Restrictions on roaming Until the past century or so, the movement of wild animals was relatively unrestricted, and their travels contributed substantially to ecological processes. As humans have increasingly altered natural habitats, natural animal movements have been restricted. Tucker et al. examined GPS locations for more than 50 species. In general, animal movements were shorter in areas with high human impact, likely owing to changed behaviors and physical limitations. Besides affecting the species themselves, such changes could have wider effects by limiting the movement of nutrients and altering ecological interactions. Science , this issue p. 466

In this paper, we investigate the formation and function of the multilevelled, fissionfusion social structure in a free-ranging African elephant, Loxodonta africana, population. We quantitatively identified the existence of four social tiers by using cluster analysis on individual association data. We assessed the effects of season and study period on social structuring and levels of cohesion within and among social units. We found that second-tier units, potentially the equivalent of the 'family', were stable across seasonal periods but the number of units increased as the study progressed and the population grew. It appears that these units were sufficiently small not to be influenced by ecologically related factors, such as resource competition, that might otherwise lead to them splitting. On the other hand, third-and fourth-tier units were significantly affected by season in a way that suggests a trade-off between ecological costs (e.g. from resource competition) and different social and ecological benefits (e.g. from predator defence, territoriality, knowledge sharing and rearing of young). Age structure also appeared to influence this multitiered social organization. The size of second-tier social units was significantly affected by the age of matriarchs: units lead by matriarchs likely to be grandmothers (i.e. females 35 years and older) were significantly larger than those lead by younger matriarchs. We present a conceptual framework for understanding the openUP (June 2007) emergence of multiple-tier social structure from interactions driven by socioecological processes. This study is the first to use rigorous quantitative methods to statistically show the existence of four hierarchical tiers of social organization in a nonhuman animal. Additionally, our results elucidate the role that ecological processes play in producing complex social structures.

Illegal wildlife trade has reached alarming levels globally, extirpating populations of commercially valuable species. As a driver of biodiversity loss, quantifying illegal harvest is essential for conservation and sociopolitical affairs but notoriously difficult. Here we combine field-based carcass monitoring with fine-scale demographic data from an intensively studied wild African elephant population in Samburu, Kenya, to partition mortality into natural and illegal causes. We then expand our analytical framework to model illegal killing rates and population trends of elephants at regional and continental scales using carcass data collected by a Convention on International Trade in Endangered Species program. At the intensively monitored site, illegal killing increased markedly after 2008 and was correlated strongly with the local black market ivory price and increased seizures of ivory destined for China. More broadly, results from application to continental data indicated illegal killing levels were unsustainable for the species between 2010 and 2012, peaking to ∼8% in 2011 which extrapolates to ∼40,000 elephants illegally killed and a probable species reduction of ∼3% that year. Preliminary data from 2013 indicate overharvesting continued. In contrast to the rest of Africa, our analysis corroborates that Central African forest elephants experienced decline throughout the last decade. These results provide the most comprehensive assessment of illegal ivory harvest to date and confirm that current ivory consumption is not sustainable. Further, our approach provides a powerful basis to determine cryptic mortality and gain understanding of the demography of at-risk species.poaching | overharvest | population estimation | extinction | endangered species consumption

Land outside of gazetted protected areas is increasingly seen as important to the future of elephant persistence in Africa. However, other than inferential studies on crop raiding, very little is understood about how elephants Loxodonta africana use and are affected by human-occupied landscapes. This is largely a result of restrictions in technology, which made detailed assessments of elephant movement outside of protected areas challenging. Recent advances in radio telemetry have changed this, enabling researchers to establish over a 24-h period where tagged animals spend their time. We assessed the movement of 13 elephants outside of gazetted protected areas across a range of land-use types on the Laikipia plateau in north-central Kenya. The elephants monitored spent more time at night than during the day in areas under land use that presented a risk of mortality associated with human occupants. The opposite pattern was found on large-scale ranches where elephants were tolerated. Furthermore, speed of movement was found to be higher where elephants were at risk. These results demonstrate that elephants facultatively alter their behaviour to avoid risk in human-dominated landscapes. This helps them to maintain connectivity between habitat refugia in fragmented land-use mosaics, possibly alleviating some of the potential negative impacts of fragmentation. At the same time, however, it allows elephants to penetrate smallholder farmland to raid crops. The greater the amount of smallholder land within an elephant's range, the more it was utilized, with consequent implications for conflict. These findings underscore the importance of (1) land-use planning to maintain refugia; (2) incentives to prevent further habitat fragmentation; (3) the testing and application of conflict mitigation measures where fragmentation has already taken place

Understanding how mammals satisfy their need for space in fragmenting ecosystems is crucial for ecosystem conservation. Using state-of-the-art global positioning system (GPS) technology we tracked 11 focal African elephants (Loxodonta africana) in Kenya at 3-hourly fix intervals and collected between 34 and 406 days per individual. Our recordings gave a high spatio-temporal resolution compared to previous studies and allowed novel insights into range use. The actual ranges of the tracked elephants are smaller than usually represented. Moreover, the ranges in our sample were complex and not confined to officially designated protected areas, except where fenced. All the unfenced elephants in our sample had distinct 'home sectors' linked by 'travel' corridors. Within each home sector the elephants concentrated in favourite 'core zones'. Such core zones tended to lie in protected areas whereas corridors typically crossed unprotected range. Elephants moved significantly faster along corridors than elsewhere in their range, which suggests awareness of danger outside the protected area. We conclude that understanding the complex use of an animal's range is crucial for conservation planning aiming to balance animal interests with those of human beings that co-habit in their range.

A dramatic expansion of road building is underway in the Congo Basin fuelled by private enterprise, international aid, and government aspirations. Among the great wilderness areas on earth, the Congo Basin is outstanding for its high biodiversity, particularly mobile megafauna including forest elephants (Loxodonta africana cyclotis). The abundance of many mammal species in the Basin increases with distance from roads due to hunting pressure, but the impacts of road proliferation on the movements of individuals are unknown. We investigated the ranging behaviour of forest elephants in relation to roads and roadless wilderness by fitting GPS telemetry collars onto a sample of 28 forest elephants living in six priority conservation areas. We show that the size of roadless wilderness is a strong determinant of home range size in this species. Though our study sites included the largest wilderness areas in central African forests, none of 4 home range metrics we calculated, including core area, tended toward an asymptote with increasing wilderness size, suggesting that uninhibited ranging in forest elephants no longer exists. Furthermore we show that roads outside protected areas which are not protected from hunting are a formidable barrier to movement while roads inside protected areas are not. Only 1 elephant from our sample crossed an unprotected road. During crossings her mean speed increased 14-fold compared to normal movements. Forest elephants are increasingly confined and constrained by roads across the Congo Basin which is reducing effective habitat availability and isolating populations, significantly threatening long term conservation efforts. If the current road development trajectory continues, forest wildernesses and the forest elephants they contain will collapse.

We use chronologies of stable isotopes measured from elephant (Loxodonta africana) hair to determine migration patterns and seasonal diet changes in elephants in and near Samburu National Reserve in northern Kenya. Stable carbon isotopes record diet changes, principally enabling differentiation between browse and tropical grasses, which use the C3 and C4 photosynthetic pathways, respectively; stable nitrogen isotopes record regional patterns related to aridity, offering insight into localized ranging behavior. Isotopically identified range shifts were corroborated by global positioning system radio tracking data of the studied individuals. Comparison of the stable isotope record in the hair of one migrant individual with that of a resident population shows important differences in feeding and ranging behavior over time. Our analysis indicates that differences are the result of excursions into mesic environments coupled with intermittent crop raiding by the migrant individual. Variation in diet, quantified by using stable isotopes, can offer insight into diet-related wildlife behavior.13-carbon ͉ 15-nitrogen ͉ chronology ͉ human-elephant conflict T he stable isotope ratios of 13 C͞ 12 C in hair records the diet of mammals (1-4). It is particularly useful in distinguishing diets of C 3 browse versus C 4 grass in tropical regions (5-7) because of the large difference in 13 C͞ 12 C ratios between plants using the C 3 and C 4 photosynthetic pathways, respectively. In tropical regions, the C 3 pathway is used primarily by trees and shrubs, whereas plants using the C 4 pathway are principally grasses (8,9).Hair is a particularly useful indicator of diet change (3, 4) because the isotope turnover of mammal tissues is high enough to resolve short-term diet changes. Recent advances in methodology, progressed through the study of large mammals with controlled diet changes (10, 11), allows detailed reconstruction of the diet history of individual large mammals in wild populations (12,13).In this study, we determine the growth rates and stable 13 C͞ 12 C and 15 N͞ 14 N ratios in elephant hair collected between 2001 and 2004. We focus on the behavior of a resident population of Samburu National Reserve, Northern Kenya, for the time period of 2000 to 2002. We compare stable isotope results of this resident population with a migrant elephant (B1013) that visited Samburu Reserve up to several times each year. Differences in isotope ratios between the resident individuals and the migrant indicate different behaviors, including rapid migration across long distances by the migrant individual and differences in the fraction C 4 biomass in the diet. The latter may be related to seasonal crop raiding, which can be quantified by using stable isotope ratios. Materials and MethodsGlobal Positioning System (GPS) radio collars were fitted to elephants in Samburu National Reserve, Northern Kenya, between 2001 and 2004 (14). Collars were programmed to record positions at hourly intervals, offering detailed records of movement. Tail hairs fro...

The internal state of an individual-as it relates to thirst, hunger, fear, or reproductive drive-can be inferred by referencing points on its movement path to external environmental and sociological variables. Using time-series approaches to characterize autocorrelative properties of step-length movements collated every 3 h for seven free-ranging African elephants, we examined the influence of social rank, predation risk, and seasonal variation in resource abundance on periodic properties of movement. The frequency domain methods of Fourier and wavelet analyses provide compact summaries of temporal autocorrelation and show both strong diurnal and seasonal based periodicities in the step-length time series. This autocorrelation is weaker during the wet season, indicating random movements are more common when ecological conditions are good. Periodograms of socially dominant individuals are consistent across seasons, whereas subordinate individuals show distinct differences diverging from that of dominants during the dry season. We link temporally localized statistical properties of movement to landscape features and find that diurnal movement correlation is more common within protected wildlife areas, and multiday movement correlations found among lower ranked individuals are typically outside of protected areas where predation risks are greatest. A frequency-related spatial analysis of movement-step lengths reveal that rest cycles related to the spatial distribution of critical resources (i.e., forage and water) are responsible for creating the observed patterns. Our approach generates unique information regarding the spatial-temporal interplay between environmental and individual characteristics, providing an original approach for understanding the movement ecology of individual animals and the spatial organization of animal populations.behavior ͉ foraging ͉ GPS data ͉ predation ͉ movement ecology C haracteristics of an animal's movement path offer insights into the external influences and internal states of an individual across time and space (1, 2). Scaled across a population or ecosystem, such information can offer new understanding of the salient factors driving the spatiotemporal structure of populations (3, 4). The methods used to garner this information have been the focus of research in the field of ecology for some time (5-7). Perhaps the most common of these methods involves analyzing correlations among consecutive displacements (i.e., step length or approximate velocity) and turning angles in the context of random walk models (4, 7). The strength of first-order autocorrelations in the directionality of movement has been used to categorize foraging strategy (8-10), whereas potential information contained in the autocorrelative structure of step lengths has largely been ignored. Given the existence of such structure in our data, as revealed by the analyses presented here, simple random walk models are clearly inadequate for characterizing movement in some animals. Our approach provides unique insight in...