Testing a key assumption of wildlife buffer zones: is flight initiation distance a species-specific trait?

The distance at which animals flee an approaching predator is known as the ‘flight initiation distance’ (FID). Wildlife managers use FID to develop buffer zones to reduce human impacts on wildlife. Many variables have been demonstrated to influence FID leading one to question whether it can be viewed as a species-specific trait. We tested this critical assumption for developing buffer zones by experimentally approaching eight species of shorebirds found at six sites around Botany Bay, 15 km south of Sydney, Australia. Botany Bay encompasses a range of human impacted areas, from urban developments with high levels of human presence, through to National Parks and wildlife protection areas where human presence is significantly lower. We found that both species and site influenced the distance birds flew away from an approaching human. Importantly, however, there was no significant statistical interaction between site and species demonstrating that ‘flighty’ species were consistently flighty while more tolerant species were consistently tolerant. Taken together, these results suggest that FID can therefore be viewed as a species-specific trait for these shorebirds. The great variability in FID suggests that wildlife managers should be somewhat conservative in developing buffer zones, but they can use previously published FID data for a given species as guidelines for setting buffer zones.     

Spatial and temporal responses of forest birds to human approaches in a protected area and implications for two management strategies

Little is known about the responses of forest birds, other than raptors, to human disturbance despite their being highly sensitive to habitat disruption. We tested five hypotheses about the spatial and temporal interactions between recreationists and wildlife by using five South American birds as model species. We measured two components of flight distance, horizontal (distance between a visitor and the base of a perching tree) and vertical (distance between the base of a tree and the perch), and estimated the third diagonal component (distance between a visitor and the perch). We found that (a) the relationships between horizontal and vertical components of flight distance were negative (rufus-collared sparrow, golden-billed saltator, chiguanco thrush), positive (spot-winged pigeon), and neutral (bay-winged cowbird), suggesting that different components are used as clues to the proximity of people when birds vacated the perching tree; (b) the distance to the nearest pathway correlated positively with flight distance components of two species (bay-winged cowbird and spot-winged pigeon); and (c) physical structure of the vegetation affected differently flight distances of all species. Interspecific comparisons showed that (d) all flight distance components increased linearly with body size, and (e) species differed in landing distances but not in response duration: large-bodied species tended to land farther than smaller-bodied ones. By implementing buffer zones (areas without access to visitors), spatial restrictions would conflict with recreational activities. We recommend re-distributing (but not restricting) human visitation by varying the number of visitors and area of visitation according to the spatial requirements of differently sized species.     

Do fences protect birds from human disturbance?

Outdoor recreation and ecotourism are becoming increasingly popular, but such human activities are not entirely benign to birds. One way to manage wildlife habitats is to restrict public access with a fence or some similar barrier, under the assumption that this provides wildlife with a refuge from human activities. We tested this assumption by measuring the responses of 10 species of birds at a site containing a fence with a relatively large number of visitors on only one side. We compared these responses to those at a less-visited, control site. Responses were measured by quantifying flight initiation distance (FID), the distance birds would allow a human to approach before fleeing. Overall, we found birds on the protected side of the fence responded similarly to birds at the low visitation control site, and significantly differently from birds at the high visitation site. Our results suggest that by reducing the number of humans and providing areas of refuge within highly visited habitats, protective barriers allow birds to behave as they would in an undisturbed environment.     

The effect of recreational disturbance on an upland breeding bird, the golden plover Pluvialis apricaria

The use of the countryside for recreation has increased dramatically in recent years. This has led to concern amongst conservationists about the effects increased human disturbance may have on important wild animal populations. In the UK, recent legislation has widened the level of access to upland habitats, which support internationally important breeding bird populations. Determining the extent to which recreational disturbance affects upland breeding birds is therefore a conservation priority. We used data collected over 13 years to investigate the impact of recreational disturbance on the distribution and reproductive performance of golden plovers breeding in close proximity to the Pennine Way, an intensively used long-distance footpath. Importantly, the Pennine Way was resurfaced in 1994 to prevent further erosion of the surrounding vegetation. We were therefore able to examine if the response of golden plovers to recreational disturbance was influenced by changes in the intensity and extent of human activity resulting from the resurfacing work. Before the Pennine Way was resurfaced, golden plovers avoided areas within 200 m of the footpath during the chick-rearing period. At this time over 30% of people strayed from the footpath and the movement of people across the moorland was therefore widespread and unpredictable. Following resurfacing, over 96% of walkers remained on the Pennine Way, which significantly reduced the impact of recreational disturbance on golden plover distribution; golden plovers only avoided areas within 50 m of the footpath at this time. Despite the clear behavioural responses of golden plovers to the presence of visitors, there was no detectable impact of disturbance on reproductive performance. In many countries, a conflict arises between the use of the countryside for recreational purposes and the protection of habitats or species of high conservation value. However, this study suggests that the implementation of simple measures to influence visitor behaviour can dramatically reduce the impact of recreational disturbance on wild animal populations.     

Patterns of species change in anthropogenically disturbed forests of Madagascar

Five main conclusions arise from this review of the responses of species to anthropogenic disturbance in Madagascar: First, species’ reactions to anthropogenic disturbance are generally negative, but remain poorly known. Our knowledge is patchy among and within higher taxonomic groups; we are still largely gathering case studies. Second, taxonomic groups vary considerably in which proximate factors are most important. Third, several groups show differing responses within different ecoregions. Whether these differences are consistent across groups requires further testing. Fourth, related species often have divergent reactions to disturbance, even within lower taxonomic groupings (families or genera). Thus, we cannot rely on phylogenetic relatedness or even ecological similarity to infer similarity in responses. Finally, disturbance typically reduces species diversity (especially of native and/or endemic species), but also causes species turnover, typically with forest specialists replaced by grassland generalists, and endemics replaced by non-endemics (including invasives). Given these knowledge gaps, we stress the urgency of applied studies that assess species’ ecology, behaviour and health across disturbance gradients, including purely anthropogenic landscapes. Remaining natural vegetation and protected areas will be unable to preserve Madagascar’s biodiversity under the impact of climatic change; we must understand responses of plants and animals to disturbance in order to create buffer zones and corridors combining secondary, degraded and natural habitats.     

Exposure to ecotourism reduces survival and affects stress response in hoatzin chicks (Opisthocomus hoazin)

Ecotourism helps to protect many habitats, but may also have negative impacts on wildlife. We investigated effects of ecotourists on reproductive success of hoatzins (Opisthocomus hoazin) and on hormonal status of their chicks in Amazonian rainforest lakes by comparing birds from undisturbed and from tourist-exposed nests. Hatching success was similar in both groups but chick survival was much lower at tourist-exposed nests than at undisturbed nests. This effect was due to an increased mortality of juveniles prior to fledging whereas small nestlings seemed largely unaffected. Juveniles, but not nestlings, living at tourist-exposed sites had a lower body mass and showed a stronger hormonal response to experimental stress compared to individuals at undisturbed sites. These data suggest that juvenile hoatzins were susceptible to tourist-induced stress which in turn may be responsible for the lower survival. In contrast, adult hoatzins that were incubating had apparently habituated to tourist presence because their flush distances at tourist-exposed nests were 50% lower than at undisturbed sites. Our findings demonstrate that individuals in different life stages show different susceptibilities to tourism. We suggest that even just watching animals during breeding can threaten their survival, but a proper scientific management of off-limit zones and area-specific guidelines for wildlife observation could reduce harmful effects.     

The buffer effect of non-breeding birds and the timing of farmland bird declines

Breeding territoriality in birds means that a certain proportion of the adult population may breed in suboptimal habitats or not breed at all. Non-breeding birds, or floaters, can have a `buffer effect' on breeding population size. Farmland bird declines in Britain are measured and analysed in terms of numbers of territory-holding birds. We use a population dynamics model to illustrate how, because of the buffer effect of floaters, there can be a time lag between the start of total population decline and detectable breeding population decline. Differences in the timing of declines of closely related farmland birds could therefore be due to slight differences in their population dynamics, rather than differences in their response to agricultural change.     

Modelling the efficacy of proposed mitigation areas for shorebirds: a case study on the Seine estuary, France

A behaviour-based model was used to explore the effect of an extension of the port at Le Havre (Port 2000), and the effect of proposed mitigation measures, on the mortality and body condition of the three main shorebird species that overwinter in the estuary of the river Seine, France. In the model, a 20% reduction in the area of mudflats on the north side of the estuary had little effect on curlew Numenius arquata mortality and body condition but significantly increased mortality and decreased body condition in dunlin Calidris alpina and oystercatchers Haematopus ostralegus. Disturbance of feeding birds both day and night had a significant effect on the mortality and body condition of all three shorebird species, as did disturbance of roosting birds. Disturbance of feeding birds in the daytime only had a significant effect on dunlin mortality and body condition, but not that of curlew and oystercatchers. In the model, the creation of a buffer zone to reduce disturbance of feeding birds restored shorebird mortality and body condition to pre-disturbance levels. A new mudflat area was also effective in mitigating the effect of habitat loss on all three shorebirds and in mitigating the effect of roost disturbance on dunlin and curlew. However, a new mudflat area was not effective in mitigating the effect of roost disturbance on oystercatcher mortality and body condition. The effectiveness of the mitigating mudflat depended as much on its size as its quality. We believe that this is the first time that anyone has been able to forecast the efficacy of proposed mitigation measures.     

The impacts of roads and other infrastructure on mammal and bird populations: A meta-analysis

Biodiversity is being lost at an increased rate as a result of human activities. One of the major threats to biodiversity is infrastructural development. We used meta-analyses to study the effects of infrastructure proximity on mammal and bird populations. Data were gathered from 49 studies on 234 mammal and bird species. The main response by mammals and birds in the vicinity of infrastructure was either avoidance or a reduced population density. The mean species abundance, relative to non-disturbed distances (MSA), was used as the effect size measure. The impact of infrastructure distance on MSA was studied using meta-analyses. Possible sources of heterogeneity in the results of the meta-analysis were explored with meta-regression.Mammal and bird population densities declined with their proximity to infrastructure. The effect of infrastructure on bird populations extended over distances up to about 1 km, and for mammal populations up to about 5 km. Mammals and birds seemed to avoid infrastructure in open areas over larger distances compared to forested areas, which could be related to the reduced visibility of the infrastructure in forested areas. We did not find a significant effect of traffic intensity on the MSA of birds. Species varied in their response to infrastructure. Raptors were found to be more abundant in the proximity of infrastructure whereas other bird taxa tended to avoid it. Abundances were affected at variable distances from infrastructure: within a few meters for small-sized mammals and up to several hundred meters for large-sized mammals.Our findings show the importance of minimizing infrastructure development for wildlife conservation in relatively undisturbed areas. By combining actual species distributions with the effect distance functions we developed, regions sensitive to infrastructure development may be identified. Additionally, the effect distance functions can be used in models in support of decision making on infrastructure planning.     

Savanna herbivore dynamics in a livestock-dominated landscape: I. Dependence on land use, rainfall, density, and time

Ecosystem processes in African savannas can be better conserved if management is based on a mechanistic understanding of wildlife dynamics in livestock-dominated landscapes. For Laikipia District, a non-protected savanna region in northern Kenya, we used spatially explicit estimates of density to characterize factors influencing the dynamics of large herbivores on three land-use types: commercial ranches that favor wildlife, communal ‘group ranches’ practicing pastoralism, and the remainder (‘transitional’ properties). For 21-year time series of nine wild and two domestic species, linear model selection was used to ascribe between 45% (Grant’s gazelle) and 95% (plains zebra) of observed variation in biomass density to land use, rainfall-dependence, density-dependence, and trends over time.Strongly opposing patterns of variation across the landscape in wildlife and livestock densities affirmed the primacy of land use among factors influencing wildlife abundance in non-protected areas. Rainfall limited densities of only the dominant grazing species throughout the monitoring period (plains zebra and cattle), and of most other species while their densities were high. Regulating effects of density were detected only for the dominant wild grazing and browsing species (zebra and giraffe). All but two wild species (zebra and Grant’s gazelle) declined on at least one land-use type, for reasons that varied among land uses.Where favored, diverse and abundant wild herbivores (mean of 1.7 t km⁻² on pro-wildlife ranches) can thrive even when sharing the landscape with a slightly higher biomass density of livestock (mean of 2.7 t km⁻²). Where not favored, only a few resilient wild species (e.g. gazelles and plains zebra) persist with high densities of livestock (mean of 4.6 t km⁻² on transitional ranches). Maintaining higher wild species diversity in the landscape will depend on the creation of a network of unfenced conservation areas in which livestock densities are persistently low or zero, which are sufficiently large to act as ‘sources’ of wild species that are prone to displacement by humans and livestock, and which generate benefits to community members that exceed opportunity costs.     

Effects of disturbance and habitat loss on papyrus-dwelling passerines

Throughout the world, but particularly in tropical regions habitat loss and disturbance are considered detrimental to biodiversity. We examine the effects of disturbance by harvesting, burning and habitat fragmentation on six bird species associated with papyrus (Cyperus papyrus) swamps. The presence or absence of these species was verified in 93 wetlands in southern Uganda between June and August 2003. Disturbance was estimated directly by observation and indirectly from examining vegetation structure. Habitat fragmentation was quantified by delineating swamps on Landsat ETM images and applying ‘Fragstats’ to calculate relevant patch metrics. The occurrence of all six species was affected positively by increased swamp size, but birds were more likely to occur in small swamps at high altitude. The shape and proximity of swamps to neighbouring wetlands had little effect on occurrence. As altitude, habitat fragmentation and disturbance were correlated, determining avian responses to disturbance was problematic. However, the occurrence of all species was affected by disturbance, often showing unimodal responses, if disturbance was not considered in conjunction with habitat fragmentation. When the effects of habitat fragmentation and disturbance were analysed together, only the occurrences of papyrus yellow warbler and white-winged warbler were affected (positively) by disturbance. Results suggest that papyrus-dwelling passerines, except papyrus gonolek, are tolerant of low intensities of disturbance, a novel finding in the tropics. We recommend that policy-makers do not prohibit harvesting, thus generating good will and encouraging rural householders to comply with additional conservation policy.     

Dampened behavioral and physiological responses mediate birds’ association with humans

The study of the mechanisms involved in phenotypic adaptations or plasticity to environmental stressors may serve to evaluate contemporary ecological and evolutionary dynamics in threatened or growing wildlife populations. By addressing human disturbance as a main stressor agent, we investigated antipredator response (flushing distance), social behavior (degree of gregarism), health (fecal pathogens), and endocrine status (fecal corticosterone levels) as potential mechanisms that permit birds to cope with anthropogenic impact in tourist areas. Our study models were species that are neither urbanized nor invasive, and that are typical of the high mountains of the Palearctic: the Alpine and the Red-billed chough (Pyrrhocorax graculus and P. pyrrhocorax), and the surrounding alpine bird community. Both chough species showed habituation, reduced flushing distances, fewer parasites and lower corticosterone levels in tourist than in non-tourist sites, and a significant variation in gregarious behavior. Behavioral and physiological adjustments permitted these corvids to optimize resource exploitation depending on habitat quality and in conditions of continuous disturbance. Human-induced changes in chough social behavior had no effect on local avian species richness, a surrogate measure for community diversity. Irrespectively of chough aggregations, bird diversity decreased significantly in recreational areas, thus not all community members tolerated human impact as choughs did. Our findings suggest that species that manage to coexist with humans primarily adjust traits that influence ecological interactions (social, foraging or antipredatory responses) and that vary rapidly (learned behaviors and physiological mechanisms that promote homeostasis).     

The distribution,population size,movements and conservation of the African black oystercatcher Haematopus moquini

The African black oystercatcher Haematopus moquini is endemic to southern Africa and occurs as a breeding species between Seal Island, South West Africa/Namibia and Mazeppa Bay, Transkei. Vagrants outside this area are rare further north than the Hoanib Estuary, South West Africa/Namibia and further east than Transkei. The present adult population is estimated to be 4781 birds of which 53% are found in the southwestern Cape Province, South Africa. Offshore islands support 29% of the world population and are identified as important localities for African black oystercatchers with a high breeding production. These birds face threats from human disturbance, expecially from off-road vehicles on the mainland, from introduced mammalian predators at islands and from competition with man for food and bait organisms. Ten islands and seven mainland sites are isolated as supporting more than 1% of the world population, and, together with their intertidal zones, are recommended for Nature Reserve status.     

Sustainable management of wildlife habitat and risk of extinction

Whether land management planning provides for sufficient habitat to sustain viable populations of indigenous wildlife is one of the greatest challenges confronting resource managers. Analyses of the effects of land management on natural resources often rely on qualitative assessments that focus on single species to reflect the risk of wildlife extinction across a planning area. We propose a conceptual framework for sustainable management of wildlife habitat that explicitly acknowledges the greater risk of an extinction event when considering the viability of multiple species, e.g., an indigenous vertebrate fauna. This concept is based on the principle that the likelihood of at least one event (i.e., species extinction) is the joint probability of the extinction probabilities of individual species, assuming independence among species’ responses to disturbance. We present an ecological rationale to support the view that, at a spatial scale of 10⁴-10⁶ ha (i.e., planning area) and a temporal scale of 10² years (i.e., planning horizon), wildlife species operating at varying ecological scales respond relatively independently to disturbances typically associated with land management. We use a hypothetical scenario of a wildlife viability assessment and Monte Carlo simulation to demonstrate that the probability of ‘any extinction’ is consistently higher than the probability of the ‘single most likely’ extinction, and that the difference between these values increases as more disturbance-sensitive species (i.e., species at risk) are analyzed. We conclude that risk assessments that rely upon the most sensitive single species may substantially underestimate the risk of wildlife extinction across a planning area. Furthermore, the selection of a planning alternative based on relative threat of local extinction of wildlife populations can vary depending on which paradigm is used to estimate risk to viability across the planning area.     

Map of bird sensitivities to wind farms in Scotland: A tool to aid planning and conservation

Government targets for renewable energy have led to a huge increase in wind farm proposals. Because of its high wind resource, Scotland has more proposed wind farms than any other UK country. Scotland’s upland habitats support many birds of conservation concern, leading to potential conflict with wind farms.To help reduce this conflict, a map of bird sensitivities has been created to guide the location of onshore wind farms in Scotland, based on distributions of 16 bird species of conservation priority and statutory Special Protection Areas. The likely sensitivity of each species to wind farms was assessed from literature, based on foraging ranges, collision risk and sensitivity to disturbance. This information was used to buffer species’ locations to identify areas of ‘high’ or ‘medium’ sensitivity. Individual species maps were converted to 1-km square resolution, and a composite map for all species created by selecting the highest sensitivity rating for each square.The map indicates greater bird sensitivity in northwest Scotland, particularly the Highlands, Western and Northern Isles. Overall, 37% of Scotland is classified as ‘high’, 25% as ‘medium’ and 38% as ‘low/unknown’ sensitivity. The overlap of the mapped species with proposed and existing wind farm developments was assessed and species for which cumulative effects of multiple wind farms are of particular concern identified. Within a Scottish context, bean goose, red kite and hen harrier showed the greatest overlap. Applications and limitations of the approach are discussed.     

Birds and green lanes: Breeding season bird abundance, territories and species richness

This study examines bird occurrence on 20 green lanes (farmland tracks with un-sealed surfaces, bordered on each side by hedgerows) and on 20 paired single hedgerows during the 2002 breeding season in Cheshire, UK. Measures of bird abundance, territories and species richness were all found to be significantly greater on green lanes than on single hedgerows. Significantly more birds were also found to occur on the inside rather than on the outside of green lanes in terms of abundance and species richness. All measures of bird occurrence on green lanes were influenced by hedge width and surrounding land use. Bird abundance on single hedgerows was influenced by the number of trees and amount of hawthorn in the hedge. Seventeen of the 18 most abundant bird species recorded in this study were found to occur in greater numbers on green lanes rather than in single hedgerows; for half of these species the difference was significant. The importance of green lanes for birds, and other wildlife is discussed, as well as their possible future within the UK landscape.     

Grassland nature reserves for breeding wading birds in England and the implications for the ESA agri-environment scheme

The two principal mechanisms available to conserve breeding wading birds on lowland wet grassland in England are nature reserves and agri-environment schemes, particularly the Environmentally Sensitive Area (ESA) scheme. ESAs offer payments to landowners in return for maintaining (‘low tiers’) or enhancing (‘high tiers’) the landscape, biodiversity and historic value of the land. Lowland wet grassland nature reserves managed by the Royal Society for the Protection of Birds (RSPB) have been successful at conserving breeding lapwings and redshank but habitat management usually only results in short-term increases in numbers of breeding snipe. Within ESAs, populations of breeding wading birds fare better on land managed under high tiers than on land managed under low tiers. Land managed under high tiers also supports more pairs of breeding wading birds per £ of ESA grant received than land managed under low tiers. Means of increasing the cost-effectiveness of the ESA scheme in conserving breeding wading birds are discussed in the light of these results.     

Partnering with local communities to identify conservation priorities for endangered Grevy’s zebra

Setting wildlife conservation priorities and determining how to meet them is challenging, particularly when policy decisions made at large scales need to be informed by a diversity of local conditions. The persistence of species that range widely demands that they coexist with people both within and outside formally protected areas. It is often politically and financially infeasible for one central body, such as a government wildlife agency, to monitor an entire population. Therefore, conservation planners are increasingly turning to local knowledge to inform conservation decisions. Here, we show the scientific and conservation benefits of recruiting and training local community members to collect data on an endangered species, the Grevy’s zebra (Equus grevyi). We recruited 18 scouts from six community-held ranches in Samburu District, Kenya. The scouts record the location, group structure and habitat of all Grevy’s zebra herds seen in walking surveys. Kernel analyses of scout herd observations indicate areas heavily used by Grevy’s zebra, and the subset of these areas favored by females with young foals. The important areas identified by the scouts closely match those inferred from analyses of GPS radiocollar data. Further, scout data reveals extensive spatial and temporal overlap between livestock and Grevy’s zebra. This overlap suggests the potential for competition between Grevy’s zebra and domestic animals. We argue that scout programs such as ours can generate valuable insights for conservation planning. In addition, such programs have the potential to improve local attitudes toward wildlife conservation.     

Micro-hotspot determination and buffer zone value for Odonata in a globally significant biosphere reserve

Reserves are frequently constrained in design and size by various financial, social or political factors. Maintenance of existing reserves must therefore rely on strategic management practices, and prioritization of conservation activities within them. Identification of global and regional hotspots have been effective for prioritizing conservation activities. Yet, identification of micro-hotspots, or overlapping areas of endemic and rare species that are under threat at the landscape scale, have largely been ignored. From a reserve management point of view, knowledge of critical micro-hotspots within a reserve, are focal points for directing cost effective, conservation initiatives, especially removal of invasive alien plants which are a major threat to biodiversity.Using diversity patterns of dragonfly assemblages, many endemic and threatened, within a biosphere reserve located in the core of a global biodiversity hotspot, we investigated the concept of micro-hotspots. As biosphere reserves contain zones with varying degrees of anthropogenic impact, we also investigated the value of buffer and transition zones for complementing the dragonfly fauna of the reserve core. We found a distinct micro-hotspot within the protected core zone which shows concordance for both endemism and species richness. We conclude that focused conservation actions to remove invasive alien plants within this micro-hotspot would help insure its continued integrity. Furthermore, while there is greater habitat degradation within the buffer and transition zones, they support many additional species, but not those necessarily endemic or threatened. The complementary value of buffer and transition zones therefore lies in increasing habitat heterogeneity and species richness of the whole reserve.     

Nature-based tourism as a form of predation risk affects body condition and health state of Podarcis muralis lizards

Nature-based tourism has experienced a greater increase, and, even although it might have deleterious effects for many wildlife species, its consequences remains little explored. Many lizard species are endangered and tourism has been proposed as a potential factor responsible of the decline of several lizards’ populations, but no study has examined the effect of tourism on lizards’ behaviour, body condition and health state. Many lizards respond to people as if they were predators, by readily escaping to refuges. However, an increase in the frequency of these antipredatory strategies can lead to a loss of body condition, which may have important consequences for short and long term fitness. We analysed the effects of tourism on escape behaviour of common wall lizards, Podarcis muralis, as well as on their body condition and health state (ectoparasites, blood parasites, and cell mediated immune response). Results showed that lizards did not modify their escape behaviour in response to tourism. Lizards had similar approach and flight distances, and escape strategies regardless of the level of tourism pressure. However, lizards inhabiting areas with high tourism levels, where they presumably needed to perform antipredatory behaviours more often, showed higher intensity of infection by ticks and lower body condition at the end of the breeding period. Moreover, lizards with poorer body condition had lower cell mediated immune responses. Therefore, tourism seems to have deleterious effects on body condition and on host-parasite relationships in this lizard species. These effects should be taking into account when designing walking paths in protected areas. Our study reports evidence that regardless lizards showed similar escape behaviour in tourist than in natural areas, their body condition and health state should be also examined to accurately assess the actual effects of tourism on lizards’ populations.