Professional Documents
Culture Documents
Introduction
Termites are social insects of the order Isoptera which enjoy a nearly worldwide
distribution. Considered pests in North America, their colonies are numerous especially
in the plains and western states. Their existence is of special relevance to archaeologists
working in the tropics and subtropics, where their mounds comprise such conspicuous
features of the landscape (See Figure 1).
Drummond (1888) remarks on the extent of termite activity in the regions of East Africa
bordering Lake Victoria, whose outlet at Jinja is the source of the White Nile:
Herodotus, with equal poetic and scientific truth, describes Egypt as “the gift of the Nile”.
Possibly had he lived today he might have carried his vision farther back still, and referred some
of it to the labours of the humble termites in the forest slopes about Victoria Nyanza
(Drummond, 1888: 158).
“Departmentof Anthropology, BrandeisUniversity,Waltham, MA 02254,U.S.A.
111
0305-4403/90/020111+33 SO3.00/0 Q 1990AcademicPressLimited
112 S. McBREARTY
This paper investigates the proposition that termites are important in altering
sedimentary profiles in tropical Africa. Where are termites found? Exactly what effect do
they have on geological deposits? Are they numerous enough to have significant impact at
archaeological and paleontological sites? Termites are considered here as one among
many post-depositional factors affecting tropical terrestrial deposits, and specific case
studies that document termite activities in archaeological contexts are presented.
Archaeological Importance
The role of termites in African archaeological contexts has occasionally been noted.
Fossilized termitaria have been recognized in an early Pleistocene occupation at Koobi
Fora in northern Kenya (Isaac, pers. comm.), in Pliocene deposits at Laetoli, northern
Tanzania (Sands, 1987) and in Plio-Pleistocene sediments in the Middle Awash region of
Ethiopia (Clark & Harris, 1985).
Cahen (1976, 1978a) and Moeyersons (1978) observe the disturbing effects of termite
burrowing in Pleistocene deposits at Gombe Point, Zaire. Similar conditions have been
noted for the site of Dinga, Kwango, Zaire (Cahen & Moeyersons, 1977) and at the
Acheulian site of Kamoa, Shaba, Zaire (Cahen, 1975). Convincing evidence of termite
disturbance at the Lupemban site of Masango, Burundi, has been presented by Van Noten
et al. (1972). Sampson (1972) suggests that the stratigraphy of the Fauresmith occurrence
at the South African site of Muirton on the Vaal River may have been affected by termites.
Crossley (1986) suggests that termites are most likely to have been responsible for the
accumulation of the upper member of the Chitimwe Formation, which overlies Middle
Stone Age artifacts at and near the archaeological site of Chaminade in northeastern
Malawi. At Muguruk, western Kenya, I have attributed some of the observed features of
the Middle Stone Age archaeological profile to termite activities (McBrearty, 1986,1988).
Watson (1967) describes the effects of termites on Iron Age interments.
TERMITES AS POST-DEPOSITIONAL AGENTS 113
(0) (b)
Amazonia (Dufour, 1987). They are particularly rich in protein and lipids which may be
lacking in a diet high in carbohydrates (Hladik, 1977); 100 g of fried termites have a
nutritive value of 561 kilocalories (Bodenheimer, 1951). African termite alates swarm at
the end of the dry season, and are thus available at a critical time when cultivated food
supplies may be running low (Thon, 1946; De Schlippe, 1956; Mielke, 1978; Freedman,
1983; and pers. obs.).
To human observers, the most striking behaviour of termites is mound building,
primarily because it takes place above ground. The mound building Macroterminae
are the most abundant group in East African savannas (e.g. Macrotermes and
Trinervirermes); other species are primarily subterranean and do not construct mounds
(e.g. Cubitermes and Odontotermes, but see Darlington, 1985). The metabolic activity of
the colony’s cultivated fungi, plus the respiration of the insects themselves, generates
considerable heat (Sands, 1987), and mounds and the air shafts they contain act as a
ventilation system which regulates the internal temperature of the colony (See Figure 2).
Mounds also improve drainage and may function secondarily in defense (Ruelle, 1964;
Wood & Sands, 1978; Collins, 1979; Darlington, 1985). In sub-Saharan Africa, large
uninhabited termite mounds are occasionally put to economic use. They may be hollowed
out and transformed into cooking ovens (Koeune, 1977) or iron smelting furnaces
(Killick, pers. comm.).
The persistence of termite mounds varies among species and with local conditions.
Mean lifespan for mounds of Trinervifermes in Upper Volta is estimated at 7 years (Roose,
1976). For Macrotermes bellicosus and Pseudacanthotermes mounds, Pomeroy (1976b)
estimates lifespans of 10 and 5 years, respectively, although colonies of other species of
Macrotermes may remain active for 80 years or more (Grasse, 1949), and colonies of
Nasutitermes may live for 50 years (Lee & Wood, 1971a). On the basis of radiocarbon
dates and the relative states of bone weathering, Watson (1967) estimates a termite mound
in northeastern Zimbabwe occupied by both Macrotermes and Odontotermes to be as
much as 700 years old, and Darlington (1985) points out that repeated recolonization of
the same site may allow some mounds to survive for centuries.
Densities of termite mounds vary with habitat (Goodland, 1965; Baroni-Urbani et al.,
1978; Pomeroy, 1978, 1983~; Golley, 1983; Josens, 1983; Salick et al., 1983) and their
116 S. McBREARTY
distribution is in part dependent upon availability of water. In an early study Kellogg &
Daval(1949) estimated that in parts of Zaire termitaria cover 30% of the ground. Sands
(1965) has computed an average density of 14.8 occupied mounds per ha for Mucrotermes
bellicosus in undisturbed woodland savanna in northern Nigeria, where the smaller
mounds of Trinervitermes reach densities of 108.6 ha-’ in the same study area.
In savanna vegetation on laterite, occupied Trinervitermes mounds may be an order of
magnitude less frequent, only 12.4 ha-’ having been observed by Bodot (1967) in Ivory
Coast. Pomeroy (1977) reports an average density for Ugandan Macrotermes mounds of
lo-40 ha-‘. Mound densities of two species of Mucrotermes in semiarid Kajiado District,
southern Kenya, range up to 9 ha-’ (Pomeroy, 1983~). In the more arid savanna of Tsavo
East National Park, southern Kenya, Mucrotermes mounds may number fewer than one
per ha, but a five- to six-fold increase in the density of mounds in 1982 over that of 1976 in
Tsavo is related by Pomeroy (1983~) to the end of drought conditions. Sands (1987)
tentatively estimates that numbers of active Macrotermes colonies ranged between one
and 3.93 ha-’ under the arid conditions prevailing at Laetoli during the Pliocene.
Termite numbers can be appreciated by contemplating the fact that each active colony
contains at least one queen, and that mature queens of some species can produce 20,000 to
30,000 eggs per day (Ruelle, 1985). Estimates of the densities of individual termites range
up to 4000 rnp2 (Wood, 1975; Wood 8~Johnson, 1978; Wood & Sands, 1978; Wood et al.,
1982), but the fumigation technique of Darlington (1979) has increased population
estimates for termite colonies by several orders of magnitude over previous live digging
methods. She has recovered over 2 million individuals from a single nest of Mucrotermes
subhyalinus less than 10 m3 in area near Kajiado, southern Kenya. Sands (1987) estimates
that a mature Mucrotermes colony may contain up to 5 million termites. Using Pomeroy’s
estimate of 9 mounds ha-’ for the Kajiado region, and Darlington’s more conservative
estimate of 2 million individuals per nest, we may assume that this semiarid habitat
supports as many as 27,000,OOOtermites ha-‘; greater numbers would be expected in more
humid regions.
The sediment used for the construction of mounds, sheets, and runways may be brought
up from considerable depth in the soil. Estimates range up to 2 or 3 m, but Pomeroy
(1976~) estimates that Ugandan Macrotermes bellicosus and Pseudacanthotermes sp.
construct their mounds primarily of topsoil from depths of 0.5 to 1.Om below the surface,
and this is probably the more common range. Termites may delve deeply in search of
water, and they have been recorded at depths of 23 m (Watson, 1972), 30 m (Marais,
1937), and even 70m (Noirot, 1970; Lee & Wood, 1971~; Wood & Sands, 1978; Josens,
1983). Their effective downward mobility is limited by the nature of substrate and the
location of the water table.
Not only do termites excavate to great depths, but also their galleries may be large, and
the total network dense and extensive (See Figure 3). Macrotermes subhyalinus creates a
network of several kilometres of branching tunnels within its 1 ha foraging area (Sands,
1987). Darlington (1979) describes subsurface foraging passages for the related species M.
michaefseni as large as 7 cm in diameter which radiate at depths of 50-60 cm from the
118 S. McBREARTY
surface to distances up to 10 m from the nest. At this distance they branch at l&25 cm
below the surface to form a dense network of smaller underground passages about 34 cm
in diameter [See Figure 4(a)]. These may be interspersed with elliptical storage pits
(Darlington, 1982). Small intersecting passages may terminate at distances up to 50 m
from the nest [Darlington, 1982; see figure 4(b)].
Obviously, stone artifacts or other objects may easily migrate downward through such
cavities. This situation is clearly described for the site of Masango, Burundi, by Van Noten
et al. (1972).
Abandoned termite galleries may become infilled with surface soil washed in from
above, and Watson (1960) has observed the resulting tube-shaped features filled with light
coloured, fine grained, less compact infill at depths of up to 8.5 m in West African soils.
Robinson (1958) documents a zone of termite galleries filled in with dark coloured,
nutrient rich, less compact soil at depths of 60-120 cm in a coffee plantation in Kenya.
Ratcliffe el al. (1952) note that infilled termite galleries penetrate to substantial depths,
often into limestone bedrock, and that plant roots preferentially follow the course of the
less densely filled galleries. In older deposits passages may persist as mineralized casts.
Vertical cylindrical structures cemented by calcite within the tuffaceous sediments at Lot.
9 at Laetoli, Tanzania, may be the fossilized remains of Pliocene termite water collecting
routes (Sands, 1987).
Mound
Inl
5-10 m
A
30-40 m
I
50 m
I
Y” Abandoned neat
(b)
I I I
D
0 IO 20 \
m
Figure 4. (a) Schematic cross-section of nest and foraging passage system of Macro-
termes michaelseni (not to scale; after Darlington, 1982: 238, figure 1); (b) Map
of underground passages excavated by a single colony of M. michuelseni (after
Darlington, 1982: 239, figure 2).
120 S. McBREARTY
Angular grOVeI,
uron plsolitha,
ESA ond MSA artifacts
LSA artifacts
Gravel, pisoliths,
Modern fermitario
0 800 1600
m
Figure 5. (a) Transverse section of the terrace of the Middle Kwilu near Thysville,
Zaire, showing stone line at the base of termite-altered yellow clayey sand (after de
Ploey, 1964: 405, figure 4); (b) transverse section of Chaminade, northern Malawi,
showing the stone line underlying termite altered red clayey sand, and their relation
to modern and fossil termitaria (after Crossley, 1986: 193, figure 2).
particles to the surface, gravel sized objects, which may include stone artifacts, accumulate
as a lag deposit at the effective base of termite activity (See Figures 5 and 6). Burrowing
rodents produce similar effects at temperate latitudes (Johnson, 1989). Termite-produced
stone lines usually have sharp upper and lower boundaries and often correspond to the
position of the water table. These mechanisms for burial of coarse debris have been
proposed for the quartz banded soils of East Africa (Ollier, 1959), for the stone horizons
near Umtali, Zimbabwe (Watson, 1962), and for Sudanese gravel and loam deposits
(Folster, 1964). Stone artifacts may become concentrated at a consistent depth in this way,
giving a false impression of an ancient land surface or living floor.
Extensive termite activity may thus produce a widespread uniform mantle of topsoil
with characteristics similar to those of the soil in a termite mound. De Ploey (1964) notes
that particle size distributions of sediments thought to be reworked by termites resemble
those of samples from nearby mounds of Macrotermes, and Pomeroy (1976~) reports that
his Ugandan termite species produces a stone free topsoil with properties closer to loam
than the average subsoil. Crossley (1986) estimates that the upper member of the
Chitimwe Formation in northeastern Malawi represents such a termite produced soil
TERMITES AS POST-DEPOSITIONAL AGENTS 121
0
2
(01 E
4
6
IO 20 30
IO u
mantle which accumulated at a rate of between 50 and 240 mm per 1000 years. I have
applied the estimates of Lee, Wood, and others in the stratigraphic interpretation of a
Middle Stone Age occupation at the site of Muguruk, western Kenya (McBrearty, 1986,
1988).
Textural Properties
Certain textural properties are characteristic of soils reworked by termites, primarily
because they select sediment for transport according to particle size. Clay size particles
may be ingested and later egested during nest building; silt particles may be grasped in the
mandibles and carried to the nest site (Lee & Wood, 1971~). The fungus growing
Macroterminae cement these soil particles together with saliva to build nests and galleries;
among the soil feeders, faeces containing the soil particles are the construction medium.
The hard outer shell of the mounds of some wood and grass eating termites, as well as the
lining of the inner nests, may be composed of orally transported soil particles mixed with
excreta (Wood & Sands, 1978).
The particle size distribution of the resulting sediment thus varies with the animals’
carrying capabilities, which is largely a function of mean body size within the colony. The
maximum particle size transported by Mucrotermes, the largest of termite genera, is 4 mm;
for the smaller Cubitermes it is probably about 1.5 mm (Darlington, 1985; Crossley, 1986).
122 S. McBREARTY
4
IO 0 6 4 2 0 -2 -4
99.9
99
95
90
00
s
50
20
IO
5
/-
.’
0.001 0.004 0.016 0.062 0,25 I 4 16
mm
Figure 7. Cumulative grain size frequencies for samples from Karonga, Malawi,
and Muguruk, Kenya. Mounds are those of Mucrotermes fulciger. [Sources: (1)
Crossley, 1986: 195, figure 4; (2) McBrearty, 1986: 122, figure 2.81.
Most species preferentially select the fine soil fractions for nest construction, with the
result that mounds have a higher proportion of fine particles than the soil from which they
are constructed (Lee&Wood, 1971a,6; Wood & Sands, 1978).
Lee & Wood (197 la) report that the workings of Australian termites result in a deposit
that is at least 20% clay. The Macroterminae of sub-Saharan Africa construct their
mounds from precisely selected clay-rich subsoil horizons (Boyer, 1948; Grasse, 1950;
Hesse, 1955; Nye, 1955; Harris, 1956; Watson, 1962, 1975; Stoops, 1964, 1968; Arshad,
198 1; Pomeroy, 1983b), and selection of clay for nest building has also been documented
for African Cubitermes and Procubitermes by Kemp (1955) Maldague (1959), Stoops
(1964), Wood et al. (1983), and Anderson & Wood (1984).
Thus surface soils in areas where termites are active are characteristically high in clay
content. The activities of Macrotermes in the Karonga area of Malawi has resulted in a soil
mantle that is 73-82% sand, 5-9% silt, and 12-20% clay (See Figure 7). In some areas of
intense erosion, sheet wash may remove the clay-rich mantle, exposing the impoverished
sandy soil beneath (Roose, 1976).
When burned, the natural concentrations of clay in old termitaria may mimic hearths
or fireplaces in the archaeological record (Clark & Harris, 1985). The clayey soils of
TERMITES AS POST-DEPOSITIONAL AGENTS 123
abandoned termite mounds are exploited for construction purposes throughout much of
sub-Saharan Africa (Milne, 1947; Lee & Wood, 1971~; and pers. obs.).
Mineralogical Properties
Termites affect soil mineralogy. It is consistently observed that termite mounds exhibit
high concentrations of certain minerals, including calcium, magnesium, phosphate,
potassium, and iron (Ap Griffeth, 1938; Adamson, 1943; Pendleton, 1943; Milne, 1947;
Sys, 1955; Robinson, 1958; Stoops, 1964; Lee &Wood, 19710; Watson, 1974u,b, 1975;
Pomeroy, 19760; Wood & Sands, 1978; Bagine, 1984). Calcium, magnesium, and potas-
sium are constituents of plant tissue, and the incorporation of saliva or faeces into the
building fabric may be the source of enrichment of these soil minerals (Bagine, 1984). In
addition Breznak et al. (1973) have demonstrated that termites of the species Coptotermes
formosunus are capable of fixing nitrogen. The mineral-rich soil of termite mounds is
widely eaten by pregnant and lactating women in sub-Saharan Africa, and Watson (1972,
1974b) uses the concentration of certain precious metals in termite earths as an aid in
prospecting.
Anderson & Wood (1984) suggest that the high mineral content of soil in the mounds of
soil feeding termites is due to the very high pH conditions to which the soil is subjected as it
passes through the termite gut. Calcium carbonate may be present as hardpan or nodules
at the base of termite mounds; Pomeroy (1976~) correlates this feature with a fluctuating
water table. Watson (1974u,b) suggests that calcium bicarbonate is carried in solution in
acid groundwater; calcium carbonate will precipitate when this charged groundwater
encounters the locally basic conditions in the soils within and beneath an active termite
nest. Calcification of nests above the water table may result from the migration of calcium-
rich groundwater upward through vertical water collecting tunnels (Sands, 1987).
The partially digested cellulose found in many termite nests contains tannins,
polyphenols, and lignin that can react with metallic ions in soil water (Sands, 1987).
Raunet (1979) stresses the importance of termites in the formation of the red friable iron-
rich soils in western Kenya. Whether they also contribute to the formation of true lateritic
duricrust has long been the subject of debate, some authors supporting this idea
(Nazaroff, 1931; Grasse, 1949, 1950; Ehrhart, 1951; Sys, 1955; Taltasse, 1957; Conrad,
1959; Tessier, 1959; Sands, 1987), others refuting it (Ap Griffeth, 1953; Boyer, 1958,1959;
Grasse & Noirot, 1959; Sillans, 1959; Lee & Wood, 1971~). Termites of the species
Anucunthotermes are thought to contribute to the formation of saline duricrusts in central
Asia (Valiakhmedov, 1979).
Maignien (1966) suggests that termites may contribute to the decomposition of lateric
crust, rather than to its formation, an idea supported by Lee & Wood (1971~). Boyer
(1959) outlines mechanisms whereby termites may be responsible for their burial, whereas
Josens (1983) notes that lateritic crusts provide suitable microenvironments for termites in
otherwise unsuitably arid habitats.
acidity is known to contribute to the dissolution of bone (Hare, 1980; White & Hannus,
1983), the action of these termites might be expected to slightly accelerate the already
formidable chemical weathering of bone found in much of the humid tropics.
Some termite species have no effect on local soil acidity (Lee & Wood, 1971a), but
others are known to raise the pH in their nests (Pendleton, 1941; Sen, 1944; Boyer, 1955,
1956; Hesse, 1955, Kemp, 1955; Nye, 1955; Sys, 1955; Gokhale et al., 1958; Robinson,
1958; Watson, 1962, 1967, 1975; Stoops, 1964; Lee & Woods, 1971a,b; Trapnell et al.,
1976; Arshad, 1977; Wood & Sands, 1978). An elevated pH contributes to an enhanced
microenvironment for bone preservation (Hare, 1980). Watson (1967) reports a mound in
an Iron Age graveyard in northeastern Zimbabwe which is presently inhabited by mem-
bers of both Macrotermes goliath and Odontotermes latericius. Human skeletons are
preserved in the alkaline soil beneath the mound, but do not survive in the surrounding
acidic soil.
Local chemical conditions favouring bone preservation may be short lived, however, as
they are dependent upon the lifespan of the colony and the mound. They are probably
offset by the physical effects of termite burrowing which contribute to bone dissolution.
Although termites may closely pack particles in nest walls, leaving little pore space, the
excavation of galleries greatly enhances overall soil porosity (Adamson, 1943; Boyer,
1958; Robinson, 1958; Ghilarov, 1962; Maldague, 1964; Lee & Wood, 1971~). Galleries
increase air circulation and permit the introduction of acid groundwater, accelerating
bone destruction.
Termites have also been observed to feed on carrion and to cause superficial bone
damage (Thorne & Kimsey, 1983). Behrensmeyer (1978) attributes damage to modern
bone in areas of nonalkaline soils north of the Amboseli basin in southern Kenya to
termites, although Hill (1987) notes that the minute surface indentations on the bone
observed by Behrensmeyer might have been produced by subterranean safari ants of the
genus Dorylus or by rootlets and soil acid.
However, it is unlikely that the termitensavannen landscape existed prior to the modern
regime of regular grass firing, which is not thought to have been established prior to
600,000 years ago (Cerling, pers. comm.). Furthermore, it is not clear how the effects of
controlled use of fire are to be distinguished from the baked clay lumps, mounds, and clay
lined depressions which may result from natural burning of termite architecture.
Case Studies
The following sites show stratigraphic and sedimentary features which may best be
explained by the action of termites.
Laetoh’, Tanzania
Fossilized termitaria are reported from both the Upper and Lower Laetolil Beds at the
Pliocene site of Laetoli in northern Tanzania (Sands, 1987). They are abundant and
remarkably well preserved at five of the 23 fauna1 localities in the Upper Laetolil Beds,
occurring as both cast and matrix fossils of hard calcite and softer clay, sometimes
cemented within the eolian volcanic deposits. Branching, parallel-sided tubular structures
with oval cross sections from a few millimetres to several centimetres in diameter are
interpreted as Macrotermes foraging tunnels, although they lack the foraging pits
described by Darlington (1982) for M. michaelseni.
Ovoid masses are thought to represent Macrotermes queen cells. They consist of either
hard calcite or softer granular sediment, resting in a usually close-fitting cavity within the
surrounding matrix. Mean widths for two samples of ovoids in the lower part of the Upper
Laetolil Beds are given as 46.9 f 14.4 cm (N= 21) and 34.5 k 16.4 cm (N= 46); heights are
22.5 & 8.6 cm and 14.6 & 6.3 cm. They were found in some instances to rest on concave
circular baseplates about twice as large in diameter as the ovoids themselves, sometimes
with columnar structures around their rims.
Two well preserved shafted chambers resembling Macrotermes nests are described from
fauna1 locality 10 W. The first is a bell-shaped structure about 60 cm high and 1-Oto 1.2 m
in diameter encased in a calcite crust and topped by two vertical ventilation shafts about
65 cm high. The second, slightly smaller, is crowned with four ventilation shafts.
The same site yielded columnar masses of hard brittle calcite up to two cubic metres in
volume and 0.5 m thick and pierced by pores and tunnels. These bear a superficial struc-
tural resemblance to vesicular laterite, but are of course of entirely different chemical
composition. These resemble repair material found on live Macrotermes nests, which is
normally built only in small amounts. Studies by Ruelle (1964) and Weir (1973) have
TERMITES AS POST-DEPOSITIONAL AGENTS 127
shown that when Macrotermes colonies are covered with plastic sheets, they respond to
the imposed heat stress by rapidly coverting the upper portions of their mounds from
closed to open ventilation systems [Sands, 1987; compare figures 2(a) and 2(b)]. Sands
(1987) postulates that at Pliocene Laetoli the natrocarbonatite ash falls, rapidly cemented
by rainfall, would effectively suffocate a termite colony, cause heat stress, and stimulate
colonies to the hectic repair activity which would result in such masses of repair material.
Using wet dust and plastic sheets, he attempted to replicate these conditions on several live
Macrotermes nests in Laetoli. Because the animals’ response was slight, Sands postulates
that the climate at Laetoli in the Pliocene was warmer than that of the present day.
Termite burrowing activity at Lot. 1OW about 1.0 to 2.5 m below marker tuBI, dated at
3.7 Ma (Hay, 1981), was quantified by observing the densities of tunnels and ovoid struc-
tures in vertical exposures; when plotted against depth they are found to have a bimodal
distribution, representing two peaks of activity beneath landsurfaces formed by successive
accumulations of airfall tuff. Using Hay’s (1987) sedimentation rates for the Upper
Laetolil Beds of 15-20 cm per 1000 years, Sands (1987) estimates that these two peaks of
termite activity span 4000-6000 years and 5000-7000 years, respectively, separated by a
period of 8000 to 10,000 years. Due to the high degree of temporal control provided by the
Laetoli chronology, Sands (1987) is able to compute densities of termitaria per 1000 year
slice of sediment; when corrected for expected longevity of a Macrotermes colony, his
estimate of 3.93 mounds ha-’ approximates densities in modern semiarid savanna
environments.
Bodo, Ethiopia
A number of mounds of burned clay have been described from Plio-Pleistocene contexts
in the Middle Awash Valley of Ethiopia (Clark et al., 1984; Clark & Harris, 1985). In some
cases, stone artifacts and fractured bone are found in the immediate vicinity of the struc-
tures; in other instances, there are no associated artifacts or bone. The question naturally
arises as to the respective roles of termites and hominids in their formation.
The features consist of cone-shaped concentrations of reddened clay between 40 and
80 cm in diameter and about 20 to 30 cm high. The clay forms irregular pellicules in some
instances; in others, particularly near the base of the cones, it is more massive. Small
discrete pellicules are found at distances up to 1.5 m downslope from the features.
Analysis of samples from Oldowan locality Bodo A-4 and Acheulian locality HAR-A3
confirm that the material is burned. The minimum temperature at which the clay had been
fired is estimated at 200” C on the basis of its remnant magnetism (Clark & Harris, 1985).
From the Middle Pleistocene locality at Bodo which has yielded cranial fragments of
Homo erectus (BOD-VP- l/l), about 400 m from the initial Bodo hominid find (Conroy ef
al., 1978; Kalb et al., 1980, 1982), Clark et al. (1984) report excavation of two of these
baked clay mounds. The features lie within dark clayey sediment which is interpreted as
overbank deposits. The concave bases of the excavated cones seem to resemble the fossil
Macrotermes baseplates described from Laetoli by Sands (1987), and interior “channels”
suggest termite activity. Colour varied from strongly reddened in the centre to yellow at
the periphery of the structures. Neither contained artifacts or bone, although both
materials occur on the surface in the vicinity.
These concentrations are interpreted as termite earth which surrounded a tree stump as
it burned (Clark et al., 1984; Clark & Harris, 1985). Clark & Harris (1985) suggest that the
fire may have been conserved or used by early hominids. This is an interesting idea, but one
that is difficult to test. There is no compelling reason to implicate hominids in the burning
of the sediments. Termite mounds are focal points of activity for primates as well as other
animals, and these islands of woody vegetation are subject to periodic natural bush fires.
128 S. McBREARTY
As James (1989) points out, the bones, stone artifacts, and baked clay at such localities
may not be in meaningful association.
Karonga, Malawi
In the region of Karonga, near the western shore of Lake Malawi, a number of
archaeological occurrences have been described which are attributed to the Sangoan and
Middle Stone Age (Clark et al., 1970, 1972). The Chiwondo Formation here consists of
sandstones, siltstones, mudstones, and marls containing Plio-Pleistocene to Middle
Pleistocene vertebrate and invertebrate fossils (Haynes, 1970; Kaufulu et al., 1981). It is
overlain unconformably by the Chitimwe Formation, which is made up of sands and
gravels containing Middle and Later Stone Age artifacts. At the site of Mwanganda’s
Village the remains of an elephant were found together with stone artifacts on the weath-
ered surface of the Chiwondo Formation at its contact with the Chitimwe Formation
(Clark & Haynes, 1970).
Crossley (1986) has described laterally discontinuous sheets of red clayey sand which
cap the Karonga sequence. The sand sheets cover a very considerable area of about
8800 km2, and are found upon a variety of substrates. Usually they overlie gravels of the
Chitimwe Formation, but in some cases they cover Mesozoic or basement rocks [See
Figures 5(b), 9(b)]. These deposits are between 3 and 6 m thick in areas with slopes of less
than 3”, and they are clearly of terrestrial origin as in many cases they drape the local
topography [See Figure 5(b)]. They lack internal structure and are composed of 73-82%
sand, 12-20% clay, and about 7% silt (See Figure 7). Near horizontal lines of scattered
stone artifacts occur within these clayey sands [Figures 7 and 9(b)]; unabraded flakes from
the Chaminade locality have been found to be conjoinable (Clark et al., 1970; Crossley,
1986).
In the Chaminade area modern Macrotermes falciger mounds occur at a density of
24 haa2, and in active mounds Crossley has observed sediment particles of the Chitimwe
Formation which has been brought up from depths of at least 5 m. From the same locality
Crossley describes dome-shaped piles of clayey sand and carbonate concretions up to 3 m
high and 6 m across, which he interprets as fossil termitaria of the genus Macrotermes.
From these observations and the character of the sediments, Crossley reasonably
infers that these sand sheets are the product of termite deposition and are derived from
the underying levels. This clearly implies a lack of stratigraphic integrity for the
archaeological contents of the biogenic mantle, and suggests some disturbance of the
underlying deposits.
TERMITES AS POST-DEPOSITIONAL AGENTS 129
Figure 8. Structureless red clayey soil with stone line (arrowed) at Ober Awach,
South Nyanza District, Kenya. Artifacts eroding from the sediment have accumu-
lated as the lag deposit in the foreground.
MSA artifacts -
Structureless
Loterised sands - red clayey sand
LSA artifacts
Mudcracks -
Artifacts
MSA artifacts
Brown clayey sand
Grey clayey sands Carbonate nodules
E E
Iron pisaliths
ESA artifacts
Coarse sands 6 Gravel
Sangoan-Lupemban
artifacts
Grey sandy clay
Cloy lens with
artifacts
Conglomerate
Phonolite
(b)
(a)
Figure 9. Stratigraphic sections. (a) Muguruk, western Kenya (after McBrearty,
1988: 394, figure 3). Circled numbers refer to members of the Muguruk Formation;
(b) Chaminade site 2, Malawi (after Crossley, 1986: 193, figure 2).
130 S. McBREARTY
and perhaps differences in the amount of time represented. Most important is the fact
that the Muguruk sediments are probably not the exclusive result of termite sediment
turnover. At Muguruk, unlike Karonga, where the sediments are terrestrial in origin,
successive fluvially aggraded landsurfaces have been subject to reworking by termites.
In spite of this, it is interesting to consider that if the sediment mantle at Muguruk were
accumulated exclusively by termites, the time elapsed could be calculated from known
termite sedimentation rates. The most common genus of termites in the region is
Macrotermes; Nye (1955) estimates that M. bellicosus accumulates about 0.3 m of
sediment in 12,000 years. From studying photographs of Muguruk taken in the late 1930s
by the site’s discoverer, the Archdeacon Owen, I estimate that before removal of tree cover
initiated the current regime of acute erosion, the occupation layer in member 4 was
overlain by about 1.5 m of uniform clay- and iron-rich sediment. It would take termites of
the species M. bellicosus about 60,000 years to accumulate this thicknes of sediment. By
contrast, using sedimentation rates for rift valley margins calculated by Schwab (1976),
fluvial accretion alone could result in this depth of deposit in less than 9000 years.
Masango, Burundi
A Lupemban assemblage excavated at the site of Masango, in central Burundi, has been
described by Van Noten et al. (1972). The occupation lies at the summit of an isolated hill,
where decomposed granite is overlain by about 1.5 m of sands, silts, and clays which are
judged to be the product of in situ weathering. In the main excavation area (9 m*) an
artifact horizon, about 5-7 cm thick, was encountered at a depth of about 10 cm. This
horizon is overlain in other parts of the site by 15560 cm of silty sand.
The upper surface of the occupation layer was irregular, but its lower surface horizontal
and fairly regular. There was no preferred artifact orientation observed for the sample of
the excavation (1 m*) for which data were recorded. However, a number of circular,
artifact-free areas about l&20 cm in diameter were encountered in excavation. Artifacts
at their perimeters showed clear inclination toward the centres of the empty areas.
Artifact densities in this unit were extremely high, exceeding 800 pieces larger that 2 cm
in maximum dimension per m*. The artifacts include classic Lupemban points made from
quartzite and Lupemban debitage, artifacts that would be expected in a Lupemban con-
text. In addition, however, small numbers of quartz Later Stone Age artifacts were
encountered, together with small quantities of charcoal, which yielded radiocarbon dates
of 5270 and 4400 f 40 BP. No bone was preserved.
The presence of termites at modern Masango is well attested by direct observation.
Termitaria are found on the summit of the hill today; their density is estimated at 25
mounds lOOOm-*. In addition, large termite burrows were encountered within the
excavation itself. Artifacts rested obliquely at the margins of these cavities and on
occasion tumbled into galleries below /Van Noten et al., 1972, figure l(c)].
Not surprisingly, Van Noten et al. suggest that termites are responsible for the
disturbance of the archaeological and stratigraphic record at Masango. Disturbance of
the material is suggested by the mixture of Lupemban and Later Stone Age artifacts, and
by radiocarbon dates in keeping with a Later Stone Age occupation on charcoal
associated with Lupemban artifacts.
Termites are implicated as the disturbing agent by a number of lines of sedimentological
evidence. The thickness and density of the archaeological occurrence, as well as the
sharply demarcated upper and lower boundaries of the archaeological layer, are con-
sistent with those of “stone lines” created by termites. Although a detailed sedimentary
description is not given, it appears that the nature and thickness of the deposit overlying
the artifact horizon is in keeping with this interpretation. Additional support is lent by the
132 S. McBREARTY
clear presence of termites at the site today, and by the circular, artifact free areas in the
occupation layer which are surrounded by inclined artifacts; these appear to be fossil
termite galleries.
Diagnostic Features
The following features may alert the archaeologist that termites have been active: (1)
modern or fossil termitaria; (2) fragments of compact or vesicular construction material;
(3) baked clay lumps or clay lined depressions; (4) excavated foraging passages; (5) com-
plete absence of bone; (6) localized patches of bone preservation; (7) complete lack of
microstratigraphy; (8) a uniform, porous, clay rich soil mantle; and (9) vertically dispersed
artifacts; or (10) vertically concentrated artifacts.
Termitaria may be complete or fragmentary bell- or cone-shaped mounds constructed
of mineralized clayey sediment. They may be underlain by a basin-like depression and
crowned by a superstructure of tubular ventilation shafts. Internally, they are likely to be
pierced by passageways and nursery galleries; they may contain fragments of fungus comb
and a central oval cavity, the remains of the royal chamber. All of these excavated features
may be infilled with sediment, and the structure and its infill may be differentially
mineralized. The soil of the mound may have a high organic content, high pH, and may be
rich in compounds of Ca, Ph, Mg, K, N, and Fe. The mound may be underlain by a patch
of localized bone preservation, and this bone may exhibit superficial damage. The deposits
may show traces of woody vegetation and burning.
Layers underlying a termite nest can be expected to exhibit excavated cavities,
chambers, and galleries in various stages of collapse, and perhaps wholly or partially filled
with matrix from an overlying layer. Excavated foraging passages may extend for great
distances and may be open, infilled with sediment, or mineralized. Artifacts derived from
higher layers may be found within them. These passages can be distinguished from root
casts by their oval, rather than round cross-sections, their branching pattern, in which
passages may be directed upward, and by the fact that they do not taper toward their ends
(Sands, 1987).
Although relic termitaria may not be preserved in all cases, their cumulative effects can
be readily discerned. The classic profile of an ancient site in which termites have artificially
concentrated archaeological material can be described as follows: the artifacts are con-
fined to a uniform depth beneath a landsurface, either ancient or modern. This layer has
sharp upper and lower contacts and lies at the junction of a structureless, porous, clay-rich
layer with underlying, coarser, more compact sediment. It may be underlain by a layer of
calcrete nodules or hardpan. There may be traces of a localized woody vegetation, patches
of well preserved bone, or remnants of fossil or modern termite galleries. These features
may be associated with lateritic duricrusts. Where termites have dispersed artifacts
through a depth of unconsolidated deposit, they are frequently reconcentrated as lag
deposits by erosive forces. I have observed these conditions in the deep red lateritic soils at
many of the sites in the Lake Victoria Basin which were described by Leakey & Owen
(1945) in their early archaeological assessment of the region (See Figure 8).
regarded as evidence for the controlled use of fire (e.g. Gowlett et al., 198 1). Scattered fired
clay lumps might easily be eroded from burned termite mounds such as those described by
Clark & Harris (1985), to be redeposited elsewhere.
Second, termites destroy or alter stratigraphy. They create an unstable landsurface
subject to collapse (Wood & Sands, 1978), and a biomantle through which artifacts may
migrate. Thus archaeological chronologies for equatorial Africa must be re-evaluated.
The work of Cahen (1976,1978a) at Gombe Point suggests that material from throughout
the Zaire basin which has been divided into chronological stages by workers such as Breuil
(1944), Breuil& Janmart (1950), Collette (1929, 1931a,b, 1935), Van Moorsel(1968), and
Mortelmans (1957a,b, 1962a,b) is not in reliable archaeological context. Crossley’s
analysis (Crossley, 1986) suggests that archaeological sites in the Malawi rift must perhaps
be looked at more critically before inferences about living floors and butchery sites can be
made.
Furthermore, these are not isolated phenomena. Crossley’s observations encompass
nearly 9000 km*, and the Kalahari sands that Cahen & Moeyersons (1977) conclude are
a reconsolidating sedimentary mantle continuously reworked by termites cover a vast
region of the southern Zaire basin. Therefore the Sangoan material from northeast
Angola described by Leakey (1949), Janmart (1947, 1953), and Clark (1963, 1966) may
owe the circumstances of its preservation to the action of termites. With the exception of a
few possible flaking floors described by Janmart (1953), the Angolan material is found
scattered as isolated specimens through many metres of fluvial and colluvial Kalahari
sands. From my own observations I know that many of the “Tumbian” (Sangoan-
Lupemban) artifacts from sites in western Kenya such as Yala Alego and Ober Awach
(Leakey & Owen, 1945) occur in similar contexts (See Figure 8).
Cahen (19786) has suggested that in light of the poor quality of our data, all artifactual
material postdating the Acheulian in Central Africa should simply be attributed to
“post-Acheulian industrial complex”. Cahen’s scheme would collapse several hundred
thousand years of human prehistory, from the disappearance of handaxes to the appear-
ance of iron working technology, into a single archaeological stage. His evaluation of
the state of Paleolithic archaeology in Central Africa seems overly pessimistic, but it is
undeniable that much informed careful work is required before even the outlines of
prehistory in this vast region of tropical Africa are clear to us.
Conclusions
Wood & Johnson (1978) observe that the use of the term in situ to denote undisturbed
artifacts may be more optimistic than realistic in many contexts. This is especially true for
tropical soils, where termites have been active for tens of millions of years. They produce
effects which may be interpreted by the unwary as the products of human behaviour or of
depositional or environmental circumstances at the time of occupation.
Clearly archaeologists have assumed a level of temporal resolution in their inter-
pretations of tropical Pleistocene remains which is rarely justified. Living floors which we
presume to represent brief instants of time may exist in the tropics, but they must be
carefully distinguished from the products of post-depositional concentrations by termites.
Living floors may be more rare in the archaeological record than archaeologists would
care to admit, but this does not mean that archaeological evidence can supply no
meaningful information simply because it does not conform to the narrow in situ model.
The method outlined by Schick (1986) for scoring sites according to degree of disturbance
seems a fruitful approach to the problem.
Perhaps in return for their wholesale destruction of stratigraphic evidence, termites
furnish new directions for future research. They selectively concentrate clay in locations
more likely to be subject to prolonged burning than other spots on the landscape.
134 S. McBREARTY
Acknowledgements
I would like to thank Charlie Vogt (Department of Entomology, Museum of
Comparative Zoology, Harvard University), Andrew Hill (Department of Anthropology,
Yale University) and Ian Deshmukh (Associates for Rural Development and USAID,
Mogadishu), for reading and commenting on earlier drafts of this paper. I thank Bambi
Thorne (Department of Entomology, Harvard University), Jo Darlington (International
Center for Insect Physiology and Ecology, Nairobi), Robert Crossley (Robertson
Research), Dave Killick (Department of Anthropology, Harvard University), Mellisa
Remis (Department of Anthropology, Yale University), and Eleanor Sterling (Depart-
ment of Anthropology, Yale University) for useful discussion of termites and their habits
and for providing useful references. Richard Hay called the Laetoli fossil termitaria to my
attention, and Mary Leakey and Andrew Hill very kindly provided me with proof copies
of sections of the Laetoli monograph before publication. Research at Muguruk was
carried out with primary funding from the L.S.B. Leakey Foundation and the National
Science Foundation. Permission to carry out the research was granted by the Office of the
President of the Republic of Kenya, and sedimentary analyses were carried out at the
National Museums of Kenya, the University of Illinois, and Harvard University. For
kindly facilitating this aspect of the project I thank Richard Leakey, David Pilbeam,
Richard Hay, John Alexander, and Ann Kosobud.
References
Adamson, A. M. (1943). Termites and the fertility of soils. Tropical Agriculture 20, 107-l 12.
Anderson, J. M. & Swift, M. J. (1983). Decomposition in tropical forests. In (S. L. Sutton, T. C.
Whitmore & A. C. Chadwick, Eds) Tropical Rainforest: Ecology & Management. Oxford:
Blackwell, pp. 287-309.
Anderson, J. M. & Wood, T. G. (1984). Mound composition and soil modification by two
soil-feeding termites (Termitinae: Termitidae) in a riparian Nigerian forest. Pedobiologia 26,
77-82.
TERMITES AS POST-DEPOSITIONAL AGENTS 135
Ap Griffeth, G. (1938). A note on termite hills. East African Agricultural Journal 4,7&7 1.
Ap Griffeth, G. (1953). Vesicular laterite. Nature 171,530.
Arshad, M. A. (1977). The role of Macrotermes in soils. International Center of Insect Physiology
and Ecology Annual Report 5,39-40.
Arshad, M. A. (198 1). Physical and chemical properties of termite mounds of two speciesof
Macrotermes (Isoptera: Termitidae) and the surrounding soils of the semiarid savanna of
Kenya. Soil Science 132, 161-174.
Badgeley, C., Tauxe, L. & Bookstein, F. L. (1986). Estimating the error of age interpolation in
sedimentary rocks. Nature 319,13914 1.
Bagine, R. N. (1984). Soil translocation by termites of the genus Odontotermes (Holmgren)
(Isoptera: Macrotermitinae) in an arid ara of northern Kenya. Oecologia (Berlin) 64,263-266.
Baroni-Urbani, C., Josens,G. & Peakin, G. J. (1978). Empirical data and demographic
parameters. In (M. V. Brian, Ed.) Production Ecology of Ants and Termites. Cambridge:
University Press,pp. 544.
Barton, R. N. E. & Bergman, C. A. (1982). Hunters at Hengistbury: some evidence from
experimental archaeology. World Archaeology 14,237-248.
Behrensmeyer, A. K. (1978). Taphonomic and ecologic information from bone weathering.
Paleobiology 4,135-149.
Behrensmeyer, A. K. (1982). Time resolution in fluvial vertebrate assemblages.Paleobiology 8,
21 l-227.
Bocek, B. (1986). Rodent ecology and burrowing behaviour: predicted effects on archaeological
site formation. American Antiquity 51,589-602.
Bodenheimer, R. s. (1951). Insects as Food: A Chapter ofthe Ecology of Man. The Hague: Junk.
Bodot, P. (1967). Etudes ecologiques des termites des savannesde BasseCote d’Ivoire. Insects
Sociaux 14,229-258.
Bouillon, A. (1970). Termites of the Ethiopian region. In (K. Krishna & F. M. Wessner, Eds)
Biology of Termites 2, 153-280.
Boyer, P. (1948). Sur lesmateriaux composant la termitihe geante de Bellicositermes rex. Comptes
Rendus des Seances de I’Academie des Sciences 227,488490.
Boyer, P. (1955). Premiere etudes pedologiques et bacteriologiques des termitiires. Comptes
Rendus Hebdomadaires des Seances de I’Academie des Sciences 240,569-571.
Boyer, P. (1956). Action des termites constructeurs sur certain solsd’Afrique Tropicale.
Proceedings of the Sixth International Congress of Soil Scientists 3,95-103.
Boyer, P. (1958). Influence des remaniements par le termite et de l’erosion sur l’evolution
pedogenetique de la termitiere epigee de Bellicositermes rex. Comptes Rendus des Seances de
I’Academie de Sciences 247,749-75 1.
Boyer, P. (1959). De I’influence des termites de la zone intertropicale sur la configuration de
certains ~01s.Revue de Geomorphologie Dynamique 10,41&44.
Brain, C. K. (198 1). Hunters or the Hunted? An Introduction to African Cave Taphonomy. Chicago:
Chicago University Press.
Breuil, H. (1944). La paleolithique au Congo Belge aprb les recherchesde Docteur Cabu; les
industries paleolithiques de la terrace de 15 metres d’un chenal secondaire compliplaine de
Piedmont de Leopoldville, apt-esles fouilles et les photographies du Docteur Cabu.
Transactions of the Royal Society of South Africa 30, 143-167.
Breuil, H. & Janmart, J. (1950). Les Limons et Graviers de 1’Angola de Nord-Est et leur Contenu
Archaeologique. Lisbon: Museuo do Dundo Publicacoes Culturais 5.
Breznak, J. A., Brill, W. J., Mertins, J. W. & Coppel, G. C. (1973). Nitrogen fixation in termites.
Nature 244,577-580.
Bullard, R. (1978). Geology in field archaeology. In (W. G. Dever & H. D. Lance, Eds) A Manual
of Field Excavation. New York: Hebrew University College, pp. 196-235.
Bunn, H. (1981). Archaeological evidence for meat-eating by Plio-Pleistocene hominids from
Koobi Fora and Olduvai Gorge. Nature 291,574577.
Bunn, H., Harris, J. K. W., Isaac, G. Ll., Kaufulu, Z., Kroll, E., Schick, K., Toth, N. &
Behrensmeyer, A. K. (1980). FxJj50: an early Pleistocenesite in northern Kenya. World
Archaeology 12,109-136.
Burnham, L. (1978). Survey of social insectsin the fossil record. Psyche 85,83-133.
136 S. McBREARTY
Milne, G. (1947). A soil reconnaissance journey through parts of Tanganyika Territory. Journal of
Ecology 35,192-265.
Moeyersons, J. (1978). The behaviour of stones and stone implements, buried in consolidating
and creeping Kalahari sands. Earth Surface Processes 3, 115-128.
Mortelmans, G. (1957a). Le Cenozoic du Congo Belge. In (J. D. Clark, Ed.) Proceedings ofthe
Third Panafrican Congress on Prehistory, Livingstone, 19.55. London: Chatto and Windus,
pp. 23-50.
Mortelmans, G. (19576). Le prehistoire du Congo Belge. Revue de I’Universitk de Bruxelles 9,
l-53.
Mortelmans, G. (1962a). Le Quaternaire du Congo occidentale et sa chronologie. In (G.
Mortelmans & J. Nenquin, Eds) Actes du Zip Congres Panafricain de Prehistoire et de l’ktude du
Quaternaire. Tervuren: MusCe Royale de I’Afrique Centrale. Series in-8”, Sciences Humaines
40(l), 97-132.
Mortelmans, G. (19626). Vue d’ensemble sur la prehistoire due Congo occidentale. In (G.
Mortelmans & J. Nenquin, Eds) Actes du IV Congres Panafricain de Prehistoire et de I’Etude du
Quaternaire. Tervuren: MusCe Royale de l’Afrique Centrale. Series in-8”, Sciences Humaines
40( 1), 129-164.
Nazaroff, P. S. (1931). Note on the spongy ironstone of Angola. Geological Magazine 68,
4433446.
Noirot, C. (1970). The nests of termites. In (K. Krishna & F. M. Weesner, Eds) The Biology of
Termites. New York: Academic Press 2,73-125.
Nye, P. H. (1955). Some soil-forming processes in the humid tropics. IV. The action of the soil
fauna. Journal of Soil Science 6,73-83.
Ollier, C. D. (1959). A two-cycle theory of tropical pedology. Journal of Soil Science 10, 137-148.
Owen, D. F. (1976). Animal Ecology in Tropical Africa. London: Longman.
Pathak, A. N. & Lehri, L. K. (1959). Studies on termite nests. I. Chemical, physical, and biological
characteristics of a termitarium in relation to its surroundings. Journal of the Indian Society of
Soil Science 7,87-90.
Pendleton, R. L. (1941). Some results of termite activity in Thailand soils. Thailand Science
Bulletin 3,29-53.
Pendleton, R. L. (1943). Importance of termites in modifying certain Thailand soils. Journal of the
American Society of Agronomists 34,340-344.
Pomeroy, D. E. (1976a). Studies on a population of large termite mounds in Uganda. Ecological
Entomology 1,49961.
Pomeroy, D. E. (19766). Some effects of mound-building termites on soils in Uganda. Journal of
Soil Science 27,377-394.
Pomeroy, D. E. (1977). The distribution and abundance of large termite mounds in Uganda.
Journal of Applied Ecology 14,465475.
Pomeroy, D. E. (1978). The abundance of large termite mounds in Uganda in relation to their
environment. Journal of Applied Ecology 15,51-63.
Pomeroy, D. E. (1983a). A striking increase in a population of termite mounds in eastern Kenya.
Kenya Journal of Science and Technology, Series B, 4,89-96.
Pomeroy, D. E. (19836). Some effects of mound-building termites on the soils of a semi-arid area
of Kenya. Journal of Soil Science 34,555-570.
Pomeroy, D. E. (1983~). Distribution and abundance of large termite mounds in a semiarid area
of southern Kenya. Kenya Journal of Science and Technology, Series B, 4,77-87.
Potts, R. (1984). Hominid hunters? In (R. Foley, Ed.) Hominid Evolution and Community
Ecology: Prehistoric Human Adaptation in Biological Perspective. London: Academic Press,
pp. 129-166.
Potts, R. (1988). Early Hominid Activities at Olduvai. New York: Aldine de Gruyter.
Potts, R. & Shipman, P. (1981). Cutmarks made by stone tools on bones from Olduvai Gorge,
Tanzania. Nature 291,577-579.
Rapp, G. & Gifford, J. A. (1982). Archaeological geology. American Scientist 70,45-53.
Rapp, G. & Gifford, J. A. (Eds) (1985). Archaeological Geology. New Haven: Yale University
Press.
TERMITES AS POST-DEPOSITIONAL AGENTS 141
Ratcliffe, F. N., Gay, F. H. & Greaves, T. (1952). Australian Termites. Melbourne: CSIRO.
Raunet, M. (1979). Importance et interactions des procesus geochemiques, hydrologiques, et
biologiques (termites) sur les surfaces d’applanissement tropicales granito-gneissiques. Exemple
au Kenya occidentale. Agronomie Tropicale 34,4&53.
Renfrew, C. (1976). Archaeology and the earth sciences.In (D. A. Davidson & M. L. Shackley,
Eds) Geoarchaeology: Earth Science and the Past, l-5, London: Duckworth.
Retallack, G. (1984). Completeness of the rock and fossil record: some estimates using fossil soils.
Puleobiology IO,5978.
Robinson, J. B. D. (1958). Some chemical characteristics of “termite soils” in Kenya coffee fields.
Journal of Soil Science 9,58-65.
Rolfsen, P. (1980). Disturbance of archaeological layers by processesin the soil. Norwegian
Archaeological Review 13,l l&l 18.
Roose, E. J. (1976). Contribution a l’ktude de f’lnfluence de la Mesofaune sur la Pedogenbe
Actuelle en Milieu Tropical. Ivory Coast: Centre d’Adiopodoume.
Rowlett, R. M. & Robbins, M. C. (1982). Estimating original assemblagecontent to adjust for
post-depositional vertical artifact movement. World Archaeology 14,73-83.
Ruelle, J. E. (1964). L’architecture du nid de Macrotermes natalensis et son sensfunctionel. In (A.
Bouillon, Ed.) Etudes sur les Termites africaines. Leopoldville, pp. 327-352.
Ruelle, J. E. (1985). Order Isoptera (Termites). In (H. S. Clarke & E. Holm, Eds) Insects of
Southern Africa. Durban: Butterworths, pp. 53361.
Sadler, P. M. (198 1). Sediment accumulation rates and the completeness of stratigraphic sections.
Journal of Geology 89,569-584.
Salder, P. M. & Dingus, L. W. (1982). Expected completeness of sedimentary sections: estimating
a timescale dependent, limiting factor in the resolution of the fossil record. Third North
American Paleontological Convention, Montreal 2,461464.
Salick, J., Herrara, R. & Jordan, C. F. (1983). Termitaria: nutrient patchiness in nutrient deficient
rainforest. Biotropica 15, l-7.
Sampson, C. G. (1972). The Stone Age industries of the Orange River Schemeand South Africa.
Bloemfontein: National Museum Memoir 6.
Sands, W. A. (1965). Mound population movements and fluctuation in Trinervitermes ebenerianus
Sjostedt (Isoptera, Termitidae, Nasutiterminae). Znsectes Sociaux 12,49-58.
Sands, W. A. (1987). Ichnocoenoses of probable termite origin from Laetoli. In (M. D. Leakey &
J. M. Harris, Eds) Laetoh: A Pliocene Site in Northern Tanzania Oxford: Oxford University
Press,pp. 409433.
Schick, K. A. (1986). Stone Age Sitesin the Making: Experiments in the Formation and
Transformation of Archaeological Occurrences. Oxford: B.A.R. International Series 163.
Schiffer, M. B. (1983). Toward the identification of site formation processes.American Antiquity
48,675-706.
Schindell, D. E. (1980). Microstratigraphic sampling and the limits of paleontologic resolution.
Paleobiology 6,4088426.
Schindell, D. E. (1982~). Gaps in the fossil record. Nature 297,282-284.
Schindell, D. E. (1982b). Resolution analysis: a new approach to gaps in the fossil record.
Paleobiology 8,34&353.
Schwab, F. L. (1976). Modern and ancient sedimentary basins: comparative evolution rates.
Geolgoy 4,723-727.
Sen, A. (1944). The influence of feed upon the composition of termite soils. Current Science 13,
28&28 1.
Shackley, M. L. (1975). Archaeological Sediments. London: Buttenvorths.
Siiriainen, A. (1977). Piecesin vertical movement-a model for rockshelter archaeology.
Proceedings of the Prehistoric Society 43,349-353.
Sillans, R. (1959). Les Savanes de I’Afrique Centrale. Paris: Lechevalier.
Stein, J. K. (1983). Earthworm activity: a source of potential disturbance of archaeological
sediments. American Antiquity 48,277-289.
Stockton, E. D. (1973). Shaw’s Creek Shelter: human displacement of artifacts and its
significance. Mankind9,112-117.
142 S. McBREARTY
Stoops G. (1964) Application of some pedological methods,to analysis pf. termite mounds. In (A.
Bou{llon. &I.) ,$tudes sur Ies Termites Africains. Ltopoldvllle: Umverslte de LCopol&ille,
pp. 379-i98.
Stoops, G. (1968). Micromorphology of some characteristic soils of the Lower Congo (Kinshasa).
Pedology l&1 l&149.
Sys,C. (1955). The importance of termites in the formation of latosols in the region of
Elizabethville. Soils of Africa 3,393-395.
Taltasse, P. (1957). Les cab&es et jacare et le r6le des termites. R&we de Giomorphologie
Dynamique ll-12,166170.
Tessier,F. (1959). The laterite of the Manuel Cape at Dakar and its fossil termitaries. Comptes
Rendus de l%lcademie des Sciences de Paris 248,3320-3322.
Thakur, M. L. (1981). Origin, dispersion, and distribution of Macroterminae (Isoptera:
Termitidae). Jo.wnal of Entomological Research 5,8&90.
Thon, L. (1946). A propos des termites au point de vue alimentaire. Bulletin Agricole du Congo
Beige 37,865~868.
Thorne, B. L. & Kimsey, R. B. (1983):Attraction of Neotropical Nasutitermes termites to carrion.
Biotropica 15,295-296.
Trapnell, C. G., Friend, M. T., Chamberlain, G. T. & Birch, H. F. (1976). The effects of fire and
termites on a Zambian woodland soil. Journal of Ecology 64,577-588.
Troll, C. (1936). Termitensavannen. In Landekundliche Vorshrift Festschriftfur Norbert Krebs.
Stuttgart: Engelhom, pp. 275-3 12.
Tutin, C. E. G. & Femandez, M. (1983). Gorillas feeding on termites in Gabon, West Africa.
Journal of Mammalogy 64,530-53 1.
Valiakhmedov, B. V. (1979). The influence of the termite Anacanthotermes ahngerianus Jacobs
(Isoptera: Insecta) on the development of takyr soils in Tadzhikistan. Doklady Akademii Nuuk
SSSR 247,48M82.
Van Andell, T. H. (1981). Consider the incompleteness of the fossil record. Nature 294,397-398.
Van Moorsel, H. (1968). Atlas de Prkhistoire de la Plaine de Kinshasa. Kinshasa: Publication de
I’UniversitC Lovanium.
Van Noten, F., Cahen, D., Keeley, L. H. & Moeyersons, J. (1978). Les Chasseursde Meer.
Brugge: Dissertations Archaeologicae Gandenses 17.
Van Noten, F., Cahen, D. & Keeley, L. H. (1980). A Paleolithic campsite in Belgium. Scient$c
American 242,48-55.
Van Noten, F., Haesaerts, P. 8cCahen, D. (1972). Un habitation Lupembien & Masango,
Burundi: rapport prkliminaire. Africa-Tervuren l&78-85.
Villa, P. (1982). Conjoinable piecesand site formation processes.American Antiquity 47,276-290.
Villa, P. (1983). Terra Amata and the Middle Pleistocenearchaeological record of Southern
France. Berkeley: University of California Publications in Anthropology 13.
Villa, P. & Courtin, J. (1983). The interpretation of stratified sites: a view from underground.
Journal of Archaeological Science 10,267-28 1.
Watson, J. A. L. & Gay, F. J. (1970). The role of grain-eating termites in the degradation of a
mulga ecosystem. Search 1,43.
Watson, J. P. (1960). Some observations of soil horizons and insect activity in granite soils.
Salisbury, Rhodesia: Proceedings of the First Federal Science Congress, pp. 271-276.
Watson, J. P. (1962). The soil beneath a termite mound. Journal of Soil Science 13,465 1.
Watson, J. P. (1967). A termite mound in an iron-age burial ground in Rhodesia. Journal of
Ecology 55,663-669.
Watson, J. P. (1972). The distribution of gold in termite mounds and soils at a gold anomaly in
Kalahari sand. Soil Science 113,3 17-321.
Watson, J. P. (1974a). Calcium carbonate in termite mounds. Nature 247,74.
Watson, J. P. (19746). Termites in relation to soil formation, groundwater, and geochemical
prospecting. Soils and Fertilizers 37, 11l-l 14.
Watson, J. P. (1975). The composition of termite (Mucrotermes spp.) mounds on soil derived from
basic rock in three rainfall zones of Rhodesia. Geoderma 14, 147-158.
Weir, J. S. (1973). Air flow, evaporation, and mineral accumulation in mounds of Mucrotermes
subhyalinus (Rambur). Journal of Animal Ecology 42,509-520.
TERMITES AS POST-DEPOSITIONAL AGENTS 143