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Expansion of human settlement in Kenya's Maasai Mara: What future for

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Article in Journal of Biogeography · June 2004

DOI: 10.1111/j.1365-2699.2004.01062.x

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Journal of Biogeography (J. Biogeogr.) (2004) 31, 997–1032
ORIGINAL
ARTICLE
1
Uganda Wildlife Authority, PO Box 3530,
Kampala, Uganda and 2International
Livestock Research Institute, PO Box 30709,
Nairobi, Kenya
*Correspondence: Richard H. Lamprey, Uganda
Wildlife Authority, PO Box 3530, Kampala,
Uganda. Tel.: +256 77 704596.
E-mail: lamprey@infocom.co.ug
ª 2004 Blackwell Publishing Ltd www.blackwellpublishing.com/jbi
Expansion of human settlement in
Kenya’s Maasai Mara: what future for
pastoralism and wildlife?
Richard H. Lamprey1* and Robin S. Reid2
ABSTRACT
Aim Wildlife and pastoral peoples have lived side-by-side in the Mara ecosystem
of south-western Kenya for at least 2000 years. Recent changes in human
population and landuse are jeopardizing this co-existence. The aim of the study is
to determine the viability of pastoralism and wildlife conservation in Maasai
ranches around the Maasai Mara National Reserve (MMNR).
Location A study area of 2250 km2 was selected in the northern part of the
Serengeti-Mara ecosystem, encompassing group ranches adjoining the MMNR.
Emphasis is placed on Koyake Group Ranch, a rangeland area owned by Maasai
pastoralists, and one of Kenya’s major wildlife tourism areas.
Methods Maasai settlement patterns, vegetation, livestock numbers and wildlife
numbers were analysed over a 50-year period. Settlement distributions and
vegetation changes were determined from aerial photography and aerial surveys
of 1950, 1961, 1967, 1974, 1983 and 1999. Livestock and wildlife numbers were
determined from re-analysis of systematic reconnaissance flights conducted by
the Kenya Government from 1977 to 2000, and from ground counts in 2002.
Corroborating data on livestock numbers were obtained from aerial photography
of Maasai settlements in 2001. Trends in livestock were related to rainfall, and to
vegetation production as indicated by the seasonal Normalized Difference
Vegetation Index. With these data sets, per capita livestock holdings were
determined for the period 1980–2000, a period of fluctuating rainfall and primary
production.
Results For the first half of the twentieth century, the Mara was infested with
tsetse-flies, and the Maasai were confined to the Lemek Valley area to the north of
the MMNR. During the early 1960s, active tsetse-control measures by both
government and the Maasai led to the destruction of woodlands across the Mara
and the retreat of tsetse flies. The Maasai were then able to expand their settlement
area south towards MMNR. Meanwhile, wildebeest (Connochaetes taurinus) from
the increasing Serengeti population began to spill into the Mara rangelands each
dry season, leading to direct competition between livestock and wildlife. Group
ranches were established in the area in 1970 to formalize land tenure for the
Maasai. By the late 1980s, with rapid population growth, new settlement areas had
been established at Talek and other parts adjacent to the MMNR. Over the period
1983–99, the number of Maasai bomas in Koyake has increased at 6.4% per annum
(pa), and the human population at 4.4% pa. Over the same period, cattle numbers
on Koyake varied from 20,000 to 45,000 (average 25,000), in relation to total
rainfall received over the previous 2 years. The rangelands of the Mara cannot
support a greater cattle population under current pastoral practices.
Conclusions With the rapid increase in human settlement in the Mara, and with
imminent land privatization, it is probable that wildlife populations on Koyake
997
R.H. Lamprey and R.S. Reid

will decline significantly in the next 3–5 years. Per capita livestock holdings on the
ranch have now fallen to three livestock units/reference adult, well below
minimum pastoral subsistence requirements. During the 1980s and 90s the
Maasai diversified their livelihoods to generate revenues from tourism, small-
scale agriculture and land-leases for mechanized cultivation. However, there is a
massive imbalance in tourism incomes in favour of a small elite. In 1999 the
membership of Koyake voted to subdivide the ranch into individual holdings. In
2003 the subdivision survey allocated plots of 60 ha average size to 1020 ranch
members. This land privatization may result in increased cultivation and fencing,
the exclusion of wildlife, and the decline of tourism as a revenue generator. This
unique pastoral/wildlife system will shortly be lost unless land holdings can be
managed to maintain the free movement of livestock and wildlife.
Keywords
Kenya, Maasai Mara, pastoralism, rangelands, bomas, livestock, wildlife.

wildlife (IIED, 1994; Western, 1994; Berger, 1996; Igoe &

INTRODUCTION
It has become a conservation dictum in Africa that the
survival of the continent’s wildlife, and particularly of its
‘megafauna’, into the twenty-first century will depend on the
goodwill of local communities. It is argued that the support
of communities for wildlife can be enhanced through
community-based wildlife management (CBWM), a process
in which local people participate in, and benefit from, the
conservation and management of wildlife on their land (Kiss,
1990; IIED, 1994; McNeely, 1995). Throughout Africa,
integrated conservation and development projects (ICDPs)
have been established by donors and non-governmental
organizations (NGOs) to encourage and promote CWBM,
and the sustainable use of natural resources (Gibson &
Marks, 1995; IIED, 2000; Barrow & Murphree, 2001).
Wildlife-based ICDPs cover a range of interventions, from
the development of local craft markets near wildlife tourism
areas, to the full ownership of wildlife and the sharing of
benefits from tourism or sport hunting (Ashley & Roe, 1998;
Hulme & Murphree, 2001).
The CBWM approach has now become enshrined in national
wildlife policies in eastern Africa (KWS, 1990; Wildlife Division,
1998; TANAPA 1994; UWA, 2002), where particular attention
has been focussed on conserving wildlife on land owned by
pastoralists such as the Maasai (Western, 1984; Homewood &
Rodgers, 1991; Igoe & Brockington, 1999). It is asserted that in
the pastoral savannas, wildlife has coexisted with man and his
stock for millenia (Parkipuny, 1997), and that there is little
competition between the two. Therefore CBWM initiatives in
savannas, with benefits to communities through game-viewing
tourism or sport hunting, will have a higher chance of success
than in agro-pastoral or agricultural areas where wildlife and
people may be mutually exclusive. Critical factors limiting the
development of CBWM in pastoral areas include insecurity of
land tenure, inequitable distribution of benefits within com-
munities, outdated legislation and lack of expertise in managing
Brockington, 1999; Barrow et al., 2000; Metcalfe, 2000).
Another factor limiting the effectiveness of ICDPs, often entirely
ignored by community conservation initiatives, is human
population growth and privatization of land. In examining the
prospects of sustainable wildlife harvest on community land,
Barrett & Arcese (1995, p. 1076) contend that ‘if human
populations grow past the point where a sustainable harvest can
provide satisfactory benefits on a per capita basis, ICDPs should
be expected to fail for the same reasons that local cooperation
may have been lacking prior to their initiation’.
Since the 1980s, CBWM has been put forward as a new
paradigm of conservation thinking (Adams & Hulme, 2001).
However, as long ago as the 1950s wildlife managers in Kenya
were making efforts to ensure that local people benefited from
wildlife, using approaches that many conservationists today
still regard as innovative. For example in 1951, the Kenya
Game Department initiated a scheme by which sport hunters
in ‘controlled areas’ paid a game fee for each animal shot to the
local district council (Game Department Annual Report,
1952). The scheme ran for 26 years1 and generated revenues
equal to those from game-viewing tourism (Strickland, 1973;
Casebeer, 1975; R.H. Lamprey, unpubl. data). A second
approach, proposed by the Game Policy Committee of 1956,
and implemented in 1961 prior to independence, was to
transfer game reserves to local government councils. District
councils, in their control of ‘Trust land’, were encouraged to
adopt game reserves as ‘African District Council Game
Reserves’, where ‘human affairs are regulated by… council
by-laws and faunal matters by the Wild Animal Protection
Order (the legislation of the Game Department)’ (Game
Department Annual Report, 1962, p. 4). After independence,
1
For a short period, from 1952 to 1958, hunting was prohibited
during the independence struggle (referred to by the colonial gov-
ernment as the ‘emergency’). Sport hunting was banned in Kenya in
1977.

998 Journal of Biogeography 31, 997–1032, ª 2004 Blackwell Publishing Ltd


these reserves were designated ‘county council reserves’2 and
legally gazetted, the principal areas being Maasai Mara,
Amboseli and Samburu. District councils3 received substantial
grants from Government in the early 1960s to develop these
areas for tourism, and to collect revenues from entry fees,
camping sites and lodge concessions. Maasai Mara and
Samburu have become prime tourism areas in Kenya
(Douglas-Hamilton et al., 1989; Lamprey, in prep.).4
Meanwhile, beginning in the mid-1960s, the issue of land
tenure on Maasai rangelands was being addressed by the
establishment of ‘group ranches’. As part of Kenya Govern-
ment’s land adjudication programme, groups of Maasai were
given title deeds to tracts of grazing land which they had
traditionally used over a long period (Lawrance Report, 1966).
There were three closely linked objectives to the group ranch
programme (Lewis, 1965; Davis, 1970; Hedlund, 1971; Oxby,
1982; Rutten, 1992). First, it was anticipated that meat produc-
tion in rangeland areas would be increased, in line with national
livestock development policies. Secondly, it was felt that if the
Maasai were given their own land they would manage it in an
ecologically sound way by designing their own grazing manage-
ment schemes, selling surplus stock, and controlling livestock
diseases effectively. Thirdly, by settling people permanently on
their own land, the state could more easily meet their needs for
basic health care and education. The concept of group mem-
bership was enshrined in the Land (Group Representatives) Act,
1968, which stated that ‘each member shall be deemed to share in
the ownership of the group ranch in undivided shares.’ Whilst
the ranch is owned by a group of ‘members’ acting as a corporate
body, the livestock remain the property of individual members.
By organizing themselves into groups, adopting constitutions,
and electing group committees, the members could acquire
loans from the Agricultural Finance Corporation for the
development of their land, for such projects as the construction
of dips, dispensaries and schools.
It was also recognized that group ranches could earn
revenues from tourism. In certain areas, such as in Narok
District, the potential for tourism was seen as being so high that
ranch planners in the 1970s were concerned that ‘because of
such a lucrative source of funds (from tourism), these ranches
have not yet had to apply for a development loan. This worries
the Ranch Planning Office in the District because… without a
loan to repay, there will be no significant offtake of cattle, but
rather, ranch finances will be used to purchase more cattle
thereby leading to a chronic overstocking problem’ (Doherty,
1979, p. 7). These concerns aside, by the 1970s many of the
critical elements were in place to ensure that the Maasai could
2
Initially established in the late 1940s as ‘national reserves’ under
Kenya Royal National Parks, these county-council ‘game reserves’ were
again redesignated as ‘national reserves’ under the 1976 Wildlife
(Conservation and Management) Act. However, they continued to be
managed by local councils.
3
In Kenya, districts are managed by ‘county councils’.
4
Amboseli was designated a National Park in 1974.
Journal of Biogeography 31, 997–1032, ª 2004 Blackwell Publishing Ltd
Expansion of human settlement in Kenya’s Maasai Mara
successfully manage and derive benefits from wildlife on their
rangelands. Maasai District Councils were managing their game
reserves, and earning revenues that could, in theory, be put
towards improving public services for their electorate. On
rangelands adjacent to the reserves, groups of Maasai had land
tenure and were in a position to directly benefit from tourism.
A critical test area for the success of wildlife conservation on
community land is the Maasai Mara ecosystem in south-
western Kenya. The area has a long history of pastoral–wildlife
interactions as indicated by the detailed palaeontological
research of Marshall (1990). As early as 2000 years ago, in
the Neolithic period, the Mara was occupied by advanced
pastoralists, who managed their cattle, sheep and goat herds
for maximized production and who did not hunt wildlife.
Today, the Mara supports an exceptionally large and diverse
population of savanna wildlife (Lamprey, 1984; Broten & Said,
1995; Ottichilo, 2000), and in the wildlife dispersal area around
the core Maasai Mara National Reserve (MMNR), Maasai
pastoralists have group or individual land tenure. Tourism in
the MMNR, and in adjacent Maasai ranches, has increased 10-
fold since the mid-1980s, with many millions of dollars being
generated for local government councils and for the Kenya
economy in general (Douglas-Hamilton et al., 1989; Lamprey,
in prep.). CBWM is considered to be highly successful in the
Mara, with direct involvement of the Maasai and local
government in wildlife conservation and in the generation of
revenue from tourism (Talbot & Olindo, 1990).
However, hopes for integrating pastoralism with conserva-
tion in the Mara are fading. Over the last 20 years, large tracts of
grazing land on the Loita plains in the north-east of the
ecosystem, used seasonally by livestock and wildlife, have been
leased out by Maasai for commercial wheat cultivation. Due to
habitat loss, poaching and other disturbances, resident wildlife
populations in the Mara have declined by over 70% over the last
20 years (Ottichilo, 2000; Homewood et al., 2001; Serneels &
Lambin, 2001). Meanwhile, in group-owned ranches adjacent
to the MMNR, the human population is increasing very rapidly
(Lamprey, 1984; Reid et al., 2003). As pressure builds for land,
ranch members are diversifying their livelihoods through
cultivation and income-generation from tourism, but there is
a dire inequality in earnings between the ranch leadership and
ordinary members (Homewood et al., 2001). During recent
planning meetings for the Mara, many Maasai indicated that
after ranch subdivision, they would prefer to cultivate (ACC,
2001). If this were to occur on a large scale, it is inevitable that
fences will be erected, and wildlife movements will be disrupted;
cultivation and large herds of wildlife are incompatible.
Subdivided ranches in the Mara ecosystem that receive
sufficient rainfall for agriculture, such as Lemek GR, now have
large areas under cultivation (Thompson, 2002). In his
economic analysis of landuse on the Mara plains, Norton-
Griffiths (1995) argues that the Maasai forgo opportunities for
more profitable forms of landuse such as cultivation, and
should now be compensated to maintain their land for wildlife.
A recent body of research presented by Homewood et al.
(2001) presents a more optimistic scenario for the Mara, in
999
R.H. Lamprey and R.S. Reid
which the Maasai have a choice of landuse options. They assert
that the Maasai remain relatively wealthy in terms of cattle
ownership, and that sustainable pastoralism can be balanced
with other ‘conservation-compatible’ livelihoods to absorb
future population growth. They maintain (Homewood et al.,
2001, p. 12547) that ‘the size of the livestock population is linked
to pastoralists decisions and their wealth… A possible tradeoff
exists for pastoralists between increasing livestock holdings and
maintaining tourist-related incomes through wildlife conserva-
tion. Similar tradeoffs have to be made by pastoralists concern-
ing the leasing of their land for mechanized agriculture and the
expansion of small-scale cultivation.’ The study concludes (p.
12549) that ‘the (Mara) ecosystem could accommodate future
population growth at low ecological cost provided land zoning
manages settlement, subsistence agriculture and their access to
and impact on key resources (swamps, water holes, wildlife
migration routes).’ The study also recognizes the urgent need
for a more equitable distribution of tourism revenues between
the elite and ordinary ranch members.
Our paper describes changes in human, livestock and
wildlife populations in the Mara over the last 50 years, to re-
assess whether pastoralism can remain a viable livelihood
option for the Maasai, and whether wildlife conservation has a
future on the group ranches. In a selected study area
encompassing key Maasai ‘group ranches’, temporal changes
in settlement distribution were determined from an analysis of
1000
Figure 1 The study area (boxed) in the
Serengeti Mara ecosystem. The dashed line
indicates the limit of the seasonal movements
of migratory wildebeest. SNP, Serengeti
National Park; MMNR, Maasai Mara
National Reserve; NCA, Ngorongoro
Conservation Area; MGR, Maswa Game
Reserve; IGR, Ikorongo Game Reserve;
GMR, Grumeti Game Reserve.
aerial photographs taken over the period 1950–74, and from
ground and aerial surveys in 1983 and 1999. Livestock and
wildlife populations were determined by aerial censuses
(‘systematic reconnaissance flights’, SRFs) conducted by the
Kenya Government over the last 20 years. Livestock units per
capita were calculated to assess whether pastoralism can
remain the principal livelihood for Maasai in the Mara
ranches. The consequences of these findings are discussed in
the context of other livelihood options, and of the imminent
fragmentation of group ranches into individual holdings.
THE STUDY AREA
The study area of 2250 km2 lies in the far north of the
Serengeti-Mara ecosystem (SME)5 (see Fig. 1), and includes
the northern part of the MMNR, the adjacent Maasai ranch of
Koyake (‘Koyake Dagurugurueti’) (971 km2) and parts of
Lemek, Ol Chorro Oirowa (‘Olchoro Oiroua’) and Ol Kinyie
Group Ranches. Place names for the study area are shown in
Fig. 2. The area is bordered in the north by the Mara River and
5
The Serengeti-Mara ecosystem encompasses the seasonal movements
of the migratory wildebeest (Gwynne & Croze, 1975; Pennycuick, 1975;
Sinclair, 1979), and includes the Serengeti National Park and Maswa,
Grumeti and Ikorongo Game Reserves in Tanzania, and the Maasai
Mara National Reserve and adjacent rangeland areas in Kenya.
Journal of Biogeography 31, 997–1032, ª 2004 Blackwell Publishing Ltd

Figure 2 The study area in the Mara
(square), and location of Maasai Mara
National Reserve, group ranches (GRs) and
names referred to in the text.
the land unit of the agricultural Kipsigis people (elevation
> 2100 m), in the east by the Loita plains (1800 m), in the
west by the steep Siria escarpment in Trans-Mara (2000 m),
and in the south by the MMNR (1700 m).
The Mara study area receives 600 mm of rainfall in eastern
areas, rising to 1000 mm at the western side where climate is
strongly influenced by the Lake Victoria weather system
(Glover, 1966; Norton-Griffiths et al., 1975; Ministry of
Agriculture, 1977; Epp & Agatsiva, 1980; Ottichilo, 2000).
The seasonal north–south shift of the inter-tropical conver-
gence zone brings the ‘short rains’ in October–November
and the ‘long rains’ in March–May (Brown & Cocheme, 1973).
Soil moisture is generally sufficient to maintain a single
growing season from November–June (Lamprey, 1984)6 the
6
Calculated using the potential evapo-transpiration (PET) data of
Woodhead (1968) modified according to Brown & Cocheme (1973),
and on the basis that the ‘growing season’ begins when rainfall exceeds
PET/2 (FAO, 1978).
Journal of Biogeography 31, 997–1032, ª 2004 Blackwell Publishing Ltd
Expansion of human settlement in Kenya’s Maasai Mara
July–October period being the ‘dry-season.’ The climate and
vegetation of the study area are characteristic of eco-climatic
zone IV, the ‘semi-arid to subhumid zone’ defined by Pratt &
Gwynne (1977) and Sombroek et al. (1980).
The soils in the Mara are ‘phonolitic tuffs’ derived from
volcanic ash (Glover, 1966), and havemoderate-high fertility
(Jaetzold & Schmidt, 1983). These blanket a geological
basement composed of Miocene quartzites, gneisses and
schists (Williams, 1964; Wright, 1967). The exposure of the
basement is represented in the stony hills in the Lemek and
Siana areas, and in the west by the prominent Siria escarpment,
a major fault line in the basement (Saggerson, 1972). The Mara
river, the most prominent river of the region, flows southwards
along the base of the escarpment. Numerous streams and
watercourses drain the central plains into the Mara River, the
largest being the Talek and its tributaries the Olare Orok,
Ntiakitiak (‘Jagartiek’), Kaimurunya and Ol Sabukiai
(Lamprey, 1984; Reid et al., 2003). Aside from these perennial
watercourses, the most important sources of permanent
1001
R.H. Lamprey and R.S. Reid
freshwater are springs arising from the hills in the Lemek/
Aitong area.
The vegetation of the study area comprises tall and short
grass plains dominated by Themeda triandra and Pennisetum
spp., interspersed by a patchwork of Acacia woodlands and
bushlands, thicket and riverine forest (Lewis, 1934; Betts,
1953; Darling, 1960; Heady, 1960; Trapnell et al., 1969;
Trump, 1972; Taiti, 1973; Herlocker, 1976; Epp & Agatsiva,
1980; Lamprey, 1984). During the 1960s previously wooded
areas were radically transformed into grassland by the
actions of fire and elephants (Lamprey, 1984; Dublin,
1995). Hillslopes in the north remain colonized by extensive
tracts of Tarchonanthus camphoratus (Maa: ol-leleshwa)
bushland, impalatable to stock (Ivens, 1967; Pratt &
Gwynne, 1977).
MAASAI SETTLEMENTS AND LAND TENURE
The Maasai are pastoral people of Nilotic origin who moved
southwards into Kenya’s central rangelands in the seven-
teenth century7 (Berntsen, 1979; Waller, 1979; Ehret, 1984).
The Kenya Maasai are traditionally divided into 11 sections
(Maa: iloshon) occupying specific areas, and defined by
differences in dialect, culture and ceremonial procedures.
The Mara study area falls within the land unit of the il-
Purko section of the Maasai, who, since their relocation
from Laikipia by the colonial authorities in 1913 (Sandford,
1919; Lamprey & Waller, 1990), occupy northern and
central Narok District.
In common with other sections, the traditional il-Purko
Maasai homestead or boma8 (Maa sing.: enkang), comprises a
ring of low huts surrounding a central livestock coral
(Lamprey, 1984; Thompson, 2002). One or more families or
‘households’ (Maa pl: olmarei) form a semi-permanent core
group within this residential association, often bound together
by age-set and clan relationships (Thompson, 2002). Each
household head, usually a married man, ‘owns’ a gate in the
boma fence.9 The huts, consisting of a frame of poles covered
7
The reader is referred to Hollis (1905); Sandford (1919); Fosbrooke
(1948); Jacobs (1965) and Spear & Waller (1993) for a general de-
scription of the Maasai and their social structure, and to Berntsen
(1979); Waller (1979) and Ehret (1986) for analyses of the Maasai
‘migrations’ and resettlements. Nestel (1984), Bekure et al. (1991);
Homewood & Rodgers (1991) and Rutten (1992) describe the nutri-
tion, economics and ecology of Maasai pastoralism.
8
Bomas are often erroneously referred to as ‘manyattas’. ‘Manyatta’ is
the Maasai word for the (generally unfenced) settlements of the war-
riors (Maa pl.: il-murrani), but has reached common usage to refer to
any Maasai settlements with permanent or semi-permanent huts.
Jacobs (1965) prefers the terms ‘kraal camp’ for permanent or semi-
permanent settlements. We use the term ‘boma’ to remain consistent
with other academic studies of Maasai pastoralism (Bekure et al., 1991;
Homewood & Rodgers, 1991; Rutten, 1992).
9
A small number of households are headed by women. Data provided
by Rutten (1992) indicate that in six group ranches in eastern Maa-
sailand, 22 of 478 households, or 4.6%, were headed by women.
1002
with mud and dung daub, are located within the boma to left
and right of the gate. The huts (Maa sing: enkaji), which may
be considered as ‘sub-households’ (Thompson, 2002), are built
and maintained by the husband’s wives (Lamprey, 1984;
Homewood & Rodgers, 1991; Coast, 2000). Each night the
households bring their livestock through their respective gates
into central compartments of the boma, to protect them from
predators and stock rustlers. The pattern of the ‘traditional’
il-Purko Maasai boma, still common in the remote areas of
Narok District, is shown in Fig. 3.
The typical il-Purko boma in the Mara is occupied for
4–5 years (Lamprey, 1984), after which the settlement
becomes uninhabitable due to the build-up of dung and
parasites. Before evacuation, a new boma is constructed
nearby, in which some of the more durable building
materials of the old boma are reused. As soon as the old
boma has dried out, it is burned.10 Unlike transhumant
Maasai such as the il-Kisongo of Amboseli, who have wet
and dry-season bomas (Western, 1973), the il-Purko Maasai
of the Mara construct permanent bomas that are used year-
round, with little transhumance (Lamprey, 1984; Coast, 2000;
Thompson, 2002). Some small-scale maize cultivation may
be practised immediately around bomas, but this is not
generally a feature of ranch areas close to MMNR, due to the
risks of crop-raiding by wildlife.11 When grazing areas far
from the bomas are used, the Maasai herders construct
temporary livestock camps (Maa pl.: ormwati) as bases to
contain stock at night. These simply comprise a circular
fence and internal stock compartments, made from Acacia
bushes cut locally. Livestock camps are generally abandoned
after a single season, and the fence is soon destroyed by
termites or grass fires.
In the Mara, as elsewhere in Maasailand, bomas are
generally clustered in favoured localities (Maa pl: in-kutot),
where soil and drainage conditions are suitable (Western &
Dunne, 1979). As these ‘neighborhoods’ expand, trading
centres, schools, dispensaries and cattle dips are established,
and eventually these areas may become small townships.
Group ranches were established in the Mara in 1970. In the
northern plains, Koyake Dagurugurueti, Ol Kinyie and Lemek
Group Ranches were registered as ranches with group title,
whilst Ol Chorro Oirowa was registered as a private ranch (see
Fig. 2). As described later in this report, the Mara ranches are
now being subdivided into private land plots. Currently, the
main physical signs of this process are the fragmentation of
bomas into family units (see Fig. 4), and the placing of
boundary markers around land holdings.
10
A boma may also be abandoned if an important occupant dies.
11
Further from the MMNR both small-scale and large-scale cultiva-
tion constitute a major source of revenue to the Maasai, especially in
Lemek and on the northern Loita Plains. Groups of Maasai pool their
land to establish ‘farming associations’, whilst local elites controlling
much larger areas lease their land to commercial wheat farmers
(Lamprey, 1984; Norton-Griffiths, 1995; Ottichilo, 2000; Serneels &
Lambin, 2001; Thompson & Homewood, 2002).
Journal of Biogeography 31, 997–1032, ª 2004 Blackwell Publishing Ltd
 Expansion of human settlement in Kenya’s Maasai Mara

Figure 3 A Maasai boma at Talek in 1986,

based on the former communal pattern, with
central enclosure for cattle, and peripheral
enclosures for sheep and goats. Photograph
taken in the early morning with herds
departing for grazing through separate
household gates.

Figure 4 A Maasai boma at Oltorotua in

2002, with cattle departing for grazing. This
boma shows the present trend towards frag-
mentation into separate household units,
with permanent corrugated-iron huts (left
and centre).

145/1/2/3/4).12 The airphotos were interpreted using a stere-

METHODS
Human settlement distribution
Table 1 summarizes the sources of data for this study. In order
to determine changes in pastoral use of the area, settlements
were mapped over a period of 50 years using a variety of
methods. Early descriptions of settlement distributions in
southern Narok before 1960 are given in district and provincial
reports of the colonial administration. Within the study area,
Maasai bomas were mapped by interpretation of complete-
coverage aerial photography taken in 1950, 1961, 1967 and
1974. These photographs, of scales varying from 1:30,000 to
1:70,000, were originally taken to produce and update 1:50,000
scale topographic maps for the area (Y731 series, numbers
osope (·4 magnification) or, where stereo-pairs were not
available, using a binocular microscope (·4 and ·8 magnifi-
cation).
In all aerial photographs, resolution and scale was suffi-
cient to detect Maasai bomas, and to count the huts (Maa pl:
enkajijik) within them. No boma was located in the same
12
Negatives and prints of each set of photography were originally
stored at the Survey of Kenya in Nairobi. For this study, new prints
were produced for the years 1961, 1967 and 1974 for lab interpretation,
whilst interpretation of the 1950 photos took place directly at the
Survey of Kenya. In the late 1980s, negatives and prints of most sets of
photography were moved from the Survey of Kenya to the UK’s
Ordinance Survey in Southampton, England.

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R.H. Lamprey and R.S. Reid

Table 1 A chronology of data sources used for the study

Huts/ People/ Human Sheep/

Year(s) Bomas boma hut population Wildlife Cattle goats Woodlands NDVI Details

1950 TAP TAP TAP Aerial photography scale 1:30,000

1961 TAP TAP TC TAP Stewart & Talbot (1962) Aerial
photography scale 1:50,000
1967 TAP TAP TAP Aerial photography scale 1:57,000
1968–72 S(O) SRF Programme of Serengeti
Research Institute begins
(Pennycuick, 1975)
1974 TAP TAP TAP Aerial photography scale 1:68,000
1977 S(K) S(K) S(K) Start of SRF Programme by
KREMU/DRSRS (Ottichilo, 2000)
1978 S(K) S(K) S(K)
1979 CEN CEN S(K) · 11 S(K) · 11 S(K) · 11 1979 Population Census
1980 S(K) S(K) S(K)
1981 S(K) · 3 S(K) · 3 S(K) · 3
1982 S(K) · 3 S(K) · 3 S(K) · 3
1983 SGR SAP SGR S(K) S(K) S(K) SGR, MVC Sample ground survey (Lamprey,
SAP 1984) Sample aerial photography,
scale 1:6000, 21% systematic
coverage
1984 S(K) S(K) S(K) MVC
1985 S(K) · 2 S(K) · 2 S(K) · 2 MVC
1986 S(K) · 3 S(K) · 3 S(K) · 3 MVC
1987 S(K) S(K) S(K) MVC
1988 MVC
1989 CEN CEN S(K) S(K) S(K) MVC 1989 Population Census
1990–97 S(K) S(K) S(K) MVC
1999 REC, SAP CEN CEN MVC Aerial survey,
distribution all bomas.
1999 Population Census
2000 S(K) S(K) S(K) MVC
2001 SAP/B SAP/B MVC Sample aerial
photography of bomas
2002 TGR TGR TGR TGR TGR

TAP, total count from aerial photography; TC, total count; S(O), systematic reconnaissance flight (SRF) by Serengeti Research Institute, analysis
based on ‘occupance’ of grid cells (Pennycuick, 1975); S(K), systematic reconnaissance flight (SRF) by Kenya Rangeland Ecological Monitoring Unit,
now DRSRS (see text); S(K), SRF carried out but data not available; SGR, sample ground count; SAP, sample, derived from strip-sample aerial
photography; SAP/B, sample photography bomas only; REC, aerial reconnaissance flight; TGR, total ground count; MVC, maximum value composite
NOAA-AVHRR NDVI data; CEN, national population census.

spot in consecutive years of photography, indicating that the
period of occupancy is less than 6 years (the minimum inter-
photo period). Abandoned il-Purko bomas are burned after a
3–4-month period of drying-out. Assuming a mean period of
occupancy of 4 years, some 6–7% of bomas in the airphotos
are unoccupied prior to burning. With their distinctive circle
of flat-roofed huts, the bomas were easily distinguished on
airphotos from the seasonal livestock camps which have no
huts, or only small shelters. From the interpretation, bomas
were mapped onto 1:50,000 base maps for digitizing.
After 1974 there is no complete coverage aerial photography.
From 1983 onwards (the start of our fieldwork period)
settlement distributions were determined from aerial observa-
tions, oblique aerial photography and systematic ground
surveys. As part of our study, the Kenya Rangeland Ecological
Monitoring Unit (KREMU)13 conducted a photo-mission in
1983 to acquire vertical panchromatic and infra-red photo-
graphy at scale 1:6000 scale along sample strips spaced 10 km
apart across the study area. This survey constituted a 21%
systematic sample of Koyake Group Ranch. Bomas ‘captured’
in the strips were used as the sample for determining huts/
boma in 1983.
In January and August 1999, systematic aerial surveys were
carried out by light aircraft at 13,500 m a.s.l. (approximately
8000 feet above ground level) to map all bomas onto 1:50,000
maps, and to obtain estimates of huts/boma. In September and
13
In the late 1980s, KREMU became the Department of Resource
Surveys and Remote Sensing (DRSRS), which was recently integrated
into the National Environment Management Authority (NEMA).

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December 2001, a sample of bomas was photographed from
the air at first light to estimate livestock/hut ratios (see next
section). As part of this exercise huts were counted from the
photos to crosscheck hut/boma estimates. Meanwhile, in
separate exercises, detailed ground counts were made of
settlements, wildlife and livestock in November 1999 for part
of Koyake and the MMNR (Reid et al., 2001), and in
November 2002 for a larger area including most of Koyake,
other adjacent group ranches and MMNR (Reid et al., 2003).
These data supplemented those of this study.
Livestock camps are clearly visible in vertical aerial photo-
graphy, and they were interpreted and mapped from all of the
complete-coverage airphoto series (1950–74). The 1983 verti-
cal aerial photography provided a 21% systematic sample of
livestock camps for estimating total camp numbers across
Koyake Group Ranch. However, mapping livestock camps
from oblique views on the ground or air proved problematic,
as the camps blend with adjacent thickets and bushlands. By
1983 the density of camps had significantly declined across the
ranch (see section ‘Expansion of Maasai settlement’), and with
the mapping problems described above, they were not
systematically counted after this year. However, as a subsidiary
exercise of the 2002 count, an intensive aerial search was made
for camps in a 96 km2 sample area in the Ilbaan woodlands
near Talek. Camp densities for this specific area could then be
compared with all previous years of photography to determine
trends.
In our study, the seven years of settlement distribution data
(1950, 1961, 1967, 1974, 1983, 1999 and 2002), serve as
‘reference years’ for the analysis of change in the Mara
ecosystem over the last half-century. Population estimates
derived from the airphotos and ground surveys were compared
with national census data of 1962, 1966, 1969, 1979, 1989 and
1999 (Morgan and Shaffer, 1966; CBS, 1979, 1989, 1999).
Whilst we recognize that the Maasai are notoriously difficult to
census, and that enumeration areas change from one census to
the next, these census data were useful for comparison with
data from the settlement counts.
Estimates of hut occupancy (people/hut) were derived
from interviews in 64 huts at Talek in 1983 (Lamprey, 1984).
These estimates were compared with those obtained in more
recent studies in the Mara and elsewhere in Maasailand
(Coast, 2000).
Wildlife and livestock populations and distribution
The first estimates of wildlife numbers in the ecosystem were
obtained from aerial total counts on either side of the Kenya–
Tanganyika border in the 1950s and early 60s (Pearsall, 1957;
Swynnerton, 1958; Grzimek & Grzimek, 1960; Stewart &
Talbot, 1962). Following the development of the SRF
methodology at the Serengeti Research Institute (SRI) (Jolly,
1969; Norton-Griffiths, 1978, 1986; ILCA, 1981), monthly or
seasonal wildlife censuses covering virtually the entire ecosys-
tem were conducted by SRI from 1969–73 (Pennycuick, 1975).
After 1973, SRFs of the Serengeti were conducted on a more
Journal of Biogeography 31, 997–1032, ª 2004 Blackwell Publishing Ltd
Expansion of human settlement in Kenya’s Maasai Mara
opportunistic time-frame (Campbell & Borner, 1995), and
cross-border counts could not easily be coordinated.
Early livestock counts in the Mara (for example from a
veterinary survey of 1960) require cautious interpretation, as
livestock numbers cannot easily be referenced to particular
localities. Useful livestock distribution data for the Mara
rangelands are only available from 1977 onwards, when
KREMU began a programme of seasonal SRF surveys of
wildlife and livestock in Narok District. The KREMU census
zone covered the entire rangeland areas of Narok District and
(later) Trans-Mara District (Stelfox & Peden, 1981; Broten &
Said, 1995; Ottichilo, 2000). Previous analyses of these data sets
have been stratified on the basis of ‘ecological units’ (Stelfox
et al., 1980)14 or for all group ranches combined (Broten &
Said, 1995; Ottichilo, 2000). We analysed raw KREMU SRF data
from 1979 to 2000 using Jolly’s ‘Method II’ (Norton-Griffiths,
1978) to obtain population estimates specifically for Koyake
Group Ranch (see Table 1). To check consistency of our Jolly II
analysis with other studies, population estimates were also re-
calculated for all ranches. We compare the Koyake data with
total ground counts of wildlife and livestock conducted in 1999
and 2002 that covered most of the MMNR and Koyake Group
Ranch (Reid et al., 2001, 2003).
To further reassess the accuracy of KREMU data, aerial
photographic surveys were conducted of a systematic sample
of bomas in Talek, Olare Orok and Mara Rienda in September
2001 (dry season) and December 2001 (wet season). The
purpose was to count cattle in bomas before they had left for
grazing, and, by extrapolation of cattle/hut ratios, to recalcu-
late the cattle population for the entire Koyake ranch as a
comparison with SRF data.
Rainfall and primary production
Rainfall has been monitored sporadically at a number of sites
throughout the Mara, but the most reliable and long-term
records are gathered at Narok, with monthly rainfall recorded
since 1914. These data were obtained from the Kenya
Meteorological Department.
Vegetation growth conditions were determined using Nor-
malized Difference Vegetation Index (NDVI) data, derived
from the Advanced Very High Resolution Radiometer (AV-
HRR) sensor on board the NOAA (National Oceanic and
Atmospheric Administration) series of satellites (Justice et al.,
1986). NDVI data are highly correlated with rainfall (Richard
& Poccard, 1998) and resultant vegetation production (Prince,
1991). For this study, NDVI data were obtained as decadal
(10 days) ‘maximum value composite’ data from sources on
the Internet, for the period 1983 to present.15 For assessment
14
In the analysis of Stelfox et al. (1980), the ‘Mara ecological unit’
combined the MMNR with Koyake; data for Koyake could not be
disaggregated from their data sets.
15
There is however, a critical gap in 1991/92, where NDVI values were
suppressed by atmospheric aerosols deriving from the eruption of Mt
Pinatubo (the ‘Pinatubo effect’).
1005
R.H. Lamprey and R.S. Reid

1.100

1.000
5-Year average of rainfall (mm)

900

800

700

600
Figure 5 Long-term rainfall at Narok,
1919–99, indicated as a 5-year cumulative
500 average. Redrawn from Lamprey (1984) with
1919 1924 1929 1934 1939 1944 1949 1954 1959 1964 1969 1974 1979 1984 1989 1994 1999
recent data from Kenya Meteorological
Year Department.

of growing conditions in Koyake group ranch, a cluster of nine 40

pixels were averaged covering the central grazing area of the 35 Enkikwe

ranch. 30 Talek

25
Cover (%)
Ol Keju
Ronkai
RESULTS 20

15
Expansion of Maasai settlement 10

Lamprey & Waller (1990) describe pastoral movements and 5

ecological changes in the Mara in the years following the great
rinderpest pandemic of the 1890s, which devastated both wild
and domestic ruminants. Their livestock herds decimated,
many Maasai died or took refuge with agricultural neighbours.
In the early years of the twentieth century, the Mara was only
used by Okiek (‘Dorobo’) hunter-gatherers. Without regular
burning and browsing, Acacia bushland began to regenerate
across the area, bringing tsetse-flies and trypanosomiasis
(Woosnam, 1913; Lewis, 1934; Ford, 1971).
In 1913 il-Purko Maasai living in the ‘Northern Maasai
Reserve’ in Laikipia were resettled by the colonial administra-
tion in an enlarged Southern Maasai Reserve that included the
rangeland and highland areas of Narok (Sandford, 1919;
Lamprey & Waller, 1990). Some il-Purko Maasai moved into
the Mara settled at Talek – there are accounts of German cattle
raids on Maasai bomas at Talek during the First World War
(Sandford, 1919, p. 121) – but they were soon forced
northwards into the Lemek Valley by the tsetse expansion
(Lewis, 1934; Beaumont, 1945). By the mid-1920s, Maasai
sectional areas had become more firmly established. The il-
Purko inhabited the Lemek Valley (the only part of the Mara
free of ‘fly’), the Loita Plains and parts of the Mau foothills.
The Siria Maasai returned to occupy the area above the Siria
escarpment16 and the Loita, weakened by conflict, withdrew
into the Loita hills to the east (Lamprey & Waller, 1990).
16
Following the rinderpest epidemic, many of the Siria Maasai had
taken refuge with Luo communities to the west, or with Okiek hunter-
gatherers in the forests.
0
1945 1955 1965 1975 1985
Year
Figure 6 Changes in woodland cover in three areas of the Mara
from 1950–83, measured by dot-grid analysis over five sample
points in each area (from Lamprey, 1984): These areas are (a)
Enkikwe (Acacia gerrardii woodland) – note woodland recovery
during the 1980s, (b) Talek ( Acacia–Commiphora woodland) and
(c) Ol Keju Ronkai inside the reserve (assumed formerly to be
Acacia woodland, now completely cleared to grassland).
The main factors affecting the pattern of Maasai occupation
of the Mara over the twentieth century were climatic
conditions and the presence of tsetse-flies (Lamprey & Waller,
1990). Tsetse-infested bushlands extended as the ‘fly-belt’
through much of the area corresponding to the MMNR. The
following discussion makes constant reference to rainfall and
trends in bushland cover. Figure 5 shows a 5-year cumulative
average of rainfall at Narok, for the period 1914–99. The figure
shows a period of particularly high rainfall during the late
1950s and early 1960s, the effect of which is discussed below.
Figure 6 shows trends in woodland cover in the Mara from
1950 to 1983 (Lamprey, 1984). The removal of bushland and
tsetse opened up the Mara to a rapid expansion of human
settlement.
The 1950 aerial photography revealed the continued
confinement of the Mara il-Purko Maasai to the Lemek
Valley (see Fig. 7). Although all bomas were located in the
valley, the livestock camp distribution indicates that the

1006 Journal of Biogeography 31, 997–1032, ª 2004 Blackwell Publishing Ltd


Figure 7 Bomas (black circles) and
temporary livestock camps (open circles) in
1950, mapped from aerial photography. At
this time the Mara National Reserve,
managed by Kenya Royal National Parks,
was located to the west of the Mara River,
in the Mara triangle.
Maasai dispersed some stock into the plains south and west
of Aitong, presumably on a seasonal basis. Meanwhile, in the
late 1950s, as pressure on grazing resources increased in the
Lemek Valley, the administrative and veterinary authorities in
Narok District embarked on a programme to expand the
available grazing area further south into the Mara woodlands.
Research was conducted on tsetse, game and pasture at
Aitong and Talek17 (Langridge et al., 1970), and bush was
cleared experimentally in trial areas (Narok District Devel-
opment Plan, 1955).
In the late 1950s, the Maasai intensified their own efforts
to remove tsetse, by systematically burning the woodlands.
Conditions over the period 1955–61 were particularly
favourable for this activity, as rainfall had greatly increased
(see Fig. 5) and grass production was high. During this
period, the grasslands could be burned as often as twice
within a season (Langridge et al., 1970). As a result,
woodlands were opened up across the Mara (Lamprey,
1984; Dublin, 1995). The decline was accelerated by elephant
activity. As areas outside the Serengeti-Mara became increas-
ingly settled in the 1950s and 60s, elephants moved into the
ecosystem, in large numbers, inflicting heavy damage on
woodlands (Game Department Annual Report, 1951; Lam-
prey et al., 1967; Glover & Trump, 1970; Langridge et al.,
1970).
17
Tsetse species found in the Mara are Glossina swynnertoni and
G. pallidipes (Lewis, 1934; Beaumont, 1945; Langridge et al., 1970). At
Talek in 1959, 4% of the tsetse flies caught were found to be carriers of
Trypanosoma congolense and T. vivax (Whiteside & Langridge, 1959).
Journal of Biogeography 31, 997–1032, ª 2004 Blackwell Publishing Ltd
Expansion of human settlement in Kenya’s Maasai Mara
In the northern Mara the destruction of vegetation was
monitored by the Tsetse Survey and Control Programme
(Langridge et al., 1970). In 1959 the programme marked 100
trees (referred to as ‘tsetse resting places’) in each of four
vegetation plots at Talek to assess change. The main species
marked were Grewia spp., Cordia ovalis, Acacia spp. and
Commiphora spp. They returned to assess the plots in 1963,
and reported that ‘between 1961 and 1963 rainfall was very
heavy throughout the district and produced a luxuriant grass
cover, which allowed bi- and tri-annual burning…. The very
fierce fires caused accelerated destruction of the vegetation
with marked changes in numbers of trees and shrubs. In plot
no. 1 only 53 of the original resting places were found again,
and of these most had suffered extensive damage. In plot no. 2,
only 31 of the original marked places remained. In plot no. 3,
only six of the original resting places were found… The
vegetation changes in the (Ilbaan) escarpment woodland were
due more to the actions of elephants than fire. In plot no. 4, 24
resting places had been smashed down to the ground and 16
had been burnt’ (Langridge et al., 1970, pp. 208–209). Thus
between 1959 and 1963, 47–94% of trees and shrubs were lost
in the study plots.
As woodlands were cleared, tsetse disappeared from much
of the Mara (Darling, 1960; Talbot & Talbot, 1963; Glover &
Trump, 1970; Langridge et al., 1970), and in 1963 the Tsetse
Survey and Control Programme was closed (Lamprey &
Waller, 1990). By the early 1960s the Maasai were able to
extend their livestock camps as far south as the Talek river,
Ol Keju Ronkai stream and the Posee plain, whilst bomas
were established at Aitong and Bardamat (see Fig. 8). The
1007
R.H. Lamprey and R.S. Reid
southwards expansion of the Maasai that began in the late
1950s was spearheaded by a wider dispersal of smallstock.18
Of the Meta Plains between Keekorok and Talek, Talbot
et al. (1961, pp. 30–31) reported that ‘there have been
increasing numbers of goats and sheep grazed in the area in
tightly packed herds numbering over 2000 animals. The fine
condition of this area is due primarily to the tsetse fly which
has kept Masai cattle out for 50 years’.19 Grimwood (1960,
p. 26) recorded that ‘sheep and goats were being grazed
further and further into the fly zone, and grass burning,
which had been carried out to improve grazing was
progressively destroying the bush, pushing back the tsetse
fly and allowing cattle to follow on the heels of the smaller
stock.’
In 1961 the MMNR, formally comprising only the ‘Mara
Triangle’20 was re-declared a ‘District Council Game Reserve’,
and was greatly enlarged to take in the plains east of the Mara
River. The new Maasai Mara Game Reserve (MMGR) now
extended eastwards to the Sianna hills, and southwards to the
18
Sheep and goats are more tolerant to trypanosomiasis than cattle
(Whitelaw, 1983), and may therefore be taken into tsetse areas during
the dry season, when fly densities are lower (Challier, 1982).
19
Lee and Martha Talbot pioneered the first wildlife research in the
Mara in the early 1960s, and conducted a landuse survey of the district
in the late 1950s.
20
The Mara Triangle is the area of MMNR bordered on the east by the
Mara River, to the west by the Siria escarpment, and to the south by
the Tanzania border.
1008
Figure 8 Bomas (black circles) and
temporary livestock camps (open circles) in
1961, mapped from aerial photography. In
the 1950s the Mara National Reserve was
managed by the Game Department as the
Maasai Mara Game Reserve (MMGR), and
in 1961 it was ‘transferred’ to Narok
District Council and designated a
‘county-council reserve’.
Tanzanian border. Under early agreements between the Game
Department and the Maasai District Council, Talek was
designated a ‘grazing area’ within the MMGR (Game Depart-
ment Annual Report, 1961; Lamprey, in prep.). Presumably,
this definition did not exclude settlement, for the 1967 aerial
photographs show a single large boma at Talek, the first in this
area for 50 years (see Fig. 9).
The Maasai population in the Mara increased rapidly after
group ranch registration in 1970. By 1974, there were 10 bomas
at Talek (see Fig. 10), and the settlement localities of Lemek,
Aitong, Koyage, Ol Kinyie, Talek and Ol Doinyo Narasha were
now well-established. As large areas in the Narok highlands
were sold or leased to outsiders for wheat cultivation during the
1970s and 80s, many Maasai families were displaced into the
more marginal ranches closer to MMNR (Duraiappah et al.,
2000). By 1983 there were some 27 bomas at Talek, and a new
locality had been established at Mara Rienda by immigrants
from Mau Narok (Fig. 11). By 1999, virtually every part of
Koyake within reach of permanent water had been occupied
(Fig. 12).
Figure 13 shows changes in the density of livestock camps
in Koyake as a whole, and in a sample area in the Ilbaan
woodlands near Talek. A significant feature is the great
increase in camps in the 1960s, associated with the southward
wave of dispersal, followed by a decline in camp density after
1974. With increasing sedentarization of the Maasai, most
grazing areas were now within daily grazing range of the
localities of Aitong, Talek and Mara Rienda. Furthermore, in
many parts of the central and northern plains, little woody
Journal of Biogeography 31, 997–1032, ª 2004 Blackwell Publishing Ltd
 Expansion of human settlement in Kenya’s Maasai Mara

Figure 9 Bomas (black circles) and tem-

porary livestock camps (open circles) in 1967,
mapped from aerial photography. The first
boma was established at Talek in 1966–67.

Figure 10 Bomas (black circles) and tem-

porary livestock camps (open circles) in 1974,
mapped from aerial photography. Note the
increasing number of bomas at Talek, and
the reduced number of temporary livestock
camps, particularly in the western areas.
The map shows the boundaries of the
group ranches (GRs), newly established in
1970. In 1973 the Maasai Mara Game Reserve
was re-gazetted under the Wildlife Animals
Protection Act, 1964, but continued to be
managed by the Narok County Council.

vegetation remained with which to build temporary camps. wildebeest. It is only in Ilbaan and Olare Sambu in the south
Western areas towards the Mara River were avoided by the that livestock camps remain a significant feature. Some
Maasai in the dry season, as these were favoured by migratory Acacia–Commiphora bushland (and tsetse) remained here,

Journal of Biogeography 31, 997–1032, ª 2004 Blackwell Publishing Ltd 1009

R.H. Lamprey and R.S. Reid

Figure 11 Bomas in 1983, mapped from the

ground and from aerial reconnaissance
flights. Temporary livestock camps were
scattered at very low density by this time,
and were not mapped (see text). In 1976,
the ‘Maasai Mara Game Reserve’ was redes-
ignated the ‘Maasai Mara National Reserve’
(MMNR) under the Wildlife (Conservation
and Management) Act, 1976; the Act speci-
fied that districts shall continue to manage
their former ‘county-council’ game reserves.
The map shows the excisions from MMNR
at Talek and Mara Rienda; these sections
were officially degazetted in 1984.

Figure 12 Bomas in 1999, mapped from

aerial reconnaissance flights. Temporary
livestock camps were not mapped (see text).

1010 Journal of Biogeography 31, 997–1032, ª 2004 Blackwell Publishing Ltd


0.25
Koyake
0.20 Ilbaan
Livestock camps km–2
0.15
0.10
0.05
0.00
1945 1955 1965 1975 1985 1995
Year
Figure 13 Changes in density of livestock camps in Koyake
GR from 1950 to 1983, and in a sample area at Ilbaan (96 km2)
from 1950 to 2002.
wildebeest avoided the more rocky terrain, and the area
remained useful for the seasonal dispersal of sheep and
goats.21
In general, bushlands in the Mara have continued to decline,
the canopy cover being reduced from 30–40% cover in 1961 to
0–5% cover in 1983 (see Fig. 6). However, in 1983 in Enkikwe
there was some recovery of woodlands, dominated by single-
species stands of Acacia gerrardii. This recovery can be
attributed to increased grazing in the area by Maasai livestock,
and the resulting suppression of fire due to reduction in
herbaceous biomass. Inside the MMNR frequent fires continue
to open up remaining thickets.
Maasai population and distribution
Bomas and huts can be clearly interpreted on aerial photo-
graphs at the scale available for this study. However, airphotos
provide no useful information on households, the primary
unit for socio-economic analysis (Coast, 2000; Thompson,
2002), which may comprise several huts.22 Therefore, we
determine human population trends in Koyake on the basis of
people per hut, huts per boma, and total bomas in the group
ranch.
Table 2 shows estimates for people per hut, people per
household and population age structure from various esti-
mates in Maasailand, as recorded through field assessments
and population censuses. In our study, the number of people
21
It might also be suggested that as livestock camps were used pri-
marily for the dispersal of sheep and goats, the decline in camps reflects
a decline in sheep and goat numbers. Talbot et al. (1961) record the
presence of many herds numbering over 2000 animals on the Meta
plains and at Talek. Herds of this size are very rarely encountered
today. However, there is no reliable information available on small-
stock numbers before KREMU SRFs began in 1977.
22
The main physical representation of the ‘household’, the gate in the
boma fence, could only just be resolved on the 1950s photos of scale
1:30,000 (the largest scale of complete-coverage airphoto series), but
interpretation was not consistent due to variations in image quality.
Journal of Biogeography 31, 997–1032, ª 2004 Blackwell Publishing Ltd
Expansion of human settlement in Kenya’s Maasai Mara
per hut was determined from a sample size of 64 huts in four
bomas at Talek in 1983 (see Fig. 14). This gave a mean of
4.61 people/hut which is close to the estimate of 4.64 people/
hut given in the 1999 national census for Koyake (see Table 2),
which carefully specified the hut as the ‘household’, whilst
recognizing the potential absence of the household head on
Census Night.23 Thus in 1999 the average boma of 7.78 huts
will accommodate an estimated 35.9 people, closely compar-
able with the estimate of 36.1 calculated from the data of Coast
(2001). Figure 13 also shows ‘reference adults’ (RA) per hut,
2005
on the crude estimation that children under 15 are measured
as 0.7 RA. Using this estimate (see section ‘Pastoral Require-
ments on Koyake’), total RA may be calculated as 0.8 · pop-
ulation, as observed in examples from Rutten (1992) and
McCabe et al. (1997).
Figure 15 presents frequency histograms of huts per boma
over the period 1950–99. In the Mara boma size remained
relatively stable at 10–12 huts from 1950 to 1983. The marginal
increase in boma size in the 1974 photography is probably due
to the immigration of Maasai families immediately after group
ranch registration, and their inclusion into new bomas.
However, by 1999 bomas had decreased in size to a mean of
7.78 ± 1.168 huts/boma (95% CL, n ¼ 79), with 1–4 huts/
boma as the class with the highest frequency.24 Figure 16 shows
the trend in boma size from 1950 to 1999. Today in the Mara,
communal bomas are disappearing, and the family ‘home-
stead’ of 1–2 households is emerging as the new form of
settlement. By November 2002, the number of huts per boma
in Koyake had fallen further to an average of 6.3 (Reid et al.,
2003).
In this study, as in others in Maasailand, the ‘household’
(Maa sing: olmarei) is defined as all those who use the gate of
the household head (usually the husband), including wives,
children, elderly dependents and any dependent brothers and
wives on the husbands side (Bekure et al., 1991; Homewood &
Rodgers, 1991; Coast, 2000). An analysis of hut/gate ratios at
Talek in 1982 gave an estimate of 10.42 people per olmarei, or
3.4 olmarei per boma (Lamprey, 1984). This may be compared
23
In the Kenya Population Censuses of 1989 and 1999, the term
‘household’ is directly analogous to people/hut in pastoral polygynous
societies such as the Maasai, as the census instructions to enumerators
clearly indicate: ‘In a polygamous marriage, if the wives are living in
separate dwelling units, and cook and eat separately, treat the wives as
separate households. Each wife with her children will therefore
constitute a separate household. The husband will be listed in the
household where he will have spent the Census Night’ (see CBS, 1989,
1999, Enumerators Instruction Manual, Section 32, p. app1–8). In
censuses before 1989, it was clear that each wife would automatically
count the male household head as part of her household, even if he was
elsewhere. Thus people/hut estimates in the 1979 census are higher
than estimates from other sources (see Table 2). People/hut estimates
were still high in 1989, but it is reasonable to assume that as this was
the first census in which the new rules above were applied, this was due
to enumeration difficulties. Apparently greater care was taken in the
1999 census to avoid this bias (see CBS, 1999).
24
The 2001 boma photography gives an estimate of 7.43 ± 1.589 huts/
boma (95% CL) for a smaller sample of 21 bomas.
1011
R.H. Lamprey and R.S. Reid

Table 2 Estimates of hut occupancy, household size and population parameters in Maasailand from various sources

People per household Population

Area/ranch/section Source People per hut (household ¼ hh) £ 15 years old (%)

Ngorongoro (Tanzania) Arhem et al. (1981) 4.14 (n ¼ 28 huts) 52.6

Ngorongoro NCAA (1999) 4.55 (n ¼ 11,354 huts) 7.70 (n ¼ 6708 hh) 53.0
Samburu (Kenya) Spencer (1973) 6.75
Kaputei high potential Nestel (1986) 4.8
(Kenya)
Kaputei low potential Nestel (1986) 4.4
Kaputei Bekure et al. (1991) – 10.78 (n ¼ 326 hh)
Kaputei Rutten (1992) 6.66 (n ¼ 221 hh) 47.0
Loodokilani Rutten (1992) 10.51 (n ¼ 75 hh) 55.1
Matapato Rutten (1992) 10.02 (n¼204 hh) 51.0
Shompole (Kenya) Coast (2001) 9.6
Lemek/Osopuko Narok District – – 52.5
Annual Report (1950)
Koyake/Lemek CBS (1979) 5.6* – –
Koyake/Lemek CBS (1989) 5.52* (n ¼ 1675 huts) – 54.7
Koyake/Lemek CBS (1999) 4.64* (n ¼ 1485 huts) – 53.4
Lemek Lamprey (1984) 7.69 ± 0.853 (95% CL)
(n ¼ 54 gates)
Talek Lamprey (1984) 4.61 ± 0.422 10.52 ± 0.973 (95% CL) 56.0
(95% CL) (n ¼ 64 huts) (n ¼ 34 gates)
Talek Coast (2001) 12.9

*Huts (Maa sing: enkaji) are defined as ‘households’ in the national censuses of 1989 and 1999 (see text).
Data recalculated from Rutten’s (1992) Table 8.2, averaged for three Kaputei group ranches (Olkinos, Embolioi, Kiboko), one Loodokilani group
ranch (Elang’ata Wuas) and two Matapato group ranches (Lorngosua, Meto).

Table 3 indicates the number of bomas, huts, households

Figure 14 Histograms of persons and reference adults/hut in
64 huts at Talek in 1983 (Lamprey, 1984). A crude estimate of
‘reference adults’ (RA) was derived by recording children under
15 years as 0.7 · RA (see text). This gives a mean of 3.84 ±
0.338 RA per hut (95% CL).
with the 12.9 people per olmarei and 2.8 olmarei per boma
specifically for the Talek area indicated in recent research by
Coast (2001).25 Again it is inferred that at Talek boma size
(olmarei per boma) is decreasing, whilst household size is
increasing.
25
Lamprey (1984) recorded that in the crowded environment of the
Lemek Valley, the average household comprised 7.69 ± 0.853 people
(95% CL).
and estimated population of the study area and Koyake, from
1950 to 1999, whilst Fig. 17 graphically indicates the increase in
huts for Koyake only, and includes the 2002 ground count
estimate. Applying the equation for mean annualized growth
rate (Preston et al., 2001), we calculated that the human
population in Koyake has increased at the rate of 7.6% per
annum over the period 1950–99. From 1983 to 1999 the
annual growth rate has been 4.4%, resulting in a doubling of
the population every 15 years.
In general, the human population estimates obtained in our
aerial reconnaissance in 1999 are supported by other sources.
Our estimates of 1828 huts in 235 bomas are similar to those
obtained in the ground counts of 1999 and 2002 (Reid et al.,
2001, 2003), making allowance for the different extents of the
ground counting areas. The 1999 ground count omitted the
eastern part of Koyake, but instead included Ol Chorro Oirowa
and parts of Lemek GR. Within this census zone 1507 huts
were counted in 246 bomas. A possible explanation for the
small difference with the 1999 aerial counts is that observers
may have had difficulty in interpreting the boma as a distinct
unit, when many bomas are breaking up into smaller, but
contiguous units (see Fig. 4). The 2002 ground count
estimated 2234 huts in 295 bomas in the 90% of Koyake that
was counted (Reid et al., 2003).
Our population estimates are also supported by those
obtained in national population censuses, although careful

1012 Journal of Biogeography 31, 997–1032, ª 2004 Blackwell Publishing Ltd

 Expansion of human settlement in Kenya’s Maasai Mara

35 35

30 Year 1950 30 Year 1961

25 Mean huts/boma = 10.97 25 Mean huts/boma = 10.66

S D = 6.12 S D = 5.32
Frequency (%)

Frequency (%)
20 n = 33 20 n = 71

15 15

10 10

5 5

0 0
1–4 5–8 9–12 13–16 17–20 21–24 25–28 29–32 33–36 1–4 5–8 9–12 13–16 17–20 21–24 25–28 29–32 33–36
Huts per boma Huts per boma

35 35

30 Year 1967 30 Year 1974

Mean huts/boma = 10.27 Mean huts/boma = 11.78

25 25
S D = 6.76 S D = 6.09
n = 94 n = 124

Frequency (%)
Frequency (%)

20 20

15 15

10 10

5 5

0 0
1–4 5–8 9–12 13–16 17–20 21–24 25–28 29–32 33–36 1–4 5–8 9–12 13–16 17–20 21–24 25–28 29–32 33–36
Huts per boma Huts per boma

30 40

Year 1983 35 Year 1999

25
Mean huts/boma = 10.72 30 Mean huts/boma = 7.78
S D = 6.94 S D = 5.22
20
n = 29 (sample) 25 n = 79 (sample)
Frequency (%)
Frequency (%)

15 20

15
10
10
5
5

0 0
1–4 5–8 9–12 13–16 17–20 21–24 25–28 29–32 33–36 1–4 5–8 9–12 13–16 17–20 21–24 25–28 29–32 33–36
Huts per boma Huts per boma

Figure 15 Frequency histograms of huts/boma in the Mara study area, for all years of settlement counts. The trend is towards bomas
with fewer houses.

14 interpretation is required due to changes in the enumeration

Mean
13 areas between census years. A census in 1950 revealed that the
Mode
12 ‘Lemek subsection of the Purko’ totalled some 3900 people
11 (Narok District Annual Report, 1950). However, as the Purko
10
settlement area extended as far east as Ololunga (well outside
Huts/boma

the study area), probably half of this total, or 1900 Maasai

9
actually lived in the Lemek Valley. In a ground count in 1945,
8
Beaumont (1945) estimated 50 bomas in the Lemek Valley,
7 including that part lying outside the study area to the east.
6 From the 1950 airphotos, 32 bomas were counted in the Lemek
5 Valley (study area part), giving an estimate of 2100 people. In
4 the 1999 population census, Koyake Group Ranch incorpor-
1940 1950 1960 1970 1980 1990 2000 2010 ated two enumeration areas, Koyake and ‘Mararianda’ (Mara
Year Rienda), totalling 6889 people, and part of the Aitong
enumeration area with 2905 people (CBS, 1999). The 1999
Figure 16 Decline in boma size (huts per boma) from 1950 to estimate of 8500 people for Koyake Group Ranch, derived
1999. The error bars give the 95% CL of the mean, and the from the aerial settlement count, is therefore representative
dotted line the mode. The 2002 data point is from the 2002
(Table 3).
ground count (Reid et al., 2003).

Journal of Biogeography 31, 997–1032, ª 2004 Blackwell Publishing Ltd 1013

R.H. Lamprey and R.S. Reid

Table 3 Bomas, huts, households and population estimate for Koyake, 1950–99

Year Bomas in Bomas in Mean huts Total huts, Households Estimated

study area Koyake* per boma ± 95% CL§ Koyake Koyake populationà

1950 32 4 10.97 ± 2.169 44 19 202

1961 62 25 10.66 ± 1.259 267 118 1229
1967 90 27 10.31 ± 1.391 278 123 1283
1974 107 50 11.78 ± 1.082 589 261 2715
1983 195 84 10.72 ± 2.639 900 398 4151
1999 531 235 7.78 ± 1.168 1828 809 8428

*Koyake did not exist before 1970; this refers to the corresponding area.
Based on people/household (‘family’) at Talek ¼ 10.42 ± 0.605 (95% CL) (see text).
àBased on people/hut ¼ 4.61 ± 0.422 (95% CL) (see Table 2).
§Although 95% CL are indicated, all huts per boma distributions are skewed (see Fig. 15). Differences in years are best tested by the non-parametric
Kruskal–Wallis test, where X2 ¼ 26.3799, d.f. ¼ 5, P < 0.001.

2400 2,234
of Trans-Mara was so tsetse-infested as to be unusable for cattle
grazing (Humphrey, 1947). The next veterinary census con-
2000
1,828 ducted in 1960 by the Agricultural and Veterinary Department
(see Simon, 1963) gives an estimate of 191,000 cattle for the
Number of Huts

1600
1200
900
800 589
267
400
44 278
0
1945 1955 1965 1975 1985
Year
Figure 17 Increase in number of huts in Koyake Group Ranch
(or equivalent area before ranch registration in 1970), for the
period 1950–2002. The 2002 data point is from the ground survey.
Trends in livestock populations
The earliest cattle population estimate in the study area is
provided by Sandford (1919, p. 103), who describes the
development of dams in Lemek Valley in 1914–16, at which
‘10,000 cattle were watered daily’.26 It is known that many cattle
died during catastrophic droughts and locust ‘visitations’ during
the early 1930s (Lamprey & Waller, 1990), but a cattle census in
1939 revealed that across Narok District the cattle population
had increased to 180,000, of which 87,000 were owned by the il-
Purko (Narok District Annual Report 1939).27 At this time much
26
According to Sandford (1919), in 1917 the il-Purko Maasai, with a
population of 16,500 people, owned 375,000 cattle and over 1,000,000
sheep/goats. These estimates, if they can be relied on (see Broch-Due &
Anderson, 1999), indicate that with 22 cattle per capita, the il-Purko
were remarkably wealthy in comparison with the situation today (see
Section ‘Pastoralism as a livelihood on Koyake’). It is extraordinary
that cattle herds had increased to this extent just 27 years after the
rinderpest pandemic. Cattle raiding must have been a major factor in
il-Purko restocking.
27
Given that the il-Purko population was estimated at 10,500 in 1950
(Narok District Annual Report, 1950), it would appear that, with eight
cattle per capita, the il-Purko were still moderately affluent (see Section
‘Pastoralism as a livelihood on Koyake’).
census areas of ‘Sianna-Talek’ and the ‘Loita Plains’, which
correspond with the ranches of Koyake, Ol Chorro Oirowa,
Lemek, Ol Kinyie and Siana now surrounding MMNR.
It is only after KREMU SRFs began in 1977 that cattle
numbers can be determined at the scale of the group ranch. We
re-analysed KREMU data for the period 1980–2000, using Jolly’s
Method II, to estimate cattle, sheep/goats, wildebeest (Conno-
1995 2005
chaetes taurinus mearnsi) and zebra (Equus quagga boehmi)
populations on Koyake. These population estimates are shown
in Table 4. Jolly II estimates were also calculated for cattle on all
Mara ranches (Koyake, Lemek, Ol Kinyie and Siana). Figure 18
shows livestock populations for all ranches and for Koyake from
1980 to 2000, rainfall at Narok for 1977–2000, and the NDVI for
Koyake from 1983 to 2000 to indicate vegetation production.
Cattle numbers on all ranches vary from 80,000 to 180,000, as
confirmed by Broten & Said (1995) for their analysis of 1977–91
KREMU data, but there has been no marked long-term increase
or decrease (Serneels & Lambin, 2001).
Livestock estimates for Koyake reflect the general trend for
all ranches. The Koyake cattle population increased to about
40,000 head in the late 1980s, following good rainfall and
increased primary production. In the early 90s, poor rainfall
resulted in a decline to about 25,000 head. A further period of
good rainfall in 1997/98, associated with ‘El Nino’, led to an
increase again to 40,000; this was followed by a massive ‘crash’
in the catastrophic ‘La Nina’ drought of 1999/2000 that
reduced cattle numbers to about 14,000 head (Reid et al.,
2001). Ground counts in 2002 showed that cattle populations
recovered rapidly again, with an estimated 27,500 head on
Koyake (Reid et al., 2003). Meanwhile, over the 1990s the
Maasai have increased their sheep and goat herds, usually a
strategy in times of drought or hardship (Grandin, 1988;
Homewood & Rodgers, 1991). The contribution of sheep and
goats to total livestock biomass increased from 7.1% in 1979 to
9.9% in the late 1990s (see Table 5). With current trends this
proportion may increase further.

1014 Journal of Biogeography 31, 997–1032, ª 2004 Blackwell Publishing Ltd

 Expansion of human settlement in Kenya’s Maasai Mara

Table 4 Summary of cattle, sheep/goat, wildebeest and zebra estimates with standard errors (SE) for Koyake, derived from KREMU SRFs of
1980–2000 by Jolly Method II analysis

Cattle Sheep and goats Wildebeest Zebra

Date of survey Estimate SE Estimate SE Estimate SE Estimate SE

01-Jul-80 16,880 8656 19,200 7693 1540 850 – –

09-Mar-81 6640 2718 9340 2541 10,140 6627 – –
27-Aug-81 20,840 7171 21960 9888 105,500 31,130 – –
27-Oct-81 11,300 5275 18,120 6708 79,960 21,262 – –
22-May-82 27,460 5277 11,980 7566 2280 748 – –
30-Aug-82 17,788 2420 21,788 9717 157,192 27,471 – –
06-Dec-82 54,774 23,591 31,830 14,981 1736 707 – –
24-Apr-85 49,408 16,071 38,586 11,344 3109 1247 4967 1084
28-Nov-85 20,822 7859 30,609 13,347 4688 1729 8257 1248
08-May-86 37,977 9013 31,480 13,864 1546 908 5872 1512
10-Aug-86 35,806 21,453 16,941 6826 32,664 8251 4293 1189
08-Nov-86 25,987 5181 25,592 10,129 13,668 4919 8306 2385
24-Apr-87 22,589 9389 22,660 14,318 2287 1825 6702 1779
12-May-89 40,745 15,030 20,213 10,840 1507 791 1560 325
13-Aug-90 44,858 24,736 16,312 10,716 127,252 75,401 12,482 2430
20-Apr-91 38,067 8302 24,858 10,237 3156 921 2199 427
12-Aug-91 38,651 7876 23,240 6358 55,987 11,629 6135 1468
14-Mar-92 19,581 4780 17,451 4760 6924 1776 22,401 3101
19-Aug-92 27,089 6390 23,067 9897 50,831 13,710 6965 2242
03-Nov-93 33,200 6348 17,367 5063 9530 1131 4628 884
15-May-94 18,165 8490 12,172 3838 1791 522 1995 949
27-Jul-96 8227 3108 10,346 4184 69,291 15,901 7934 1116
28-May-97 32,518 5857 24,149 4000 1073 467 3085 1610
19-Aug-97 46,392 9335 32,261 9713 24,273 15,041 8626 2606
21-Oct-00 7748 2679 60,603 16,589 16,507 5041 10,674 6294

Figure 19 shows that the Koyake cattle population lags
1–2 years behind the peaks and troughs of rainfall and primary
production, and demonstrates that 75% of the variation in cattle
numbers on Koyake (wet-season population) is explained by
rainfall in the previous 2 years. This confirms that, as with resident
wildebeest (Serneels & Lambin, 2001), rainfall is the limiting factor
for cattle in the Mara, and that the Maasai continue to maximize
their herds opportunistically in response to favourable conditions.
Dry-season cattle estimates show no relationship at all with
rainfall, presumably as there is considerable on/off-ranch cattle
movement during the wildebeest migration.
During the 1970s, much of the dry-season stock movement
was into Trans-Mara, where Koyake members negotiated
reciprocal grazing arrangements with their Siria neighbours on
Oloirien and Kimentet group ranches (Lamprey, 1984). Today,
with grazing land becoming scarcer, such agreements are no
longer easily reached with the Siria Maasai, except in years of
extreme drought such as in 1999/2000. Movements mainly
take place between il-Purko ranches (Koyake, Ol Kinyie,
Lemek and Siana) in an opportunistic manner, as herders take
advantage of localized ‘green flushes’ and also of the availab-
ility of crop residues on commercial farms. Thus, from the
mid-1980s onwards there is no statistically significant differ-
ence in wet- and dry-season cattle numbers on Koyake.
Available SRF data for Trans-Mara indicate the cattle
numbers fluctuate from 150,000 to 285,000, roughly in response
to rainfall conditions. However, the SRF during the drought year
of 2000 gives a high estimate of 244,000; during the prevailing
dire conditions Maasai of Koyake and other Narok ranches had
no choice but to take some of their cattle to Trans-Mara.
Trends in migratory wildlife
In the early 1970s, Maasai moving southwards into the Mara
plains following the eradication of tsetse met head-on with the
Serengeti-Mara’s expanding population of migratory wilde-
beest. Prior to 1963, the wildebeest population of the SME had
been held in check by rinderpest, which prevailed endemically
in the ruminant populations with low-level transmission
between wildlife and livestock28 (Talbot & Talbot, 1963;
28
The wildebeest herds of the Serengeti-Mara ecosystem appear to
have recovered quite rapidly from the rinderpest pandemic of the
1890s. In 1927 the Game Department reported that ‘wildebeest in the
south-western portion of the Maasai Reserve occasioned complaint…
in view of the amount of grass and water, which the vast herds con-
sume and also of the fact that they foul the ground when they calve
down, causing snotsiekte (malignant catarrhal fever) in the Maasai
cattle. If any attempt is to be made to reduce the numbers of these
animals in that area, it will have to be in concert with the Tanganyika
authorities, and on a huge scale, as there is over our borders an
apparently inexhaustible wildebeest reservoir’ (Game Department
Annual Report, 1927, p. 13).

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R.H. Lamprey and R.S. Reid

200,000 Cattle koyake

Cattle all ranches
180,000 Sheep/Goats koyake

160,000

140,000
Population estimate

120,000

100,000

80,000

60,000

40,000

20,000

0.550
NDVI (a)
0.500 NDVI (b)

0.450
Mean NDVI

0.400

0.350

0.300

0.250

1200

1000
Figure 18 Top: 1980–2000 cattle popula-
Rainfall (mm)

800
tion on all ranches (Koyake, Lemek, Ol
600 Kinyie, Siana and Ol Choro Oirowa)
400 (Broten & Said, 1995), cattle on Koyake
200 only, and sheep/goats on Koyake only.
Middle: NDVI(a) central Koyake 1983–2000,
0
and NDVI(b) ‘Mara eco-unit’ 1983–94
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
00

(Ottichilo, 2000). Bottom: Rainfall at

19
19
19
19
19
19
19
19
19
19
19
19
19
19
19
19
19
19
19
19
19
19
19
20

Y e ar Narok 1977–2000.

Table 5 Seasonal biomass (kg km)2) of cattle, sheep/goats,

Sinclair, 1979). Early aerial counts conducted in 1961 estima-
wildebeest and zebra on Koyake, for three SRF periods, based on a
ted a total Serengeti wildebeest population of 220,000 animals,
wet season of November–June, and a dry season of July–October
which calved on the Serengeti plains in the wet season and
Total moved northwards into the central Serengeti woodlands in the
biomass dry season (Grzimek & Grzimek, 1960).
Cattle Smallstock Wildebeest Zebra (kg km)2) Sinclair (1979) describes the rapid increase in the Serengeti
wildebeest population that followed the eradication of
1979
rinderpest by the ‘JP-15 vaccination campaign’ in 1963 (Atang
Wet season 6990 461 292 1004 8748
Dry season 5253 544 7359 2554 15,710 & Plowright, 1969), and favourable dry-season rainfall in the
Annual mean 6411 489 2648 1521 11,068 late 1960s. From 1969 onwards, with a population increasing
1985–91 beyond a million, the Serengeti ‘migration’ annually spilled
Wet season 6058 637 526 1082 8304 over into the Mara in Kenya (Pennycuick, 1975). By 1975 the
Dry season 7159 433 8852 1527 17,971 Serengeti wildebeest population had stabilized at approxi-
Annual mean 6425 569 3302 1230 11,526 mately 1.4 million animals, regulated by the dry-season food
1992–2000 supply (Sinclair, 1979), and subsequent surveys during the
Wet season 5141 420 594 1605 7761 1980s showed little significant change from this number (see
Dry season 4511 791 7078 2107 14,486
Serneels and Lambin 2001). During the drought of the
Annual mean 4931 544 2755 1773 10,003
mid-1990s the wildebeest population declined to 900,000,
Conversion to biomass for different species as follows (Stelfox et al., but recovery has been rapid, the most recent count in
1980): cattle, 180 kg; sheep/goat, 23 kg; wildebeest, 123 kg; zebra, 1999 giving an estimate of 1.3 million (Mduma et al., 1999;
200 kg. TWCM, 1999).

1016 Journal of Biogeography 31, 997–1032, ª 2004 Blackwell Publishing Ltd

 Expansion of human settlement in Kenya’s Maasai Mara

55,000

50,000

45,000

40,000

Cattle population estimate

35,000

30,000

25,000
Wet season
20,000 Dry season
2
15,000 Linear (Wet season) R = 0.7467
2
Linear (Wet season) R = 0.0003
10,000
Figure 19 Relationship between wet and
dry season cattle estimates on Koyake, and 5000

the cumulative rainfall of the previous 900 1100 1300 1500 1700 1900 2100

2 years (1987–2000 data). Total rainfall previous 2 years (mm)

Figure 20 Density and distribution of wildebeest in the Mara ecosystem, from 11 SRF’s of 1979 (after Lamprey, 1984). The northern
Loita plains wildebeest on Ol Kinyie are defined as the ‘resident population’; these move south-west into the Mara in the dry season to
join the much greater population of Serengeti migrants. This seasonal pattern has prevailed (Serneels & Lambin 2001), the essential
difference being the drastic decline in the Mara’s resident wildebeest from 100,000 in 1979 to just 20,000 today (Ottichilo, 2000).

In the late 1970s, the wildebeest seasonal distribution in the
Mara followed the pattern indicated in Fig. 20 (Lamprey,
1984). Today, of the 300,000–750,000 wildebeest now entering
the Mara each year from the Serengeti, some 50,000–150,000
move into the four group ranches of Lemek, Koyake, Ol Kinyie
and Siana (Broten & Said, 1995; Ottichilo, 2000). Our analysis
of monthly KREMU SRF data of 1979 shows the influx of
120,000 wildebeest into Koyake Group Ranch in June/July,
followed by their gradual recession back into the MMNR in
August–October (see Fig. 21).
Table 4 presents population estimates of wildebeest and
zebra on Koyake for 1980–2000. Ottichilo (2000) has shown
that the influx of migratory wildebeest in the Mara in the dry
season is strongly correlated with vegetation conditions, as
indicated by the NDVI of the rangelands.
Trends in resident wildlife
The Mara has its own ‘resident’ wildlife populations that move
locally within the area. The Mara’s resident wildebeest calve on
the Loita plains and move into the Mara Reserve and adjacent
ranches to mix with the Serengeti populations during the dry
season (see Fig. 20) (Talbot & Talbot, 1963; Stelfox et al., 1980;
Lamprey, 1984; Ottichilo, 2000; Serneels & Lambin, 2001). In
the early 1960s this resident population was estimated at
20,000 animals (Darling, 1960; Stewart & Talbot, 1962). As

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R.H. Lamprey and R.S. Reid
160,000
140,000 Wildebeest
Cattle
120,000
Animal population
100,000
80,000
60,000
40,000
20,000
0
16- 15- 14- 15- 14- 14- 13- 13-
Dec Jan Feb Mar Apr May Jun Jul
Survey date
with the Serengeti wildebeest, the Mara population increased
fivefold to 100,000 animals over the period 1961–79 (Stelfox
et al., 1980; Ottichilo, 2000).
Since 1980, resident wildlife populations have declined by
over 70% across the Mara rangelands, both inside and outside
the MMNR (Ottichilo, 2000). The resident wildebeest popu-
lation is now reduced from 100,000 animals in the late 70s to
just 20,000 today. Ottichilo recorded similar declines in the
other major species of the Mara plains, zebra (Equus quagga
boehmii), buffalo (Syncerus caffer brachyceros), giraffe (Giraffa
camelopardalis tippelskirchi), eland (Taurotragus oryx patter-
soni), waterbuck (Kobus ellipsiprymnus defassa), topi (Damal-
iscus lunatus jimela), kongoni (Alcelaphus buselaphus cokei),
Thomson’s gazelle (Gazella thomsoni) and Grant’s gazelle
(Gazella granti robertsi). This reduction is attributed to the
conversion of grazing land on the Loita plains to wheat, and to
increased poaching (Ottichilo, 2000; Serneels & Lambin, 2001).
In the wet season (November–June), fewer than 2000
wildebeest are found in Koyake, these animals being essentially
resident. However, wet-season wildebeest and zebra numbers
show a slight increasing trend, possibly due to displacement
from the Loita plains. In the 1990s, these species contribute
7.7% and 20.7% respectively to total herbivore biomass on
Koyake. In the ground census in November 2002 when the
migration was late returning to the Serengeti, there were at
least 10,400 zebra and 18,100 wildebeest on Koyake (Reid
et al., 2003).
Wildlife/livestock interactions, stocking densities
and ‘carrying capacity’
Wildebeest and zebra compete directly with cattle for forage
(Casebeer & Koss, 1970; Bourn & Blench, 1999; Voeten, 1999);
in the same grazing area they eat the same grass species, and
any differences relate to selection of leaf/stem or sward
components. Both cattle and wildebeest require a dry-matter
intake of about 2.5% to maintain condition (Sinclair, 1977;
Lamprey & Field, 1983; Bekure et al., 1991). On this basis, a
herbaceous dry-matter (DM) production of nearly
1018
Figure 21 Monthly estimates (error
12- 11- 11- 10- 10-
bars ± SE) of cattle and wildebeest on
Aug Sep Oct Nov Dec
Koyake, from KREMU SRFs of 1979. Derived
from Lamprey (1984).
3.2 kg ha)1 day)1 will be required on Koyake to support
25,000 cattle and 60,000 wildebeest (a mean estimate for the
early 80s) during dry-season months. Using the rainfall-
production equation of Sinclair (1979), the average dry-season
rainfall of 204 mm at Aitong (Lamprey, 1984) will stimulate a
DM production of 3.5 kg ha)1 day)1, just sufficient to main-
tain the observed herbivore numbers.
An analysis of data presented in Stelfox et al. (1980) and
Broten & Said (1995) indicates that cattle, sheep/goats,
wildebeest and zebra contribute 88–92% of herbivore biomass
on the ranches. Buffalo contribute some 17% to herbivore
biomass in MMNR/Koyake combined as the ‘Mara eco-unit’,
but in Koyake alone, only 2% of the biomass. Table 5 shows
the seasonal biomass of livestock (cattle and sheep/goats) and
wildlife (wildebeest/zebra) for Koyake, clustered into three
separate time periods: 1979, 1985–91 and 1992–97. These data
suggest that Koyake Group Ranch supports a slightly lower
biomass of herbivores in 1992–2000 than in 1979, presumably
due to the reduced rainfall during the 1990s. Table 5 also
shows that at the height of the dry season when migratory
wildebeest are present, the biomass on Koyake may exceed
17,000 kg km)2.
In assessing ‘carrying capacity’, early studies related veget-
ation production and large mammal biomass to rainfall, in
protected areas or pastoral rangelands (Coe et al., 1976;
Le Houerou & Hoste, 1977; Pratt & Gwynne, 1977; Bourn,
1978; Jewell, 1980; Bell, 1982; Deshmukh, 1984; East, 1984;
Bekure et al., 1991). Collectively, this research demonstrated
that an area such as the Mara, which has volcanic soils and an
annual rainfall of 700–900 mm, will support a herbivore
biomass of 10,000–15,000 kg km)2, a biomass range well
supported by our data in Table 5.
In the late 1980s, the relevance of ‘traditional’ models of
carrying capacity was questioned. It was suggested that in
semi-arid and arid rangelands, where rainfall is very variable,
livestock populations never achieve the equilibrium implicit in
the concept of a fixed carrying capacity (Westoby et al., 1989;
Behnke et al., 1993; Sandford, 1995). In such ‘non-equilib-
rium’ systems, livestock herders closely track grazing resources
Journal of Biogeography 31, 997–1032, ª 2004 Blackwell Publishing Ltd

spatially and temporally to maximize production. More
recently, Illius & O’Connor (2000), in considering the impact
of grazing on vegetation, have focussed the debate once again
on density-dependence in grazing systems. Through model-
ling, they contend that even in arid and semi-arid rangelands
‘the animal population is in long-term equilibrium with dry-
season resources, on which it depends for survival; that dry-
season resource areas and outlying areas thus operate in a
source-sink manner; and that the ratio of these areas deter-
mines the strength of consumer-resource coupling outside the
dry-season range’ (Illius & O’Connor, 2000, p. 283). In
vigorous defence of the non-equilibrium concept, Sullivan &
Rohde (2002) assert that it reflects pastoral strategies in arid
environments where herders are responsive to the vagaries of
an unpredictable climate.
Our study shows that livestock populations exhibit the high
variability associated with an unpredictable rainfall and
opportunistic herding practices, with numbers varying from
25,000 after 1–2 years of below-average rainfall, to some
45,000 in good years. Although a large part of the Mara
rangeland area lies in the subhumid zone, rainfall variability is
high, with a coefficient of variation of 40–60% (Ottichilo,
2000). During droughts, such as those of 1984, 1991 and 2000,
the cattle population crashes as animals die or are sold off.
Estimated over the long-term, through the peaks and troughs
of ‘normal’ rainfall variability, the domestic and wild herbivore
biomass on Koyake falls within the range 10,000–
11,500 kg km)2 annum)1, in general agreement with the
‘classic’ estimates of ‘carrying capacity’.
The rangelands of the Mara show extraordinary resilience in
maintaining huge populations of domestic and migratory wild
herbivores year after year. However, the long-term effect
impact of this heavy grazing on the Mara rangelands has never
been rigorously investigated. A few early studies showed that
there is local deterioration in range conditions close to Maasai
settlement, manifested by a loss of vegetation cover, change in
vegetation composition and increased soil erosion (Heady,
1966; Skovlin, 1980). Using LANDSAT imagery, Lamprey
(1984) determined a significant inverse relationship between
herbaceous vegetation cover and livestock density at the scale
of the group ranch. The application of more recent research in
soil degradation in pastoral rangelands (Biot, 1993) may shed
further light on the ability of the Mara grasslands to sustain
such a vast herbivore population.
The Koyake Maasai refer to the annual wildebeest migration
as their ‘yearly famine’, as the wildebeest ‘finish the grass’ and
bring disease (Lamprey, 1984). The Maasai describe ‘turning
disease’ as a particular problem; wildebeest herds bring Oestrid
bot-flies, the larvae of which cause brain damage to their
wildebeest and cattle hosts and cause them to circle endlessly
(Sachs & Debbie, 1969). Malignant catarrhal fever (MCF), a
viral disease transmissible to cattle during the wildebeest calving
season (Rossiter et al., 1983; Mushi, 1985) is not a problem on
Koyake, as the ranch is not a wildebeest calving area. MCF is
factor in livestock grazing in Ol Kinyie Group Ranch, as the
Mara’s resident wildebeest calve on the Loita Plains.
Journal of Biogeography 31, 997–1032, ª 2004 Blackwell Publishing Ltd
Expansion of human settlement in Kenya’s Maasai Mara
Recent research in the Maasai Mara has shown that, to a
certain degree, some species of wildlife are attracted to Maasai
areas (Reid et al., 2001). Moderately grazed ‘lawns’ in the
general vicinity of bomas often attract small to moderate-sized
wildlife species that prefer short grass, such as Thomson’s
gazelle. The close proximity of settlements may also afford
additional protection from predators, whilst the nutrient-rich
sites of old bomas may provide improved grazing. However,
many species of high tourism value, such as elephants, rhino
and the big carnivores avoid settlement areas, so this potential
‘synergism’ is limited.
Pastoralism as a livelihood on Koyake
Various authors have estimated the number of livestock
required to support pastoral households. These estimates differ
according to prevailing rangeland conditions, and the nature
of the pastoralism itself (whether ‘pure’ or agro-pastoral).
Early research indicated that subsistence pastoralism could be
maintained with 2.5–3.5 standard stock units (SSU ¼ 450 kg),
equivalent to 6–9 cattle, per capita of the human population
(Brown, 1971; Pratt & Gwynne, 1977; Jewell, 1980). Kjaerby in
Homewood & Rodgers (1991) calculates that where livestock
are exchanged for grain, five cows per capita are needed to
meet subsistence requirements. Bekure et al. (1991, p. 27), in
estimating the needs of Maasai in Kaputei ranches, simply state
that ‘there must be at least 10 cattle for each person if the
population is to subsist on a diet of milk and meat alone’, but
this clearly applies to pure pastoralism, now a rare phenom-
enon. Herd composition is critical to this determination, but
available data indicate that herds in eastern Maasailand are
similar to those of the Mara area, with mature cows making up
58% of the herd in Kajiado ranches (Bekure et al., 1991),
compared with 57% in the Mara area (Thompson 2002). In
both areas, about 25% of mature cattle are lactating at any one
time.
Comparisons of more recent estimates of pastoral needs
have been complicated by the use of different units.29 Table 6
attempts to standardize these estimates according to cattle per
person, and ‘livestock units’30 (LU ¼ 250 kg) per ‘reference
29
For example livestock holdings may be determined as cattle alone
(Bekure et al., 1991); the mix of animals (whether smallstock, cattle or
others) (Jewell, 1980); ‘standard stock units’ (SSUs) of 450 kg (Brown,
1971; Pratt & Gwynne, 1977; Jewell, 1980); ‘livestock equivalents’ (LE)
of liveweight0.75 (Nestel, 1986), closely equivalent to the ‘tropical
livestock unit’ (TLU) or ‘livestock units’ (LU) of 250 kg (Boudet &
Riviere, 1968; Bekure et al., 1991; McCabe et al., 1997; ACC, 2001).
Human needs may be considered in terms of the ‘family’ (Brown,
1971; Dahl & Hjort, 1976; Pratt & Gwynne, 1977; Jewell, 1980); the
‘household’ (Bekure et al., 1991; ACC, 2001); the individual (adults
and children together) (Bekure et al., 1991); the ‘reference adult’ (RA)
derived from the ‘active adult male equivalent’ (AAME) based on
energy requirements of different age classes (FAO, 1974; Little, 1980;
Nestel, 1986; McCabe et al., 1997; ACC, 2001); or in terms of human/
livestock ‘metabolic ratios’ (Prins, 1992).
30
Originally derived from the Tropical Livestock Unit (TLU) of
250 kg of Boudet & Riviere (1968).
1019
R.H. Lamprey and R.S. Reid

Table 6 Minimum subsistence requirements for pastoralism, as indicated by various studies

LU/capita LU/RA LU/RA

Author Unit of assessment Cattle/capita (cattle only) (cattle only) (all stock)

Brown* in Pratt & Cattle and smallstock, per family of eight persons 5.5 4.0 5.0 5.6
Gwynne (1977)
Dahl & Hjort (1976) Cattle per family of eight persons 10.0 7.2 9.0
Jewell (1980) Cattle and smallstock per family of eight persons 5.5 4.0 5.0 7.5
McCabe et al. (1997) Cattle and smallstock, per family of eight persons 5.5 4.0 5.0 8.7
Nestel (1986) Cattle and smallstock, per AAME (intermediate wealth) 5–13
Bekure et al. (1991) Cattle, per capita 10.0 7.2 9.0

*See also Brown (1971).

18
LU/RA (cattle only)
16 LU/RA (cattle and sheep/goats)
14

12

10
LU/RA

4
2 Figure 22 The decline in livestock hold-
0 ings per capita on Koyake, expressed as
1978 1980 1982 1984 1986 1988 1990 1992 1994 1996 1998 2000 Livestock Units/Reference Adult (LU per RA)
Year (see text).

adult’ (RA) (Little, 1980; McCabe et al., 1997). The RA closely Table 7 Estimation of livestock units per reference adult on
approximates, but simplifies, the ‘active adult male equivalent’ Koyake Group Ranch (971 km)2), for livestock data averaged
(AAME) (FAO, 1974; Nestel, 1986; Bekure et al., 1991), and for 1992–2000, and human population estimate of 1999.
both may be estimated as persons · 0.80 (where Maasai
Parameter Cattle Sheep/goat All livestock
populations have a similar age structure).31 Nestel (1986)
stratified Kaputei Maasai into the poor, with 0–5 LU/AAME, Population 27,395 23,645 51,040
the intermediate, with 5–13 LU/AAME, and the wealthy with kg km)2 4931 544 5475
>13 LU/AAME. Biomass entire 4,788,115 528,064 5,316,178
On the basis that the Maasai population is increasing at the ranch (kg)
rate of 4.4% per annum (see section ‘Maasai population and TLU entire ranch 19,152 2112 21,265
LU/capita (8500 people) 2.25 0.25 2.50
distribution’), Fig. 22 shows the LU/RA ratio for Koyake over
LU/RA 2.82 0.31 3.13
the period 1979–2000. In the early 1980s, LU per RA in Koyake
declined to very low levels due to decreased rainfall (just
512 mm in 1981), and the movement of many cattle into
Trans-Mara. By 1985, cattle numbers had increased in minimum subsistence levels, despite the good rainfall of the
response to favourable rainfall (see Figs 11 & 18), and per late 1990s. Today on Koyake there are fewer than three cattle
capita holdings increased to 16 LU/RA. Since 1990, however, per person, equivalent to 2.82 LU/RA. If sheep and goats are
per capita livestock holdings have steadily declined to below also included, this estimate increases to 3.13 LU/RA. Table 7
presents a general summary of livestock holdings for Koyake
for the period 1979–2000.
31
The RA is calculated as follows: adult male ¼ 1: adult These estimates are corroborated by the aerial photographic
female ¼ 0.86; children 0–5 ¼ 0.52; children 6–10 ¼ 0.85; male child survey of cattle in bomas, conducted in September 2001 (dry
11–15 ¼ 0.96; female child 11–15 ¼ 0.86 (FAO, 1974). In the absence season, wildebeest present), and December 2001 (typical ‘short
of detailed population age structures on Koyake, it is assumed that rains’ wet season, wildebeest absent). This analysis, shown in
population age structures are similar to those in Ngorongoro and
eastern Maasailand (with 52–55% of the population under 15 years –
Fig. 23, gives an estimate of 16.3 cattle/hut (3.5 cattle/person
see Table 2). The conversion of 0.8 · population is calculated from or 3.2 LU/RA) for the dry season, and 13 cattle/hut (2.8 cattle/
data of Rutten (1992) and McCabe et al. (1997). person or 2.5 LU/RA) for the wet season. By extrapolation on

1020 Journal of Biogeography 31, 997–1032, ª 2004 Blackwell Publishing Ltd


100
80
Figure 23 Histograms of cattle/hut, from
Frequency
aerial photographic surveys of September 60
2001 (dry season) and December 2001 (wet
season). Weighting the mean on huts per
40
boma, this analysis gives the following sea-
sonal estimates: dry season, 16.3 cattle per
20
hut (SD ¼ 14.39, n ¼ 174), equivalent to
3.5 cattle per person or 3.2 LU per RA; wet
0
season, 13 cattle per hut (SD ¼ 5.97,
2.5
n ¼ 238), equivalent to 2.8 cattle per person
or 2.5 LU per RA.
huts numbers, this provides an estimate of 25,000–30,000
cattle for the ranch, well in agreement with SRFs and with the
ground count estimates of 27,500 cattle for Koyake in 2002
(Reid et al., 2003). The dry-season histogram in Fig. 20 is
heavily skewed, due to the herding of cattle surplus to
immediate requirements into small bomas in the central
plains, probably to remove them from the main wildebeest
concentration. Such herding commonly takes place under
herding agreements between Maasai families.
The December 2001 photographic survey indicated that
during the wet season, cattle were distributed evenly amongst
the households of the ranch; there was no significant difference
in cattle/hut ratios for the affluent area of Talek
(13.411 ± 1.1674 cattle/hut, 95% CL, n ¼ 120 huts), and the
central plains area of Oltorotua/Olare Orok (13.962 ±
1.054 cattle/hut, 95% CL, n ¼ 80 huts).
Group ranch subdivision
In the Mara, Lemek GR was subdivided in 1999, and Koyake
ranch members then voted to follow suit (M. McCartney, pers.
comm.). Whilst group ranch subdivision in Narok District has
not been analysed in detail, the causal factors are generally
similar to those that brought about the first wave of
subdivisions in Kajiado District in eastern Maasailand (ole
Pasha, 1986; Bekure et al., 1991; Galaty, 1992; Rutten, 1992;
Campbell, 1993; Kimani & Pickard, 1998). According to these
authors, group ranches have largely failed because they cannot
serve as economically viable units, and because ranch
committees have mismanaged group resources for their own
ends. Increasingly disillusioned with their committees, a
younger generation of ranch members pressed to subdivide
their ranches, to gain individual security of land tenure, and
advance their livelihoods on their own land. In Kajiado,
subdivision began in the late 1970s, and accelerated in the
1980s.32 The process was often chaotic. In ranches close to the
nation’s capital Nairobi, ‘land committees’, generally com-
32
The process became increasingly politicized when in 1989 President
Moi ‘advised all Kajiado residents to have their group ranches subdi-
vided and each member gets his title deed to avoid differences in the
future’ (Kenya Times, 15 April 1989).
Journal of Biogeography 31, 997–1032, ª 2004 Blackwell Publishing Ltd
Expansion of human settlement in Kenya’s Maasai Mara
Sep 2001 (dry season)
Dec 2001 (wet season)
7.5 12.5 17.5 22.5 27.5 32.5 37.5 62.5 67.5 72.5 77.5
Cattle/hut
posed of ranch chairman and other key individuals, allocated
large tracts to themselves, prompting a subdivision scramble as
ordinary members tried to convert their intangible ‘shares’ into
land (Galaty, 1992). In his analysis of group ranch subdivision
in Kajiado, Rutten (1995) considered the overall impact of
subdivision to be catastrophic for the Maasai: ‘Land subdivi-
sion has been most favourable to the most powerful. Whole
groups of Maasai pastoralists have been excluded (and) only a
mere 10% of the households have a sufficient large parcel
during the dry season period. Land sales to outsiders worsened
this situation by a reduction of the area available and by
buying of animals, using the proceeds of the land sales. These
outsiders fence off their land for cultivation or act as absentee
landowners interested in speculation only.’
Despite the sedenterization brought on by land privatization,
Rutten (1992) noted that ranch subdivision in Kajiado did not
automatically result in fencing, for three reasons. First, fencing is
very expensive, a typical Kaputei plot of 40 ha costing KShs
150,000 (equivalent to US$ 6550) to fence in 1990.33 Such
costs could clearly be halved if shared with neighbours, but in
the Mara this would still amount to more than double the
annual income of an ordinary Maasai household. Secondly, in
1990 many plots in Kaputei had not been occupied by their
owners, and thus had not been developed. Thirdly, Rutten
(1992, p. 363) questions whether ‘livestock owners are really
interested in turning away from the traditional communal
use of the pastures forced on them by climatic conditions’.34
However, fencing is mainly carried out by outsiders, who buy
land for cultivation and speculation (Rutten, 1992). Kimani
& Pickard (1998) record that in some ranches in 1996, over
50% of plots created at original subdivision were purchased
by non-Maasai, and that Maasai holdings are less than half of
their original size. Finding themselves on plots too small for
33
In 1999 the fence for the 40 ha plot would cost KShs 490,000 (c. US$
7000), calculated using the GPD Deflator (Local Currency Series) and
Official Exchange Rates given by World Bank (2001), where US$
1 ¼ KShs 22.9 in 1990, and US$ 1 ¼ KShs 70.4 in 1999.
34
We note, however, that at the time of subdivision in Kaputei, Rutten
records that livestock holdings exceeded 11 LE/AAME. Clearly,
pastoralism remained a viable livelihood for the Kaputei Maasai in the
late 1980s, and there was relatively little incentive to explore alternative
livelihoods.
1021
R.H. Lamprey and R.S. Reid
pastoralism, and without the know-how or resources to
develop these areas for other livelihoods, more Maasai famil-
ies are now cultivating, or dividing their plots for further sales.
In the Mara, as in eastern Maasailand, the concept of
‘membership’ has become a critical aspect of the subdivision
process. At the time of group ranch adjudication, the
membership essentially comprised household heads who
applied to the Adjudication Committee for group title of the
ranch (Bekure et al., 1991; Rutten, 1992). Their names were
added to the ‘Adjudication Register’, and following the
establishment of the ranch new members could only be added
by a 60% vote of the membership. In some ranches in eastern
Maasailand, membership has been stable and closely con-
trolled, whilst in others the membership has shown a
significant increase.35 As subdivision approached in eastern
ranches in the mid-1980s, membership registers were carefully
scrutinized to remove members whose claims were thought to
be invalid.
Ranches in Narok District have not been studied in such
detail as those in Kajiado, but there are indications that
membership rose rapidly after adjudication. For example,
Strickland, (1973, sec. 4.2) stressed that ‘Maasai leaders (in the
Mara) must be consulted on the feasibility of restricting the
number of shares in the ranches…, which would prevent the
groups from continually taking on more members than they
can support’. Our data indicate that on Koyake Group Ranch,
established in 1970, the number of households (the early
members) increased from 123 in 1967 to 364 households in
1974 (see Table 3). By 1999, the Koyake GR membership stood
at 1020 (ACC, 2001), drawn from an estimated 800 households.
A land survey completed in 2002 divided Koyake into plots
of an average size of about 150 acres (60 ha), to be allocated to
the 1020 members (M. McCartney, pers. comm.). As indicated
in our study and by Coast (2000), an early manifestation of
ranch subdivision was the fragmentation of the traditional
boma into one- or two-family units, and the proliferation of
these new ‘homesteads’ within their localities. As subdivision is
implemented, there is vigorous lobbying amongst the Koyake
membership to acquire plots with permanent water. Dams
built by conservation NGOs for the benefit of the community
may soon fall within the plots of influential Maasai. Land
allocation meetings have been acrimonious. It is alleged that
non-members (perhaps 300 of the 800 households) are not
being given land, and may be evicted from Koyake. By October
2003, many households were ‘deconstructing’ their bomas and
relocating to communal areas at Aitong and Talek whilst
decisions are made about their land allocation. Trading centres
serving surrounding neighbourhoods, such as that at Maili
Tano (‘Five miles’) between Aitong and Mara Rienda have
35
For example the membership of Olkinos Group Ranch, a ‘Phase I’
Kaputei ranch, rose from 90 to 116 over the period 1973–90. In
Oldoinyo Nyokie Group Ranch, a ‘Phase II’ section, the ranch of the
Loodokilani membership rose from 162–354 over the period 1982–90
(Rutten, 1992). Some of the other ranches show surprising stability in
membership.
1022
now been demolished. Meanwhile, those Maasai already
confident in their land claim have constructed permanent
homesteads incorporating brick-and-mabaati houses, wells
and fenced stock enclosures.
In the Mara, livestock holdings have dropped below the
subsistence minimum as subdivision approaches. Thompson &
Homewood (2002) report that interviewed households in the
Mara would allocate 38% of their plot area (about 23 ha) to
cultivation, and 60% (36 ha) to livestock. Assuming that all
wildlife were excluded, this latter area would support about 25
cattle for each household of 10.5 people, equivalent to just
2.0 LU/RA.
DISCUSSION
Over the last 30 years, the clearing of tsetse-infested bushlands
in the Mara enabled il-Purko Maasai to settle throughout the
northern Mara plains. In the early 1970s the group ranch system
formalized their land tenure. On Koyake, the group ranch
selected for our detailed study, 56% of the population is below
the age of 15, and the population is increasing at 4.4% per
annum. Many Maasai have also moved into the Mara from the
higher potential areas of Mau Narok as more areas are leased for
commercial wheat production (Duraiappah et al., 2000).
Our temporal analysis of settlement pattern indicates that
since 1967, Maasai have occupied the more outlying areas of
Koyake through ‘pulses’ of expansion. The primary factor
influencing the establishment of bomas is the distribution of
permanent water sources; analysis of the 2002 ground count
data revealed that 75% of bomas were within 2 km of water
(Reid et al., 2003). The first wave of expansion out of Lemek
Valley in 1961–67 was to the Talek river, which holds water
throughout the year. The second wave occurring in the early
1970s saw an increase in settlement at Talek, and the new
occupation of Mara Rienda close to the Mara River. In the
third expansion phase from the early 1980s to present, suitable
areas in Enkikwe and along the Olare Orok and Ntiakitiak
streams were occupied.
If settlement expansion continues, it is probable that wildlife
populations will decline drastically in the rangelands around
MMNR. We base this conclusion on two bodies of evidence. The
first relates to direct observation of wildlife behaviour with
respect to settlement distribution. Analysis of data gathered
during the 2002 ground count indicate that wildlife density
declines significantly beyond a boma density of 0.5 bomas km)2
(Reid et al., 2003). Currently, the density of bomas on Koyake is
about 0.3 bomas km)2. However, as ranch subdivision is
implemented and bomas disaggregate into 1–2 household units
(see Fig. 16 and Table 3), we anticipate a rapid increase in boma
density beyond 0.5 bomas km)2, and a more even distribution
of bomas across the landscape. Modelling of these changes using
GIS predicts that wildlife populations will fall by at least 40%
(Reid et al., 2003), and that certain species attractive for
tourism, such as the carnivores, rhino and eland may disappear
entirely from the Maasai ranches. Tourism associations now
report that predators are now absent from large parts of
Journal of Biogeography 31, 997–1032, ª 2004 Blackwell Publishing Ltd

Enkikwe and along Olare Orok, and that much of the area is
worthless for game-viewing tourism (Campfire, 2002).
The second body of evidence is more inferential, and relates to
the necessary changes in livelihoods that will take place as pas-
toralism declines. If pastoralism is to continue supporting the
increasing human population, livestock numbers must be
increased correspondingly. However, this study shows that
although cattle numbers in the Mara ecosystem oscillate in res-
ponse to rainfall and primary production, there has been no long-
term increase in the cattle population since SRF surveys were
started in 1980 (see also Broten & Said, 1995; Serneels & Lambin,
2001). Indeed, the SRF estimates of 80,000–180,000 obtained in
the SRFs, are similar to (if not slightly less than) the estimate of
191,000 obtained in the first reliable cattle census of 1960.
Over the period 1980–2000, Koyake Group Ranch has
supported 25,000–45,000 cattle, with a mean of about 27,000.
Livestock numbers obtained from SRFs are well corroborated
by the 2001 aerial photographic study, and from the 2002
ground counts. On the basis that the ranch supports 8500
people in 800 households, livestock holdings are estimated at
2.82 LU/RA, well below the minimum subsistence of about
5 LU/RA. The inclusion of sheep and goats increases this ratio
to 3.1 LU/RA. With the onset of ranch subdivision, the Maasai
of the Mara are not ‘buffered’ by an excess of livestock, and
may shortly have to cultivate, or take up other livelihoods that
are not compatible with conservation.
The results of our study cannot easily be reconciled with
those of Thompson (2002) and Homewood et al. (2001), who
calculated a ratio of 8.2 LU/RA at Talek and Aitong36 and
assert that in the Mara as whole the Maasai are relatively
wealthy. There are a number of possible reasons for this
discrepancy. First, Homewood et al. (2001)conducted their
ground sampling in 1999, when aerial counts (in 1997)
indicated a cattle population of about 45,000 (see Table 4),
following 3 years of above-average rainfall (see Fig. 18). At this
time, livestock holdings were estimated at 5.5 LU/RA (see Fig.
22). Secondly, it is possible that their sampling frame included
the more wealthy and influential Maasai, who must endorse
household research at the local level, and possibly ‘steer’
researchers towards the households for survey. Thirdly, at the
time of their assessment, there may have been local clustering
of stock at Talek due to extraneous factors such as local
rainfall, or the presence of migratory wildebeest in key grazing
areas in the ranch. We suggest that livestock counting
techniques based largely on household questionnaires at the
local level are subject to bias, no matter how carefully they are
designed (Bekure et al., 1991).37 As Thompson (2002, p. 82)
concedes, ‘mobility and extended networks of entitlements to
36
Extrapolation of the livestock estimates of Homewood et al. (2001)
to the ranch as a whole indicates a Koyake cattle population of some
130,000 (cf. maximum of 45,000 as calculated in this study).
37
Thompson (2002) describes how questions on livestock numbers
were carefully couched in terms of family requirements, to which
families may have replied with optimum, rather than actual numbers.
Journal of Biogeography 31, 997–1032, ª 2004 Blackwell Publishing Ltd
Expansion of human settlement in Kenya’s Maasai Mara
livestock and livestock products confuse simple measures
based on numbers of stock present at any one time.’
The accumulation and re-distribution of cattle by wealthy
Maasai, through the networks indicated above, is well known
in Maasailand (Homewood, 1992; Rutten, 1992). In their
detailed study in eastern Maasailand group ranches, Bekure
et al. (1991) recorded that 20% of the households own 60% of
the cattle. Lamprey (1984) estimated from cattle dip records
that 25% of households owned 52% of the cattle38 whilst
Thompson (2002) records that at Talek 20% of the households
own 49% of the cattle. By means of cattle loans, wealthy
Maasai help their extended family, whilst at the same time
extending their political influence throughout the ranch.39
Despite the decline in household livestock holdings in
Koyake, Coast (2000) records that some 70% of Maasai at
Talek still view pastoralism as their main occupation, but she
accepts that ‘(interview) questions… referred to the amount of
time an individual spends on a particular occupation, rather
than any consideration of the amount of income derived from
that activity.’ Actual income from livestock is small. According
to Thompson (2002) and ACC (2001), at Talek only about 4%
of bulls/steers and 0.4% of cows (heifers, milk-cows) are sold
at market. These sources indicate that income from livestock
production at Talek is about US$ 200–400 per household per
annum, about 10–20% of household income. Recalculated as
returns to land, livestock sales generate about US$ 5–13 ha)1
per annum (Douglas-Hamilton et al., 1989; Norton-Griffiths,
1995; Thompson & Homewood, 2002), a relatively insignif-
icant amount compared with small-scale cultivation, which in
Lemek may generate upwards of US$ 120 ha)1 per annum
(Thompson & Homewood, 2002).
In pastoral terms, the ordinary Maasai of the Mara find
themselves in a position similar to those of Ngorongoro in
Tanzania, where livestock holdings have dropped from 6.8 LU/
RA in 1987 (Homewood & Rodgers, 1991; McCabe et al., 1997) to
2.7 LU/RA in 1998 (NCAA, 1999). In Ngorongoro land use is
controlled by the Ngorongoro Conservation Area Authority
(NCAA), which collects all tourist entry and lodge franchise fees.
Until 1995 cultivation was banned, and the Maasai were largely
dependent on famine relief. With the cultivation ban lifted, some
85% of Ngorongoro Maasai households now cultivate, compared
with fewer than 5% 10 years ago (Coast, 2001; McCabe, 2002).40
38
On the basis of dip records, Lamprey (1984) calculated the average
household herd to be 129.9 ± 9.8 cattle (SE, n ¼ 144), but these herds
also included loaned cattle distributed amongst many poorer house-
holds. This is likely to be an overestimate as this method only recorded
those who could afford to dip their cattle, and therefore excluded the
poor.
39
As there has been no change in the leadership of Koyake Ranch
Committee for over 20 years (Thompson, 2002), it may be suggested
that committee members are continuously re-elected by a constituency
made dependent by cattle loans.
40
Cultivation in Ngorongoro was prohibited to maintain the ‘natural
environment’. The change in landscape as a result of cultivation has
prompted some conservationists to request UNESCO to re-evaluate
the status of Ngorongoro as a World Heritage Site.
1023
R.H. Lamprey and R.S. Reid
In the face of declining LU/RA ratios, and without significant
returns from other livelihoods, the Maasai in Ngorongoro are
‘commoditizing’ their livestock economy by selling more stock.
The percentage ‘turnover’ of their herds is about 8%, compared
with 4% in Loliondo (Tanzania) (Galvin et al., 2002) and
2–4% on Koyake (ACC, 2001; Thompson 2002).41 With this
high turnover, livestock sales in Ngorongoro generate 83% of
household income, compared with just 10–20% of household
income at Talek in the Mara. Crop sales in Ngorongoro, whilst
generating just 20% of household income, are nonetheless
gaining in importance. Maasai households in Ngorongoro are
thus diversifying their livelihoods away from subsistence
pastoralism, a strategic necessity when per capita livestock
holdings are falling (Zaal & Dietz, 1999; Little et al., 2001).
In contrast to Ngorongoro, land use in the Mara ranches is
under the control of the Maasai themselves, and there are more
livelihood options. Tourism has become an important source
of revenue for some ranch members who live close to MMNR.
Household incomes are now supplemented by tourist campsite
rents, ‘dividends’ from local wildlife trusts, employment in
lodges, revenues from ‘cultural manyattas’, cultivation leases
and remittances from family members now living in the cities.
This diversification has brought some wealth for family
economic improvement; Coast (2000) noted that at Talek
over 47% of families have at least one dwelling with a
corrugated iron (‘mabaati’) roof (an indicator of household
improvement), compared with just 6% in Ngorongoro.42
However, although there are many tourism developments
and activities in the Mara ranches, the majority of Maasai
households have benefited little from them. In the 1980s no
mechanism was in place for ranches to charge tourists for entry
on to their land, and the Narok County Council could not
consistently honour an agreement to remit a proportion of
MMNR earnings to the ranches. Tourism revenues on Koyake
were generated from bed-night tariffs on two lodges and from
camping fees, and amounted to <1% of the total revenue
generation in the Mara (Douglas-Hamilton et al., 1989).43
During the 1990s the situation improved, as ‘wildlife associ-
ations’ were established on group ranches adjacent to the
reserve to collect and disburse game-viewing fees. However,
influential leaders gained control of key tourist campsites and
now capture a high proportion of tourism revenues, averaging
41
In his cattle sales analysis, Thompson (2002) omits two ‘outliers’
from Talek that skew his percentage estimates upwards.
42
The 2001 aerial photographs indicated that across Koyake GR, 15%
of all huts were tin-roofed, with no significant difference between
ranch areas. This can be compared with estimates of 20% and 24% for
the 1999 and 2002 ground counts respectively. On the 2001 photos,
tin-roofed and brick-walled ‘prestige’ houses’, used for day-living and
welcoming of guests, could clearly be discerned from simpler ordinary
tin-roofed huts (enkajijik). About 2.5% of all buildings, or 13% of
tin-roofed houses, were prestige houses, and the majority of these
(40%) were located at Talek.
43
Douglas-Hamilton et al. (1989) did not assess the value of tourism
wage-labour to Mara Maasai.
1024
some US $3000 per household per annum (Thompson &
Homewood, 2002). Maasai households immediately adjacent
to MMNR remain at a distinct advantage in their employment
opportunities in lodges inside the reserve, a livelihood that
contributes over 30% to household income (ACC, 2001). For
ranch members who live further from MMNR, ‘tourism
dividends’ from the wildlife associations are their only source
of tourism revenues. These dividends amount to just US$ 30
per household per annum, representing about 1% of house-
hold income (ACC, 2001; Thompson & Homewood, 2002;
R.H. Lamprey, unpubl. data).
Wildlife associations in the Mara are now being dissolved
by their dissatisfied memberships. Ironically, one of the last
activities of the main association, the Koyake-Lemek Wildlife
Trust, was to fund the group-ranch subdivision survey out of
its wildlife earnings. Subsequent survey costs remain a
significant budget item of the associations, with 50% of
revenues to be paid to a ‘Land Demarcation Account.’
The wildlife associations may be compared with two other
models of management in the Mara. The first is the Ol
Chorro Oirowa Wildlife Management and Conservation
Association (OOWMCA), established in the northern Mara
in 1992. Here, a group of wealthy Maasai amalgamated their
private ranches of 5–20 km2 to create an 86 km2 sanctuary for
wildlife, free of bomas, from which they earn high revenues
from an exclusive lodge concession. Even under these ‘ideal’
conditions, however, splits have occurred in the membership
over the division of tourism revenues, to the extent that in
1999 some members voted to form a splinter group, and to
draw in other group ranches in blocking the entry of
OOWMCA tourists.
The second model is the ‘Mara Conservancy’, an NGO
established in 2000 to manage the Mara Triangle, the part of
MMNR lying within Trans-Mara District (Mara Conservancy,
2000). The conservancy operates under a long-term manage-
ment lease from the Trans-Mara County Council. The aim of
this endeavour is to ensure transparency of revenue returns to
the Council, and proper management of the reserve. Imme-
diately after its establishment, annual revenues increased from
US$ 200,000 to US$ 1,000,000 as embezzlement at Trans-Mara
MMNR entry gates was curtailed (McCartney, pers. comm.;
Walpole & Leader-Williams, 2001). Potentially, the Maasai in
Koyake and other ranches can enter into similar management
contracts with outside agencies.
Despite these possibilities, tourism revenues in the Mara
appear to have peaked in the mid-1990s (Lamprey, in prep.).
By 1999 the ranch areas were so saturated with lodges and
campsites that a moratorium was placed on further develop-
ments by the Minister responsible for tourism. During the late
1990s, tourism in the Mara, and throughout the country as a
whole, declined in the face of a deteriorating political climate
in Kenya. Meanwhile, the key attraction, the wildlife, is also
declining (Ottichilo, 2000), and as more bomas and permanent
homesteads are established in the rangelands, the tourism
value of the area will decrease further. In 2003 a long-
established lodge on Koyake closed permanently, for the
Journal of Biogeography 31, 997–1032, ª 2004 Blackwell Publishing Ltd

reason that they found themselves in the expanding village of
Mara Rienda. Further closures are to be expected following the
subdivision of Koyake.
After subdivision the following principal livelihood options,
implemented singly or in combination, will be available to
landowners:
• ‘Commoditization’ of livestock production: this may develop in
the same way as in Kajiado (Zaal & Dietz, 1999)44 and
Ngorongoro (Galvin et al., 2002). With high costs, and the
need for access to permanent water, plots will remain
unfenced for some time, and herders will graze their cattle
freely over a number of holdings (Rutten, 1992). Currently
livestock sales in the Mara generate about US $5 ha)1
(Norton-Griffiths, 1995). At a later stage, some landowners
will orientate themselves towards fully commercialized meat
and/or dairy production, both systems involving fencing, the
installation of dams and boreholes, and possibly stall-feeding
(ACC, 2001).
• Tourism: after subdivision, it will be challenging for
landowners to maintain the elements of an ‘unspoiled’
wilderness landscape necessary for wildlife-based tourism. On
the basis of current practices, tourism generates about US
$10 ha)1, but this will only be maintained if land is unfenced
and undeveloped. Landowners will need to amalgamate
adjacent land holdings to create viable game-viewing or
(possibly) sport hunting areas. However, with an average plot
size of just 0.6 km2, it will require the cooperation of many
landowners for this to be a realistic option. Furthermore, it will
be important for the resulting ‘conservancies’ to be established
adjacent to MMNR, which will act as wildlife reservoir.
Campsites at Talek are likely to continue, as they overlook the
MMNR and give a wilderness vista; some new landowners
along Talek river have already fenced their private campsites.
However, campsites along rivers such as Olare Orok are
unlikely to remain viable, as plots on both sides of the river will
be quickly occupied by new landowners (Lamprey, in prep.).
Some tourism-based livelihoods, such as employment in
lodges, will continue, but this is unlikely to contribute
significantly to household income except in areas immediately
adjacent to MMNR (ACC, 2001).
• Small-scale cultivation: we have noted the rapid increase in
cultivation in Ngorongoro with the decline in per capita
livestock holdings. With climatic conditions similar to those
of Ngorongoro (volcanic soils, rainfall greater than
700 mm), the Mara area is suitable for maize and millet
cultivation (Jaetzold & Schmidt, 1983). At US $50–100 ha)1,
returns to land from cultivation are expected to exceed
returns from livestock and tourism by at least 300%
(Norton-Griffiths, 1995; Thompson & Homewood, 2002).
On the basis of current practices, small-scale cultivation may
44
Zaal & Dietz (1999) consider this commoditization in terms of
‘Calorific Terms of Trade’ (CtoT), and provide the example from
Kajiado in which one goat with calorific value of 29,250 calories might
be bartered for 67 kg of maizemeal of 230,000 calories, giving a CToT
advantage of 8 : 1 in favour of the pastoralist.
Journal of Biogeography 31, 997–1032, ª 2004 Blackwell Publishing Ltd
Expansion of human settlement in Kenya’s Maasai Mara
be carried out by individual landowners, or by an
association of landowners. Initially cultivation plots will be
small, limited by the cost of labour and fencing. At a later
stage, more associations may be established, and larger areas
will come under cultivation.
• Mechanised cultivation: current practice involves the leasing
out of large tracts on the northern Loita plains for commercial
wheat production. Again, on Koyake land holdings this will
only be viable if plots can be amalgamated. Expected returns
will be approximately US $60 ha)1 (Norton-Griffiths, 1995;
Thompson & Homewood, 2002).
• Land sales: landowners may choose to sell their land to
outsiders. As has been shown in Kajiado, fencing has rapidly
accelerated as new landowners consolidate and protect their
agricultural or livestock developments (Rutten, 1992).
On the critical land pressures building in the Mara, the
Narok County Council is silent. Whilst the council receives
massive revenues from wildlife-based tourism (Lamprey, in
prep.), its recent development plans merely state that tourism
is beneficial, agriculture should be promoted, and problem
animals should be controlled (Narok District Development
Plans, 1994, 1997). Meanwhile, conservation NGOs have
hastily assisted in the preparation of a Natural Resources
Management Plan 2001–05 for the Mara ranches (ACC, 2001).
This plan prescribes the zonation of Koyake into different
landuses, with interventions such as improvements in live-
stock, disease control, and extension of better farming
methods. However, subdivision is proceeding, and land-zoning
cannot easily be implemented when new landowners embark
on their own initiatives for self-advancement45. In an effort to
maintain Koyake as a wildlife area after subdivision, tour
operator associations are proposing to directly compensate
landowners to keep their land open and unfenced. Spatial
planning, so difficult to implement in the Mara, will be central
to this scheme. In a simpler zonation plan, tourism associa-
tions propose that Koyake be divided into just two manage-
ment blocks. Block I will cover all prime game-viewing land
holdings from the Mara River eastwards to the Aitong-Talek
road, and landowners will be actively encouraged to maintain
their areas for tourism through compensation. Block II lying
east of the Aitong-Talek road, has lower tourism potential, and
is targeted for settlement and water-development projects
(M. McCartney, pers. comm.; Lane, 2002).46
Classical approaches to ‘community-based wildlife manage-
ment’, involving the sharing of tourism revenues amongst
pastoralists with communal land tenure, have largely failed in
the Mara. Whilst the group ranches harbour a world-famous
wildlife resource, it has not been in the interests of leaders to
45
On a practical note, this plan has designated the entire Enkikwe area,
where immigration and settlement is occurring most rapidly, as the
‘conservation and tourism zone’ where settlement is supposedly dis-
couraged.
46
The District Land Office has now conceptually divided the ranch
into five blocks, the occupation of which will depend on ranch
membership status.
1025
R.H. Lamprey and R.S. Reid
distribute benefits equitably. An affluent minority has done
very well, whilst the main costs of keeping land open for
wildlife have been borne by ordinary Maasai. There are strong
parallels with the Kajiado ranches, where, according to
Campbell (1993, p. 268), ‘the political and economic elites
who have conspired to prevent the development of policies
which would have expanded the economic potential of these
areas for the majority of their inhabitants now seek to acquire
these areas for alternative land uses which will yield significant
wealth for the few.’ Out of such arguments, a more pragmatic
conservation narrative holds that CBWM may have little role
to play in conserving wildlife in communal areas outside
protected areas, where human needs are increasing rapidly and
other livelihoods are more rewarding (Barrett & Arcese, 1995;
Neumann, 1997; Songorwa, 1999). As exemplified by the Mara
ranches, where government wildlife agencies have little control
over landuse, the last chance for wildlife will involve the
creation of conservancies on private land, with adequate
compensation paid to individual landowners to forgo cultiva-
tion. This process of ‘set aside’ will not be driven by
governments or donors (Norton-Griffiths, 1995), but by a
private sector anxious to conserve one of Kenya’s most
important tourism areas.
ACKNOWLEDGMENTS
We thank Mike Rainy and Cathy Wilson for their assistance in
collecting data during the January 1999 aerial survey, and staff
of the Department of Resource Surveys and Remote Sensing for
providing SRF data. This research was funded in 1982/3 through
grants from the Natural Environment Research Council in UK,
the New York Zoological Society, the UK Overseas Develop-
ment Administration (now DfID), The East African Wildlife
Society and partially in 1999 through a grant from USAID
through the International Livestock Research Institute.
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BIOSKETCHES

Richard Lamprey is an ecologist specializing in wildlife conservation and management, with experience in Kenya, Tanzania,
Uganda, Ethiopia and Eritrea. His particular focus is to train wildlife agency staff in ecological monitoring, biodiversity assessment
and protected area design. Since 1995 he has assisted the Uganda Wildlife Authority in preparing and implementing a protected area
system plan for Uganda.

Robin Reid is a systems ecologist who leads research on livestock and environment issues at ILRI, focusing on the causes of land-use
change and how such changes affect ecosystems in Africa. She currently leads a global pastoral systems project at ILRI, which
attempts to balance pastoral development and ecosystem conservation.

1032 Journal of Biogeography 31, 997–1032, ª 2004 Blackwell Publishing Ltd

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