Hotter deserts and the impending challenges for the Spiny-tailed Lizard in India.

Ectotherms are particularly vulnerable to climate change, especially those living in extreme areas, such as deserts, where species are already thermally constrained. Using the vulnerable herbivorous lizard Saara hardwickii as a model system, we used a multi-pronged approach to understand the thermal ecology of a desert agamid and potential impacts of rising temperatures. Our data included field-based measures of operative temperatures, body temperatures, and activity, as well as lab-based measures of thermal limits, preferences, and sprint speed. As expected, the temperature dependence of locomotor performance and foraging activity were different, and in the worst-case global warming scenario (SSP5-8.5), potential sprint speed may decrease by up to 14.5% and foraging activity may decrease by up to 43.5% by 2099. Burrows are essential thermal refuges, and global warming projections suggest that S. hardwickii may be restricted to burrows for up to 9 h per day by 2099, which would greatly limit critical activities, like foraging and seeking mating opportunities. Overall, we show that key information on thermal ecology, including temperature-sensitive behaviours in the wild, is necessary to understand the multiple ways in which increasing temperatures may influence ectothermic vertebrates, especially for species like S. hardwickii that are already vulnerable to environmental change.

Too much information? Males convey parasite levels using more signal modalities than females utilise.

Elaborate sexual signals are thought to have evolved and be maintained to serve as honest indicators of signaller quality. One measure of quality is health, which can be affected by parasite infection. Cnemaspis mysoriensis is a diurnal gecko that is often infested with ectoparasites in the wild, and males of this species express visual (coloured gular patches) and chemical (femoral gland secretions) traits that receivers could assess during social interactions. In this paper, we tested whether ectoparasites affect individual health, and whether signal quality is an indicator of ectoparasite levels. In wild lizards, we found that ectoparasite level was negatively correlated with body condition in both sexes. Moreover, some characteristics of both visual and chemical traits in males were strongly associated with ectoparasite levels. Specifically, males with higher ectoparasite levels had yellow gular patches with lower brightness and chroma, and chemical secretions with a lower proportion of aromatic compounds. We then determined whether ectoparasite levels in males influence female behaviour. Using sequential choice trials, wherein females were provided with either the visual or the chemical signals of wild-caught males that varied in ectoparasite level, we found that only chemical secretions evoked an elevated female response towards less parasitised males. Simultaneous choice trials in which females were exposed to the chemical secretions from males that varied in parasite level further confirmed a preference for males with lower parasites loads. Overall, we find that although health (body condition) or ectoparasite load can be honestly advertised through multiple modalities, the parasite-mediated female response is exclusively driven by chemical signals.

The extent of rapid colour change in male agamid lizards is unrelated to overall sexual dichromatism.

Dynamic colour change is widespread in ectothermic animals, but has primarily been studied in the context of background matching. For most species, we lack quantitative data on the extent of colour change across different contexts. It is also unclear whether and how colour change varies across body regions, and how overall sexual dichromatism relates to the extent of individual colour change. In this study, we obtained reflectance measures in response to different stimuli for males and females of six species of agamid lizards (Agamidae, sister family to Chameleonidae) comprising three closely related species pairs. We computed the colour volume in a lizard-vision colour space occupied by males and females of each species and estimated overall sexual dichromatism based on the area of non-overlapping male and female colour volumes. As expected, males had larger colour volumes than females, but the extent of colour change in males differed between species and between body regions. Notably, species that were most sexually dichromatic were not necessarily those in which males showed the greatest individual colour change. Our results indicate that the extent of colour change is independent of the degree of sexual dichromatism and demonstrate that colour change on different body regions can vary substantially even between pairs of closely related species.

Animal movement ecology in India: insights from 2011-2021 and prospective for the future.

The field of animal movement ecology has advanced by leaps and bounds in the past few decades with the advent of sophisticated technology, advanced analytical tools, and multiple frameworks and paradigms to address key ecological problems. Unlike the longer history and faster growth of the field in North America, Europe, and Africa, movement ecology in Asia has only recently been gaining momentum. Here, we provide a review of the field from studies based in India over the last 11 years (2011-2021) curated from the database, Scopus, and search engine, Google Scholar. We identify current directions in the research objectives, taxa studied, tracking technology and the biogeographic regions in which animals were tracked, considering the years since the last systematic review of movement ecology research in the country. As an indication of the growing interest in this field, there has been a rapid increase in the number of publications over the last decade. Class Mammalia continues to dominate the taxa tracked, with tiger and leopard being the most common species studied across publications. Invertebrates and other small and medium-sized animals, as well as aquatic animals, in comparison, are understudied and remain among the important target taxa for tracking in future studies. As in the previous three decades, researchers have focussed on characterising home ranges and habitat use of animals. There is, however, a notable shift to examine the movement decision of animals in human-modified landscapes, although efforts to use movement ecology to understand impacts of climate change remain missing. Given the biogeographic and taxonomic diversity of India, and the fact that the interface between anthropogenic activity and wildlife interactions is increasing, we suggest ways in which the field of movement ecology can be expanded to facilitate ecological insights and conservation efforts. With the advancement of affordable technologies and the availability of analytical tools, the potential to expand the field of movement ecology, shift research foci, and gain new insights is now prime.

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.

Colours of stress in male Indian rock agamas predict testosterone levels but not performance.

Rapid physiological colour change offers dynamic signalling opportunities that can reveal distinct information to receivers in different contexts. Information content in dynamic colours, however, is largely unexplored. In males of the Indian rock agama (Psammophilus dorsalis), stressful events, including male-male agonistic interactions, induce a colour change, wherein the dorsal band turns yellow and the lateral bands turn orange. We aimed to determine whether these pigment-based dynamic colours convey information about individual quality. Using an agamid-specific visual model, we first quantified the chromatic and achromatic contrasts of each colour component displayed by males during handling stress, which induces the maximal response of aggression-typical colours. We then measured baseline testosterone levels, morphology (body mass and size), and performance measures (bite force and sprint speed) of these lizards. Chromatic contrasts of the dorsal yellow and lateral orange bands, individually and relative to each other (internal pair), were negatively correlated with testosterone levels, while the chromatic contrast of the internal pair was positively correlated with body condition. The lack of an association between colour contrasts and both bite force and sprint speed indicate that the conspicuousness of colours expressed during stressful events, such as agonistic interactions, do not reveal male performance ability. Despite our expectations of a positive relationship with testosterone, morphology (body condition), and performance (bite force, sprint speed), we find that for P. dorsalis, the conspicuousness of stress-induced colours provide only some information about individual quality. We speculate that the dynamicity of physiological colours may influence their function as content-containing signals in social interactions.

Correlated evolution of parental care with dichromatism, colors, and patterns in anurans.

Parental care is widespread and has fitness benefits. But caregiving parents incur costs including higher predation, and this may lead to selection for body colors or patterns that help mitigate the risks of caring. The evolution of coloration, including sexual dichromatism, however, can be driven by other factors, such as sexual selection. Therefore, examining the associations between parental care and color patterns may provide key insights into evolutionary patterns and selection pressures for parental care. Our comparative analysis of 988 anuran species reveals that dichromatic species are less likely to provide parental care, irrespective of the caregiving sex, and are more likely to breed in aquatic habitats. We then examined whether dorsal colors and patterns that enhance crypticity or function as aposematic signals are associated with the caregiving sex, and the modality of care (transport or stationary). Only caregiving males are more likely to have dorsal Stripes, but none of the colors (Green-Brown, Red, Yellow, Blue-Black) and other patterns (Plain, Bands, Spots, Mottled-Patches) were associated with caregiving females or the modality of care. Overall, sexual dichromatism, breeding ecology, and parental care are associated, but the evolution of caregiving behavior does not appear to influence the myriad colors and patterns characteristic of anurans globally.

The ecology of sleep in non-avian reptiles.

Sleep is ubiquitous in the animal kingdom and yet displays considerable variation in its extent and form in the wild. Ecological factors, such as predation, competition, and microclimate, therefore are likely to play a strong role in shaping characteristics of sleep. Despite the potential for ecological factors to influence various aspects of sleep, the ecological context of sleep in non-avian reptiles remains understudied and without systematic direction. In this review, we examine multiple aspects of reptilian sleep, including (i) habitat selection (sleep sites and their spatio-temporal distribution), (ii) individual-level traits, such as behaviour (sleep postures), morphology (limb morphometrics and body colour), and physiology (sleep architecture), as well as (iii) inter-individual interactions (intra- and inter-specific). Throughout, we discuss the evidence of predation, competition, and thermoregulation in influencing sleep traits and the possible evolutionary consequences of these sleep traits for reptile sociality, morphological specialisation, and habitat partitioning. We also review the ways in which sleep ecology interacts with urbanisation, biological invasions, and climate change. Overall, we not only provide a systematic evaluation of the conceptual and taxonomic biases in the existing literature on reptilian sleep, but also use this opportunity to organise the various ecological hypotheses for sleep characteristics. By highlighting the gaps and providing a prospectus of research directions, our review sets the stage for understanding sleep ecology in the natural world.

An expanding cityscape and its multi-scale effects on lizard distribution.

Urbanization results in complex and variable changes to environmental conditions, which translate to shifts in selection pressures for organisms. Size of a city as well as the intensity and extent of urbanization can synergistically influence how organisms are impacted. However, less is known about how landscape heterogeneity, rate of land-use change, and scale of urbanization affect species persistence. We evaluate the ways in which urbanization changes the environment and examine how some of these environmental factors influence the presence of the lizard Psammophilus dorsalis (Peninsular rock agama), in Bengaluru, India. Variability in environmental factors across the study area was characterised by measures of habitat composition and diversity, habitat connectivity, rate of habitat change, predation pressure, land surface temperature (LST) and artificial light at night (ALAN), that were derived from remotely sensed and citizen science data. Most of these factors showed high variance across two measures of urbanization: distance from city center and proportion of built-up area. Habitat diversity and ALAN were the only two factors that changed predictably and in a non-linear way, with distance from the city center and proportion of built-up area. We then used a multi-scale approach to examine the relative importance of some these environmental factors at the landscape scale, as well as additional factors at the microhabitat-scale, in predicting the presence and relative abundance of P. dorsalis respectively. At the landscape scale, LST, which is positively correlated with proportion of cropland, predicted lizard presence; whereas at the microhabitat scale, P. dorsalis was more likely to be found in sites with higher proportions of rocks. Overall, we demonstrate that urbanization can result in environmental predictors that do not vary linearly across the urbanization gradient. For the iconic rock agama, many of these environmental factors do not seem to be strong selection pressures that influence their distribution in the expanding cityscape. Whether this urban utilizer can continue to persist with increasing anthropogenic development is uncertain. To better understand drivers of species persistence, we emphasize the importance of quantifying urbanization across multiple axes, considering environmental factors that are relevant to species at different spatial and temporal scales.

A longitudinal study of perceived stress and cortisol responses in an undergraduate student population from India.

Young adults entering college experience immense shifts in personal and professional environments. Such a potentially stressful event may trigger multiple psychological and physiological effects. In a repeated-measures longitudinal survey (N = 6 time-points) of first year cohort of residential undergraduate students in India, this study evaluates multiple psychological parameters: PSS14 (Perceived Stress Scale), K10 (distress scale) and positive mood measures, along with salivary cortisol levels. We find that compared to women, men showed significantly lower levels of salivary cortisol and also a decrease in perceived stress (PSS14) and distress (K10) with time. By contrast, women reported similar perceived stress and distress levels over time but had higher cortisol levels at the end of the academic year. Academic stress was reported by the students to be the most important stressor. This study highlights notable gender-/sex-differences in psychological and physiological stress responses and adds a valuable longitudinal dataset from the Indian undergraduate student cohort which is lacking in literature.

Diet influences latitudinal gradients in life-history traits, but not reproductive output, in ectotherms.

Aim: Latitudinal gradients in life-history traits are apparent in many taxa and are expected to be strong for ectotherms that have temperature-driven constraints on performance and fitness. The strength of these gradients, however, should also be affected by diet. Because diet type (carnivory, omnivory, herbivory) influences accessibility to nutrition and assimilation efficiency, we aim to study how diet affects latitudinal gradients in lifetime reproductive output and the underlying life-history traits in ectotherms. Location: Global. Time period: Recent. Major taxa studied: Lizards (Reptilia, Squamata, Sauria). Methods: We used empirical (352 species) and phylogenetically imputed data (563 species) to analyse the interactive effects of latitude and diet on life-history traits (longevity, age at maturity, reproductive life span, hatchling mass, clutch/brood size, clutch/brood frequency, female mass) and lifetime reproductive output of lizards. Results: Lifetime reproductive output does not significantly differ in lizards across diet types, and only carnivores exhibit a small increase at higher latitudes. Diet type, however, influences latitudinal patterns of individual life-history traits. Carnivores exhibit a shift towards 'slower-paced' life histories at higher latitudes for most traits (increased longevity, age at maturity, reproductive life span, and decreased clutch frequency). By contrast, herbivores either display 'faster-paced' life histories (reduction in reproductive life span, hatchling mass, female mass) or no change (clutch frequency, clutch size, age at maturity) at higher latitudes. Omnivores exhibit intermediate and muted latitudinal patterns. Main conclusions: We suggest that the nutritional challenges of herbivory, compounded by thermal constraints at higher latitudes, may explain differences in life-history characteristics of herbivorous ectotherms. Intermediate patterns exhibited by omnivores highlight how flexibility in diet can buffer environmental challenges at higher latitudes. Our results indicate that lizards with different diet types display various trends in their life histories across latitudes, which eventually balance out to result in similar reproductive outputs throughout their lifetime, with little benefits to carnivory.

Asymmetric interspecific competition drives shifts in signalling traits in fan-throated lizards.

Interspecific competition can occur when species are unable to distinguish between conspecific and heterospecific mates or competitors when they occur in sympatry. Selection in response to interspecific competition can lead to shifts in signalling traits-a process called agonistic character displacement. In two fan-throated lizard species-Sitana laticeps and Sarada darwini-females are morphologically indistinguishable and male agonistic signalling behaviour is similar. Consequently, in areas where these species overlap, males engage in interspecific aggressive interactions. To test whether interspecific male aggression between Si. laticeps and Sa. darwini results in agonistic character displacement, we quantified species recognition and signalling behaviour using staged encounter assays with both conspecifics and heterospecifics across sympatric and allopatric populations of both species. We found an asymmetric pattern, wherein males of Si. laticeps but not Sa. darwini showed differences in competitor recognition and agonistic signalling traits (morphology and behaviour) in sympatry compared with allopatry. This asymmetric shift in traits is probably due to differences in competitive abilities between species and can minimize competitive interactions in zones of sympatry. Overall, our results support agonistic character displacement, and highlight the role of asymmetric interspecific competition in driving shifts in social signals.

Mate-searching context of prey influences the predator-prey space race.

Predation risk is a strong driver of prey distribution and movement. However, fitness-influencing behaviours, such as mating, can alter risk and influence predator-prey space-use dynamics. In tree crickets, Oecanthus henryi, mate searching involves acoustic signalling by immobile males and phonotactic movement by females. Space-use patterns in tree crickets relative to their primary predators, green lynx spiders (Peucetia viridans), should therefore depend on their current mate-searching state; whether males are calling or non-calling and whether females are phonotactic or non-phonotactic. We first measured the degree of spatial anchoring of crickets to specific bushes in the field and determined whether that influenced the probability of broad-scale spatial overlap with spiders. In the absence of spiders, all crickets, independent of sex or male calling status, were found to be spatially anchored to specific types of bushes and not uniformly distributed on the landscape. At the broad spatial scale, spiders were more likely to be found on bushes with female crickets and, to a lesser degree, calling male crickets. At a finer spatial scale within a bush, movement strategies of crickets not only varied depending on the presence or absence of a spider, but also on their current mate-searching state. Phonotactic females showed clear predator avoidance, whereas calling and non-calling males moved towards the spider instead of away, similar to predator inspection behaviour seen in many taxa. As the strongly selected sex, males are more likely to undertake risky mate-searching activities, which includes inspection of predator positions. Overall, we found that all crickets were predictably anchored at the landscape scale, but their sex and mate-seeking behaviour influenced the degree of overlap with predators and their antipredator movement strategies. Reproductive strategies within a prey species, therefore, can alter predator-prey space race at multiple spatial scales.

Non-consumptive predator effects on prey population size: A dearth of evidence.

There is a large and growing interest in non-consumptive effects (NCEs) of predators. Diverse and extensive evidence shows that predation risk directly influences prey traits, such as behaviour, morphology and physiology, which in turn, may cause a reduction in prey fitness components (i.e. growth rate, survival and reproduction). An intuitive expectation is that NCEs that reduce prey fitness will extend to alter population growth rate and therefore population size. However, our intensive literature search yielded only 10 studies that examined how predator-induced changes in prey traits translate to changes in prey population size. Further, the scant evidence for risk-induced changes on prey population size have been generated from studies that were performed in very controlled systems (mesocosm and laboratory), which do not have the complexity and feedbacks of natural settings. Thus, although likely that predation risk alone can alter prey population size, there is little direct empirical evidence that demonstrates that it does. There are also clear reasons that risk effects on population size may be much smaller than the responses on phenotype and fitness components that are typically measured, magnifying the need to show, rather than infer, effects on population size. Herein we break down the process of how predation risk influences prey population size into a chain of events (predation risk affects prey traits, which affect prey fitness components and population growth rate, which affect prey population size), and highlight the complexity of each transition. We illustrate how the outcomes of these transitions are not straightforward, and how environmental context strongly dictates the direction and magnitude of effects. Indeed, the high variance in prey responses is reflected in the variance of results reported in the few studies that have empirically quantified risk effects on population size. It is therefore a major challenge to predict population effects given the complexity of how environmental context interacts with predation risk and prey responses. We highlight the critical need to appreciate risk effects at each level in the chain of events, and that changes at one level cannot be assumed to translate into changes in the next because of the interplay between risk, prey responses, and the environment. The gaps in knowledge we illuminate underscore the need for more evidence to substantiate the claim that predation risk effects extend to prey population size. The lacunae we identify should inspire future studies on the impact of predation risk on population-level responses in free-living animals.

Risk of social colours in an agamid lizard: implications for the evolution of dynamic signals.

The forces of sexual and natural selection are typically invoked to explain variation in colour patterns of animals. Although the benefits of conspicuous colours for social signalling are well documented, evidence for their ecological cost, especially for dynamic colours, remains limited. We examined the riskiness of colour patterns of Psammophilus dorsalis, a species in which males express distinct colour combinations during social interactions. We first measured the conspicuousness of these colour patterns on different substrates based on the visual systems of conspecifics and predators (bird, snake, canid) and then quantified actual predation risk on these patterns using wax/polymer lizard models in the wild. The black and red male state exhibited during courtship was the most conspicuous to all visual systems, while the yellow and orange male aggression state and the brown female colour were least conspicuous. Models bearing the courtship colour pattern experienced the highest predator attacks, irrespective of the substrate they were placed on. Thus, social colours of males are not only conspicuous but also risky. Using physiological colours to shift in and out of conspicuous states may be an effective evolutionary solution to balance social signalling benefits with predation costs.

Lizards from suburban areas learn faster to stay safe.

Enhanced cognitive ability is beneficial in unpredictable and harsh environments, as it enables animals to respond with flexibility. For animals living in urbanized areas, local environments not only are altered but can rapidly change during their lifetime. Urban residents are therefore challenged with identifying novel dangers and safe refuges in dynamic environments. We demonstrate that the tropical agamid lizard Psammophilus dorsalis experiences dramatically different habitats not only across the rural to urban spatial scale but also over the short temporal scale of a few years in suburban areas. Differences in environmental stability are expected to affect rates of learning and reversal learning in resident lizards. In testing arenas, lizards from these populations were required to choose a designated 'safe' refuge instead of an 'unsafe' one after simulated predator attacks. The contingency for safety was switched during the reversal learning task. In general, P. dorsalis showed high rates of learning and reversal learning, but lizards from suburban areas were quicker to learn and unlearn the location of the safe refuge than those from rural areas. This demonstrates for the first time to our knowledge that suburban lizards have faster learning and reversal learning skills for a key survival-related behaviour, finding safety in unpredictable environments.

Social coping styles of lizards are reactive and not proactive in urban areas.

Animals engage in social interactions with changes in their behaviour and physiology. Environmental challenges, however, can influence social interactions by adding additional stressors. Here, we investigated the effects of urbanisation on the behaviour and hormonal responses of a tropical lizard species, Psammophilus dorsalis, during social interactions. We recorded behaviour of males from suburban and rural areas during controlled encounters with other males and females. We then measured corticosterone and testosterone levels of individuals at 10 min intervals, from immediately after the social encounter to 30 min later and then at 120 min after the interaction period. We found that differences in social behaviours and subsequent hormone levels were largely driven by habitat, and not social context. Overall, we found that fewer suburban males showed behavioural displays compared to rural males during social encounters. For those that displayed, intensity of aggression was similar across populations, but courtship intensity was lower for suburban males compared to rural males. Suburban males also had significantly elevated levels of corticosterone both under control conditions (no social encounter) and following intra- and intersexual interactions, while rural males retained low levels of corticosterone across contexts. Social interactions were associated with an increase in testosterone levels in all males, but only rural males maintained elevated levels for up to 120 min after interactions with females. Thus, lizards from these suburban and rural populations showed key differences in responsiveness to and recovery from social challenges, a pattern that suggests alternative coping styles ('proactive' vs. 'reactive'). These differences in social coping styles could influence consequences of sexual selection in an urbanised world.

Wind farms have cascading impacts on ecosystems across trophic levels.

Wind farms are a cleaner alternative to fossil fuels for mitigating the effects of climate change, but they also have complex ecological consequences. In the biodiversity hotspot of the Western Ghats in India, we find that wind farms reduce the abundance and activity of predatory birds (for example, Buteo, Butastur and Elanus species), which consequently increases the density of lizards, Sarada superba. The cascading effects of wind turbines on lizards include changes in behaviour, physiology and morphology that reflect a combination of predator release and density-dependent competition. By adding an effective trophic level to the top of food webs, we find that wind farms have emerging impacts that are greatly underestimated. There is thus a strong need for an ecosystem-wide view when aligning green-energy goals with environment protection.

Lizards assess complex social signals by lateralizing colour but not motion detection.

Vertebrates lateralize many behaviours including social interactions. Social displays typically comprise multiple components, yet our understanding of how these are processed comes from studies that typically examine responses to the dominant component or the complex signal as a whole. Here, we examined laterality in lizard responses to determine whether receivers separate the processing of motion and colour signal components in different brain hemispheres. In Psammophilus dorsalis, males display colours that dynamically change during courtship and aggressive interactions. We tested the visual grasp reflex of both sexes using robotic stimuli that mimicked two signal components: (1) multiple speeds of head-bobbing behaviour and (2) multiple colours. We found no laterality in response to different motion stimuli, indicating that motion similarly attracts attention from the two visual fields across sexes. Notably, receivers showed left visual field dominance to colours, especially when males were exposed to 'aggression-specific' colours and females to 'courtship-specific' colours.

Change in Mesoherbivore Browsing Is Mediated by Elephant and Hillslope Position.

Elephant are considered major drivers of ecosystems, but their effects within small-scale landscape features and on other herbivores still remain unclear. Elephant impact on vegetation has been widely studied in areas where elephant have been present for many years. We therefore examined the combined effect of short-term elephant presence (< 4 years) and hillslope position on tree species assemblages, resource availability, browsing intensity and soil properties. Short-term elephant presence did not affect woody species assemblages, but did affect height distribution, with greater sapling densities in elephant access areas. Overall tree and stem densities were also not affected by elephant. By contrast, slope position affected woody species assemblages, but not height distributions and densities. Variation in species assemblages was statistically best explained by levels of total cations, Zinc, sand and clay. Although elephant and mesoherbivore browsing intensities were unaffected by slope position, we found lower mesoherbivore browsing intensity on crests with high elephant browsing intensity. Thus, elephant appear to indirectly facilitate the survival of saplings, via the displacement of mesoherbivores, providing a window of opportunity for saplings to grow into taller trees. In the short-term, effects of elephant can be minor and in the opposite direction of expectation. In addition, such behavioural displacement promotes recruitment of saplings into larger height classes. The interaction between slope position and elephant effect found here is in contrast with other studies, and illustrates the importance of examining ecosystem complexity as a function of variation in species presence and topography. The absence of a direct effect of elephant on vegetation, but the presence of an effect on mesoherbivore browsing, is relevant for conservation areas especially where both herbivore groups are actively managed.