Possible causes of decreasing migratory ungulate populations in an East African savannah after restrictions in their seasonal movements

In many areas in Africa, seasonal movements of migratory ungulates are restricted and their population numbers decline, for example in the Tarangire region, Tanzania. Here, agriculture restricts migration of ungulates to their wet season ranges. We investigated whether low forage quality or supply are possible causes of population decline of wildebeest and zebra when access to these wet season ranges is restricted and migratory herds have to reside in the dry season range year-round. We simulated grazing through a clipping experiment in the dry season range during the wet season. Clipping negatively affected forage supply and had a positive effect on forage quality by increasing proportions of live and leaf biomass as well as nutrient concentrations in the leaves. However, increase in forage quality in the dry season range due to grazing was not as such that requirements of wildebeest during the wet season, when females are lactating, could be met. We conclude that low forage quality in the dry season range during the wet season could cause the decrease in migratory ungulate populations in the Tarangire region. With this study, the necessity of protecting wet season ranges from expanding human activities to safeguard migratory systems is supported.     

Elephant survival,rainfall and the confounding effects of water provision and fences

Elephant are increasing across some areas of Africa leading to concerns that they may reduce woodlands through their feeding. Droughts may help limit elephant numbers, but they are generally both episodic and local. To explore more general impacts of rainfall, we examine how its annual variation influences elephant survival across ten sites. These sites span an almost coast-to-coast transect of southern Africa that holds the majority of the ~500,000 remaining savanna elephants. Elephants born in high rainfall years survive better than elephants born in low rainfall years. The relationship is generally weak, except at the two fenced sites, where rainfall greatly influenced juvenile survival. In these two sites, there are also extensive networks of artificial water. Rainfall likely affects elephant survival through its influence on food. The provision of artificial water opens new areas for elephants in the dry season, while fencing restricts their movements in the wet season. We conclude that the combination of these factors makes elephant survival more susceptible to reductions in rainfall. As a result, elephants living in enclosed reserves may be the first populations to feel the impacts of global warming which will decrease average rainfall and increase the frequency of droughts. A way to prevent these elephants from damaging the vegetation within these enclosed parks is for managers to reduce artificial water sources or, whenever practical, to remove fences.     

Reversal of breeding season by lowland birds at higher altitudes in western Cameroon

Evidence of breeding during the dry season is given for 31 species of lowland birds in montane and semi-montane areas in western Cameroon. At least 17 of these species are shown to breed in the wet season at lower altitudes in West Africa. This reversal of breeding season may be due to the unusually heavy rainfall and high humidities which cause temperatures to decrease more rapidly with altitude than on mountains with drier climates. Low temperatures and heavy rainfall during the wet season prevent almost all montane species from breeding then, and affect similarly the lowland birds whose ranges overlap with those of montane birds. Breeding seasons of some lowland species in Cameroon have previously been considered prolonged, but separation of breeding records by altitude reveals clear seasonality.     

The role of wetlands in wildlife migration in the Tarangire ecosystem,Tanzania

Twenty-two years of rainfall data from six sites, 5 years of animal migration data and 2 years of water quality at 13 sites were explored to quantify the role of water in the Tarangire ecosystem. Inter-annual fluctuations in rainfall were large and not predictable solely from the Southern Oscillation Index. Seasonal fluctuations of rainfall were pronounced, with marked wet and dry seasons. In the dry season, the only drinking water available for wildlife was the Tarangire River and a number of small, scattered wetland-fringed water holes. Their salinity was often high (>8 ppt) and was higher in dry years than in wet years, as well as at the start of the wet season. Water quantity and quality may control the annual migration of wildebeest, zebra, elephants and buffaloes. These animals aggregate in the dry season in areas with the least salty water. The timing of seasonal variations in rainfall is largely predictable and controls annual migration. All wildebeest and most zebras migrated out of Tarangire National Park and into the wider Tarangire ecosystem at the start of the wet season, and they returned into the park in the dry season. Some elephants and buffaloes also migrated in out of the park and a larger resident population remained, whose size may vary inter-annually depending on surface water quantity and quality. The extent of the migration zone may also vary inter-annually.This revised version wa published online in March 2005 with corrections to the issue cover date.     

THE BREEDING SEASON IN S.W. ECUADOR

The breeding season on the Santa Elena peninsula in S.W. Ecuador is described on the basis of 1761 nests found in the four years 1955 to 1958. The environment and climate are outlined—a cool, dry season from about May to November and a warm, variable wet season from December to April, when alone rain may fall. It is shown that each year the general breeding season is closely correlated with the rainfall. The peak of breeding varies from year to year by at least a month and its length from about six weeks to three and a half months.
Although all species for which there are enough data, are stimulated to breed after important falls of rain, most of those species of small land-birds which are completely resident, attempt to nest before the rain and often continue for some time afterwards. Some specific differences, between seed- and insect-eating finches, seem to be adaptations to food supply and the availability of nest sites and building material. Such evidence as there is suggests that raptors, waders and waterfowl also nest in the wet season (often late), rather than in the dry season. (Unfortunately herons and sea-birds do not nest in the area.)
Annual differences in the amount of breeding by the same species are thought not to be entirely due to differences in weather between years.
It is suggested that the late (August-September) breeding in the Galápagos Islands described by Lack (1950) and the apparent dry-season nesting of raptors, waders and waterfowl there might be found to be linked with variable or abnormally wet weather, if records of rainfall from the islands were available.     

Response of African Elephants (Loxodonta africana) to Seasonal Changes in Rainfall

The factors that trigger sudden, seasonal movements of elephants are uncertain. We hypothesized that savannah elephant movements at the end of the dry season may be a response to their detection of distant thunderstorms. Nine elephants carrying Global Positioning System (GPS) receivers were tracked over seven years in the extremely dry and rugged region of northwestern Namibia. The transition date from dry to wet season conditions was determined annually from surface- and satellite-derived rainfall. The distance, location, and timing of rain events relative to the elephants were determined using the Tropical Rainfall Measurement Mission (TRMM) satellite precipitation observations. Behavioral Change Point Analysis (BCPA) was applied to four of these seven years demonstrating a response in movement of these elephants to intra- and inter-seasonal occurrences of rainfall. Statistically significant changes in movement were found prior to or near the time of onset of the wet season and before the occurrence of wet episodes within the dry season, although the characteristics of the movement changes are not consistent between elephants and years. Elephants in overlapping ranges, but following separate tracks, exhibited statistically valid non-random near-simultaneous changes in movements when rainfall was occurring more than 100 km from their location. While the environmental trigger that causes these excursions remains uncertain, rain-system generated infrasound, which can travel such distances and be detected by elephants, is a possible trigger for such changes in movement.     

Scales of spatiotemporal variability in macroinvertebrate abundance and diversity in monsoonal streams: detecting environmental change

1. We quantified spatial and temporal variability in benthic macroinvertebrate species richness, diversity and abundance in six unpolluted streams in monsoonal Hong Kong at different scales using a nested sampling design. The spatial scales were regions, stream sites and stream sections within sites; temporal scales were years (1997–99), seasons (dry versus wet seasons) and days within seasons. 2. Spatiotemporal variability in total abundance and species richness was greater during the wet season, especially at small scales, and tended to obscure site‐ and region‐scale differences, which were more conspicuous during the dry season. Total abundance and richness were greater in the dry season, reflecting the effects of spate‐induced disturbance during the wet season. Species diversity showed little variation at the seasonal scale, but variability at the site scale was apparent during both seasons. 3. Despite marked variations in monsoonal rainfall, inter‐year differences in macroinvertebrate richness and abundance at the site scale during the wet season were minor. Inter‐year differences were only evident during the dry season when streams were at base flow and biotic interactions may structure assemblages. 4. Small‐scale patchiness within riffles was the dominant spatial scale of variation in macroinvertebrate richness, total abundance and densities of common species, although site or region was important for some species. The proportion of total variance contributed by small‐scale spatial variability increased during the dry season, whereas temporal variability associated with days was greater during the wet season. 5. The observed patterns of spatiotemporal variation have implications for detection of environmental change or biomonitoring using macroinvertebrate indicators in streams in monsoonal regions. Sampling should be confined to the dry season or, in cases where more resources are available, make use of data from both dry and wet seasons. Sampling in more than one dry season is required to avoid the potentially confounding effects of inter‐year variation, although variability at that scale was relatively small.     

Multi‐century tree‐ring precipitation record reveals increasing frequency of extreme dry events in the upper Blue Nile River catchment

Climate‐related environmental and humanitarian crisis are important challenges in the Great Horn of Africa (GHA). In the absence of long‐term past climate records in the region, tree‐rings are valuable climate proxies, reflecting past climate variations and complementing climate records prior to the instrumental era. We established annually resolved multi‐century tree‐ring chronology from Juniperus procera trees in northern Ethiopia, the longest series yet for the GHA. The chronology correlates significantly with wet‐season (r = .64, p < .01) and annual (r = .68, p < .01) regional rainfall. Reconstructed rainfall since A.D. 1811 revealed significant interannual variations between 2.2 and 3.8 year periodicity, with significant decadal and multidecadal variations during 1855–1900 and 1960–1990. The duration of negative and positive rainfall anomalies varied between 1–7 years and 1–8 years. Approximately 78.4% (95%) of reconstructed dry (extreme dry) and 85.4% (95%) of wet (extreme wet) events lasted for 1 year only and corresponded to historical records of famine and flooding, suggesting that future climate change studies should be both trend and extreme event focused. The average return periods for dry (extreme dry) and wet (extreme wet) events were 4.1 (8.8) years and 4.1 (9.5) years. Extreme‐dry conditions during the 19th century were concurrent with drought episodes in equatorial eastern Africa that occurred at the end of the Little Ice Age. El Niño and La Niña events matched with 38.5% and 50% of extreme‐dry and extreme‐wet events. Equivalent matches for positive and negative Indian Ocean Dipole events were weaker, reaching 23.1 and 25%, respectively. Spatial correlations revealed that reconstructed rainfall represents wet‐season rainfall variations over northern Ethiopia and large parts of the Sahel belt. The data presented are useful for backcasting climate and hydrological models and for developing regional strategic plans to manage scarce and contested water resources. Historical perspectives on long‐term regional rainfall variability improve the interpretation of recent climate trends.     

Trade-off between resource seasonality and predation risk explains reproductive chronology in impala

We investigated the variation in birth synchrony displayed by impala Aepyceros melampus populations across their distribution from southern to eastern Africa. Our analysis was based on field data from Chobe National Park in Botswana and Mala Mala Private Game Reserve in South Africa (4 and 13 years of monitoring, respectively). We compared our results with those from other studies conducted across the impala species range. Impala lambing was highly synchronized in Chobe with 90% of lambs born within 2 weeks in mid-November. Variation in rainfall in the preceding wet season explained 74% of variation in the dates of the first lamb observation in Mala Mala. In Chobe, the earliest birth peak occurred after the highest rainfall and the body condition of lambs in that cohort was also best for both males and females. No association was found between the lunar cycle and the estimated onset of the conception period, despite previous studies having found an association between the lunar cycle and the rutting behaviour in males. On a regional scale, impalas in areas with a marked dry season (several months with no rain) tend to synchronize births with the onset of the rains, when grass quality is highest. Number of months with rain explained 78% of the regional variation in birth synchrony. Neither latitude nor total rainfall contributed significantly to a stepwise multiple regression model. These data support the theory that impalas synchronize births in areas with a highly seasonal food supply, and temporally space births in less seasonal (equatorial) areas to reduce predation risks.     

The natural regulation of buffalo populations in East Africa: The food supply as a regulating factor,and competition*

Evidence is presented to show that both the quality and quantity of food available to a buffalo population falls below the minimum maintenance requirements of that population at certain times of the year. In the Serengeti grasslands there was a shortage of the only good quality component, grass leaf, in the dry season, with the result that the animals consumed an increasing proportion of poor quality food such as grass stem. By the end of the dry season the diet had dropped in quality below the minimum maintenance level. In areas such as Mt. Meru where there was a more continuous growing season, the high density of animals kept the standing crop of leaf at a low level. During the cooler dry season the growth of leaf became insufficient in quantity for the maintenance requirements of the population. These two quite different situations suggested that food shortage was a more general phenomenon in eastern Africa. Various measurements of feeding behaviour were made. Total grazing time per 24 h did not differ between seasons but ruminating time may have increased as the season became drier and could have been a response to the more fibrous food. Analysis for cycles of activity showed that there was more temporal organization during the dry season. These changes in activity cycles appeared to be related to the increase in energy expenditure produced by heat stress and sweating. Old animals with poor teeth did not compensate for the poor food supply by changing their feeding behaviour. There was a positive relationship between annual rainfall and mean crude density in different areas of eastern Africa, indicating that regulation was taking place. Since rainfall determined the amount of available food, it could have operated through the food supply. On a finer scale it was found that the extent of the preferred riverine habitats was also related to density. Thus rainfall, the extent of riverine habitat and perhaps soil moisture were three limiting factors that determined mean density and all could have taken effect through the food supply. As a result of initial selective grazing the amount of available leaf declined as the dry season progressed to the extent that by the end of the season the proportion of this component in the diet fell to a very low level. The impact of the population on its limited food supply indicated that intra-specific competition was acting as the cause of regulation. Measurements of wildebeest eating the same food in the same habitats as buffalo showed that inter-specific competition was also taking place. A small proportion (7“) of the wildebeest population could have reduced the buffalo population by approximately 18^o,‘, from its potential population size. The buffalo population in the Serengeti was regulated by adult mortality which was caused by undernutrition as a result of food shortage. This food shortage was caused by intra- and inter-specific competition. The effect of predation and disease was to hasten the response of the population to changes in the food supply. The limiting factors determining the mean level of the available food were, amongst others, rainfall, soil moisture and the extent of the preferred riverine habitat. The effect of interspecific competition could result in a complex regulation of populations through their food supply. There appears to be no foundation for hypotheses which invoke over-utilization or damage as a consequence of regulation through food.     

Climate variability and change over southern Africa: impacts and challenges

In this paper, the influence of climate variability and change on the environment was studied over southern Africa using ground-based and remotely sensed data. A time series analysis of rainfall and temperature anomalies indicated that there was a high rainfall and temperature variability in the region. The influence of global teleconnections on rainfall patterns over southern Africa showed that in some areas there was a spatial variation in their strength, increasing from west to east. Maps of NDVI, from 1982 to 2004, showed that changes in vegetation cover were more apparent during the dry season than during the wet season. The study also revealed that climate variability and change are linked to decreasing rainfall and hence, decreasing regional water resources and biodiversity and increasing environmental degradation. With the regional population expected increase, this depletion of resources poses the greatest regional environmental challenge to humankind.     

Plant compositional change over time increases with rainfall in Serengeti grasslands

Previous work from the Plains of Serengeti National Park (SNP) suggested that plant species composition (i.e. the composite measure of species' identities and abundances) is unchanged on the temporal scale of decades ( Belsky 1985 ), raising the possibility that African grazing ecosystems may be resistant to climate change. In this study, grassland plant composition was measured three times between 2000 and 2007 at eight permanent sites spanning the SNP environmental gradient. Confirming previous work, plots in the Serengeti Plains were relatively stable with respect to plant composition through time. In contrast, sites in all other regions of the ecosystem showed greater change in plant species composition during the study. Three factors, annual rainfall, dry season rainfall, and fire frequency, were linearly correlated (all r>0.73) with the amount of composition change at the sites. A path model, which accounted for the strong correlation between annual and dry season rainfall and for the known effect of annual rainfall on fire frequency, was used to elucidate which factors were associated with composition change and which were spuriously correlated. The path model demonstrated that change in plant composition was positively related to annual rainfall, but not dry season rainfall or fire frequency. However, dry season rainfall was positively associated with changes in plant species identity, one component of compositional change, across sites over time. Because climate models predict both increased wet season and decreased dry season rainfall in East Africa, these findings raise uncertainty about how these grasslands will respond to future changes. This study and others ( Belsky 1985 ) suggests that plant assemblages adapted to low rainfall in Serengeti may exhibit compositionally stability, while the same may not be true in regions of higher average rainfall and larger species pools.     

Nesting biology and mating system in an alpine population of Temminck's Stint Calidris temminckii

A colour-marked population of Temminck's Stint Calidris temminckii was studied in an alpine area of southern Norway over 3 years. Birds arrived from late May until late June, and egg-laying occurred over approximately 1 month. Each year, one to three males out of about 15–17 males still displayed after the last recorded clutch-completion date.
There were six documented cases of double-clutching. Inter-clutch intervals varied (range 2–8-9-1 days), longer intervals probably being caused by weather-induced food shortage. A successively bigamous mating pattern was recorded in both sexes. A polygynously (bigamously) mated male invariably incubated its first mate's clutch. Females consistently changed mates between layings of successive clutches, the last one incubated by themselves.
An excess of nesting females was found, particularly in parts of the study area where nesting started late. This appears to have been caused mainly by a considerable annual immigration of late-arriving females, having probably laid their first clutch(es) elsewhere, in nesting habitats that were available earlier. I suggest that male availability is relatively unimportant compared with other factors governing female movements between layings of successive clutches; females may increase their reproductive success, either by achieving a longer egg-laying season (i.e. by moving from nesting habitats/areas available early in the season to those available later) or by being capable of utilizing favourable feeding habitats/conditions in different areas.     

Seasonal polyphenism in Bicyclus (Lepidoptera: Satyridae) butterflies: different climates need different cues

A comparison is made between northern and southern hemisphere populations of Bicyclus butterflies in Africa regarding their responses in wing pattern polyphenism to seasonal change in rainfall and temperature. In southern habitats where temperature and rainfall are often positively correlated, a high temperature during the larval period induces conspicuous wet season forms whereas a fall in temperature elicits cryptic dry season forms. In northern habitats, however, where temperature and rainfall usually are negatively correlated, a rise in temperature should not induce a wet season form because such a rise is correlated with the onset of the dry season. Here, wing pattern plasticity, as measured using museum material, was regressed on mean monthly values for rainfall and temperature. Rainfall appeared to be a frequent determinant of wing pattern plasticity whereas temperature was much less often a significant independent variate. We conclude that the wing pattern may only respond to seasonal change in temperature if rainfall and temperature are positively correlated; in other situations rainfall remains the only significant determinant for wing pattern plasticity.     

THE MIGRATION PATTERNS OF QUELEA QUELEA IN AFRICA

Red-billed Queleas migrate, at the beginning of the wet season, away from their dry-season concentration areas and towards areas where rain started several weeks earlier. Considerable fat deposits are accumulated for this “early-rains migration”. The direction taken by the migrants, the distance they must fly, and the timing of the movement are dependent upon the timing of the rains and the way the rain-front moves. On the return “breeding migration” individuals in breeding condition stop to breed (in large aggregations) wherever they find conditions suitable for the founding of colonies. The location of the colonies can vary greatly from year to year. The migrations performed by several populations, in different parts of Africa, are predicted on the basis of general rainfall patterns, and the predictions tested against the facts available. There is evidence that individual females are able to produce a succession of broods in the same breeding season, at colonies which may be very far apart, and probably with different mates. The adaptive value of this “itinerant breeding” is discussed. Many other bird species, which breed at a particular phase of the wet-season/dry-season cycle, are expected to perform similar “itinerant breeding”.     

Dry season watering alters the significance of climate factors influencing phenology and growth of saplings of savanna woody species in central Zambia,southern Africa

Although tree growth in southern African savannas is correlated with rainfall in the wet season, some studies have shown that tree growth is controlled more by rainfall in the dry season. If more rainfall occurred in the dry season in future climates, it would affect the growth of savanna trees, especially saplings that have shallower roots which limit access to subsoil water during the dry season when leaf flush and shoot extension occur. Recent paleobotanical evidence has revealed that there was relatively more precipitation in the dry season in eastern Africa in the Eocene than under the current climate. Saplings therefore can be expected to respond more to water addition during the dry season than mature trees that have more stored water and deeper roots that access subsoil water. Accordingly, I hypothesized that irrigation in the dry season should (i) advance the onset of the growing season, (ii) increase growth rates and (iii) alter the growth responses of saplings to climate factors. To test these hypotheses saplings of five savanna woody species were irrigated during the hot‐dry season at a site in central Zambia and their monthly and annual growth rates compared to those of conspecifics growing under control conditions. Although the responses among the species were variable, all irrigated saplings had significantly higher monthly and annual growth rates than control plants. In addition, dry season watering significantly altered the climatic determinants of sapling growth by either strengthening the role of the same climatic factors that were important under control conditions or displacing them altogether. In conclusion, more precipitation during the hot‐dry season is likely to have significant positive effects on sapling growth and consequently reduce the sapling‐tree transition periods and promote future tree population recruitment in some southern African savanna tree species.     

FACTORS AFFECTING THE FOOD SUPPLY AND BREEDING SEASON OF RESIDENT BIRDS AND MOVEMENTS OF PALAEARCTIC MIGRANTS IN A TROPICAL AFRICAN SAVANNAH

The fact that Palaearctic migrants arrive in the northern tropical savannah of Africa during the dry season suggests potential competition for food with African species. Moreover, in the southern tropical savannah African species breed during the rainy seasons, when Palaearctic migrants are present. In the equatorial area of Serengeti, East Africa, an index of the food supply for insectivorous birds was obtained from 3 years of light-trap measurements and sweep net samples. Adults of Lepidoptera, Coleoptera, Orthoptera and Isoptera are sparse in the dry season but become locally abundant after the first rainstorms that mark its close. They are apparently blown by converging winds ahead of the inter-tropical front and settle to lay eggs where rain has fallen. These early storms therefore produce localized superabundances of food. In the ensuing rainy seasons insect abundance remains high. African insectivorous birds breed during the wet period, reaching a peak two months after the insect increase. It is suggested that this lag is due to the need to recover body condition, build up reserves for eggs, develop gonads and wait for vegetation and insect larvae to develop. In the samples available, breeding records of above-ground nesters peaked in the first rains, while ground-nesters peaked in the second (main) rains. Predators bred towards the end of the rains, when there is an abundance of fledglings and small mammals. Thus the food supply could act as the ultimate factor determining the timing of the breeding season in this area. Palaearctic migrants arrive in the Serengeti 4–10 weeks ahead of the main rain front. However, most species are only found where rain has fallen recently. When conditions dry up they move on to other wet areas. Thus they overlap with African species only where there is a superabundance of insects. When the rains become widespread Palaearctic migrants disperse into their usual habitats, and therefore appear not to compete for available resources with closely related species of African birds. The situation in West Africa, where residents and migrants overlap throughout the dry season, cannot be explained in the same terms.     

The sensitivity of annual grassland carbon cycling to the quantity and timing of rainfall

Global climate models predict significant changes to the rainfall regimes of the grassland biome, where C cycling is particularly sensitive to the amount and timing of precipitation. We explored the effects of both natural interannual rainfall variability and experimental rainfall additions on net C storage and loss in annual grasslands. Soil respiration and net primary productivity (NPP) were measured in treatment and control plots over four growing seasons (water years, or WYs) that varied in wet‐season length and the quantity of rainfall. In treatment plots, we increased total rainfall by 50% above ambient levels and simulated one early‐ and one late‐season storm. The early‐ and late‐season rain events significantly increased soil respiration for 2–4 weeks after wetting, while augmentation of wet‐season rainfall had no significant effect. Interannual variability in precipitation had large and significant effects on C cycling. We observed a significant positive relationship between annual rainfall and aboveground NPP across the study (P=0.01, r²=0.69). Changes in the seasonal timing of rainfall significantly affected soil respiration. Abundant rainfall late in the wet season in WY 2004, a year with average total rainfall, led to greater net ecosystem C losses due to a ～50% increase in soil respiration relative to other years. Our results suggest that C cycling in annual grasslands will be less sensitive to changes in rainfall quantity and more affected by altered seasonal timing of rainfall, with a longer or later wet season resulting in significant C losses from annual grasslands.     

Nesting Behavior of <Emphasis Type="Italic">Gorilla gorilla diehli</Emphasis> at Kagwene Mountain,Cameroon: Implications for Assessing Group Size and Density

We recorded nesting data at 569 fresh night nest sites, comprising 7032 individual nests, of Cross River gorillas inhabiting the Kagwene Mountain in western Cameroon. The mean night nest group size was 12.4. Overall, 55% of night nests were constructed on the ground and 45% in trees. Significantly more arboreal nests were constructed in the wet season (69%), vs. the dry season (19%). Day nest construction was common at Kagwene (n = 260 nest sites, mean nest group size = 5.98) and we encountered significantly more day nest sites in the wet season. Nest site reuse was also common (35%), though not related to season. Our results of nesting habits concur with those from other western gorilla studies, in which rainfall influences arboreal nesting. However, we encountered wet season arboreal nesting, day nest construction, and overall nest site reuse more frequently than reported for other sites. Our observations have considerable implications when estimating group size and density using traditional nest count data. The gorillas at Kagwene inhabit the highest altitudinal range of all Cross River gorilla subpopulations and rainfall is also high; therefore other subpopulations may demonstrate different nesting characteristics. However, one should consider our findings when attempting to estimate Cross River gorilla density at other localities through nest site data.     

Movements of the migratory wildebeest population in the Serengeti area between 1960 and 1973

Data on the monthly distributions of the Serengeti migratory wildebeest population between June 1960 and May 1973 have been analysed. It has been shown that the population performs an annual migration, spending the wet season on the Serengeti plains, then moving west towards Lake Victoria at the beginning of the dry season, and later north to the northern extension of the Serengeti National Park or into the Masai Mara Game Reserve in Kenya. The wildebeest return to the plains as soon as the rain begins, but if this is delayed they may move south-west from the northern areas into the corridor first. Annual differences in the pattern of the migration can be correlated with differences in rainfall. The rainfall affects particularly the timing of the movements of the wildebeest on and off the plains, and the extent to which they utilize the northern areas. A large increase in the size of the population over the years studied has resulted in a greater utilization of the northern areas. It has been shown that about half the total range of the wildebeest population is outside the boundaries of the Serengeti National Park, and it is therefore important that these areas should be protected if such a large population is to survive.