Landscape-scale habitat response of African elephants shows strong selection for foraging opportunities in a human dominated ecosystem

Balancing trade-offs between foraging and risk factors is a fundamental behavior that structures the spatial distribution of species. For African elephants Loxodonta africana, human pressures from poaching and conflict are primary drivers of species decline, but little is known about how elephants structure their spatial behavior in the face of human occupancy and predation. We seek to understand how elephants balance trade-offs between resource access, human presence and human predatory risk factors (poaching and conflict killing) in an unfenced, dynamic ecosystem where elephants persist primarily outside protected areas in community rangelands. We used tracking data from 101 elephants collected between 2001 and 2016. We investigated elephant behavior in response to landcover, topography, productivity, water, human features and human predation risk using third-order resource selection functions. We extended this analysis by employing a mixed-effects multinomial regression to identify temporal shifts in habitat use, and evaluated temporal shifts in movement patterns by estimating mean squared displacement across different productivity periods. Across periods, elephants displayed strong selection for productive areas and areas near water. Temporal shifts in habitat use showed that, during the dry period, elephants were clustered around permanent water sources where humans also congregated. At the onset of the wet period, a shift occurred where elephants moved away from permanent water and from permanent settlements towards seasonal water sources and seasonal settlements. Our findings indicate that foraging and water access are important limiting factors affecting elephants that potentially restrain their spatial responses to humans at the scale of our analysis. Given that pastoralists and elephants rely on the same resources, increasing human and livestock populations enhance pressure on shared resources and space in Africa's drylands. The long-term conservation of elephants will require approaches that reduce poaching as well as landscape level planning to prevent negative impacts from increasing competition for preferred resources.     

A population survey of forest elephants (Loxodonta africana cyclotis) in northern Congo

From February to April of 1989 transects were run in four forested areas of northern Congo (Brazzaville) to census elephant populations using dung counts. A total of 401.0 km of line transects was completed during the survey. An overall extrapolated density of 0.7 elephants/km² was obtained with a variation from 0.3 to 0.9 elephants/km². This study shows for the first time that elephants occur in high densities in a number of sites in northern Congo. When elephant dung density is plotted against distance from the nearest village for each transect a significant positive linear correlation results. Based on this correlation we hypothesize that the population of elephants in much of northern Congo is high. Local reports and our own observations indicate that elephant poaching is heavy in northern Congo and that the CITES ban on trade in African elephant ivory has had a limited effect on the level of poaching.     

Effective population size dynamics reveal impacts of historic climatic events and recent anthropogenic pressure in African elephants

Two hundred years of elephant hunting for ivory, peaking in 1970–1980s, caused local extirpations and massive population declines across Africa. The resulting genetic impacts on surviving populations have not been studied, despite the importance of understanding the evolutionary repercussions of such human-mediated events on this keystone species. Using Bayesian coalescent-based genetic methods to evaluate time-specific changes in effective population size, we analysed genetic variation in 20 highly polymorphic microsatellite loci from 400 elephants inhabiting the greater Samburu-Laikipia region of northern Kenya. This area experienced a decline of between 80% and 90% in the last few decades when ivory harvesting was rampant. The most significant change in effective population size, however, occurred approximately 2500 years ago during a mid–Holocene period of climatic drying in tropical Africa. Contrary to expectations, detailed analyses of four contemporary age-based cohorts showed that the peak poaching epidemic in the 1970s caused detectable temporary genetic impacts, with genetic diversity rebounding as juveniles surviving the poaching era became reproductively mature. This study demonstrates the importance of climatic history in shaping the distribution and genetic history of a keystone species and highlights the utility of coalescent-based demographic approaches in unravelling ancestral demographic events despite a lack of ancient samples. Unique insights into the genetic signature of mid-Holocene climatic change in Africa and effects of recent poaching pressure on elephants are discussed.     

Fine-scale population genetic structure in a fission-fusion society

Nonrandom patterns of mating and dispersal create fine-scale genetic structure in natural populations — especially of social mammals — with important evolutionary and conservation genetic consequences. Such structure is well-characterized for typical mammalian societies; that is, societies where social group composition is stable, dispersal is male-biased, and males form permanent breeding associations in just one or a few social groups over the course of their lives. However, genetic structure is not well understood for social mammals that differ from this pattern, including elephants. In elephant societies, social groups fission and fuse, and males never form permanent breeding associations with female groups. Here, we combine 33 years of behavioural observations with genetic information for 545 African elephants ( Loxodonta africana ), to investigate how mating and dispersal behaviours structure genetic variation between social groups and across age classes. We found that, like most social mammals, female matrilocality in elephants creates co-ancestry within core social groups and significant genetic differentiation between groups (Φ_ST = 0.058). However, unlike typical social mammals, male elephants do not bias reproduction towards a limited subset of social groups, and instead breed randomly across the population. As a result, reproductively dominant males mediate gene flow between core groups, which creates cohorts of similar-aged paternal relatives across the population. Because poaching tends to eliminate the oldest elephants from populations, illegal hunting and poaching are likely to erode fine-scale genetic structure. We discuss our results and their evolutionary and conservation genetic implications in the context of other social mammals.     

Movement patterns of forest elephants (Loxodonta cyclotis Matschie, 1900) in the Odzala-Kokoua National Park,Republic of Congo

African forest elephants (Loxodonta cyclotis Matschie, 1900) are ecological engineers that play a fundamental role in vegetation dynamics. The species is of immediate conservation concern, yet it is relatively understudied. To narrow this knowledge gap, we studied the drivers of daily movement patterns (linear displacements) of forest elephants—characterised by a set of geographical, meteorological and anthropogenic variables—in the Odzala-Kokoua National Park, Republic of Congo. Explicitly, we used conditional random forest to model and disentangle the main environmental factors governing the displacements of six forest elephants, fitted with GPS collars and tracked over 16 months. Results indicated that females moved further distances than males, while the presence of roads or human settlements disrupted elephant behaviour resulting in faster displacements. Forest elephants moved faster along watercourses and through forest with understory dominated by Marantaceae forests and bais, but moved slower in savannahs. Finally, flood-prone areas—described by elevation and accumulated precipitation—and higher temperatures prevented longer displacements. We expect these results to improve the knowledge on the species movements through different habitats, which would benefit its conservation management.     

Genetic relatedness and disrupted social structure in a poached population of African elephants

We use genetic measures of relatedness and observations of female bonding to examine the demographic signature of historically heavy poaching of a population of free-ranging African elephants. We collected dung samples to obtain DNA and observed behaviour from 102 elephant families over a 25-month period in 2003–2005 in Mikumi National Park, Tanzania. Poaching reduced the population by 75% in the decade prior to the 1989 ivory trade ban; park records indicate that poaching dropped significantly in Mikumi following the ban. Using 10 microsatellite loci, DNA was genotyped in 203 elephants and pair-wise relatedness was calculated among adult females within and between groups. The Mikumi population is characterized by small group size, considerable variation in group relatedness, females with no first-order adult relatives and females that form only weak social bonds. We used gene-drop analysis and a model of a genetically intact pedigree to compare our observed Mikumi group relatedness to a simulated genetically intact unpoached expectation. The majority of groups in Mikumi contain 2 to 3 adults; of these, 45% were classified as genetically disrupted. Bonding, quantified with a pair-wise association index, was significantly correlated with relatedness; however only half of the females formed strong bonds with other females, and relatedness was substantially lower for a given bond strength as compared to an unpoached population. Female African elephants without kin demonstrated considerable behavioural plasticity in this disturbed environment, grouping with other females lacking kin, with established groups, or remaining alone, unable to form any stable adult female-bonds. We interpret these findings as the remaining effect of poaching disturbance in Mikumi, despite a drop in the level of poaching since the commercial trade in ivory was banned 15 years ago.     

Animal movement in dynamic landscapes: interaction between behavioural strategies and resource distributions

Most ecological and evolutionary processes are thought to critically depend on dispersal and individual movement but there is little empirical information on the movement strategies used by animals to find resources. In particular, it is unclear whether behavioural variation exists at all scales, or whether behavioural decisions are primarily made at small spatial scales and thus broad-scale patterns of movement simply reflect underlying resource distributions. We evaluated animal movement responses to variable resource distributions using the grey teal (Anas gracilis) in agricultural and desert landscapes in Australia as a model system. Birds in the two landscapes differed in the fractal dimension of their movement paths, with teal in the desert landscape moving less tortuously overall than their counterparts in the agricultural landscape. However, the most striking result was the high levels of individual variability in movement strategies, with different animals exhibiting different responses to the same resources. Teal in the agricultural basin moved with both high and low tortuosity, while teal in the desert basin primarily moved using low levels of tortuosity. These results call into question the idea that broad-scale movement patterns simply reflect underlying resource distributions, and suggest that movement responses in some animals may be behaviourally complex regardless of the spatial scale over which movement occurs.     

Effect of Colony, Patch Distance, And Trajectory Sense on Movement Complexity in Foraging Ants

Animals display foraging trajectories when exploiting food patches. These displacements have been studied, particularly in organisms that forage from a central place. The complexity of a movement path may be analyzed by means of the fractal dimension, an index that estimates the tortuosity of a trajectory. In field experiments we studied the effect of trajectory sense, distance to the resource patch and colony on the movement complexity in a common ant of central Chile. We found that these factors and their interactions significantly affected the complexity of movement paths. We discuss whether mortality risk determines more linear trajectories when the resource patch is distant from the nest, and whether the information acquired from the environment by workers determines less complex return trips.     

Forest elephant paths and movements at the Odzala National Park, Congo: the role of clearings and Marantaceae forests

The role of forest clearings and Marantaceae forest on elephant movement at the Odzala National Park was studied by describing and mapping elephant paths. Three types of paths were distinguished based on their direction, length, width, elephant activity and the types of forests they cross. They were called boulevards, foraging paths, and clearing alleys. All paths explored were heavily used. Boulevards are used for travelling over long distances (up to 34 km) and for linking up rapidly favourite sites such as clearings. Foraging paths mainly run through the medium-density Marantaceae forests which provide elephants with both herbaceous foods and tree fruit. Clearing alleys form a dense network of tracks around clearings. This network could be a result of an anti-predatory monitoring behaviour of elephants before entering the clearings where they have long experienced heavy poaching. The heavy trampling which results from the elephant activity could contribute to the maintenance of clearings by preventing germination and seedling survival of the numerous seeds dispersed in their dung.     

Estimating elephant density using motion-sensitive cameras: challenges,opportunities, and parameters for consideration

With extinction rates far exceeding the natural background rate, reliable monitoring of wildlife populations has become crucial for adaptive management and conservation. Robust monitoring is often labor intensive with high economic costs, particularly in the case of those species that are subject to illegal poaching, such as elephants, which require frequent and accurate population estimates over large spatial scales. Dung counting methods are commonly employed to estimate the density of elephants; however, in the absence of a full survey calibration, these can be unreliable in heterogeneous habitats where dung decay rates may be highly variable. We explored whether motion-sensitive cameras offer a simple, lower cost, and reliable alternative for monitoring in challenging forest environments. We estimated the density of African savanna elephants (Loxodanta africana) in a montane forest using the random encounter model and assessed the importance of surveying parameters for future survey design. We deployed motion-sensitive cameras in 65 locations in the Aberdare Conservation Area in Kenya during June to August in 2015 to 2017, for a survey effort of 967 days, and a mean encounter rate of 0.09 ± 0.29 (SD) images/day. Elephants were captured in 16 locations. Density estimates varied between vegetation types, with estimates ranging from 6.27/km² in shrub, 1.1/km² in forest, 0.53/km² in bamboo (Yushania alpine), and 0.44/km² in the moorlands. The average speed of animal movement and the camera detection zone had the strongest linear associations with density estimates (R = −0.97). The random encounter model has the potential to offer an alternative, or complementary method within the active management framework for monitoring elephant populations in forests at a relatively low cost.     

Modelling large herbivore movement decisions: Beyond food availability as a predictor of ranging patterns

The ability of animals to adapt to their changing environment will depend in part on shifts in their ranging patterns, but when and why individuals choose to move requires detailed understanding of their decision‐making processes. We develop a simple decision‐making model accounting for resource availability in habitually used ranges. We suggest that disparities between model predictions and animal tracking data indicate additional factors influencing movement decisions, which may be identified given detailed system‐specific knowledge. The model was evaluated using movement data from satellite‐tracked elephants (Loxodonta africana) inhabiting the Amboseli basin in Kenya, moving from savannah areas with low quality but constant resource availability, to areas with temporally constrained higher nutrient availability. Overall, the model fits the data well: There was a good correlation between predicted and observed locations for the combined data from all elephants, but variation between individuals in how well the model fits. For those elephants where model predictions were less successful, additional factors likely to affect movement decisions, including reproduction, anthropogenic threats, memory and perception are suggested. This protocol for building and testing decision‐making models should contribute to success in attempts to preserve sufficient space for large herbivores in their increasingly human‐dominated ecosystems.     

Demography of a population with a long history of poaching and the utility of the individual identification technique as a tool for monitoring intermittently studied elephant populations

In many elephant populations in Africa, adverse effects of poaching including altered age and sex structure and long calving intervals continue to negatively impact the rate of recovery. The study reported here characterized the demographic status of the elephant population in Meru National Park, Kenya, so as to determine the size, age and sex structure, and calving interval of the population and to compare its demographic parameters against those from the relatively stable elephant population at Amboseli. Additionally, a demographic study conducted 7 years earlier enabled us to determine the demographic performance of the population and to explicitly test the utility of the individual identification technique in monitoring intermittently studied populations. We found 392 elephants in the park. The proportion of older individuals was lower, and the calving interval was longer than estimates from Amboseli. We recognized 16% of elephants indexed in the previous study. Our results suggest that the individual identification technique may be useful in monitoring demographic changes including poaching in intermittently studied elephant populations. Overall, our results suggest that although the Meru elephant population showed signs of recovery, poaching continues to imperil its recovery.     

A stakeholder perspective into wildlife policy in India

We investigated perceptions of wildlife policy and issues through questionnaires (n = 148) administered to policy makers, conservation scientists, individuals representing non-government organizations (NGOs), and field officials, who implemented government policies and enforced laws. We found significant differences among attitudes of stakeholders identifying major threats to wildlife, the use of science, the role of poaching in conservation, and the composition of species illegally traded. Policy makers and field officials differed in their views with NGOs and scientists on community response to wildlife policies and the varying threat perceptions to different species due to poaching and illegal trade. We noted ambiguity among stakeholders about sustainable use principles in India. Policies must be more effective in conservation and the process of making policy must be broad-based and participatory if wildlife conservation is to advance on the subcontinent. © 2011 The Wildlife Society.     

Hotspot elephant‐poaching areas in the Eastern Selous Game Reserve,Tanzania

Elephant poaching is a significant problem in many parts of Africa, including Tanzania. This study identifies areas within the Eastern Selous Game Reserve (ESGR) that are at a higher risk of elephant poaching for the illegal ivory trade. We obtained data on the distribution of poached elephants and the seasons of poaching from 2008 to 2013 from the elephant mortality database of the Selous Game Reserve in the eastern and north‐eastern sectors. The incidences of poaching were higher during the wet season. The hot spots of elephant poaching were observed on the edges of the ESGR. This finding was attributed to the involvement of local people adjacent to the ESGR in poaching activities due to a lack of economic opportunities. This study will serve as a tool to guide and inform reserve managers involved in wildlife conservation in Tanzania. Recommended solutions to address the problem of elephant poaching are improved economic opportunities for local people, enhanced conservation education and research and improved governance and law enforcement.     

How do African elephants utilize the landscape during wet season? A habitat connectivity analysis for Sioma Ngwezi landscape in Zambia

1. The influence of environmental factors on the distribution and persistence of African elephants (Loxodonta africana) is pertinent to policy makers and managers to formulate balanced plans for different land‐use types.
2. The study focuses on movement of elephants and how they utilize foraging areas in Sioma Ngwezi landscape in Zambia by answering the following questions: (1) Which environmental variables and land‐cover class predict the movement of elephants during the wet season in Sioma Ngwezi landscape? (2) What is the wet season suitable habitat for elephants in Sioma Ngwezi landscape? (3) What are the major wet season movement corridors for elephants in Sioma Ngwezi landscape?
3. We used GPS telemetry data from the collared elephants to assess habitat connectivity. Maximum entropy (MaxEnt) and linkage mapper were the tools used to predict habitat suitability, movement corridors, and barriers in the landscape during the wet season.
4. The study identified elevation, land cover, and NDVI as the most important environmental predictors that modify the dispersal of elephants in the landscape during the wet season. Additionally, a total of 36 potential wet season corridors were identified connecting 15 core areas mainly used for foraging and protection from poachers in the landscape. Of these, 24 corridors were highly utilized and are suggested as priority corridors for elephant movement in the landscape.
5. The identified wet season habitats and functional corridors may help to combat elephant poaching by patrolling areas with high relative probability of elephant presence. The findings may also help abate human–elephant conflict such as crop‐raiding by managing identified corridors that run into agriculture zones in the game management area.

     

Human–elephant coexistence challenges in Myanmar: An analysis of fatal elephant attacks on humans and elephant mortality

Understanding the underlying causes behind human–elephant conflict (HEC)-driven mortality of humans and elephants will help improve both parties’ wellbeing. The objective of this study was to examine the temporal and spatial mortality patterns of humans and elephants and the influence of local attitudes, conflict factors and habitat factors on elephant poaching. We used the Myanmar Forest Department data from 2001 to 2020 for humans and 2011 to 2020 for elephants together with explanatory data on human attitudes, habitat, and conflict factors. Approximately seven persons were killed annually in elephant attacks, with a bias towards men. The annual mortality of elephants during the study period was on average 16 individuals, and most elephants were killed by humans. There was a significant relationship between the number of killed humans and human-killed elephants around HEC villages. Villages with more property damage exhibited a higher rate of human mortality, which also correlated with negative feelings of local people towards elephants. Elephant poaching was higher in villages with less suitable habitat available for elephant use. Human encroachment is an important cause of HEC, leading to human loss and forming the main threat to the survival of wild elephants. We suggest local involvement to ensure good governance in conflict resolution and mitigation strategies and to strengthen law enforcement.     

Using Poaching Levels and Elephant Distribution to Assess the Conservation Efficacy of Private,Communal and Government Land in Northern Kenya

Efforts to curb elephant poaching have focused on reducing demand, confiscating ivory and boosting security patrols in elephant range. Where land is under multiple uses and ownership, determining the local poaching dynamics is important for identifying successful conservation models. Using 2,403 verified elephant, Loxodonta africana, mortality records collected from 2002 to 2012 and the results of aerial total counts of elephants conducted in 2002, 2008 and 2012 for the Laikipia-Samburu ecosystem of northern Kenya, we sought to determine the influence of land ownership and use on diurnal elephant distribution and on poaching levels. We show that the annual proportions of illegally killed (i.e., poached) elephants increased over the 11 years of the study, peaking at 70% of all recorded deaths in 2012. The type of land use was more strongly related to levels of poaching than was the type of ownership. Private ranches, comprising only 13% of land area, hosted almost half of the elephant population and had significantly lower levels of poaching than other land use types except for the officially designated national reserves (covering only 1.6% of elephant range in the ecosystem). Communal grazing lands hosted significantly fewer elephants than expected, but community areas set aside for wildlife demonstrated significantly higher numbers of elephants and lower illegal killing levels relative to non-designated community lands. While private lands had lower illegal killing levels than community conservancies, the success of the latter relative to other community-held lands shows the importance of this model of land use for conservation. This work highlights the relationship between illegal killing and various land ownership and use models, which can help focus anti-poaching activities.     

What if it gets crowded? Density‐dependent tortuosity in individual movements of a Neotropical mammal

Effects of density dependence on animal movements have received much attention in ecology, but it is still debated to what extent dispersal and movements in general are density dependent, and their potential contribution to population regulation processes. Here, we determine the occurrence and nature of density dependence in the movements of a Neotropical marsupial, the black‐eared opossum Didelphis aurita Wied‐Neuwied 1826. Using spool‐and‐line tracking devices, we estimated the tortuosity of fine‐scale movements of 149 individuals by their fractal dimension D. We evaluated the relative importance of population size, reproductive or climatic seasons and reproductive maturity of individuals as determinants of movement tortuosity, using a model selection approach. Population size was the most important determinant of movement tortuosity, with season (climatic seasons for females, reproductive seasons for males) and reproductive maturity as secondary but also important variables. We detected a positive density‐dependent effect on movement tortuosity, resulting in more intensive use of areas by individuals during periods of high population size. This positive association between movement tortuosity and population size is more likely to result from intraspecific competition, which forces individuals to explore their environment more intensively during high‐density periods. Therefore, despite being density dependent, movements in D. aurita apparently do not contribute to population regulation mechanisms.     

Leveraging multidimensional heterogeneity in resource selection to define movement tactics of animals

Increasing interest in the complexity, variation and drivers of movement‐related behaviours promise new insight into fundamental components of ecology. Resolving the multidimensionality of spatially explicit behaviour remains a challenge for investigating tactics and their relation to niche construction, but high‐resolution movement data are providing unprecedented understanding of the diversity of spatially explicit behaviours. We introduce a framework for investigating individual variation in movement‐defined resource selection that integrates the behavioural and ecological niche concepts. We apply it to long‐term tracking data of 115 African elephants (Loxodonta africana), illustrating how a behavioural hypervolume can be defined based on differences between individuals and their ecological settings, and applied to explore population heterogeneity. While normative movement behaviour is frequently used to characterise population behaviour, we demonstrate the value of leveraging heterogeneity in the behaviour to gain greater insight into population structure and the mechanisms driving space‐use tactics.     

Wildlife crime promoted by weak governance

Wildlife crime such as illegal hunting of elephants and rhinos is currently one of the major issues in conservation. In order to combat poaching, law enforcement is essential. Here, we reviewed wildlife cases from the Northern Zone Anti‐Poaching Unit, Arusha, Tanzania (APU‐Arusha), focusing on the types of wildlife offence committed, prosecution outcomes and the sentences awarded to the offenders who were found guilty by the courts. Eighty‐two registered cases were thoroughly reviewed. The majority of wildlife offences committed were unlawful possession of, and failure to report, government trophies. It was found that only 16% of the accused were found guilty, 6% were acquitted, 30% of the charges were withdrawn, and 48% of the cases were discharged by the director of public prosecution (DPP) or magistrate. The poor conviction rate was attributed to weak evidence, failure of the prosecution to prove cases beyond reasonable doubt, lack of cooperation between the zonal game office and the DPP, and in some cases, the accused jumped their bails. Understanding the factors which allow individuals to continue to engage in poaching and the factors that promote the practice of good governance is vital in trying to stop wildlife crimes.