Graeme Shannon

Leadership in elephants: the adaptive value of age

The value of age is well recognized in human societies, where older individuals often emerge as leaders in tasks requiring specialized knowledge, but what part do such individuals play in other social species? Despite growing interest in how effective leadership might be achieved in animal social systems, the specific role that older leaders may play in decision-making has rarely been experimentally investigated. Here, we use a novel playback paradigm to demonstrate that in African elephants (Loxodonta africana), age affects the ability of matriarchs to make ecologically relevant decisions in a domain critical to survival—the assessment of predatory threat. While groups consistently adjust their defensive behaviour to the greater threat of three roaring lions versus one, families with younger matriarchs typically under-react to roars from male lions despite the severe danger they represent. Sensitivity to this key threat increases with matriarch age and is greatest for the oldest matriarchs, who are likely to have accumulated the most experience. Our study provides the first empirical evidence that individuals within a social group may derive significant benefits from the influence of an older leader because of their enhanced ability to make crucial decisions about predatory threat, generating important insights into selection for longevity in cognitively advanced social mammals.

A synthesis of two decades of research documenting the effects of noise on wildlife

Global increases in environmental noise levels – arising from expansion of human populations, transportation networks, and resource extraction – have catalysed a recent surge of research into the effects of noise on wildlife. Synthesising a coherent understanding of the biological consequences of noise from this literature is challenging. Taxonomic groups vary in auditory capabilities. A wide range of noise sources and exposure levels occur, and many kinds of biological responses have been observed, ranging from individual behaviours to changes in ecological communities. Also, noise is one of several environmental effects generated by human activities, so researchers must contend with potentially confounding explanations for biological responses. Nonetheless, it is clear that noise presents diverse threats to species and ecosystems and salient patterns are emerging to help inform future natural resource‐management decisions. We conducted a systematic and standardised review of the scientific literature published from 1990 to 2013 on the effects of anthropogenic noise on wildlife, including both terrestrial and aquatic studies. Research to date has concentrated predominantly on European and North American species that rely on vocal communication, with approximately two‐thirds of the data set focussing on songbirds and marine mammals. The majority of studies documented effects from noise, including altered vocal behaviour to mitigate masking, reduced abundance in noisy habitats, changes in vigilance and foraging behaviour, and impacts on individual fitness and the structure of ecological communities. This literature survey shows that terrestrial wildlife responses begin at noise levels of approximately 40 dBA, and 20% of papers documented impacts below 50 dBA. Our analysis highlights the utility of existing scientific information concerning the effects of anthropogenic noise on wildlife for predicting potential outcomes of noise exposure and implementing meaningful mitigation measures. Future research directions that would support more comprehensive predictions regarding the magnitude and severity of noise impacts include: broadening taxonomic and geographical scope, exploring interacting stressors, conducting larger‐scale studies, testing mitigation approaches, standardising reporting of acoustic metrics, and assessing the biological response to noise‐source removal or mitigation. The broad volume of existing information concerning the effects of anthropogenic noise on wildlife offers a valuable resource to assist scientists, industry, and natural‐resource managers in predicting potential outcomes of noise exposure.

African elephant home range and habitat selection in Pongola Game Reserve, South Africa

ABSTRACT The ranging behaviour and habitat occupancy by three elephant groups (cow herd, bulls, and an orphan group) were studied over a two-year period in a small, fenced reserve. No summer dispersal was observed. Distinct seasonal home ranges were exhibited for all groups, with the summer (wet season) ranges being smaller than the winter (dry season) ranges. Home range size was much smaller than in other locations. The dam and surrounding high density of patches of vegetation of high nutritional quality are thought to be the reasons. Habitat selection was strongly evident with all of the elephant groups selecting River Line habitats in the dry season. In the wet season the cow herd and orphans selected the more open Acacia habitats and the bulls exhibited no significant habitat preference.

A framework to assess evolutionary responses to anthropogenic light and sound

Human activities have caused a near-ubiquitous and evolutionarily-unprecedented increase in environmental sound levels and artificial night lighting. These stimuli reorganize communities by interfering with species-specific perception of time-cues, habitat features, and auditory and visual signals. Rapid evolutionary changes could occur in response to light and noise, given their magnitude, geographical extent, and degree to which they represent unprecedented environmental conditions. We present a framework for investigating anthropogenic light and noise as agents of selection, and as drivers of other evolutionary processes, to influence a range of behavioral and physiological traits such as phenological characters and sensory and signaling systems. In this context, opportunities abound for understanding contemporary and rapid evolution in response to human-caused environmental change.

The role of foraging behaviour in the sexual segregation of the African elephant

Elephants (Loxodonta africana) exhibit pronounced sexual dimorphism, and in this study we test the prediction that the differences in body size and sociality are significant enough to drive divergent foraging strategies and ultimately sexual segregation. Body size influences the foraging behaviour of herbivores through the differential scaling coefficients of metabolism and gut size, with larger bodied individuals being able to tolerate greater quantities of low-quality, fibrous vegetation, whilst having lower mass-specific energy requirements. We test two distinct theories: the scramble competition hypothesis (SCH) and the forage selection hypothesis (FSH). Comprehensive behavioural data were collected from the Pongola Game Reserve and the Phinda Private Game Reserve in South Africa over a 2.5-year period. The data were analysed using sex as the independent variable. Adult females targeted a wider range of species, adopted a more selective foraging approach and exhibited greater bite rates as predicted by the body size hypothesis and the increased demands of reproductive investment (lactation and pregnancy). Males had longer feeding bouts, displayed significantly more destructive behaviour (31% of observations, 11% for females) and ingested greater quantities of forage during each feeding bout. The independent ranging behaviour of adult males enables them to have longer foraging bouts as they experience fewer social constraints than females. The SCH was rejected as a cause of sexual segregation due to the relative abundance of low quality forage, and the fact that feeding heights were similar for both males and females. However, we conclude that the differences in the foraging strategies of the sexes are sufficient to cause spatial segregation as postulated by the FSH. Sexual dimorphism and the associated behavioural differences have important implications for the management and conservation of elephant and other dimorphic species, with the sexes effectively acting as distinct “ecological species”.

Elephants can determine ethnicity, gender, and age from acoustic cues in human voices

Significance Recognizing predators and judging the level of threat that they pose is a crucial skill for many wild animals. Human predators present a particularly interesting challenge, as different groups of humans can represent dramatically different levels of danger to animals living around them. We used playbacks of human voice stimuli to show that elephants can make subtle distinctions between language and voice characteristics to correctly identify the most threatening individuals on the basis of their ethnicity, gender, and age. Our study provides the first detailed assessment of human voice discrimination in a wild population of large-brained, long-lived mammals, and highlights the potential benefits of sophisticated mechanisms for distinguishing different subcategories within a single predator species. Animals can accrue direct fitness benefits by accurately classifying predatory threat according to the species of predator and the magnitude of risk associated with an encounter. Human predators present a particularly interesting cognitive challenge, as it is typically the case that different human subgroups pose radically different levels of danger to animals living around them. Although a number of prey species have proved able to discriminate between certain human categories on the basis of visual and olfactory cues, vocalizations potentially provide a much richer source of information. We now use controlled playback experiments to investigate whether family groups of free-ranging African elephants (Loxodonta africana) in Amboseli National Park, Kenya can use acoustic characteristics of speech to make functionally relevant distinctions between human subcategories differing not only in ethnicity but also in sex and age. Our results demonstrate that elephants can reliably discriminate between two different ethnic groups that differ in the level of threat they represent, significantly increasing their probability of defensive bunching and investigative smelling following playbacks of Maasai voices. Moreover, these responses were specific to the sex and age of Maasai presented, with the voices of Maasai women and boys, subcategories that would generally pose little threat, significantly less likely to produce these behavioral responses. Considering the long history and often pervasive predatory threat associated with humans across the globe, it is likely that abilities to precisely identify dangerous subcategories of humans on the basis of subtle voice characteristics could have been selected for in other cognitively advanced animal species.

Activity Budgets and Sexual Segregation in African Elephants (Loxodonta africana)

Abstract The activity budget hypothesis is 1 of 4 main hypotheses proposed to explain sexual segregation by large herbivores. Because of their smaller body size, females are predicted to have higher mass-specific energy requirements and lower digestive efficiency than males. As a result, females are expected to forage longer to satisfy their nutritional demands. Maintaining the cohesion of a mixed-sex group with differing activity budgets and asynchronous behavioral patterns is increasingly difficult, ultimately leading to spatial segregation of males and females. We tested this hypothesis using data (2002–2005) from 3 distinct populations of African elephants (Loxodonta africana), a species that exhibits marked sexual segregation. Group and individual behaviors were assessed at discrete points in time throughout the day, with a minimum of 10 min between consecutive records. Focal samples of individual male and female elephants also were recorded, with behavioral data logged every minute for 15 min. Data were grouped into 5 behavioral categories: drinking, resting, walking, feeding, and other. Neither activity rhythms nor feeding time varied significantly between the sexes and behavioral patterns were very similar. We propose that social and environmental factors influence behavioral rhythms to a greater extent than does body size, whereas increasing feeding time is only 1 method by which elephants can improve nutritional return. This is especially pertinent when considering their generalist foraging approach, substantial energy demands, and hindgut fermentation. We conclude that the activity budget hypothesis is unlikely to be the causal mechanism in the sexual segregation of African elephants, a finding that concurs with recent experimental and field research on a range of sexually dimorphic herbivores.

SHORT-DURATION DAYTIME MOVEMENTS OF A COW HERD OF AFRICAN ELEPHANTS

Abstract We examined daytime movements of a herd of African elephants (Loxodonta africana) at 10-min, 15-min, and 20-min intervals in Pongola Game Reserve, South Africa. This group tended to proceed in a consistent direction during consecutive movements, especially during long moves. Serial movement lengths and serial movement angles were autocorrelated at 10-min and 15-min intervals but not at 20-min intervals, indicating that 20-min intervals may be a suitable temporal scale to avoid oversampling. Herd movements followed a Lévy-modulated correlated random walk. In addition, looping movements were detected. Spatial scale of the loops averaged about 1 km. Movement strategies that include both Lévy walks and correlated random walks are thought to optimize foraging.

Effects of social disruption in elephants persist decades after culling

BackgroundMulti-level fission-fusion societies, characteristic of a number of large brained mammal species including some primates, cetaceans and elephants, are among the most complex and cognitively demanding animal social systems. Many free-ranging populations of these highly social mammals already face severe human disturbance, which is set to accelerate with projected anthropogenic environmental change. Despite this, our understanding of how such disruption affects core aspects of social functioning is still very limited.ResultsWe now use novel playback experiments to assess decision-making abilities integral to operating successfully within complex societies, and provide the first systematic evidence that fundamental social skills may be significantly impaired by anthropogenic disruption. African elephants (Loxodonta africana) that had experienced separation from family members and translocation during culling operations decades previously performed poorly on systematic tests of their social knowledge, failing to distinguish between callers on the basis of social familiarity. Moreover, elephants from the disrupted population showed no evidence of discriminating between callers when age-related cues simulated individuals on an increasing scale of social dominance, in sharp contrast to the undisturbed population where this core social ability was well developed.ConclusionsKey decision-making abilities that are fundamental to living in complex societies could be significantly altered in the long-term through exposure to severely disruptive events (e.g. culling and translocation). There is an assumption that wildlife responds to increasing pressure from human societies only in terms of demography, however our study demonstrates that the effects may be considerably more pervasive. These findings highlight the potential long-term negative consequences of acute social disruption in cognitively advanced species that live in close-knit kin-based societies, and alter our perspective on the health and functioning of populations that have been subjected to anthropogenic disturbance.

Ecological Thresholds in the Savanna Landscape: Developing a Protocol for Monitoring the Change in Composition and Utilisation of Large Trees

Background Acquiring greater understanding of the factors causing changes in vegetation structure - particularly with the potential to cause regime shifts - is important in adaptively managed conservation areas. Large trees (≥5 m in height) play an important ecosystem function, and are associated with a stable ecological state in the African savanna. There is concern that large tree densities are declining in a number of protected areas, including the Kruger National Park, South Africa. In this paper the results of a field study designed to monitor change in a savanna system are presented and discussed. Methodology/Principal Findings Developing the first phase of a monitoring protocol to measure the change in tree species composition, density and size distribution, whilst also identifying factors driving change. A central issue is the discrete spatial distribution of large trees in the landscape, making point sampling approaches relatively ineffective. Accordingly, fourteen 10 m wide transects were aligned perpendicular to large rivers (3.0–6.6 km in length) and eight transects were located at fixed-point photographic locations (1.0–1.6 km in length). Using accumulation curves, we established that the majority of tree species were sampled within 3 km. Furthermore, the key ecological drivers (e.g. fire, herbivory, drought and disease) which influence large tree use and impact were also recorded within 3 km. Conclusions/Significance The technique presented provides an effective method for monitoring changes in large tree abundance, size distribution and use by the main ecological drivers across the savanna landscape. However, the monitoring of rare tree species would require individual marking approaches due to their low densities and specific habitat requirements. Repeat sampling intervals would vary depending on the factor of concern and proposed management mitigation. Once a monitoring protocol has been identified and evaluated, the next stage is to integrate that protocol into a decision-making system, which highlights potential leading indicators of change. Frequent monitoring would be required to establish the rate and direction of change. This approach may be useful in generating monitoring protocols for other dynamic systems.