Afr. J. Ecol.

1984, Volume 22, pages 229-244

Seasonal herd dynamics of a savanna elephant population

D. WESTERN and W. K. LINDSAY Animal Research and Conservation
Centre of The New York Zoological Society, P.O. Box 481 77, Nairobi, Kenya

Summary
The ecology and behaviour of the Amboseli elephant population in southern
Kenya has, unlike most populations studied so far, been largely unaffected by
human pressures, so provides an opportunity to examine and explain natural
seasonal cycles in movement, habitat selectivity and herd dynamics. Water
distribution and forage quality largely dictate seasonal migrations. During dry
seasons elephants are confined within the vicinity of permanent water and, like
the other wild herbivores in the ecosystem, select habitats with progressively
more abundant but less digestible forage. Herd size increases spontaneously at the
onset of each rains, and declines progressively through the dry season as food
reserves diminish. Various hypotheses for the seasonal aggregations are con-
sidered, including foraging, social, reproductive and anti-predator. The evidence
is still inadequate to show conclusively whether recent changes in elephant
movement result from increased human pressures, or changes in rainfall and food
availability.

Resume
L’ecologie et le comportement de la population d’elephants de 1’Amboseli (S
Kenya) a etC, au contraite de la plupart de celles etudiees jusqu’ici, Cpargnie de
toute pression anthropique, ce qui a permis d’ktudier et de discuter les cycles
naturels de mouvements saisonniers, la selectivitk des habitats et la dynamique
des populations. La disponibilite en eau et la qualit6 du fourrage dictent mani-
festement les migrations saisonnieres. Durant la saison seche les elephants sont
confines au voisinage des points d’eau permanents et, tout comme les autres
herbivores sauvages de cet ecosysteme, choisissent progressivement des milieux
offrant une nourriture plus abondante mais moins digeste. La taille du troupeau
augmente spontanement des la venue des pluies et decroit progressivement au
long de la saison seche lorsque les reserves de nourriture s’amenuisent. Differentes
hypotheses quant aux aggregations saisonnieres sont consideries y compris celles
relatives au comportement alimentaire, social, reproducteur et defensif. I1 n’y a
pas encore d’evidence permettant de conclure que les recentes modifications des
deplacements sont dues a une pression humaine croissante, une modification des
chutes de pluies ou de la disponibilite de nourriture.

Introduction
Wherever they occur, elephants greatly affect the structure of vegetation and
perhaps animal communities (Laws, 1970; Thomson, 1975). However, there is
229
230 D. Western and W. K. Lindsay

some debate about whether their impact on the landscape occurs naturally, or
whether it also reflects human disturbance within the ecosystem. The debate, is
difficult to resolve because so little information exists on undisturbed elephant
populations. Most studies have looked at elephants compressed into sanctuaries
in the last few decades, or subjected to heavy hunting or cropping, both of which
cause unusually large aggregations and woodland depletion (Laws, Parker &
Johnstone, 1975; Barnes, 1980). It is questionable whether an elephant’s natural
ecology and behaviour can be inferred from populations in which densities,
ranging and foraging patterns, and herd dynamics, have been greatly changed by
human disturbance.
The Amboseli ecosystem in southern Kenya is one of the few areas studied
so far where elephants have not been greatly disturbed by man (Western, 1973,
1975). The animal and plant communities in Amboseli have been monitored for
the last 15 years (Western, 1973, 1982) and we draw on this information to look
at the seasonal ranging patterns, habitat use and herd dynamics of this natural
elephant population. We also consider recent changes in human activity which
might be altering the ecology of Amboseli’s elephants and discuss the results in
relation to other populations.

The Amboseli ecosystem

Amboseli refers to a Pleistocene lake basin lying at the foot of Kilimanjaro along
the Kenya-Tanzania border (Fig. 1). The ecosystem, described in detail elsewhere
(Western, 1973, 1975; Western & van Praet, 1973), need only be summarized
briefly here.
The rainfall, which averages 350 mm annually, falls in two seasons, the short
rains of November and December, and the long rains of March to May. The
vegetation is sparse, bushed-grassland (Pratt, Greenway & Gwynne, 1966) except
in the Amboseli basin, where there are extensive alkaline grasslands, woodlands,
and swamps (Western & Sindiyo, 1972). Until the 1940s water was seasonally
restricted to the basin’s swamps but subsequently, boreholes provided the tra-
ditional human occupants of Amboseli, the pastoral Maasai, permanent water
sources in surrounding areas. Elephants, other large ungulates, and Maasai and
their livestock, migrate seasonally between the bushed grasslands and the basin.
The wet-season range of most ungulates covers more than 5000 km2 whereas
their dry-season range occupies approximately 600 km2 in and around the basin
(Western, 1975). In 1977,388 km2of the basin was gazetted a national park. Else-
where the ecosystem was divided into group ranche+i.e. ranches under collective
Maasai ownership. Except for their exclusion from the national park, Maasai
seasonal movements have altered little. The only sizeable arable farmlands occur
over 20 miles away, along the NW and SE flanks of Kilimanjaro. The north-
ern slope remains uncultivated and elephants continue to migrate from the
Kilimanjaro forest to the basin. Small agricultural settlements have sprung up
east of the basin in the last few years, but have yet to displace elephants entirely
from the small swamps found there.
Elephants have occurred in Amboseli at least since the beginning of the 1900s
(P. Ole Taiko, pers. comm.). In recent years their numbers have increased from
approximately 480 in 1975 to 580 in 1980 (Moss, 1977, 1981). All individuals
have been identified and births, deaths and various social interactions have been
 Seasonal elephant-herd dynamics 23 1

monitored since 1972 (Moss, 1977, 1981, 1983). The social structure of the
Amboseli elephants appears similar to that of other elephant populations (e.g.
Douglas-Hamilton, 1972); females and juveniles occur in stable family groups
whereas bachelor bulls are found in more variable, all-male groups, and associate
temporarily with cow/calf herds (Moss, 198 1 ; Poole & Moss, 198I ; Moss & Poole,
1983).

Methods
We have used three sources of information to estimate the numbers and distri-
bution of elephants in the Amboseli ecosystem; the first a series of sample counts
over the entire ecosystem; the second a series of total counts within the Amboseli
basin; and the third, radio tracking of individual elephants. The sample counts
span the period 1973-1976, include thirteen flights over an area of 8500 km2
and cover the entire dispersal zone of the migratory ungulates (Western, 1975).
The area was divided into 5 x 5-km grids and flown at least 4 times each year
to coincide with wet and dry seasons; each flight line was 5 km apart. Observers
counted (or photographed and later counted) all animals falling within a 200 m
strip, and so sampled 8% of the study area; the aircraft was flown at 160 km h-I
and at a height of 90 m above ground. The counting technique has been detailed
elsewhere (Western, 1975).
The total aerial counts were flown monthly between 1975 and 1979 and
covered 600 km2 of the Amboseli basin. These counts were conducted in the same
way as the sample counts, except that flight lines were never more than 1 km
apart, even over open country, and far less over close cover. The location and
size of each herd was recorded on a vegetation map of the basin. A ‘herd’ was
arbitrarily defined as any aggregation showing some cohesion and separated from
other individuals or groups by a clear break in distribution (a distance greater than
the diameter of the herd). Bull herds were distinguished from cow/calf groups in
all counts starting in early 1976. Of the sixty counts in the series, fifty-three were
classified by gender. The counts, which commenced at 10.00 hours, took 60 to
90 min to complete.
Ranging patterns of some individuals were monitored by radio tracking. Six
females radio-collared between 1974 and 1979 were relocated on 275 occasions.
On every occasion observers noted the location, group composition, and activity
of each herd associated with the radio-collared female.
Rainfall was recorded at 01 Tukai, the national park’s headquarters, located
within the central basin. ‘Seasons’, as used in some of our analyses, were defined
by monthly rainfall total as follows: short rains-November-December; short
dry-January-March; long rains-March-May; early long dry-May-
June; mid long dry-July-August; late long dry-September-October. They were
adjusted slightly according to rainfall patterns in individual years.
Herb layer biomass was measured monthly at thirteen sites. The plots, each
300 m2 in area, were distributed among the five principal Amboseli habitats.
Herb layer biomass was estimated from grass height and cover according to a
relationship derived from clipped quadrats:
log,,(dry mass)= 1.02 loglo(heightxcover)-0~38(r2=0-89;n=42),
where mean height is measured in cm and ground cover as a percentage. The
232 D. Western and W. K. Lindsay

nutritional quality of herb layer vegetation (evaluated by crude protein content)

was periodically estimated from samples clipped at each plot.
We have not considered the contribution of woody vegetation to food avail-
ability in this study since most of the Amboseli basin habitats are sparsely wooded,
with the exception of the small patches of dense swamp edge. Browsing was at
all times less common than grazing (Lindsay, 1982; Poole, 1982), even in the dry
season.

Results
Annual range
The absence of adjacent populations in most directions simplifies our assessment
of how far Amboseli elephants range. To the north, elephants can move
unchecked until the settled agricultural areas north of the Mombasa/Nairobi
road, 80 km away. However, numerous reconnaissance flights and aerial counts
showed that animals from Amboseli seldom, if ever, reach this far north (Fig.
1). Westwards, there is no settlement within 150 km, except for an isolated
pocket around Namanga, 60 km away. Regular flights by one of us (D.W.) during
the late 1960s and 1970s, failed to locate any Amboseli elephants west of
Namanga. The nearest elephant population occurs 100 km beyond Namanga on
the Nguruman Escarpment, separated by an arid stretch of the Great Rift Valley.
Historical evidence and aerial counts (Fig. 1) suggest that although Amboseli
elephants had an unrestricted range, they seldom moved more than 50 km from
the basin.
The Kilimanjaro elephant population inhabits the forest tract along the
mountain’s northern slope (Afolayan, 1975). These animals periodically enter the
basin, usually during the rains (C. Moss, pers. comm.). All elephants in the basin
population have been individually identified (Moss, 1981, 1983) and can be
distinguished from the Kilimanjaro elephants, which are more timid and differ
in appearance, having smaller ears, more slender, skewed tusks, and a slighter
build (C. Moss, pers. comm.). There is no evidence from aerial or radio-tracking
records that the basin elephants use the Kilimanjaro forest.
To the east, another elephant population occurs in Tsavo West National Park
where Cobb (1976) documented a wet-season migration corresponding to that
in Amboseli. During the rains the Tsavo population extends westwards and
overlaps slightly the eastern range of the Amboseli population (Fig. 1). The
two populations can be distinguished by their reaction to low-flying aircraft. In
response to extremely heavy poaching during the early 1970s (Cobb, 1980), the
Tsavo elephants react strongly to engine noise by bunching tightly and fleeing,
whereas the Amboseli elephants, less poached and more conditioned to the
continuous tourist traffic of Amboseli, show far less response. Based on such
reactions, Amboseli elephants have rarely been located east of the Loitokitok road
(Fig. 1).
Data from the radio-collared elephants demonstrate a similar annual range
(Fig. 1). The area encompassed by 275 observations extends beyond the range
defined by the aerial surveys in only two locations. The total range of the
population based on aerial surveys is 3042 km2, the range based on radio-collared
animals is 2756 km2 and combined range from both sources is 3588 km2.
 233

- 20s

- 30s

Fig. 1. Total range occupied by the Amboseli elephant population determined from aerial counts
(-) and radio-tracking records (---). Circles represent averaged densities of all Amboseli elephant
sightings from aerial counts between 1973 and 1976, squares represent sightings of adjacent Tsavo
West elephants. Large, medium and small circles and squares represent densities > 4 km-*, 2-4 km-2
and < 2 km-*, respectively; solid circles and squares represent dry-season counts, open circles and
squares represent wet-season counts.

Seasonal migration
By partitioning aerial sample counts of the ecosystem into wet and dry seasons
we can examine the seasonal distribution of elephants over the period 1973-1979.
During the rains herds ranged more widely than during the dry season (Fig. I).
On average, 42% of the population was located more than 10 km from the basin
during wet seasons, compared to 17% during the dry season. Radio-collared
animals were always observed within 15 km of the basin during dry season,
whereas during the rains they were located up to 40 km away. Monthly total
counts of the Amboseli basin revealed regular departures each rains prior to 1977,
and a consistent concentration each dry season (Fig. 2). As suggested previously
for all migratory ungulates (Western, 1975), movement from the basin is appar-
ently triggered by the rains, and a return to it by dwindling water supplies in the
234 D. Western and W . K. Lindsay

Drought

- W el

/-Swamp -l

LGrosslonds
Bush

I 4 100

300 g
z
c
0
200 raal
-
W

n 100
h

I
1976 1977 I978 I979 I980
Fig. 2. Monthly rainfall (histogram) and elephant numbers in the Amboseli basin and the standing
mass of vegetation in the main habitats. The drought, wet and dry years referred to in the text are
indicated here.

bushlands. If this is the case with elephants, their numbers in the basin should
be negatively related to rainfall. To test this hypothesis we calculated linear
regressions of elephant numbers on cumulative rainfall over different periods
ranging from 5 to 90 days preceding each count, using both untransformed and
log-transformed data. A double-log linear model regressing elephant numbers
against rainfall gave the best fit.
The total number of elephants and the number of cows and calves (family-unit
members) in the basin, were most highly correlated with cumulative rainfall over
the preceding 15 days (total numbers; r2= 0.154, P < 0.0 1, n = 60; cow/calf
numbers: r2=0-137, < 0.0 1, n = 53). Bull numbers, however, were most highly
correlated with rainfall over the preceding 50 days r2= 0. I 57, P < 0.0 1, n = 53).
Elephants have become considerably less migratory since late 1977 (Fig. 2).
To test whether the breakdown in migration was due to climatic changes we
partitioned the counts into a pre-November 1977 period (the 'drought years'), a
November 1977-pre-November 1979 period (the 'wet years'), and a post-October
 Seasonal elephant-herd dynamics 235

Table 1. Seasonal mean numbers and densities of elephants km-* in Amboseli Basin habitats

Habitat

No. of Bushed- Open Dense

Season counts grassland Grassland woods woods Swamp Basin

Short rains 10 No. 232.6 4.6 146.2 77.9 33.8 267.4

Den. 0.035 1.227 8.287 1.374 0.668
Short dry 5 No. 125.6 99.0 85.4 87.0 83.2 3744
Den. 0.758 0.716 9.255 3.382 0.936
Long rains II No. 327.3 9.1 92.5 19.5 26.7 172.7
Den. 0.069 0.776 2.074 1.085 0432
Long dry 9 No. 225.7 4.0 854 97.5 72.2 274.3
Early Den. 0.03 1 0.716 10.372 2.935 0.685
Mid 10 No. 214.5 1.6 94.6 964 76.2 285.5
Den. 0.012 0.794 10.255 3.098 0.7 13
Late 8 No. 182.8 5.3 445 73.4 185.0 3 17.2
Den. 0.04 I 0.373 7.809 7.520 0.793
Total No. counts 53
Area of
habitat types
(km? +2000 130.6 1 19.2 9.4 24.6 400.2

1979 period (‘dry years’) (Fig. 2). We found that total, cow/calf and bull numbers
were more strongly related to rainfall during the drought years than over the whole
study period (total numbers: rz= 0.450, P < 0.00 1, n = 26; cow/calf numbers:
r2=0-539, P < O - O O l , n= 19; bull numbers: r2=0.392, P<O.Ol, n= 19). After
October 1977, linear relationships declined to insignificance for all heads. We
consider why migrations have changed in recent years in the discussion.

Habitat selection
In the course of their daily foraging in and around the central basin, some
elephants use the woodland and bushland habitats south of the basin each night.
By mid-morning, most animals are feeding within the basin where they remain
until late evening. Thus, the aerial total counts in the basin, which are made
between 10.00 and 11.30 hours can demonstrate only the daytime trends in
habitat use within the basin. However, the use made of all habitats can be
approximated by subtraction (see below).
In Table 1 we present the mean density and numbers of elephants in the basin’s
habitats for each season. A xz analysis shows that there was non-random use of
habitats in all seasons (x2 values by season: short rains=7405; short dry=
3276; long rains = 1307; early long dry = 10,8 14; mid long dry = 1 1,509; late long
dry = 13,059; P < 0.00 1). Use changed progressively between seasons. The low-
production alkaline grasslands were seldom used by elephants and we have
excluded them from later analyses. The open acacia woodlands were used most
236 D. Western and W. K. Lindsay

80 Bush

60

40
C
._ Open
c
-
0 wood
2 2c I
n
+
0 Long
$- rains
0

2c

40
Fig. 3. Use of habitats by the Amboseli elephants during the long rains and long dry season based
on fifty-three aerial counts between 1976 and 1980.

heavily early in the dry season, the denser woodlands mid to late dry season, and
the swamps and swamp edges at the end of each dry season (Table 1).
We can infer the use made of bushlands by subtracting the number in the
basin from the total number in the population, which fluctuated around 500
during the study period (C. Moss, pers. comm.). We have shown the diurnal use
of all habitats by expressing the percentage of the total population found in each
of them. The results (Fig. 3) are summarized for the long rains and late long-dry
seasons only, when differences in habitat condition were most extreme. The short
rains and short dry seasons showed a similar, though less pronounced, trend in
habitat use.
Elephants used the bushed-grasslands most during the rains and concentrated
within the basin during the dry season. Some family units, known as ‘peripheral
groups’ (Moss, 1981; Moss & Poole, 1983) continued to use the bushlands during
the dry season. During the rains, elephants within the basin used the open
woodlands most, then progressively moved to the denser woodlands and swamps
as the dry seasons progressed (Fig. 3, Table 1). By the end of the dry seasons over
one-third of the population was concentrated in the swamps, which occupy less
than 10% of the basin, and little more than 1% of the ecosystem. Males and
females showed similar movements through the dry season, though males
favoured habitats with more abundant woody vegetation (Lindsay, 1982).
In a test of the relationship between habitat selection by elephants and habitat
forage quality, we assumed that the net digestive value of forage is directly
proportional to its protein content, a relationship widely established for both
ruminant and non-ruminant herbivores (Glover & French, 1957). Our results
show that during the rains, when water imposed no distributional constraint and
all forage was abundant, elephant numbers varied positively with habitat quality
(rs= 1-00,P<0.05,n =4, Spearman’s rank correlation) and inversely with habitat
production (rs= 1.00, P<0.05, n=4). Numbers were highest in the bushed-
 Seasonal elephant-herd dynamics 237

Table 2. Seasonally averaged elephant numbers in the habitats they predominantly use in the
Amboseli ecosystem

Wet season Dry season

Bushed- Open Dense Open Dense

grassland woods woods Swamp Bush woods woods Swamp

No. of elephants 282 I18 47 30 183 45 73 I85

Pasture quality
(O/O crude protein) 18.7 14.0 12.5 7.00 9.1 9.1 10.8 5.8
Pasture mass (g m2) 227 1202 3312 4393 18 72 2010 2723

grasslands and progressively lower in the open woodlands, denser woodlands and
swamps-i.e., along the quality gradient-and the sequence selected during the
dry season (Fig. 3). The nutritional value of all pastures declined substantially by
the end of the dry season, by which time the elephants had moved along the qual-
ity gradient of the denser woodlands and swamps, the only habitats remaining
with a large standing crop of forage. The swamps were more heavily used than
all other habitats, despite having the lowest pasture quality (Table 2).

Herd dynamics
Elephant herd size varied greatly. To what extent does it vary between seasons
and habitats?
Since 1967 large numbers of elephants have been observed gathering together
during the rains (D.W., pers. observation). The largest aggregations exceeded 400
animals-more than 80% of the entire population. During the dry season, herd
size dwindled and groups larger than twenty animals were seldom observed during
the severe drought of the mid-1970s. To test whether these observations were
statistically significant, we classified the aerial total counts into one of six seasons,
as in the analysis of habitat selection (Table 1). In these analyses we have included
only counts with more than 2 10 elephants-the largest aggregations seen during
the counts.
In a one-way analysis of variance, arithmetic mean group size (MGS) varied
significantly between seasons ( F = 3.9709, P < 0.0 I , n = 40). The largest MGS were
observed during the rains, the smallest during the long dry seasons. A potential
bias in our data could affect the analysis of seasonality in MGS. The large
groups, which commonly form outside the basin during the rains (D.W., personal
observation), are inevitably excluded from the aerial surveys within it. Because
such a bias reduces the seasonal difference in MGS, our results give a conservative
picture of seasonal variation.
To test whether MGS varied systematically with rainfall, we examined all
counts of more than 2 10 animals and regressed MGS against cumulative rainfall
over the preceding 5-90 days. The strongest relationships for all herds and
cow/calf herds were found in a linear regression of MGS against cumulative rain-
fall over the previous 30 days (all groups r2=0.150,P<0.05, n=40,
238 D. Western and W . K . Lindsay

cow/calf groups: rz = 0.2 15, P < 0.0 I , n = 40). The highest significance value for
bull groups was found when MGS was regressed against cumulative rainfall over
the previous 10 days (r2=0*193,P<O.OOl, n=53). MGS for all groups was more
strongly related to rainfall in the wet years and dry years than over the whole
study period (wet: r2=0.270,P<0.05, n = 16; dry: r2=0-582,P<O.Ol, n = 12). No
significant relationship was found for the drought years because so few animals
stayed within the basin each rains. Rainfall accounted for more of the variance in
herd size during the dry years (58%) than during the wet years (27%), suggesting
that herd size responded more rapidly to seasonal changes during dry than wet
years.
Mean group size also vaned between habitats in a given season, and within
habitats between seasons. There was, however, a great deal of variation, which
we attribute to the continual formation and breakdown of herds as they encoun-
tered each other, mingled, moved on, or changed habitat. Male herds seldom
averaged more than four individuals, whereas between 1971 and 1980 female
herds averaged more than twenty-five. Perhaps due to the small herd size of males,
and a high inter-herd variance, we found no significant relationship between
group size and habitat use. Cow/calf herds did, however, show a tendency to be
larger in some habitats than others. Mean group size also changed progressively
within a habitat from wet to dry season. Though lacking data for bushlands, our
casual observations suggested that during the rains MGS was largest in this habitat
and, within the basin, highest in the open woodlands and lowest in the closed
woodlands and swamps. By late dry season herds in the bushed-grasslands were
few and small (D.W., personal observation), and within the basin, progressively
larger in the open woodlands and swamps. Throughout the dry season there was,
therefore, a progressive shift in the location of largest MGS from open woodland
to swamps. There was a significant correlation between mean numbers and mean
group size with respect to habitats (Spearman’s rank correlation, r,= 0309,
P<O.O1, ~ = 2 9 ) .

Discussion
Seasonal migration and range
The seasonal migrations and home range of Amboseli’s elephants were little
affected by human pressures until 1977. Both the timing and extent of elephant
migrations were similar to those of other migratory ungulates in the ecosystem,
including zebra, wildebeest, Coke’s hartebeest, and Maasai livestock-cattle,
sheep and goat. The reasons are probably similar in all cases (Western, 1975):
water is available during the dry season only within the basin, thus confining most
water-dependent species to within a 15 km foraging radius of the swamps. During
the rains, water is widely distributed and enables the migratory ungulates to select
any pasture. In the bushlands ungulate movements are dictated by localized
storms and green forage. Here, wet-season food availability greatly exceeds the
requirements of migrants, whose numbers are limited by food supplies in the
dry-season range (Western, 1973). The seasonal surplus gives elephants, and other
migrants, little reason to move great distances from the basin into the bushlands.
The farthest elephants were little more than 50 km away (Fig. 1). Migration often
starts within a few hours of the first rains, long before vegetation has responded to
 Seasonal elephant-herd dynamics 239

it, further supporting the hypothesis that water restricts the movement of
migrants during the dry season.
There is good evidence that dwindling water supplies force the return of
migrants to the basin before food supplies are depleted in the bushlands (Western,
1975). Support for this argument is given by the observation that boreholes, sunk
for livestock during the 1940s and 1950s, allowed them to stay longer in the
bushlands than zebra and wildebeest, which had no access to artificial water
sources. But, elephants return to the basin earlier than other species (Western,
1975). Their earlier return cannot be explained by water availability alone. We
show later that forage requirements also contribute to their return from the
bushland areas.
What accounts for the diminished elephant migrations after 1977? Changes
in both human activity and climate offer plausible explanations. Changes in
human activity include poaching, which increased sharply during the early
1970s, more agricultural settlement and, in 1977, the removal of Maasai and their
livestock from the national park.
The agricultural settlements lie east of the basin, but occupy less than 1% of
the elephant’s dry-season range and cannot explain their lessened migration. The
1977 displacement of Maasai from the basin cannot explain the change either
since, as noted earlier, the pastoralists traditionally migrated to the bushlands at
the same time as other migratory ungulates, including elephants. The bushland
areas used most frequently by elephants are no more settled today than in the
1960s. There remain two other possibilities related to human activity. First,
domestic stock, which formerly made up 65% of the basin biomass (Western,
1975), were excluded from the national in 1977, thus giving extra forage to wild
herbivores. Secondly, during the 1970s elephants were heavily poached in their
wet-season range, but little so in the basin, which therefore provided them a safe
refuge. Neither of the last two explanations can yet be distinguished from another,
based on climate. The wetter years, 1977-1979, may have sufficiently increased
forage production in the basin (Fig. 3) that elephants had no reason to migrate
during the rains. By considering the reasons for habitat choice by elephants it may
be possible to identify specific tests which could distinguish between these three
hypotheses.

Habitat selection
At the onset of the rains, elephants migrate to the bushed-grasslands, then succes-
sively select the open woodlands, denser woodlands, swamp edges and swamps
during the dry seasons (Table 2, Fig. 3). They probably do so for the same reasons
as the other wild and domestic migrants in the ecosystem which show a similar
pattern of habitat selection (Western, 1973). In the course of a season there is
a characteristic grazing succession in which elephant buffalo, zebra, wildebeest
and Thomson’s gazelle succeed each other through the habitat sequence. Vesey-
Fitzgerald (1965) and Bell (1971) described a similar grazing succession for
Serengeti and Rukwa, respectively, postulating that large herbivores using a
habitat first open the pasture to smaller migrants by removing coarser forage and
promoting secondary growth. McNaughton (1976) showed that in the Serengeti
wildebeest grazing promotes secondary grass production and attracts Thomson’s
240 D. Western and W. K. Lindsay

gazelle. However, although this hypothesis may explain habitat selection by the
following migrants, it fails to explain the selection by initiating species, such as
elephants.
There is a more parsimonious explanation for the grazing succession in
Amboseli. From the results of pasture analysis presented earlier, we can conclude
that all species select forage with the highest digestible energy whenever possible,
but are forced to use successively poorer pastures as food supplies dwindle. The
biggest species, because of their greater food requirements, abandon favoured
habitats before smaller species. This hypothesis is consistent with optimal foraging
theory (MacArthur & Pianka, 1966; Schoener, 1971), which states that an indi-
vidual will select food items in a manner which will maximize its rate of energy
assimilation. Although search time and handling costs can constrain optimality
(Charnov, 1976), in its simplest form the theory predicts that the highest-quality
foods should be consumed whenever encountered, and that as they become rarer,
diet should expand progressively to include less valuable foods (Townsend &
Hughes, 198 1).
Is it possible to distinguish, using pasture analysis, whether the post- 1977
reduction in elephant range reflects a normal foraging response to pasture changes
produced by wetter years, or a response to human activities?
During the worst drought years, peak biomass in the open woodlands (< 15
g m-2) was less than in the bushlands (>30 g m-2) (Fig. 2), which had higher
quality pastures (Table 2). Elephants, therefore obtain food higher in both quality
and quantity by migrating to bushlands during the rains. During the wet years,
biomass in the open woodlands increased greatly, often to levels in excess of
600 g m-2-far higher than in the bushlands (Fig. 2). The additional mass and
its relatively high quality, was perhaps sufficient to dissuade elephants from
migrating. The persistance of a large standing mass of vegetation in the woodlands
and swamps following the return of drier years can be attributed to the reserves
built up during the wetter years, and to the removal of livestock from the national
park in June 1977.
We cannot, therefore, presently tell whether man or climate has caused a
reduction in elephant range. Nevertheless, we can propose a test that will do
so if the surplus forage in the basin woodlands is reduced to low levels once
again. If foraging selection is responsible, elephants will resume their wet-season
migrations to bushland areas. If poaching, or some other human pressure is
responsible, elephants will not resume full seasonal migrations.

Herd-size functions
The explanation for changes in herd size between seasons and habitats can be
examined from two alternative perspectives-why herds form, and why they
fragment. We first consider why herds fragment, since we can make certain
deductions from the data already presented.
During the rains, elephants aggregate into large herds, which gradually break
up during the dry season. Because MGS differs between seasons and habitats,
we can dismiss the simple explanation that the herds form and break down at
random. A refinement of this hypothesis holds that herds randomly meet and
 Seasonal elephant-herd dynamics 24 1

aggregate in direct proportion to their use of a habitat, such that the preferred
habitats support the highest densities, and largest group sizes (Rubenstein, 1978).
The correlation previously shown between MGS and numbers of elephants in a
habitat adds some validity to this hypothesis. It fails, however, to account for the
aggregations of several hundred elephants which form immediately the rains
begin, and before there has been substantial vegetation response. These herds
usually form when elephants migrate to the bushlands, where their density is low
and their movement large (Fig. 1) -the converse of predictions derived from the
random-aggregation hypothesis. There is evidence from behavioural studies
(Douglas-Hamilton, 1972; Moss, 198 1 ; Moss & Poole, 1983) that herd fusion and
fission, far from being random, follows predictable patterns of association between
family units. Families tend to associate according to a hierarchial clustering into
bond groups and clans, and breakdown in the reverse order. The aggregation
patterns are related to spatial proximity of herds, as shown by Douglas-Hamilton
(1972) in Lake Manyara.
A plausible explanation of herd breakdown is that families which can benefit
by association when forage is abundant (see below), must divide to maintain
foraging efficiency when it is scarce. In this explanation the costs of large-herd
association increase faster through the dry season than the benefits of small-herd
association, which include less feeding interference and greater foraging efficiency
(Alexander, 1974). By moving onto the more abundant forage of the denser
woodlands and swamps, a herd might delay division, We might anticipate that
closely related individuals have most to gain through inclusive fitness by
maintaining group cohesion (Hamilton, 197 1). For any given habitat considerable
variation in herd size might result from such a compromise between feeding and
social benefits and would add greatly to herd-size variation within habitats.
Allowing for herd size, we would expect that herds comprising closely related
animals would use the abundant pastures more often than herds of less related
individuals.
If herds do divide in response to declining food resources during the dry season,
why do they aggregate during the rains? There are a number of possible, non-
exclusive explanations: (i) anti-predation (Laws et al., 1975); (ii) feeding facili-
tation, kin-group resource acquisition (Wrangham, 1980), or perhaps to exchange
information (Ward & Zahavi, 1973); (iii) social benefits, such as extended social
networds and early establishment of dominance hierarchies (Moss, 198 I); and (iv)
reproductive benefits, which include improved mating opportunities (Cox & Le
Boeuf, 1977).
It is difficult to isolate each explanation and quite likely that aggregations
benefit individuals in a number of ways, although certain factors seem more likely
than others. We now consider their relative importance.
There is evidence that large aggregations of elephants in many localities form
in response to heavy predation and disturbances by man. In Kabalega, Tsavo and
Serengeti, elephants show extreme flight response to people and vehicles, and
females form a protective circle around the young (Laws et al., 1975). In Amboseli
one of us (D.W.) observed large seasonal aggregations in the late 1960s, prior to
any substantial poaching, which showed little flight response or anti-predator
bunching, and which were generally loose herd associations, rather than tight
formations. Moreover, the herds formed and divided seasonally, where in areas
242 D. Western and W . K. Lindsay

of intensive hunting, the large herds persist year-round (I. S. C. Parker, pers.
comm.)
There is similarly no strong evidence that herd formation results in foraging
benefits, either through facilitation, or through information exchange. Indeed it
is not uncommon to see spontaneous herd fusions immediately before the rains,
before any possible benefits from feeding aggregations or information exchange.
Moss (1 98 1) has suggested that individual elephants derive various social
benefits in an aggregation, including the re-establishment of kin and non-kin
bonds. Only detailed studies can demonstrate the nature and extent of such
benefits. Until then the hypothesis remains plausible, but unsubstantiated.
Circumstantial evidence suggests that seasonal aggregations of elephants in
Amboseli constitute breeding herds which could benefit oestrous females. Oestrus
is seasonal, with peaks during and shortly after the rains when the large herds are
formed (C.J. Moss, pers. comm.). There is also direct evidence that dominant
males come into musth during and shortly after the rains, whereas the subordinate
males are in musth during the dry seasons when few females are in oestrus (Poole
& Moss, 1981; Poole, 1982). The large aggregations could therefore act as
competitive arenas in which a female increases her chance of securing a dominant
male. Oestrous females might also avoid the intense harassment of subordinate
males by the attendance of dominant males (Poole, 1982). This is similar to the
strategy found in female aggregations of elephant seals (Cox & Le Boeuf, 1977),
and might explain why elephant herds form and break down faster during dry
than wet years: during dry years oestrus is more seasonally synchronized
(C.J. Moss, pers. comm.) and would considerably restrict the period of breeding
benefits.

Comparison with other elephant populations

Few elephant populations elsewhere in Africa have been studied in as much detail
as Amboseli’s. We can, however, make some general comparisons of annual
home range, density, seasonal movements, habitat selection, and herd dynamics
between elephant populations.
The 500 elephants in the 3500-km2 total range of the Amboseli population,
occur at an overall density of 0.14 elephants km-2. Densities in the dry-season
concentration area of the basin were 0 . 4 4 9 km-2, and in specific habitats, 10
km-2 (Table 1).
Densities of elephants in other populations, reviewed by Laws et al. (1975)
and Eltringham (1 977), range from 0.18 km-2 in Serengeti (Watson & Bell, 1969),
0.88 km-2 in Tsavo National Park, 3.80 km-2 in North Bunyoro, Uganda (Laws
et al., 1975), to 5.00 km-2 in Lake Manyara National Park (Douglas-Hamilton,
1972) and generally increases with annual rainfall. Radio-tracking studies in
Tsavo East National Park (Leuthold & Sale, 1973) showed annual ranges for
individual elephants of some 1600 km2.
Seasonal migrations and habitat selection have been reported for many ele-
phant populations (Laws et al., 1975; Eltringham, 1977; Caughley & Goddard,
1975). The movements appear to coincide with changes in food and water
availability. The Tsavo population (Cobb, 1976; Leuthold, 1977) covers a similar-
sized range, and shows similar seasonal movements to those of Amboseli’s
 Seasonal elephant- herd dynamics 243

elephants. There is some evidence from radio-tracking records of Tsavo elephants

(Leuthods, 1977) that, following their wet-season movements, cow/calf groups
return to the same area each dry season, as in Amboseli’s elephants (Moss, 1977,
I98 1 ; Moss & Poole, 1983).
In areas of poaching or conflict with human populations, the overall range of
populations become restricted and large defensive aggregations of elephants are
soon evident (Laws et al., 1975). Range compression frequently leads to extensive
woodland reduction (Laws, 1970). We believe that, until the mid- to late-1970s
the Amboseli elephant population was little disturbed by human activity and that
annual patterns of migration, habitat selection and herd dynamics were natural,
seasonal events. The impact on habitats was still relatively minor (Western &
van Praet, 1973). Changes in movements since then may reflect responses to
poaching, in common with many other ecosystems (Laws, 1970), or may be
responses to changing forage availability. Until food becomes scarce, it will be
difficult to tell which factor is responsible.
The evidence from Amboseli and elsewhere suggests to us that elephants under
threat from man may exhibit similar, but exaggerated herd responses to those
found in natural populations. If so, we will need to define behavioural criteria,
and the magnitude of herd response to seasonal change, that distinguish stressed
and non-stressed populations.

Acknowledgments
We thank David Maitumo, Field Assistant to the project, and the various people
who helped with aerial counts, including Harvey Croze, Steve Cobb, Chris
Hillman, Patrick Hamilton, Chris Gakahu and Nick Georgiadis. Joe Kioko and
Bob Oguya, Wardens of Amboseli, were helpful in numerous ways. Cynthia Moss
and Joyce Poole have added greatly to our work. This study constitutes part of
David Western’s field studies funded by the New York Zoological Society.

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