E. Afr. Wildl. J.

, 1975, Volume 13, pages 265-286

Water availability and its influence on the structure

and dynamics of a savannah large mammal community

D A VI D WEST E R N New York Zoological Society

Summary
The Amboseli ecosystem is in a semi-arid savannah environment in which water
availability is highly seasonal, a feature which has an important bearing on the struc-
ture and efficiency of the large mammal community.
Data are presented to show that the seasonal movements, a wet season dispersal and
dry season concentration of water-dependent species (obligate drinkers) can be related
to the seasonality of rainfall and water availability, but that similar patterns do not
prevail in the water-independent species.
Evidence suggests that water availability is a crucial parameter in calculating the
carrying capacity of a range, and that the duration of ephemeral supplies, which
reflect rainfall seasonality, evapotranspiration, and soil water storage capacity, is
equally important. A simplified model is presented which calculates the carrying
capacity of water-bound communities-taking into account these two limiting factors.
There appears to be a physiological barrier to a heavy utilization of the low water
content grasses beyond reach of the water-dependent species, since most water-inde-
pendent species select high water content browse in order to maintain a positive water
balance. Significantly, while the water-independent community is almost exclusively
composed of browsers, most water-dependent species are grazers. The biomass
density of large mammals beyond 15 km from water, a radius which encloses 99.5 % of
the biomass of water-dependent species, is extremely low compared to areas within
10 km of water.
As a consequence of the physiological barrier to a heavy utilization of low water
content forage in water-deficient areas, the food chain efficiencies are appreciably
lower than those with permanent water sources. Livestock development programmes
have long appreciated this constraint and employed water development schemes to
increase range-carrying capacity.
Seasonality in the movement patterns of Amboseli is compared to other large
mammal communities and management implications are discussed.
Present address: Zoology Department, University of Nairobi, Box 30197, Nairobi, Kenya.

265
266 David Westem

Introduction
Seasonal changes in the distribution of large mammals in African savannahs vary in
extent from the extensive migrations of the Serengeti wildebeest, which cover an eco-
system of some 25 000 km2 (Norton-Griffiths, 1973) to localized movements within
confined areas, such as Nairobi National Park, covering 114 km2 (Foster & Coe,
1968). Within this range three categories are widely recognized, based on the patterns
of seasonal movement. These are migratory, resident and what can be coined dispersal
systems. that is wet season dispersal, dry season concentration populations. Seasonal
movements have been described for a number of ecosystems in relation to rainfall,
habitats and grazing conditions (Talbot & Talbot, 1963; Lamprey, 1964; Vesey-
Fitzgerald, 1965; Bell, 1971). It has not been obvious in most cases, however,
whether these movements are in relation to water availability, pasture conditions, or
~ o m ecombination of the two. It would seem probable that most populations will
select the best pastures available to them within the areas accessible, but in many cases
accessibility appears limited, particularly in species dependent on free water, a com-
modity limited in distribution during the dry season.
Lamprey ( I 964), noting the confined distribution of the Tarangire National Park
herbivores in the dry season, commented ‘The extent and distribution of permanent
water in the dry season throughout the Masai area of Tanganyika is probably the most
important limiting factor in the number and distribution of game animals in the
savanna of East Africa and the carrying capacity of the country as a whole is closely
related to the carrying capacity of the land within “cruising range” of the dry season
water supplies.’
If this hypothesis is correct it implies that the utilization of the savannah environ-
ments cannot be predicted simply on the basis of primary production, but that there is
a physiological barrier that limits food chain efficiency. It also implies that both the
distribution of permanent water and the duration of ephemeral sources are necessary
in calculating carrying capacity, and that the more restricted are these two parameters,
the lower will be the total biomass supported within the ecosystem. Studies on the
adaptations of large mammals to arid environments (Schmidt-Nielsen, 1964; Taylor,
1970; Macfarlane & Howard, 1972) suggest that the problems of maintaining a positive
water balance in water-deficient environments are sufficient to suspect that a physio-
logical barrier to a high food chain efficiency may well exist.
The hypothesis seems relevant to the Amboseli dispersal system and data from a
long term ecological study of the area (Western, 1973) will be used to describe the
5easonal movements and biomass distribution of the large mammal community in
response to water availability, and to compare with both migratory and resident
populations elsewhere. In this study emphasis is placed on the gross characteristics of
the large mammal community rather than the response of individual species, although
i t will be shown that autecological data is valuable and complementary where sufficient
detail can be obtained.

The study area and methods

The Amboseli ecosystem covers nearly 3000 km2 of eastern Kajiado District in
Southern Kenya, Fig. 5, of which some 600 km2 is a dry season concentration area in
and around the Amboseli basin, Fig. 1. The dry season range has been described by
Western & Sindiyo (19721, Pennycuick & Western (1972) and in more detail by
Seasonal water availability 267
268 D a d Western

Western (1973). Most of the area receives less than 350 mm of rain annually and the
vegetation is typically Acacia and Comniipliora scattered woodland and bushland,
classified by Pratt, Greenway & Gwynne (1966) in Ecological Zone V of their East
African rangeland classification.
Permanent water is restricted to the vicinity of the dried-out Pleistocene lake bed of
the Amboseli basin, the discharge zone of much of the water from the northern slopes
of Kilimanjaro. Elsewhere water is absent in the dry season except for a number of
boreholes which have provided an important source to livestock, the supplies of which
are not usually available to wildlife.
The extent of the ecosystem is more or less defined by the annual range covered by
the dispersal populations of the Amboseli basin and tends to overlap with adjacent
dispersal communities, The area defined for this study (2400 km2) is slightly smaller
than the total annual range of the dispersal community, but the limits of the area are
drawn so that permanent water supplies outside the ecosystem will not be closer than
those within it, Fig. 5. Furthermore, the area south of the basin has been omitted from
this analysis since much of i t increases rapidly in elevation to the forest zone of
Kilimanjaro and covers an environment with somewhat less extreme aridity problems
to those north of the basin.
For the purpose of the present paper, animal numbers and movements are mostly
considered in relation to the dry season concentration area. The concentration area in
the dry season is sufficiently small that a relatively high sampling fraction of the popu-
lation is possible, whereas during the rains most species are spread over an area too
large to permit intensive counts at frequent intervals. Intensive counts of the dry
season concentration area are therefore used in this study, and where necessary are
supplemented with less intensive counts over the entire ecosystem. The extensive
counts are based on systematic strip counts from an aircraft using similar methods to
those mentioned below, but using a four-seater Cessna 180 and three observers, one
making an inventory of environmental conditions, the other two counting animals
between streamers demarcating strips each 200 m wide on the ground.
On the systematic count transects were flown at 5-km intervals, and the lengths were
also subdivided into 5-km units so that the entire ecosystem is covered by a 5-km
square grid, Fig. 5.
Distance from water profiles on the extensive survey 'using 5-km square grids was
determined in 5-km categories. Intervals of 5-km radii were drawn around each water
point and each grid square was assigned the distance from water category that covered
most of the area within it.
A series of aerial and ground counts were established in the dry season concentration
area of the Amboseli ecosystem in December 1967 and formed the basis of an eco-
logical monitoring programme which is still continuing. A description of the study area
and the aerial counting techniques have been published (Western & Sindiyo, 1972;
Pennycuick & Western, 1972; Western, 1973) and the results form the basis of the
animal counts presented in this paper.
Animal distributions in relation to water are based on randomized ground counts
using a sampling system described by Western ( I 973). Before each count, all the water
sources in the study area were located, usually from an aircraft, and profiles were
drawn at 2-km intervals around each source. Using this technique, it was possible to
determine the distance of any point along the transects relative to the nearest water
source.
 M o n t h 01 count

Spccte, Apr.!h') July:69 Sept.16'J Oct.ih9 Dec.16Y Jan.170 Feh./70 Mar./70 Apr./70 May/70 Juiic;'i(I July!70

Zehra Y 2369 3720 305x 3007 285 I5 0 2034 2938 70 63X 3X3X 4299
SE 493 956 75x 55 I 94 I40 372 574 61 161 X50 695
\\I i64770 XXhXJX 729027 7 I 6XhX 67944 35760 484906 700411) I 66x8 I52090 9 I4976 I o 2 4 n I~
Wlldehec*l Y 2166 3071 2605 2759 I286 614 I840 4603 973 2212 l2XX 4270
\I 323 770 503 633 313 210 346 I ooo I27 552 440 933
\\I 35X'JOh ?OX865 431649 457166 2 I3090 I01740 ioJxxn 762717 161226 366528 544x21 707539
Kongoni Y 7x I04 I6 I I7 I13 X 0 19x 115 25 7X IY3
St 29 67 I0 68 72 7 0 X0 62 IX 36 77
\It I0623 14165 2179 15935 I 539 I 1090 0 26968 I5663 3405 I0624 262~7
T garelk Y 83 0 0 I36 I76 4nx I72 54 I48 75 9X I 86
st 32 0 0 76 62 379 76 22 47 47 39 60
\! I 1619 0 0 2652 3432 Y5 I 6 3354 I053 xnh I463 IYII 3627
G . ga,cllc Y I496 4XX 47') 75 I 90 I 1221 6X I 724 61 5 713 I60 X38
FI 292 20 I 117 226 263 391 258 257 19') 147 40 I85
\\ I 74650 24351 23'JO? 37475 449ho 6092~ 33Y81 36128 3 1 ~ 7 35579 7984 41816
IIlll)ill;l Y 719 0 79 I32 I55 1512 276 297 827 In 3 I43 600
St 29') 0 71 69 88 800 16X 204 329 I05 125 262
\\ I 32643 J 3587 5993 7037 6x645 12530 I3484 3XOOO x30x h492 27240
Or?.\ Y 260 0 26 1x1 I93 197 0 0 Ill 21 0 4
SL 143 0 17 I33 I23 IXJ 0 0 42 I2 0 4
\\ t 43420 0 4342 30227 32231 32899 0 0 1x537 3507 0 668
F-land Y 0 0 0 (I 59 0 293 IX 0 71 0 0
St tl 0 0 0 51 0 238 16 0 62 0 0
UI 0 0 0 0 2 I429 0 io64in 6538 0 257x7 0 0
i~uiraio Y I 107 21 612 577 71 95 707 724 1014 43x 33 49
SF 942 I0 403 337 41 xx 45') 460 797 244 20 in
\\I 552836 104x7 3206l 5 2XXl54 35457 47443 353056 361 566 506392 21x737 I64x0 24471
tlcphnnt Y 410 462 263 249 I n4 95 60 21 I 4 350 II 9 212
St 14X 24 I I5 2 I02 x4 XY 38 I00 4 204 75 II 9
UI I23tKJO~I I3X6000 7x9000 747000 5520M) 2XSWo I ROOOO 633000 Ixxw Io5i)ooo 3S7"Wl 636000
( i iralle Y I IY 33 I6 423 4 30 I46 99 YI 79 I39 191
St 41 25 x 336 4 77 55 35 43 35 75 I06
\II Y I 844 25469 I2349 32647 I 30x7 30100 I I2683 76408 70234 60972 imnn 147416 F
Ca1tlc Y 8735 22267 19201 I3023 I826 4096 II 9 6 19201 4730 I408 9498 5027 B
SE 3493 6244 5517 2430 Ill6 2316 891 6058 2196 931 3566 I67R 8
\It 2419595 541 X679 5318677 1607371 505802 I I34592 552892 53 I8677 1310210 390016 2630946 1392479
Shoat Y 3030 7x83 5953 54x7 482 71 24 I 5x49 62 0 352 1192
9
St 1431 1916 2764 I652 420 66 17Y 1731 54 0 306 858
Ut 48177 I 25340 94653 X7243 7664 I129 3832 Y2Y99 986 0 5597 I8953
Ostrich Y I37 I37 I74 I2n 50 39 24 I 63 181 321 131 I02
SE 3n 60 95 53 I5 20 68 44 67 117 37 36
\\ t I 5550 I 5550 I Y749 1452X 5675 4427 27354 7151 20544 36434 I4869 11577
Donkey Y 86 300 679 49n 42 0 69 445 0 0 209 I02
SE 63 217 249 I62 36 0 53 209 0 0 I34 81
wt 10790 3YOOO XX270 64740 5460 0 8970 57580 o 0 27170 I3260 h,
7.7
m
54 9-2 7-9 6-5 I 5 I 9 -- 8-1 2-2 24 47 4.2 W
270 David Western

Seasonal movements
Data on the numbers and biomass of animals within the dry season study area are
given in Table 1 for the most numerous large mammal species, Thomson’s gazelle size
and upward. Small numbers of other species occur, for example waterbuck, lesser
kudu, bushbuck, but their numbers are low and an accurate census of their numbers
was difficult using large-scale monitoring techniques.
The fluctuations in numbers of the numerically dominant species are shown in
Fig. 2 and comparing these figures with the distribution of rainfall over the period, it
is clear that there are marked seasonal fluctuations that broadly coincide with rainfall
patterns.

Fig. 2. Variation in the numbers of some principal Iarge mammal species against rainfall in the dry
season range of the Amboseli ecosystem. Shoat equals sheep and goat figurescombined. 0,wildebeest;
0 , zebra: r - , Grant’s gazelle:
1 , elephant; .,oryx; ~., ostrich; *, shoat; A,cattle.

Although the total biomass of animals within the dry season study area show a close
relationship to rainfall, it is desirable to account for these changes quantitatively.
Data for calculating biomass figures is based on values given in Table 2.
Rainfall data are derived from the 01 Tukai raingauge which has been maintained
continuously since 1967. Because records are taken from a single rainfall station (others
 Seasonal water availability 211

Table 2. Population weights used in the calculations of biomass

~

Mean
Species weight (kg) Authority*
Zebra-Equus burchelli (Gray) 238
Wildebeest-Connochaetes taurinus (Burchell) 165
Kongoni-Alcelaphus buselaphus (Pallas) 136
Thomson’s gazelle-Gazella thomsonii (Guntter) 20
Grant’s gazelle-Gazella granti (Brooke) 50
Impala-Aepyceros melampus (Lichtenstein) 45
Oryx-Oryx beisa (Ruppell) 167 Ledger (1964)
Eland-Taurotragus oryx (Pallas) 363
Buffalo-Cyncercus cafer (Sparrnann) 500 Bouliere (1965)
Elephant-Loxodonta ufricana (Blurnenbach) 3000
Giraffe-Giraffa camefeopardafis (L.) 772
Cattle-Bos indicus (L.) 277
Sheep-Chis aries (L.)
Goat-Capra hircus (L.) 16
Ostrich-Strutheo camelus (L.) 114
Donkey-Equus asinus (L.) 130 G. M. 0. Maloiy
(1972, pers. comm.)
* All sources unless indicated are from Foster and Coe (1968).

were destroyed by Maasai soon after installation), very high correlations cannot be
expected in an area with considerable rainfall variations. Moreover, rainfall will not
necessarily provide a good index of the actual availability of water to plant and animal
life, since the drying power of the air, landscape and soil characteristics, and so on,
may have an overriding effect that nullifies a 1 : 1 ratio of rainfall ( p ) to available
water.
To obtain some idea of ‘rainfall effectiveness’ the modifying influence of these
factors should be taken into account. While it is not possible to provide an assessment
of landscape and soil characteristics on water availability to plant and animal life,
some account of the drying power of the air can be made by using the ratio of p/ET.
The evaporative demand (E,) can be obtained from data given on a national scale by
Woodhead (1968); however, when the soil is covered by vegetation it is more correct
to use the term potential evapotranspiration (ET),which is simply a measure of the
potential drying power of the air assuming a saturated soil, ET can be taken as approxi-
mately 0.8 of E, in dry, low-lying areas (Brown & Cocheme, 1969). By using the ratio
p / E , as a crude measure of the drying power of the air, it should be possible to obtain
a better prediction of animal responses to available moisture than rainfall alone. Both
sets of data are given in Table 3, and because the rainfall over the last month may not
necessarily be the best index of the amount of water available freely, and in vegetation,
a moisture index over a variable period before each count has been computed, ranging
from 1 to 70 days.
The correlation of total biomass in the dry season area with rainfall is poor, the
highest correlation occurring with cumulative rainfall over the last 50 days (r =
-0.399, d.f. 10, P > O.lO), a suggestive but not statistically significant result. The
relationship is shown graphically in Fig. 2 and suggests that there is a large decline in
total large mammal biomass with a small amount of rainfall. A variety of other
correlations were tried over various time intervals before each count using both actual
 b
2
5
Table 3. Summary of rainfall ( p ) and rainfall to evapotranspiration ( p I E T )ratios used to predict biomass and animal numbers in dry season concentration area 3
___ ~~

Days prior 1

to count: I 5 10 15 20 30 40 50 60 70 3
Count
date: p p/ET p p/ET p p/ET p pIET p pIET I> pIET I) p I E T p pIET 1) p / E T p p/ET
1969 18 Mar. 14 0.07 29 0.11 29 0.09 53 0.15
19 Apr. 25 0.54 32 0.91 32 0.35 92 0.64 92 0.47 92 0.37 93 0.31 106 0.30
16 May 3 0.78 9 0.27 9 0.10 9 0.07 40 0.23 51 0.23 101 0.36 101 0.31
12 June 3 0.10 3 0.04 3 0.02 6 0.03 12 0.06 12 0.05 44 0.15
9 July 3 0.02 3 0.02 3 0.02 12 0.04
19 Aug.
30 Sept.
20 Oct. 2 0.74 2 0.04 2 0.05 5 0.05 5 0.03 5 0.02 5 0.02 5 0.02 5 0.01
I4 NOV. 2 0.52 5 0.24 8 0.18 8 0.24 18 0-20 20 0.14 20 0.10 23 0.10 23 0.08 23 0.07
7 Dec. 3 0.08 25 1.28 42 0.48 50 0.38 60 0.35 60 0.26 62 0.22 65 0.19
1970 24 Jan. 19 3.80 51 2.03 52 1.04 57 1.54 83 0.83 83 0.59 83 0.45 83 0.36 87 0.32 109 0.34
15 Feb. I 0.01 53 0.34 60 0.30 60 0.25 60 0.21 60 0.18
15 Mar. 15 0.29 15 0.38 15 0.14 15 0.10 15 0.07 16 0.06 16 0.22 68 0.28
17 Apr. 6 0.04 21 0.1 1 21 0.08 21 0.07 21 0.06
16 May 4 0.14 4 0.05 4 0.03 4 0.04 4 0.02 10 0.04 14 0.04
15 June 4 0.02 4 0.02 6 0.02
21 July
 Seasonal water availability 273

rainfall and ‘effective rainfall’ (p/E,) and a variety of transformations (square root,
semi-log and double-log). Furthermore, because there was apparently a longer lag in
the dry season return of livestock relative to wildlife, a separate set of computations
was run for each separately. Similar computations were run for ‘plains game’, zebra,
wildebeest and kongoni. The best correlations were obtained with log/log transforma-
tions, and a graph summarizing the results is presented in Fig. 3. Few correlations
were obtained using straight rainfall figures, but the value p/ET improved the relation-
ships to statistically significant levels. Because of the high degree of variability contri-
buted by elephant biomass, a species for which the sampling technique employed was
largely inadequate by virtue of a few herds with relatively large numbers, their contri-
bution to the total biomass has been omitted. There can be little doubt that with more
appropriate counting (high sampling fractions or total counts) the fluctuations of
elephant biomass would be found to follow similar rainfall-related changes.
The correlation of total biomass (less elephant contribution) with p/ET is most sig-
nificant over the last 50 days (r = -0.583, P < 0.05) falling away either side of this
time interval. Splitting the total biomass into two major components, livestock and
plains game, the former were found to correlate best with p/ET over the last 50 days,
the latter over the last 40 days. The statistical correlations for livestock were not
significant (P < 0.05) while those for plains game increased to a correlation coefficient
of 0.691 (P < 0.02), Fig. 4.

lor

I l l 1 I
. .
l l I I
10 20 30 40 50 60 70 80 90
Cumulative rainfall over preceding 50 days Preceding interval (days)
FIG. 3. FIG.4.
Fig. 3. Total large mammal biomass in the dry season range plotted against cumulative rainfall over
the preceding 50 days. The open circle is of a count following a number of rapid moves into and out
of the basin prompted by unseasonal light rains and rapid desiccation. Even a more substantial fall
was inadequate to stimulate a full emigration following this series of short-lived migrations.
Fig. 4. Correlation of herbivore biomass in the dry season range with ~ / E values
T over a preceding
time interval. The total biomass excludes elephant and buffalo; plains game includes zebra, wilde-
beest and kongoni.

The results are interpreted in the following way.

Even relatively small quantities of rain falling shortly before a count will lead to a
rapid, large scale emigration. Immigration rates after rain cessation are, by contrast,
slow, and need not reflect the total effective rainfall over a few days previously.
Similarly, since even heavy rainfalls have little residual effect after 2 months or so,
records prior to this offer little correlation between effective rainfall and animal
274 David Western

numbers. Rainfall variation over the period of 40-50 days seems to provide the best
correlations. As more data become available it will be worth improving the predictive
model by relating animal numbers to the time-specific quantity of rain, that is, given
values of effective rainfall at varied time intervals prior to the count.
Due to the high variability of counts, it is not possible to obtain significant predic-
tions for individual species numbers in the dry season area except in the case of zebra
and wildebeest. Regression equations relating animal numbers to p/ET over the last
40 days are as follows:
Zebra: logy = 1.72-0.03 Log .s ( r = -0674, d.f. 10, P :< 0.02)
Wildebeest: log 1’ = 4.32 - 1.06 log s ( r = - 0.695, d.f. 10, P < 0.02).
Species which do not show any significant seasonal fluctuation in numbers are
impala, Thomson’s gazelle, rhino and buffalo. Oryx show an inverse pattern, moving
into the basin area during the wet season and out during the dry season.
Two general features emerge from the data. Firstly, emigration is relatively spon-
taneous and can occur within a few hours of a heavy rainfall. In 1969 almost all the
animals that normally migrate had moved out of the basin within a few hours of a
rainfall of 6.9 mm. Immigration rates are by contrast slow, and it is usually a few
weeks after a reasonable rainfall before substantial numbers begin to move back into
the dry season range.
Secondly, the immigration rates of livestock are slower than plains game and are
not strongly predictable on the basis of p or p/ET. The weaker correlation is believed
to stem in large part from the high count variance in livestock. Settlement counts are a
more useful measure of settlement movements in this respect since they can be totally
counted with absolute accuracy and are highly correlated with actual livestock numbers
( r = 0.745, d.f. 10, P < 0.01). Substituting the log of settlement counts for livestock
numbers, significant correlations are obtained with log y/Er ( P < 0.05). It is apparent
that the livestock move back into the basin slightly later than plains game.
The relative rates of immigration and emigration are well demonstrated by settle-
ment counts. To reach the dry season peak numbers from a wet season minimum took
3 months in each of two dry seasons, but emigration achieved the reverse situation
within a month. In fact the emigration was complete within a few days, but counts
were not sufficiently frequent to quantify this rate of change.

Distributions from water

The results indicate that there is a progressive contraction in the range of most species
within the Amboseli ecosystem as the area dries out. Since the Amboseli basin is the
only area with natural, permanent water during the dry season, it is possible that the
concentrations are in response to a diminishing water distribution. Before passing on
to consider the evidence for this, data on the distribution of animal species in relation
to dry season water supply are required. Water is more or less ubiquitous in the rains
and animals are rarely more than a few kilometres from the nearest available water.
Data from ground counts relating the dry season density distribution of species
to the vicinity of nearest available water are given in Table 4.All species, except oryx,
and the total biomass show a significant concentration around water (P < 0.001,
using a Kolomogorov-Smirnov one-sample test, Siegel, 1956). However, more recent
data are now available, based on aerial counts over the entire ecosystem. Distance from
 Seasonal water availability 215

Table 4. Distribution of species relative to water during the dry season. Data are summed for the months
of February, August, September and October, 1968, and June and August, 1969
Distance from water (km)
Den*
Species 0-2 2 4 4-6 6-8 8-10 10-12 12-14 14-16 %t
Zebra 10.6 7.1 2.9 5.9 3.9 0.5 0.6 1.1 den
68 85 91 98 100 100 100 100 %
Wildebeest 13.9 23.2 5.4 3.4 1.6 0.1 0 0 den
56 90 97 99 100 100 100 100 %
Kongoni 0.5 0 0.4 0 0.1 0 0 0 den
81 82 I00 100 100 100 100 100 %
Thomson’s g. 2.6 1.8 0.2 0 0 0 0 0 den
78 98 100 100 100 100 100 I00 %
Grant’s g. 1.8 1.5 1.2 1.2 2.4 1.5 0 0.6 den
56 73 85 90 96 100 100 100 %
Impala 1.9 0.2 0 1.o 0 0 0 0 den
88 92 92 100 100 100 100 100 %
Oryx 0 0 I .7 0.2 0.9 0 0.8 0 den
5 6 79 84 94 94 100 100 %
Eland 0 0 0 0 0 0 0 0 den
0 0 0 0 0 0 0 0 %
Buffalo 0.7 0 0 0 0 0 0 0 den
100 100 100 100 100 100 100 100 %
Elephant 0.5 0 0 0.3 0 0 0 0 den
89 89 89 100 100 100 100 100 %
Giraffe 0.4 0.3 0.1 0.1 0.1 0 0 0 den
75 82 95 99 100 100 100 100 %
Cattle 40.0 18.4 6.4 7.0 1.o 29.3 0 0 den
71 83 87 89 89 100 100 100 %
Shoat 5.2 2.2 17.0 0 22.4 0 1.6 0 den
39 45 85 85 99 99 100 100 %
Ostrich 0.1 0.8 0.2 0.1 0 0 0.1 0 den
27 84 95 100 100 100 100 100 %
Donkey 1.6 0 0 0.2 0.2 1.9 0 0 den
87 87 87 89 90 100 100 100 %
47 64 79 81 91 95 98 100 Expected
distribution
(cumulative %)
3619 847 273 188 44 120 4 1 Biomass
distribution
(in 000’s kg)
71 87 92 97 97 99 100 100 Cumulative
distribution
of biomass (%)
* Density/km*
t Cumulative frequency distribution (%)

water profiles taken from these extensive surveys Table 6 , are given for livestock and
wildlife in Fig. 5. Because water development in the form of boreholes has made
available to livestock, but not wildlife, sources of water in areas previously dry, they
possess a different distance from water profile. Both livestock and various wildlife
276 David Western

Fig. 5. Area included in the spatial analysis of animals distribution in relation to dry season water
supply. Squares are 5 d 5 km. (a) Traditional permanent water supplies are shown by solid squares,
post-1944 water development for livestock by half squares. (b) Distance from water profiles in 5 km
categories for wildlife species. The same profiles applied to livestock pre-water development. (c)
Distance from water profiles for livestock resulting from water development.

species are only distributed within a limited distance from water, but other wildlife
species are not obviously confined to the vicinity of water. The densities of both former
categories are extremely low or absent beyond 10 km from water; the latter category
shows relatively high densities as far as 35 km from water. The distinction between
these distributions is used to separate species into two communities (Fig. 6): a water
dependent, or water bound, and water independent or non-water bound community.
Table 5 presents the list of species within each community based on data from Table 4
and recorded distributions for other species.

Causes of seasonal movements

Seasonality in the movements of large mammals between wet and dry season ranges
in Amboseli could result from water availability, pasture conditions, or a combination
of the two. The evidence seems to suggest that the dry season concentrations at least
are largely due to water availability.
Range conditions in the wet season area do not deteriorate below those of the dry
season range, based on evidence from both the standing crop biomass and quality of
vegetation. Standing crop figures during the prolonged dry season of 1969 ranged from
approximately 244 kg/ha to 320 kg/ha in a study plot typical of the wet season range,
and 102 kg/ha to 192 kg/ha on the Amboseli plains where there was still a substantial
biomass occupancy (Western, 1973).
Qualitatively, the only measurements that were made on pastures were protein
determinations and these indicated that in the wet season range during the same
 Seasonal water availability 211

Table 5. Distribution of feeding strategies between water-

bound and non-water-bound communities
Water-bound Non-water-bound
Zebra G Oryx G
Wildebeest G Eland B
Hartebeest* G Grant’s gazelle M/B
Thomson’s gazelle M/G Giraffe B
Buffalo G Ostrich B
Elephant M Lesser kudu B
Cattle G Gerenuk B
Sheep G Dik-Dik B
Goat M
Donkey G
Warthog G
Waterbuck G
Reedbuck G
Hippo G
Unclassified in terms of water dependence: impala and
rhino
* Water independent in some instances.
G = grazer.
B = browser.
M = mixed feeder.

Table 6. Density distribution (no/km2) of various species with distance from water, dry season
1973
Distance from water (km)
Species 0-5 5-10 10-15 15-20 20-25 25-30 30-35
Zebra 1.05 2.00 0.60 0.18
Wildebeest 13.85 0.87 0.16
Kongoni 0.35
Giraffe 0.63 0.03 0.14 0.50 0.42 0.70
Grant’s gazelle 1.68 3.60 0.75 0.82 0.96 0.46 0.80
Ostrich 0.23 0.03 0.22 0.50 1.oo
Cattle 15.50 24.60 5.90
Sheep & Goat 23.10 9.50 0.03

period crude protein levels were around 6.2 % while on the Amboseli plains they were
5 % . It is significant that the wet-season ranges are vacated by the water-dependent
wildlife species immediately it rains, even before the grasslands have had time to
respond. It is likely that the basin pastures-which comprise almost entirely halo-
phytic grasses-are inferior to much of the grazing in the wet-season range.
It is particularly pertinent that the two categories of animals which are not depen-
dent on water in the basin, livestock and water-independent wildlife, can remain in
the wet-season range, and do so, year round. Despite the fact that a large number of
stock do move into the basin area, 60% remain in the wet-season range where water
developments have extended their traditional dry-season range. However, even within
the newly created dry-season range, the livestock populations still show a similar
water dependence and there are still interstitial areas which are beyond access from
water, and receive little or no use. Such areas, especially those towards the llaingurun-
yeni Hills, are typically left with a good grass cover at the end of the dry season.
278 David Western

i
l
-5
259

5 J '
71
10 I5
- 48 13' I 250
I5 20 20 2 5 ' 2 5 - 3 0 ' 3 0 35
Distance from water (km)

Fig. 6. Biomass density distribution of water-bound and non-water-bound species, based on aerial
counts in October 1973. The maximum distance of any point in the ecosystem t o the nearest available
livestock watering point is less than 20 km. The large biomass recorded 35 k m from water is un-
doubtedly an artifact of a low sampling level, since very few areas are this far from water. (a) Water-
bound hildlife; (b) water-bound livestock; ( c ) non-water-bound wildlife.

Following the water development schemes initiated in and around the Amboseli
ecosystem in the 194Os, there was a three-fold increase in livestock numbers, and this
apparently was in response to the increased area of dry-season grazing (Western,
1973). There was no apparent, comparable decrease in wildlife numbers, indicating
that there was a net increase in the carrying capacity of the ecosystem. It was the very
fact that there was unutilized grazing away from the confines of water that prompted
the water development scheme.
During the 1969 drought, Maasai livestock had consumed the available pasture in
the basin area and the herders then switched to a strategy of using grasslands as far to
the north as their stock could be safely walked, since there was a large volume of
unutilized forage at the fringes of their grazing range, and beyond. The furthest
settlement from water was 1 1 km, enabling cattle to forage out to a maximum of 15
km, entailing a total of some 35 km every 2 days. The Maasai regarded water as the
absolute constraint on utilizing as yet unexhausted grazing outside the normal dry
season range, and were willing to press their cattle to the limits in order to reach it.
The foregoing evidence implies that the availability of surface water is the principal
cause of the large seasonal shifts in biomass between the wet and dry season areas.
With the onset of the rains, animals are not bound by the localized distribution of
water and spread out over a large area of wet season pasture, very little of it overlap-
ping with thedry season range. However, as the surface pools dry out with the cessation
of the rains the water-dependent species must shift back into the Amboseli basin to be
within commuter distance of permanent water. This constraint does not apply to
water-independent species, which show little seasonal shift in range.

Seasonality, water availability and food chain efficiency

Despite the recent redistribution of livestock created by water development schemes,
 Seasonal water availability 279

and the resulling coalescence of what were previously relatively discrete wet and dry
season ranges, water-dependent and water-independent communities within the
Amboseli ecosystem are still discernible as applied to wildlife rather than livestock.
Even for livestock, it is still possible to determine water-dependent profiles that limit
their dry season range and allow interstices in the ecosystem that are almost devoid
of any water-dependent species, livestock or wildlife. The fact that the water-dependent
curves for livestock apply ubiquitously, allows various assumprions to be made about
the distribution of biomass in the ecosystem prior to water development, and to calculate
the extent of the interstices that were then free of water-bound species.
Applying the water-dependence curves for both the water-bound and non-water-
bound communities (livestock are included in the former) to a map giving the distri-
bution of permanent water in the ecosystem, prior to water development, Fig. 5, a
prediction of the traditional distribution of dry season biomass, can be given. The map
is indeed no more than the present situation minus the livestock supported around the
developed water points, and although there may have been a slightly different pro-
portion of livestock to wildlife biomass, this would not radically affect the overall total.
The delineation of the two communities, water-dependent and water-independent
has important implications for the flow of energy through the large mammal eco-
system, within which 52 % of the area, the 15-km profile from water, encloses 99 % of
the dry season biomass of the water-dependent community in the dry season, the
remaining area being occupied solely by water-independent species. If the seasonal
movement patterns already outlined are applied to calculate the amount of time
that was spent outside the dry season range, this appears relatively small, 20% of
the biomass occupancy. In other words, half the ecosystem that was covered by wet
season range received only 20% of the water-dependent biomass occupancy of the
dry season area. Using the 10-km profile which enclosed 88 % of the water-dependent
biomass, the differences in use of the two seasonal ranges would have been even
greater and would not have been accounted for by differences in primary production.
By contrast, the biomass density of water-independent species shows little change
with distance from water.
The obviously large differences in food chain efficiency (consumption by trophic
level nlproduction by trophic level n - 1) that existed in the Amboseli ecosystem in the
past, and still prevail to a lesser extent, are of immediate ecological and development
significance, but before discussing these aspects it is worth considering the role of water
availability and seasonality in structuring the ecosystem into two ecological com-
ponents.
The distribution of permanent water broadly separates the Amboseli ecosystem into
two communities, water-bound and non-water-bound. The division would seem to
hold some degree of generality based on the feeding radius of the water-bound com-
munity around water. In a large ecosystem with highly localized water distribution, the
size of the water-bound community relative to the non-water-bound community
should be small, and conversely where water is well distributed or ubiquitous. If the
water dependence curves are widely applicable, then the model given below will be
valid for other areas.
By definition, the distribution of permanent water in an ecosystem can be regarded
as a constant parameter resulting from its drainage and geomorphic characteristics.
Over a number of years even so-called permanent water may dry up, but here we are
more concerned with seasonal than long term changes. Given the distribution of
280 David Westeni

permanent water and the water-dependence biomass curves, the distribution limits of
the water-dependent community in the dry season should be predictable. The spatial
limits enclosing a given percentage of the community can be calculated by:
It

A = Z w,2-d
i I
where A = spatial limits of dry season distribution; i = percentage of biomass
included; I’ = radius of circle enclosing i percent of biomass; d = total area of over-
lap between adjacent r’s.
If there is no seasonal movement out of the dry season range the carrying capacity
of the water-dependent community can be readily calculated from the carrying capa-
city of in sifu food supplies and their availability and quality. This situation only applies
to resident communities; in migratory communities the ratio of biomass occupancy
in the wet to that in the dry season ranges will proportionally increase the figure. The
duration and distribution of ephemeral water will limit the time spent out of the dry
season range and while potential ephemeral water reservoirs are determined by the
geomorphic properties of the area, their duration will vary with the quantity and
distribution of rainfall. To a large extent there is a regular seasonal pattern in the
quantity and distribution of rainfall, and this offers a large measure of predictability
in calculating the ratio of time spent In the wet season and dry season areas, allowing
for the time lags in rainfall and ephemeral water depletion.
The duration of ephemeral water can be calculated on the basis of effective rainfall,
the drying power of the atmosphere, animal consumption and a constant which is the
water storage capacity of the area. The time lag in return times of animals to the
Amboseli dry season area has been shown to have a reasonable level of predictability
on the basis of p/E,, the best fit-with cumulative values over the previous 40-50
days-presumably reflecting the water storage constant. Using these sorts of data to
calculate biomass occupancy outside the dry season area, one can roughly calculate
the biomass of the water-dependent community as follows :

Where: B = biomass of water-dependent community; li = carrying capacity in

biomass density of dry season range; b = biomass occupancy outside dry season
range: d = as in equation (1).
Presumably many further refinements are possible, such as water-dependence
curves which are related to seasonal changes in heat stress and vegetation moisture.
The main point made is that one can begin to use seasonal and environmental data to
provide rough estimates of carrying capacity and food chain efficiencies.
I t should be stressed that the seasonality in rainfall can be as important as the total
rainfall in determining the biomass levels of a water-dependent community. A given
quantity of effective rain will allow a gerater biomass occupancy outside the dry season
area if it is spread over a longer rather than shorter period, and there may be consider-
able variation in the spread of a given quantity of rain, for example the number of
dry months at Mashuru in the extreme north of the Amboseli ecosystem varies from
2 to 6 for years with a fall of 350 mm. Tyrell’s (1975) precipitation seasonality index
uhich expresses the duration of annual rainfall on a scale of 1 (equitable distribution)
to 100 (all falling in 1 month) gives a useful measure of seasonality. A high p J value
 Seasonal water availability 28 1

will restrict the wet season dispersal phase and potential biomass levels of the water-
bound community.

Physiological barriers to energy consumption

Most of the difference that apparently existed in the food chain efficiencies of the
water-dependent and water-independent communities prior to water development,
and still partially persists, can be accounted for by the poor utility of graze by the
water-independent community. Livestock increases in the Amboseli ecosystem and
possibly Kajiado District in general resulted in large part from the extension of dry
season grazing made available by water development, since there is evidence suggesting
that the traditional dry season range was fully stocked by 1915, following the recovery
of cattle from rinderpest (Sandford, 1919). A few areas still remain beyond dry season
grazing range and are frequently compared to the areas close to water to show the
effects of overgrazing by livestock. Overgrazing, however, is a relative term, unless
based on the long term food chain efficiencies of areas being compared, and in this
sense the water-dependent community is considerably more efficient, although in
absolute terms higher efficiencies may be obtainable with different stocking rates.
What seems to underlie the differences in food chain efficiencies between water-
dependent and water-independent communities is a physiological barrier: with
decreasing water availability a negative water balance becomes progressively more
difficult to avoid, but a number of strategies exist for doing so: (1) physiological
adaptations to conserve water, (2) feeding selection for a high water-content diet, (3)
behavioural adaptations which include gearing activity cycles to minimize desiccation
and to consume forage when it contains high moisture content.
Taylor (1968, 1969a, 1970) has discussed these adaptations to arid environments by
East African herbivores. Strategies 2 and 3 may be useful in extending the range of
highly water-dependent species and the actual amount will be a function of ambient
heat stress and food moisture content.
The selection of food with a high water content strongly favours browse rather than
graze intake, due to grasses in semi-arid and arid environments tending to have much
lower moisture levels in the dry season, often considerably below 10% water content
in environments close to Amboseli (Cassady, 1973). There is little data available on
browse water levels at similar times, but Taylor & Lyman (1967) showed that eland,
during a drought in northern Kenya, were consuming food with a 50-60% water
content through a selection of high moisture level browse plants. Taylor (1968, 1972)
has shown how Grant’s gazelle can maintain a positive water balance without free
water by selecting high water content food.
By classifying the water-dependent and water-independent communities in Amboseli
into browsers, grazers and mixed feeders, the division of the two communities on the
basis of feeding strategies can be tested. A classification of this sort is given in Table 5
using Hofmann’s (1973) division of ruminants on the basis of food intake. The assign-
ment of non-ruminants into similar categories is based on personal observations and
information available in the literature. The only deviation from Hofmann’s classifi-
cation is eland which is believed to be predominantly a browser (J. C. Hillman, personal
communication). The results show large differences between the two communities on
the basis of grazers and non grazers, 86 % of the water-dependent species being grazers
(or predominantly so, with no browsers) and 89 % of the water-independent community
being browsers or predominantly so with only 1 grazer. The division is in fact so
striking that they can be classified as grazing and browsing communities respectively.
Both oryx and Grant’s gazelle show a variety of physiological adaptations to
surviving without free water intake (Taylor, 1969b, 1972; J. King, personal communi-
cation) and both take in a significant proportion of graze, but even here, observations
i n Amboseli indicate that the graze to browse ratio in Grant’s falls to low levels in the
dry season, a feature noted elsewhere by Stewart & Stewart (1970).
The conclusion drawn is that there is a physiological barrier to a high level of
utilization of low moisture content grasslands in the dry season, if free water is un-
available. There does, in other words, seem to be a niche or niches available and an
alternative to the physiological barrier hypothesis is one which suggests that species
are missing through extinction, but since the environment is one which is relatively
inaccessible to man, a principal cause of much recent extinction in large mammals,
this is unlikely. Alternatively, it could be supposed that the niche has only recently
been opened up by changing climates or human activity, but arid-adapted species, the
oryx, Grant’s gazelle, and gerenuk have exploited these environments, as far as we
know, for a long period, so this too seems unlikely. Physiological barriers to food
exploitation have been shown to exist in invertebrates in similar environments, for
example in Tsavo National Park, where faecal decomposition by dung beetles is
limited by the availability of water (Anderson & Coe, 1974), and it is not unreasonable
to expect a similar situation in large mammal communities. The literature on adapta-
tions to arid environments by large mammals is voluminous (Schmidt-Nielsen, 1964;
Maloiy, 1972) and the importance of food water intake in maintaining a positive water
balance is well documented. For example, a declining appetite in livestock with
decreased water intake is well established and Macfarlane & Howard (1972) have
demonstrated that water turnover is inversely proportional to stocking rate in sheep in
the Negev, Israel, a factor which could of itself limit food chain efficiency in water-
deficient environments.

Comparison with other areas

Large mammal communities can be classified int.0 three broadly recognized categories
on the basis of seasonal movements: ( 1 ) migratory: (2) dispersal (wet season dispersal/
dry season concentration); (3) resident.
A number of other ecosystems contain communities of large mammals with seasonal
dispersals similar to those in Amboseli, for example Tarangire (Lamprey, 1963), the
Athi-Kapiti Plains (Peterson & Casebeer, 1972), southern Samburu District (Rainy,
19731, Tsavo West (S. Cobb, personal communication) and several Tanzania Maasai-
land, livestock-dominated communities (Jacobs, 1965). Few data are available for
quantitative comparison, but there is evidence to suggest that water availability does
have an important bearing on the distributional limits of water dependent species in
the dry season (Lamprey, 1964; Rainy, 1973).
The Serengeti large mammal community is regarded as largely migratory (Bell,
1971) but nevertheless, distinct wet and dry season ranges are evident, the Serengeti
Plains and woodlands respectively, and although it does seem that the main migrations
are in response to pasture conditions, the plains are coincidentally devoid of water
during the dry season. The dry season range has a welI-distributed supply of perma-
nent water and in many respects the seasonal movements are similar to those in
 Seaonal water availability 283

Amboseli. In both situations the movements from the seasonally watered wet season
range to the permanently watered dry season range is slow relative to the sudden
reverse migrations following rainfall. Within the dry season ranges there are progressive
concentrations in the moister areas of greater grass production, though of poorer
quality; in Serengeti the concentration is towards the northern high rainfall zones
(M. Norton-Griffiths, personal communication), in Amboseli onto the low-lying,
swamp pastures. In fact the outstanding differences are not so much the pattern of
movements, as in their scale; the annual movements in Serengeti cover an area some
ten times that covered in Amboseli.
Ngorongoro Crater ecosystem is regarded as supporting a resident population of
herbivores which move comparatively little during the rains. Wet and dry season
movements within the Crater are well defined, however, there being a movement onto
the higher ground during the rains and a progressive concentration onto the swamp
pastures in the dry season. In fact the similarity between the seasonal movements of
Amboseli and Ngorongoro is striking, but in the latter case the entire ecosystem is
within reach of permanent water, excluding the area outside the Crater which is used
during the rains by a small percentage of the animals. Once again the scale of move-
ments are of a different order of magnitude occupying approximately 300 km2 com-
pared to 3000 km2 in Amboseli and some 25 000 km2 in Serengeti.
A definition of communities on the basis of seasonal movements, as commonly
used, is not particularly meaningful, at least in the case of the three examples com-
pared. It is the scale of movements that has been used to distinguish between the three
categories.
Although in the context of seasonal movements the distinction between resident,
dispersal and migratory populations is not particularly meaningful or useful, a classi-
fication on the basis of scale can have some merit in for example determining the
impact of predator populations on the herbivores. Kruuk (1972) has shown that as a
consequence of migratory movements on a large scale in Serengeti, the impact of
predator populations on herbivores is small compared to Ngorongoro where the off-
take is extremely high.
The case is an interesting one, presenting another example of factors which may
limit food chain efficiency, in this instance high prey mobility as a limitation on energy
flow into the secondary consumers.

General remarks
The division of large mammals into water-dependent and water-independent species
is no doubt a simplification of the underlying situation. It is possible that certain of
the species classified as water independent only appear so within 35 km of water, the
maximum distance in the Amboseli study area, and that at distances greater than this,
or under more heat stressed environments, species such as giraffe and ostrich may
show water dependent distributions. The concept as used is nevertheless useful within
Amboseli, and in all probability, similar environments.
Much of the Amboseli basin is extremely saline and alkaline and the vegetation is
composed predominantly of halophytes (Western & Van Praet, 1973). If higher
dietary mineral salt loads are ingested, the greater water turnover needed to maintain
an ionic balance could appreciably increase the water dependence of animals feeding
on these pastures. Overall, however, this is unlikely to affect the absolute range of
284 David Western

species, since the soils immediately outside the basin, and well within the cruising
range of water-dependent species, are oxisols, derived from the basement complex,
and are non-saline.
It cannot be calculated from the Amboseli data whether the low biomass density of
browsers relative to grazers of less than 1 : 10 is related to a differential primary pro-
duction of browse and graze, but this seems unlikely. It is quite possible that the
seasonal production of browse is composed to a large degree of deciduous species
with a sharp production and leaf-fall cycle. Most of the grasses are by comparison
perennial and remain available over a full seasonal cycle. It is conceivable then that
seaonality could have yet another significant influence on energy flow through the
system, reducing food chain efficiency in browsing communities faced with a high
percentage of deciduous plants and a short rainfall season.
An increased intake of browse relative to graze as the dry season progresses has
been demonstrated in mixed feeders such as Grant’s gazelle (Stewart & Stewart,
1970), and the effect of so doing would probably be to increase the preformed water
intake, a necessary channel of supply in avoiding dehydration (Taylor, 1972), but
whether this is the reason for a dietary switch is unclear. Cassady (1973) has suggested
that there is a close correlation between grass moisture content and crude protein
level. Utilizing data from his published tables showed a highly significant correlation
between these two parameters (Panicum ma.uimum r = 0.84, P < 0.01 ; Digitaria
milanjiana r = 0.79, P < 0.01). It seems quite possible then that a switch in diet
to a greater browse intake could result from both a simultaneous decline in food
moisture and quality in graze. The quality of food alone is often adequate to reduce
animal consumption.

Management implications
Seasonal movements in large mammal communities pose difficultiesfor their conserva-
tion, especially in those areas where there is a distinct dispersal and concentration
phase. In most cases, of which Amboseli, Samburu, and Mara Game Reserves, and
Nairobi National Park are good examples, conservation action is taken before the
seasonal movements are worked out, and protection is provided for the conspicuous
phase, the dry season concentrations. The former human inhabitants are usually
displaced to the wet season range, and as a result of this artificial division, two conse-
quences arise. In the wet season areas livestock rapidly increase through water debelop-
ment and the reserve supply of forage made available, and in due course reach a new
carrying capacity.
The demand for additional livestock production has increased substantially as a
result of human population increments, to the extent that the loss of forage to
seasonal incursions by wildlife is viewed with increasing hostility. Simultaneously
wildlife area5 have become economically more valuable and conservation pressures
for the full protection of wildlife communities have increased sufficiently that pro-
tection is sought for the wet season dispersal phase. The prospect of the seasonal
migrations being severed jeopardizes the dry season populations and would in most
cases lead to a severe biomass reduction, about 30% in the case of Amboseli.
In Amboseli, and inevitably elsewhere, an imbalance is created-‘Government
animals’ (wildlife) are permitted to move onto private land and are protected by
legislation, whereas privately owned animals (livestock) are not permitted to move
 Seaonal water availability 285

onto government land, the national parks. It is too much to expect the private land
owners to accept a situation in which their cattle can be shot for occasionally trespass-
ing on government land in periods of pasture shortage, while government-owned
wildlife may consume over 50% of their available pasture. Economically the imbalance
may be even more dramatic; in the case of Amboseli wildlife revenues exceed &750000
p.a. and yet nothing is received by the landowners, whose production loss amounts to
&23000 p.a.-or 8 % of their annual subsistence production (Western & Thresher,
1973).
Unless the landowners in the dispersal paths of parks animals are provided incen-
tives to do otherwise, it is difficult to see how the seasonal migrations can continue.
In Amboseli, Western & Thresher (1 973) have formulated a development plan that
takes into account the opportunity costs of wildlife to landowners, and seeks, by the
payment of wildlife utilization fees to ranchers in the dispersal areas of the ecosystem,
to provide such incentives. The utilization fees would be drawn from revenues earned
by the Park, but would be phased out proportional to direct earnings from wildlife,
largely through tourist viewing activities. The total revenues of the Park and land-
owners would be vastly increased by an integrated land use approach, relative to
earnings from separate livestock and wildlife enterprises.
In many areas there will be insufficient revenues generated by the protected portion
of the annual wildlife range to afford compensatory payments to landowners. In such
situations a number of alternatives exist; the compensation figures could be paid
proportional to revenue earnings by alternative government funding, or direct
revenues could be encouraged through hunting, cropping or tourism, but it might be
necessary in some situations to discourage seasonal migrations altogether by game
barriers or other deterrents. Purely aesthetic reasons are not likely to play a significant
role, except perhaps in the case of non-competitive species like giraffe.

Acknowledgments
The main study was supported by a Ford Foundation Travel and Study Award and a
Leverhulme Overseas Research Award. The research programme is currently supported
by the New York Zoological Society.
I am grateful to Dr Colin Pennycuick who did most of the piloting during the
monthly aerial counts, and to Drs Harvey Croze, Stephen Cobb, and Chris Hillman
who contribute to the aerial sampling programme. Thanks are also due to Sandy
Price who assisted in manuscript preparation.

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