Inequalities in noise will affect urban wildlife.

Understanding how systemic biases influence local ecological communities is essential for developing just and equitable environmental practices that prioritize both human and wildlife well-being. With over 270 million residents inhabiting urban areas in the United States, the socioecological consequences of racially targeted zoning, such as redlining, need to be considered in urban planning. There is a growing body of literature documenting the relationships between redlining and the inequitable distribution of environmental harms and goods, green space cover and pollutant exposure. However, it remains unknown whether historical redlining affects the distribution of urban noise or whether inequitable noise drives an ecological change in urban environments. Here we conducted a spatial analysis of how urban noise corresponds to the distribution of redlining categories and a systematic literature review to summarize the effects of noise on wildlife in urban landscapes. We found strong evidence to indicate that noise is inequitably distributed in redlined urban communities across the United States, and that inequitable noise may drive complex biological responses across diverse urban wildlife, reinforcing the interrelatedness of socioecological outcomes. These findings lay a foundation for future research that advances relationships between acoustic and urban ecology through centring equity and challenging systems of oppression in wildlife studies.

Predicting the spatial expansion of an animal population with presence-only data.

Predictive models can improve the efficiency of wildlife management by guiding actions at the local, landscape and regional scales. In recent decades, a vast range of modelling techniques have been developed to predict species distributions and patterns of population spread. However, data limitations often constrain the precision and biological realism of models, which make them less useful for supporting decision-making. Complex models can also be challenging to evaluate, and the results are often difficult to interpret for wildlife management practitioners. There is therefore a need to develop techniques that are appropriately robust, but also accessible to a range of end users. We developed a hybrid species distribution model that utilises commonly available presence-only distribution data and minimal demographic information to predict the spread of roe deer (Capreolus caprelous) in Great Britain. We take a novel approach to representing the environment in the model by constraining the size of habitat patches to the home-range area of an individual. Population dynamics are then simplified to a set of generic rules describing patch occupancy. The model is constructed and evaluated using data from a populated region (England and Scotland) and applied to predict regional-scale patterns of spread in a novel region (Wales). It is used to forecast the relative timing of colonisation events and identify important areas for targeted surveillance and management. The study demonstrates the utility of presence-only data for predicting the spread of animal species and describes a method of reducing model complexity while retaining important environmental detail and biological realism. Our modelling approach provides a much-needed opportunity for users without specialist expertise in computer coding to leverage limited data and make robust, easily interpretable predictions of spread to inform proactive population management.

Horizontal viewsheds of large herbivores as a function of woodland structure.

There is great potential for the use of terrestrial laser scanning (TLS) to quantify aspects of habitat structure in the study of animal ecology and behaviour. Viewsheds-the area visible from a given position-influence an animal's perception of risk and ability to respond to potential danger. The management and conservation of large herbivores and their habitats can benefit greatly from understanding how vegetation structure shapes viewsheds and influences animal activity patterns and foraging behaviour. This study aimed to identify how woodland understory structure influenced horizontal viewsheds at deer eye height. Mobile TLS was used in August 2020 to quantify horizontal visibility-in the form of Viewshed Coefficients (VC)-and understory leaf area index (LAI) of 71 circular sample plots (15-m radius) across 10 woodland sites in North Wales (UK) where fallow deer (Dama dama) are present. The plots were also surveyed in summer for woody plant size structure, stem density and bramble (Rubus fruticosus agg.). Eight plots were re-scanned twice in winter to compare seasonal VC values and assess scan consistency. Sample plots with higher densities of small stems had significantly reduced VC 1 m from the ground. Other stem size classes, mean percentage bramble cover and understory LAI did not significantly affect VC. There was no difference in VC between summer and winter scans, or between repeated winter scans. The density of small stems influenced viewsheds at deer eye height and may alter behavioural responses to perceived risk. This study demonstrates how TLS technology can be applied to address questions in large herbivore ecology and conservation.

Capturing red squirrels (Sciurus vulgaris) on camera: A cost-effective approach for monitoring relative abundance and habitat preference.

Effective methods for monitoring animal populations are crucial for species conservation and habitat management. Motion-activated cameras provide an affordable method for passively surveying animal presence across the landscape but have mainly been used for studying large-bodied mammals. This paper explores the relative abundance and habitat preferences of red squirrels (Sciurus vulgaris) in coniferous forests using cameras and live trapping. The study was conducted in two forests (Newborough and Pentraeth) on Anglesey, North Wales, with a total of 50 sampling locations across four habitat categories. Detailed woodland structure and composition data were gathered around each sampling location. We found a strong positive correlation between the number of individual red squirrels live trapped over 10 days with the number of camera images of squirrels recorded during a previous 5-day period. The time interval between camera deployment and the first recorded image of a red squirrel showed a significant negative correlation with the number of individuals live trapped. Red squirrel relative abundance was negatively related to forest canopy openness, while the presence of Scots pine and increased tree species diversity were positively associated with the relative abundance of squirrels. There was also a strong site difference with lower relative abundance at Newborough compared with Pentraeth, which likely reflects the heavy thinning of mature forest at Newborough reducing tree crown connectivity. The results show that remotely activated cameras are an effective method for monitoring red squirrel populations across varying animal densities. The cameras also provided crucial information on red squirrel habitat preferences that can aid in woodland management and conservation efforts. Cameras have great potential to collect data on the population status of other small mammals, but it is essential that these methods are validated on a species-by-species basis.

A multifaceted framework to establish the presence of meaning in non-human communication.

Does non-human communication, like language, involve meaning? This question guides our focus through an interdisciplinary review of the theories and terminology used to study meaning across disciplines and species. Until now, it has been difficult to apply the concept of meaning to communication in non-humans. This is partly because of the varied approaches to the study of meaning. Additionally, while there is a scholarly acknowledgement of potential meaning in non-human cognition, there is also scepticism when the topic of communication arises. We organise some of the key literature into a coherent framework that can bridge disciplines and species, to ensure that aspects of meaning are accurately and fairly compared. We clarify the growing view in the literature that, rather than requiring multiple definitions or being split into different types, meaning is a multifaceted yet still unified concept. In so doing, we propose that meaning is an umbrella term. Meaning cannot be summed up with a short definition or list of features, but involves multiple complexities that are outlined in our framework. Specifically, three global facets are needed to describe meaning: a Signal Meaning Facet, an Interactant Meaning Facet, and a Resultant Meaning Facet. Most importantly, we show that such analyses are possible to apply as much to non-humans as to humans. We also emphasise that meaning nuances differ among non-human species, making a dichotomous approach to meaning questionable. Instead, we show that a multifaceted approach to meaning establishes how meaning appears within highly diverse examples of non-human communication, in ways consistent with the phenomenon's presence in human non-verbal communication and language(s). Therefore, without further recourse to 'functional' approaches that circumvent the critical question of whether any non-human meaning exists, we show that the concept of meaning is suitable for evolutionary biologists, behavioural ecologists, and others to study, to establish exactly which species exhibit meaning in their communication and in what ways.

Seasonal range fidelity of a megaherbivore in response to environmental change.

For large herbivores living in highly dynamic environments, maintaining range fidelity has the potential to facilitate the exploitation of predictable resources while minimising energy expenditure. We evaluate this expectation by examining how the seasonal range fidelity of African elephants (Loxodonta africana) in the Kruger National Park, South Africa is affected by spatiotemporal variation in environmental conditions (vegetation quality, temperature, rainfall, and fire). Eight-years of GPS collar data were used to analyse the similarity in seasonal utilisation distributions for thirteen family groups. Elephants exhibited remarkable consistency in their seasonal range fidelity across the study with rainfall emerging as a key driver of space-use. Within years, high range fidelity from summer to autumn and from autumn to winter was driven by increased rainfall and the retention of high-quality vegetation. Across years, sequential autumn seasons demonstrated the lowest levels of range fidelity due to inter-annual variability in the wet to dry season transition, resulting in unpredictable resource availability. Understanding seasonal space use is important for determining the effects of future variability in environmental conditions on elephant populations, particularly when it comes to management interventions. Indeed, over the coming decades climate change is predicted to drive greater variability in rainfall and elevated temperatures in African savanna ecosystems. The impacts of climate change also present particular challenges for elephants living in fragmented or human-transformed habitats where the opportunity for seasonal range shifts are greatly constrained.

The effects of population management on wild ungulates: A systematic map of evidence for UK species.

INTRODUCTION: Over recent decades, the abundance and geographic ranges of wild ungulate species have expanded in many parts of Europe, including the UK. Populations are managed to mitigate their ecological impacts using interventions, such as shooting, fencing and administering contraception. Predicting how target species will respond to interventions is critical for developing sustainable, effective and efficient management strategies. However, the quantity and quality of evidence of the effects of interventions on ungulate species is unclear. To address this, we systematically mapped research on the effects of population management on wild ungulate species resident in the UK. METHODS: We searched four bibliographic databases, Google Scholar and nine organisational websites using search terms tested with a library of 30 relevant articles. Worldwide published peer-reviewed articles were considered, supplemented by 'grey' literature from UK-based sources. Three reviewers identified and screened articles for eligibility at title, abstract and full-text levels, based on predefined criteria. Data and metadata were extracted and summarised in a narrative synthesis supported by structured graphical matrices. RESULTS: A total of 123 articles were included in the systematic map. Lethal interventions were better represented (85%, n = 105) than non-lethal interventions (25%, n = 25). Outcomes related to demography and behaviour were reported in 95% of articles (n = 117), whereas effects on health, physiology and morphology were studied in only 11% of articles (n = 14). Well-studied species included wild pigs (n = 58), red deer (n = 28) and roe deer (n = 23). CONCLUSIONS: Evidence for the effects of population management on wild ungulate species is growing but currently limited and unevenly distributed across intervention types, outcomes and species. Priorities for primary research include: species responses to non-lethal interventions, the side-effects of shooting and studies on sika deer and Chinese muntjac. Shooting is the only intervention for which sufficient evidence exists for systematic review or meta-analysis.

Social Disruption Impairs Predatory Threat Assessment in African Elephants.

The transmission of reliable information between individuals is crucial for group-living animals. This is particularly the case for cognitively advanced mammals with overlapping generations that acquire detailed social and ecological knowledge over long lifetimes. Here, we directly compare the ecological knowledge of elephants from two populations, with radically different developmental histories, to test whether profound social disruption affects their ability to assess predatory threat. Matriarchs (≤50 years of age) and their family groups received playbacks of three lions versus a single lion roaring. The family groups in the natural Amboseli population (Kenya) reliably assessed the greater predatory threat presented by three lions roaring versus one. However, in the socially disrupted Pilanesberg population (South Africa), no fine-scale distinctions were made between the numbers of roaring lions. Our results suggest that the removal of older and more experienced individuals in highly social species, such as elephants, is likely to impact the acquisition of ecological knowledge by younger group members, particularly through the lack of opportunity for social learning and cultural transmission of knowledge. This is likely to be exacerbated by the trauma experienced by juvenile elephants that witnessed the culling of family members and were translocated to new reserves. With increasing levels of anthropogenic disturbance, it is important that conservation practitioners consider the crucial role that population structure and knowledge transfer plays in the functioning and resilience of highly social and long-lived species.

Contrasting selection pressure on body and weapon size in a polygynous megaherbivore.

Sexual size dimorphism (SSD) is a common morphological trait in ungulates, with polygyny considered the leading driver of larger male body mass and weapon size. However, not all polygynous species exhibit SSD, while molecular evidence has revealed a more complex relationship between paternity and mating system than originally predicted. SSD is, therefore, likely to be shaped by a range of social, ecological and physiological factors. We present the first definitive analysis of SSD in the common hippopotamus (Hippopotamus amphibius) using a unique morphological dataset collected from 2994 aged individuals. The results confirm that hippos exhibit SSD, but the mean body mass differed by only 5% between the sexes, which is rather limited compared with many other polygynous ungulates. However, jaw and canine mass are significantly greater in males than females (44% and 81% heavier, respectively), highlighting the considerable selection pressure for acquiring larger weapons. A predominantly aquatic lifestyle coupled with the physiological limitations of their foregut fermenting morphology likely restricts body size differences between the sexes. Indeed, hippos appear to be a rare example among ungulates whereby sexual selection favours increased weapon size over body mass, underlining the important role that species-specific ecology and physiology have in shaping SSD.

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.

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.

Rangeland dynamics: investigating vegetation composition and structure of urban and exurban prairie dog habitat.

Rapid human population growth and habitat modification in the western United States has led to the formation of urban and exurban rangelands. Many of these rangelands are also home to populations of black-tailed prairie dogs (Cynomys ludovicianus). Our study aimed to compare the vegetation composition of an urban and exurban rangeland, and explore the role that prairie dogs play in these systems. The percent absolute canopy cover of graminoids (grasses and grass-likes), forbs, shrubs, litter, and bare ground were estimated at sampling areas located on and off prairie dog colonies at an urban and an exurban site. Herbaceous forage quality and quantity were determined on plant material collected from exclosure cages located on the colony during the entire growing season, while a relative estimate of prairie dog density was calculated using maximum counts. The exurban site had more litter and plant cover and less bare ground than the urban site. Graminoids were the dominant vegetation at the exurban plots. In contrast, mostly introduced forbs were found on the urban prairie dog colony. However, the forage quality and quantity tests demonstrated no difference between the two colonies. The relative prairie dog density was greater at the urban colony, which has the potential to drive greater vegetation utilization and reduced cover. Exurban rangeland showed lower levels of impact and retained all of the plant functional groups both on- and off-colony. These results suggest that activities of prairie dogs might further exacerbate the impacts of humans in fragmented urban rangeland habitats. Greater understanding of the drivers of these impacts and the spatial scales at which they occur are likely to prove valuable in the management and conservation of rangelands in and around urban areas.

Recommended survey designs for occupancy modelling using motion-activated cameras: insights from empirical wildlife data.

Motion-activated cameras are a versatile tool that wildlife biologists can use for sampling wild animal populations to estimate species occurrence. Occupancy modelling provides a flexible framework for the analysis of these data; explicitly recognizing that given a species occupies an area the probability of detecting it is often less than one. Despite the number of studies using camera data in an occupancy framework, there is only limited guidance from the scientific literature about survey design trade-offs when using motion-activated cameras. A fuller understanding of these trade-offs will allow researchers to maximise available resources and determine whether the objectives of a monitoring program or research study are achievable. We use an empirical dataset collected from 40 cameras deployed across 160 km(2) of the Western Slope of Colorado, USA to explore how survey effort (number of cameras deployed and the length of sampling period) affects the accuracy and precision (i.e., error) of the occupancy estimate for ten mammal and three virtual species. We do this using a simulation approach where species occupancy and detection parameters were informed by empirical data from motion-activated cameras. A total of 54 survey designs were considered by varying combinations of sites (10-120 cameras) and occasions (20-120 survey days). Our findings demonstrate that increasing total sampling effort generally decreases error associated with the occupancy estimate, but changing the number of sites or sampling duration can have very different results, depending on whether a species is spatially common or rare (occupancy = ψ) and easy or hard to detect when available (detection probability = p). For rare species with a low probability of detection (i.e., raccoon and spotted skunk) the required survey effort includes maximizing the number of sites and the number of survey days, often to a level that may be logistically unrealistic for many studies. For common species with low detection (i.e., bobcat and coyote) the most efficient sampling approach was to increase the number of occasions (survey days). However, for common species that are moderately detectable (i.e., cottontail rabbit and mule deer), occupancy could reliably be estimated with comparatively low numbers of cameras over a short sampling period. We provide general guidelines for reliably estimating occupancy across a range of terrestrial species (rare to common: ψ = 0.175-0.970, and low to moderate detectability: p = 0.003-0.200) using motion-activated cameras. Wildlife researchers/managers with limited knowledge of the relative abundance and likelihood of detection of a particular species can apply these guidelines regardless of location. We emphasize the importance of prior biological knowledge, defined objectives and detailed planning (e.g., simulating different study-design scenarios) for designing effective monitoring programs and research studies.

Behavioral responses associated with a human-mediated predator shelter.

Human activities in protected areas can affect wildlife populations in a similar manner to predation risk, causing increases in movement and vigilance, shifts in habitat use and changes in group size. Nevertheless, recent evidence indicates that in certain situations ungulate species may actually utilize areas associated with higher levels of human presence as a potential refuge from disturbance-sensitive predators. We now use four-years of behavioral activity budget data collected from pronghorn (Antilocapra americana) and elk (Cervus elephus) in Grand Teton National Park, USA to test whether predictable patterns of human presence can provide a shelter from predatory risk. Daily behavioral scans were conducted along two parallel sections of road that differed in traffic volume--with the main Teton Park Road experiencing vehicle use that was approximately thirty-fold greater than the River Road. At the busier Teton Park Road, both species of ungulate engaged in higher levels of feeding (27% increase in the proportion of pronghorn feeding and 21% increase for elk), lower levels of alert behavior (18% decrease for pronghorn and 9% decrease for elk) and formed smaller groups. These responses are commonly associated with reduced predatory threat. Pronghorn also exhibited a 30% increase in the proportion of individuals moving at the River Road as would be expected under greater exposure to predation risk. Our findings concur with the 'predator shelter hypothesis', suggesting that ungulates in GTNP use human presence as a potential refuge from predation risk, adjusting their behavior accordingly. Human activity has the potential to alter predator-prey interactions and drive trophic-mediated effects that could ultimately impact ecosystem function and biodiversity.

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

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.

Effects of social disruption in elephants persist decades after culling.

BACKGROUND: Multi-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. RESULTS: We 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. CONCLUSIONS: Key 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.

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.

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 (> or =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.

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".