that constitute such a large fraction of the African continent. Despite at-
tempts by governments t o settle, modernize, or otherwise convert pastoralists to
cultivators, some of these populations continue to exploit successfully vast land
areas and to resist conversion to a sedentary existence. Whether pastoralists can
continue to avoid indefinitely the transformation of their lifestyles is not clear.
Group ranches and restricted grazing have already brought about changes that
limit nomadism and further reduce those populations which maintain a traditional
form of pastoralism. There is, then, some urgency to study these populations before
they disappear or are dramatically transformed. Another reason to conduct re-
search on traditional pastoralists is that we need basic information in order to
judge the effects of settlement on the health and adaptability of the people. This is
especially the case when settlement is planned with the objective of improving
people’s lives.
How best can we study African pastoral populations either to understand their
successful behavioral and biological ways of coping with a harsh environment or to
provide the information needed to plan for almost inevitable change? A means that
seems most productive is a multidisciplinary and systems approach that enables
scientists from a number of fields to collaborate on one or more populations. This
approach avoids focussing on simple deterministic models of biology or behavior
and encourages integrative kinds of explanation, encourages synthesis, and stands
a better chance of providing the kinds of analytic and predictive models that will
help us to better understand these people.
ACKNOWLEDGMENTS
Thanks are due to colleagues Ann B. Stahl, who read and offered detailed sug-
gestions on one section, and longtime friend Neville Dyson-Hudson, who helped to
sort out the distribution of pastoral populations in Africa. I appreciate also the firm
but diplomatic patience of Emoke J.E. Szathmary.
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YEARBOOK OF PHYSICAL ANTHROPOLOGY 32:215-247 (1989)

Human Biology of African Pastoralists

KEY WORDS Pastoral nomads, Environmental adaptation, Biobehavior,

ABSTRACT Pastoralism in Africa is a subsistence tradition that has a

AFRICAN PASTORALISTS AND THEIR ENVIRONMENT

tant to trypanosomiasis than are domestic species, a circumstance that limits

of language and livestock practices. A variety of sources of information can be used

Fig. 2. Distribution of some pastoral populations in Africa.

standing of how conditions of behavior, within environmental contexts, influence

environmental cycles of resource availability, such buffering is always imperfect,

TABLE 1.Measures of health and adaptability

HEALTH AND ADAPTABILITY ASSESSMENT

MEASURES OF HEALTH AND ADAPTABILITY

Seasonal food availability exerts a considerable influence on the diets of most

TABLE 2. Prevalences of lactose malabsorption in some pastoral and non-pastoral populations’

“adaptation” to digest lactose seem t o be virtually absent. Further definition of

Infant, child, and adolescent growth

diet or favorable environmental conditions. This can be demonstrated by compar-

Adult size and body composition

Emerson and others (1972) estimated caloric costs of a full-term pregnancy to be

Protein Nutrition I nfection/Parasites

OPERATION AND FITNESS

in pastoralists? 3) What are the causes of seasonal variation in fertility? 4) What

Pastoralist responses to drought

This heterogeneity of fertility in pastoralists is only part of the remarkable

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