Long-term surveys of age structure in 13 ungulate and one ostrich species in the Serengeti, 1926-2018.

The Serengeti ecosystem spans an extensive network of protected areas in Tanzania, eastern Africa, and a UNESCO Wold Heritage Site. It is home to some of the largest animal migrations on the planet. Here, we describe a dataset consisting of the sample counts of three age classes (infant, juvenile and adult) of 13 ungulate and one ostrich species. Sample counts were tallied visually from the ground, or, in some instances, aerial photographs, during a period extending from 1926 to 2018. Observed animals were assigned to age classes based on specific criteria for each species. For nine of the 14 species of this dataset, the number of sampling years is over 30. This resulted in a total of 533 different records of count across age classes. By computing age-class ratios, these data can be used to measure long-term recruitment success at different ages of the tallied species. In particular, the temporal extent of these data allows comparison of patterns to other long-term processes, such as the El Niño-Southern Oscillation (ENSO).

Long-term monitoring of Serengeti bird species occurrence, abundance, and habitat.

The Serengeti ecosystem contains one of the most diverse bird assemblages in Africa. We present here a data set consisting of abundances of bird species in different habitats of the Serengeti ecosystem over a 87-yr time frame. This data set comprises 66,643 georeferenced occurrences for 568 species from 1929 to 2017. Most records contain feeding location, food source, distribution status, and observation locality. The records originate from three different but complementary methodologies: points, sites, and transects. The point method (bird species records 1929-2017) is based on ad hoc observations and includes rare species or those in special habitats. These points came from published records as well from the research program of A. R. E. Sinclair and colleagues. The site method (1966-2017) is based on structured observations at sites selected to represent specific habitats, and replicated within habitats and over time. At each site, birds were recorded by sight and sound over a radius of 50 m for 10 min. The transect method (1997-2011) is based on road transects covering different areas of the ecosystem. Road transects were traversed using a vehicle with observers travelling at 30 km/h. Bird species were those easily seen from a vehicle out to 50 m either side. As most transects were traversed multiple times, this method provides information on temporal change in abundance for a select set of species. No copyright restrictions apply to the use of this data set other than citing this publication.

Landscape-level movement patterns by lions in western Serengeti: comparing the influence of inter-specific competitors, habitat attributes and prey availability.

BACKGROUND: Where apex predators move on the landscape influences ecosystem structure and function and is therefore key to effective landscape-level management and species-specific conservation. However the factors underlying predator distribution patterns within functional ecosystems are poorly understood. Predator movement should be sensitive to the spatial patterns of inter-specific competitors, spatial variation in prey density, and landscape attributes that increase individual prey vulnerability. We investigated the relative role of these fundamental factors on seasonal resource utilization by a globally endangered apex carnivore, the African lion (Panthera leo) in Tanzania's Serengeti National Park. Lion space use was represented by novel landscape-level, modified utilization distributions (termed "localized density distributions") created from telemetry relocations of individual lions from multiple neighbouring prides. Spatial patterns of inter-specific competitors were similarly determined from telemetry re-locations of spotted hyenas (Crocuta crocuta), this system's primary competitor for lions; prey distribution was derived from 18 months of detailed census data; and remote sensing data was used to represent relevant habitat attributes. RESULTS: Lion space use was consistently influenced by landscape attributes that increase individual prey vulnerability to predation. Wet season activity, when available prey were scarce, was concentrated near embankments, which provide ambush opportunities, and dry season activity, when available prey were abundant, near remaining water sources where prey occurrence is predictable. Lion space use patterns were positively associated with areas of high prey biomass, but only in the prey abundant dry season. Finally, at the broad scale of this analysis, lion and hyena space use was positively correlated in the comparatively prey-rich dry season and unrelated in the wet season, suggesting lion movement was unconstrained by the spatial patterns of their main inter-specific competitors. CONCLUSIONS: The availability of potential prey and vulnerability of that prey to predation both motivate lion movement decisions, with their relative importance apparently mediated by overall prey abundance. With practical and theoretical implications, these results suggest that while top carnivores are consistently cognizant of how landscape features influence individual prey vulnerability, they also adopt a flexible approach to range use by adjusting spatial behaviour according to fluctuations in local prey abundance.

The predator-prey power law: Biomass scaling across terrestrial and aquatic biomes.

Ecosystems exhibit surprising regularities in structure and function across terrestrial and aquatic biomes worldwide. We assembled a global data set for 2260 communities of large mammals, invertebrates, plants, and plankton. We find that predator and prey biomass follow a general scaling law with exponents consistently near ¾. This pervasive pattern implies that the structure of the biomass pyramid becomes increasingly bottom-heavy at higher biomass. Similar exponents are obtained for community production-biomass relations, suggesting conserved links between ecosystem structure and function. These exponents are similar to many body mass allometries, and yet ecosystem scaling emerges independently from individual-level scaling, which is not fully understood. These patterns suggest a greater degree of ecosystem-level organization than previously recognized and a more predictive approach to ecological theory.

Resource wars and conflict ivory: the impact of civil conflict on elephants in the Democratic Republic of Congo--the case of the Okapi Reserve.

Human conflict generally has substantial negative impacts on wildlife and conservation. The recent civil war (1995-2006) in the Democratic Republic of Congo (DRC) resulted in a significant loss of wildlife, including elephants, due to institutional collapse, lawlessness and unbridled exploitation of natural resources such as minerals, wood, ivory and bushmeat. We used data from distance sampling surveys conducted before and after the war in a protected forest, the Okapi Faunal Reserve, to document changes in elephant abundance and distribution. We employed Generalized Additive Models to relate changes in elephant distribution to human and environmental factors. Populations declined by nearly fifty percent coinciding with a major increase in elephant poaching as indicated by reports of ivory trade during the war. Our results suggest that humans influenced elephant distribution far more than habitat, both before and after the war, but post-war models explained more of the variation. Elephant abundance declined more, closer to the park boundary and to areas of intense human activity. After the war, elephant densities were relatively higher in the centre of the park where they were better protected, suggesting that this area may have acted as a refuge. In other sites in Eastern DRC, where no protection was provided, elephants were even more decimated. Post-war dynamics, such as weakened institutions, human movements and availability of weapons, continue to affect elephants. Survival of remaining populations and recovery will be determined by these persistent factors and by new threats associated with growing human populations and exploitation of natural resources. Prioritizing wildlife protection, curbing illegal trade in ivory and bushmeat, and strengthening national institutions and organizations in charge of conservation will be crucial to counter these threats.

Trophic downgrading of planet Earth.

Until recently, large apex consumers were ubiquitous across the globe and had been for millions of years. The loss of these animals may be humankind's most pervasive influence on nature. Although such losses are widely viewed as an ethical and aesthetic problem, recent research reveals extensive cascading effects of their disappearance in marine, terrestrial, and freshwater ecosystems worldwide. This empirical work supports long-standing theory about the role of top-down forcing in ecosystems but also highlights the unanticipated impacts of trophic cascades on processes as diverse as the dynamics of disease, wildfire, carbon sequestration, invasive species, and biogeochemical cycles. These findings emphasize the urgent need for interdisciplinary research to forecast the effects of trophic downgrading on process, function, and resilience in global ecosystems.

Predicted impact of barriers to migration on the Serengeti wildebeest population.

The Serengeti wildebeest migration is a rare and spectacular example of a once-common biological phenomenon. A proposed road project threatens to bisect the Serengeti ecosystem and its integrity. The precautionary principle dictates that we consider the possible consequences of a road completely disrupting the migration. We used an existing spatially-explicit simulation model of wildebeest movement and population dynamics to explore how placing a barrier to migration across the proposed route (thus creating two disjoint but mobile subpopulations) might affect the long-term size of the wildebeest population. Our simulation results suggest that a barrier to migration--even without causing habitat loss--could cause the wildebeest population to decline by about a third. The driver of this decline is the effect of habitat fragmentation (even without habitat loss) on the ability of wildebeest to effectively track temporal shifts in high-quality forage resources across the landscape. Given the important role of the wildebeest migration for a number of key ecological processes, these findings have potentially important ramifications for ecosystem biodiversity, structure, and function in the Serengeti.

A disease-mediated trophic cascade in the Serengeti and its implications for ecosystem C.

Tree cover is a fundamental structural characteristic and driver of ecosystem processes in terrestrial ecosystems, and trees are a major global carbon (C) sink. Fire and herbivores have been hypothesized to play dominant roles in regulating trees in African savannas, but the evidence for this is conflicting. Moving up a trophic scale, the factors that regulate fire occurrence and herbivores, such as disease and predation, are poorly understood for any given ecosystem. We used a Bayesian state-space model to show that the wildebeest population eruption that followed disease (rinderpest) eradication in the Serengeti ecosystem of East Africa led to a widespread reduction in the extent of fire and an ongoing recovery of the tree population. This supports the hypothesis that disease has played a key role in the regulation of this ecosystem. We then link our state-space model with theoretical and empirical results quantifying the effects of grazing and fire on soil carbon to predict that this cascade may have led to important shifts in the size of pools of C stored in soil and biomass. Our results suggest that the dynamics of herbivores and fire are tightly coupled at landscape scales, that fire exerts clear top-down effects on tree density, and that disease outbreaks in dominant herbivores can lead to complex trophic cascades in savanna ecosystems. We propose that the long-term status of the Serengeti and other intensely grazed savannas as sources or sinks for C may be fundamentally linked to the control of disease outbreaks and poaching.

Effect of habitat area and isolation on fragmented animal populations.

Habitat destruction has driven many once-contiguous animal populations into remnant patches of varying size and isolation. The underlying framework for the conservation of fragmented populations is founded on the principles of island biogeography, wherein the probability of species occurrence in habitat patches varies as a function of patch size and isolation. Despite decades of research, the general importance of patch area and isolation as predictors of species occupancy in fragmented terrestrial systems remains unknown because of a lack of quantitative synthesis. Here, we compile occupancy data from 1,015 bird, mammal, reptile, amphibian, and invertebrate population networks on 6 continents and show that patch area and isolation are surprisingly poor predictors of occupancy for most species. We examine factors such as improper scaling and biases in species representation as explanations and find that the type of land cover separating patches most strongly affects the sensitivity of species to patch area and isolation. Our results indicate that patch area and isolation are indeed important factors affecting the occupancy of many species, but properties of the intervening matrix should not be ignored. Improving matrix quality may lead to higher conservation returns than manipulating the size and configuration of remnant patches for many of the species that persist in the aftermath of habitat destruction.

Climate extremes promote fatal co-infections during canine distemper epidemics in African lions.

Extreme climatic conditions may alter historic host-pathogen relationships and synchronize the temporal and spatial convergence of multiple infectious agents, triggering epidemics with far greater mortality than those due to single pathogens. Here we present the first data to clearly illustrate how climate extremes can promote a complex interplay between epidemic and endemic pathogens that are normally tolerated in isolation, but with co-infection, result in catastrophic mortality. A 1994 canine distemper virus (CDV) epidemic in Serengeti lions (Panthera leo) coincided with the death of a third of the population, and a second high-mortality CDV epidemic struck the nearby Ngorongoro Crater lion population in 2001. The extent of adult mortalities was unusual for CDV and prompted an investigation into contributing factors. Serological analyses indicated that at least five "silent" CDV epidemics swept through the same two lion populations between 1976 and 2006 without clinical signs or measurable mortality, indicating that CDV was not necessarily fatal. Clinical and pathology findings suggested that hemoparsitism was a major contributing factor during fatal epidemics. Using quantitative real-time PCR, we measured the magnitude of hemoparasite infections in these populations over 22 years and demonstrated significantly higher levels of Babesia during the 1994 and 2001 epidemics. Babesia levels correlated with mortalities and extent of CDV exposure within prides. The common event preceding the two high mortality CDV outbreaks was extreme drought conditions with wide-spread herbivore die-offs, most notably of Cape buffalo (Syncerus caffer). As a consequence of high tick numbers after the resumption of rains and heavy tick infestations of starving buffalo, the lions were infected by unusually high numbers of Babesia, infections that were magnified by the immunosuppressive effects of coincident CDV, leading to unprecedented mortality. Such mass mortality events may become increasingly common if climate extremes disrupt historic stable relationships between co-existing pathogens and their susceptible hosts.

Group formation stabilizes predator-prey dynamics.

Theoretical ecology is largely founded on the principle of mass action, in which uncoordinated populations of predators and prey move in a random and well-mixed fashion across a featureless landscape. The conceptual core of this body of theory is the functional response, predicting the rate of prey consumption by individual predators as a function of predator and/or prey densities. This assumption is seriously violated in many ecosystems in which predators and/or prey form social groups. Here we develop a new set of group-dependent functional responses to consider the ecological implications of sociality and apply the model to the Serengeti ecosystem. All of the prey species typically captured by Serengeti lions (Panthera leo) are gregarious, exhibiting nonlinear relationships between prey-group density and population density. The observed patterns of group formation profoundly reduce food intake rates below the levels expected under random mixing, having as strong an impact on intake rates as the seasonal migratory behaviour of the herbivores. A dynamical system model parameterized for the Serengeti ecosystem (using wildebeest (Connochaetes taurinus) as a well-studied example) shows that grouping strongly stabilizes interactions between lions and wildebeest. Our results suggest that social groups rather than individuals are the basic building blocks around which predator-prey interactions should be modelled and that group formation may provide the underlying stability of many ecosystems.

Effective enforcement in a conservation area.

Wildlife within protected areas is under increasing threat from bushmeat and illegal trophy trades, and many argue that enforcement within protected areas is not sufficient to protect wildlife. We examined 50 years of records from Serengeti National Park in Tanzania and calculated the history of illegal harvest and enforcement by park authorities. We show that a precipitous decline in enforcement in 1977 resulted in a large increase in poaching and decline of many species. Conversely, expanded budgets and antipoaching patrols since the mid-1980s have greatly reduced poaching and allowed populations of buffalo, elephants, and rhinoceros to rebuild.

The role of predation in the decline and extirpation of woodland caribou.

To select appropriate recovery strategies for endangered populations, we must understand the dynamics of small populations and distinguish between the possible causes that drive such populations to low numbers. It has been suggested that the pattern of population decline may be inversely density-dependent with population growth rates decreasing as populations become very small; however, empirical evidence of such accelerated declines at low densities is rare. Here we analyzed the pattern of decline of a threatened population of woodland caribou (Rangifer tarandus caribou) in British Columbia, Canada. Using information on the instantaneous rate of increase relative to caribou density in suitable winter foraging habitat, as well as on pregnancy rates and on causes and temporal distribution of mortalities from a sample of 349 radiocollared animals from 15 subpopulations, we tested 3 hypothesized causes of decline: (a) food regulation caused by loss of suitable winter foraging habitat, (b) predation-sensitive foraging caused by loss of suitable winter foraging habitat and (c) predation with caribou being secondary prey. Population sizes of caribou subpopulations ranged from < 5 to > 500 individuals. Our results showed that the rates of increase of these subpopulations varied from -0.1871 to 0.0496 with smaller subpopulations declining faster than larger subpopulations. Rates of increase were positively related to the density of caribou in suitable winter foraging habitat. Pregnancy rates averaged 92.4% +/-2.24 and did not differ among subpopulations. In addition, we found predation to be the primary cause of mortality in 11 of 13 subpopulations with known causes of mortality and predation predominantly occurred during summer. These results are consistent with predictions that caribou subpopulations are declining as a consequence of increased predation. Recovery of these woodland caribou will thus require a multispecies perspective and an appreciation for the influence of inverse density dependence on population trajectories.

Ecological change, group territoriality, and population dynamics in Serengeti lions.

Territorial behavior is expected to buffer populations against short-term environmental perturbations, but we have found that group living in African lions causes a complex response to long-term ecological change. Despite numerous gradual changes in prey availability and vegetative cover, regional populations of Serengeti lions remained stable for 10- to 20-year periods and only shifted to new equilibria in sudden leaps. Although gradually improving environmental conditions provided sufficient resources to permit the subdivision of preexisting territories, regional lion populations did not expand until short-term conditions supplied enough prey to generate large cohorts of surviving young. The results of a simulation model show that the observed pattern of "saltatory equilibria" results from the lions' grouping behavior.

What limits the Serengeti zebra population?

The populations of the ecologically dominant ungulates in the Serengeti ecosystem (zebra, wildebeest and buffalo) have shown markedly different trends since the 1960s: the two ruminants both irrupted after the elimination of rinderpest in 1960, while the zebras have remained stable. The ruminants are resource limited (though parts of the buffalo population have been limited by poaching since the 1980s). The zebras' resource acquisition tactics should allow them to outcompete the ruminants, but their greater spatial dispersion makes them more available to predators, and it has been suggested that this population is limited by predation. To investigate the mechanisms involved in the population dynamics of Serengeti zebra, we compared population dynamics among the three species using demographic models based on age-class-specific survival and fecundity. The only major difference between zebra and the two ruminants occurred in the first-year survival. We show that wildebeest have a higher reproductive potential than zebra (younger age at first breeding and shorter generation time). Nevertheless, these differences in reproduction cannot account for the observed differences in the population trends between the zebra and the ruminants. On the other hand, among-species differences in first-year survival are great enough to account for the constancy of zebra population size. We conclude that the very low first-year survival of zebra limits this population. We provide new data on predation in the Serengeti and show that, as in other ecosystems, predation rates on zebras are high, so predation could hold the population in a "predator pit". However, lion and hyena feed principally on adult zebras, and further work is required to discover the process involved in the high mortality of foals.