Evaluating expert-based habitat suitability information of terrestrial mammals with GPS-tracking data.

Aim: Macroecological studies that require habitat suitability data for many species often derive this information from expert opinion. However, expert-based information is inherently subjective and thus prone to errors. The increasing availability of GPS tracking data offers opportunities to evaluate and supplement expert-based information with detailed empirical evidence. Here, we compared expert-based habitat suitability information from the International Union for Conservation of Nature (IUCN) with habitat suitability information derived from GPS-tracking data of 1,498 individuals from 49 mammal species. Location: Worldwide. Time period: 1998-2021. Major taxa studied: Forty-nine terrestrial mammal species. Methods: Using GPS data, we estimated two measures of habitat suitability for each individual animal: proportional habitat use (proportion of GPS locations within a habitat type), and selection ratio (habitat use relative to its availability). For each individual we then evaluated whether the GPS-based habitat suitability measures were in agreement with the IUCN data. To that end, we calculated the probability that the ranking of empirical habitat suitability measures was in agreement with IUCN's classification into suitable, marginal and unsuitable habitat types. Results: IUCN habitat suitability data were in accordance with the GPS data (> 95% probability of agreement) for 33 out of 49 species based on proportional habitat use estimates and for 25 out of 49 species based on selection ratios. In addition, 37 and 34 species had a > 50% probability of agreement based on proportional habitat use and selection ratios, respectively. Main conclusions: We show how GPS-tracking data can be used to evaluate IUCN habitat suitability data. Our findings indicate that for the majority of species included in this study, it is appropriate to use IUCN habitat suitability data in macroecological studies. Furthermore, we show that GPS-tracking data can be used to identify and prioritize species and habitat types for re-evaluation of IUCN habitat suitability data.

Collaborative agent-based modeling for managing shrub encroachment in an Afroalpine grassland.

We co-designed an agent-based model of an Afroalpine grassland in Ethiopia that is experiencing unwanted shrub encroachment. The goal was to enable managers of a community conservation area to better understand the drivers of shrub encroachment and to test possible management actions for controlling shrubs. Due to limited site-specific data, we parameterized this model using insights from published literature, remote sensing, and expert opinion from scientists and local managers. We therefore sought to explore potential future scenarios rather than make highly accurate predictions, focusing on facilitating discussions and learning among the diverse co-management team. We evaluated three social-ecological scenarios with our model, examining: (1) the impact of changing precipitation regimes on vegetation, (2) whether changing the frequency of guassa grass harvests would improve the long-term sustainability of the grassland, and (3) whether the combination of grass harvest and shrub removal would affect shrub encroachment. We found that the model was highly sensitive to the amount of grass harvested each year for local use. Our results indicate that the guassa grass was more resilient than shrubs during persistent dry climatic conditions, whereas a reduction in only the early spring rains (known as the "belg") resulted in considerable loss of grass biomass. While our modeling results lacked the quantitative specificity desired by managers, participants in the collaborative modeling process learned new approaches to planning and management of the conservation area and expanded their knowledge of the ecological complexity of the system. Several participants used the model as a boundary object, interpreting it in ways that reinforced their cultural values and goals for the conservation area. Our work highlights the lack of detailed scientific knowledge of Afroalpine ecosystems, and urges managers to reconnect with traditional ecological management of the conservation area in their pursuit of shrub encroachment solutions. The decline or absence of the belg rains is becoming increasingly common in the Ethiopian highlands, and our results underscore the need for more widespread understanding of how this changing climatic regime impacts local environmental management. This work lays a foundation for social-ecological research to improve both understanding and management of these highly threatened ecosystems.

Body size and digestive system shape resource selection by ungulates: A cross-taxa test of the forage maturation hypothesis.

The forage maturation hypothesis (FMH) states that energy intake for ungulates is maximised when forage biomass is at intermediate levels. Nevertheless, metabolic allometry and different digestive systems suggest that resource selection should vary across ungulate species. By combining GPS relocations with remotely sensed data on forage characteristics and surface water, we quantified the effect of body size and digestive system in determining movements of 30 populations of hindgut fermenters (equids) and ruminants across biomes. Selection for intermediate forage biomass was negatively related to body size, regardless of digestive system. Selection for proximity to surface water was stronger for equids relative to ruminants, regardless of body size. To be more generalisable, we suggest that the FMH explicitly incorporate contingencies in body size and digestive system, with small-bodied ruminants selecting more strongly for potential energy intake, and hindgut fermenters selecting more strongly for surface water.

Drivers of site fidelity in ungulates.

While the tendency to return to previously visited locations-termed 'site fidelity'-is common in animals, the cause of this behaviour is not well understood. One hypothesis is that site fidelity is shaped by an animal's environment, such that animals living in landscapes with predictable resources have stronger site fidelity. Site fidelity may also be conditional on the success of animals' recent visits to that location, and it may become stronger with age as the animal accumulates experience in their landscape. Finally, differences between species, such as the way memory shapes site attractiveness, may interact with environmental drivers to modulate the strength of site fidelity. We compared inter-year site fidelity in 669 individuals across eight ungulate species fitted with GPS collars and occupying a range of environmental conditions in North America and Africa. We used a distance-based index of site fidelity and tested hypothesized drivers of site fidelity using linear mixed effects models, while accounting for variation in annual range size. Mule deer Odocoileus hemionus and moose Alces alces exhibited relatively strong site fidelity, while wildebeest Connochaetes taurinus and barren-ground caribou Rangifer tarandus granti had relatively weak fidelity. Site fidelity was strongest in predictable landscapes where vegetative greening occurred at regular intervals over time (i.e. high temporal contingency). Species differed in their response to spatial heterogeneity in greenness (i.e. spatial constancy). Site fidelity varied seasonally in some species, but remained constant over time in others. Elk employed a 'win-stay, lose-switch' strategy, in which successful resource tracking in the springtime resulted in strong site fidelity the following spring. Site fidelity did not vary with age in any species tested. Our results provide support for the environmental hypothesis, particularly that regularity in vegetative phenology shapes the strength of site fidelity at the inter-annual scale. Large unexplained differences in site fidelity suggest that other factors, possibly species-specific differences in attraction to known sites, contribute to variation in the expression of this behaviour. Understanding drivers of variation in site fidelity across groups of organisms living in different environments provides important behavioural context for predicting how animals will respond to environmental change.

Hierarchical global plant biophysical regions as potential analysis units.

Regional and global vegetation simulations can be problematic when analysis units to which parameters are assigned do not align with plant productivity and phenology. Having a suite of predefined biophysical regions at a variety of scales that correspond to differences in plant productivity and phenology would allow analysts to select a set of analysis units at the scale needed. In other cases, environmental or social responses may be hypothesized to be related to differences in plant dynamics. One may compare the discrimination in such data that biophysical regions at different scales provide to determine which best distinguishes the responses in question, such that like responses fall within the same regions to the degree possible. If those relationships are significant, the responses may then be extrapolated based on the biophysical regions. I defined hierarchical biophysical regions based on plant productivity and phenology by clustering global 0.083 degree resolution normalized difference vegetation indices (NDVI) over a 10 year period. Agglomerative average-linkage distances based on squared error between clusters were conducted using an iterative sampling approach to merge more than 2 million clusters into fewer and fewer clusters based on NDVI greenness profiles comprised of 240 values over 10 years, until all cells were in a single cluster. Greater and greater differences in greenness profiles were ignored at higher levels of the hierarchy. Using a difference increment of 0.1, 253 non-duplicative sets of clusters were created, and 107 of those were included in animations that may be used to explore differences in global plant dynamics. Differences in clusters were quantified based on comparing the focal set of cluster results with 10 other cluster sets. Analysts may use the hierarchical clusters to improve the alignment of their parameter sets that inform plant growth and other dynamics with real-world plant dynamics.

Effects of body size on estimation of mammalian area requirements.

Accurately quantifying species' area requirements is a prerequisite for effective area-based conservation. This typically involves collecting tracking data on species of interest and then conducting home-range analyses. Problematically, autocorrelation in tracking data can result in space needs being severely underestimated. Based on the previous work, we hypothesized the magnitude of underestimation varies with body mass, a relationship that could have serious conservation implications. To evaluate this hypothesis for terrestrial mammals, we estimated home-range areas with global positioning system (GPS) locations from 757 individuals across 61 globally distributed mammalian species with body masses ranging from 0.4 to 4000 kg. We then applied block cross-validation to quantify bias in empirical home-range estimates. Area requirements of mammals <10 kg were underestimated by a mean approximately15%, and species weighing approximately100 kg were underestimated by approximately50% on average. Thus, we found area estimation was subject to autocorrelation-induced bias that was worse for large species. Combined with the fact that extinction risk increases as body mass increases, the allometric scaling of bias we observed suggests the most threatened species are also likely to be those with the least accurate home-range estimates. As a correction, we tested whether data thinning or autocorrelation-informed home-range estimation minimized the scaling effect of autocorrelation on area estimates. Data thinning required an approximately93% data loss to achieve statistical independence with 95% confidence and was, therefore, not a viable solution. In contrast, autocorrelation-informed home-range estimation resulted in consistently accurate estimates irrespective of mass. When relating body mass to home range size, we detected that correcting for autocorrelation resulted in a scaling exponent significantly >1, meaning the scaling of the relationship changed substantially at the upper end of the mass spectrum.

Efectos del Tamaño Corporal sobre la Estimación de los Requerimientos de Área de Mamíferos Resumen La cuantificación precisa de los requerimientos de área de una especie es un prerrequisito para que la conservación basada en áreas sea efectiva. Esto comúnmente implica la recolección de datos de rastreo de la especie de interés para después realizar análisis de la distribución local. De manera problemática, la autocorrelación en los datos de rastreo puede resultar en una subestimación grave de las necesidades de espacio. Con base en trabajos previos, formulamos una hipótesis en la que supusimos que la magnitud de la subestimación varía con la masa corporal, una relación que podría tener implicaciones serias para la conservación. Para probar esta hipótesis en mamíferos terrestres, estimamos las áreas de distribución local con las ubicaciones en GPS de 757 individuos de 61 especies de mamíferos distribuidas mundialmente con una masa corporal entre 0.4 y 4,000 kg. Después aplicamos una validación cruzada en bloque para cuantificar el sesgo en estimaciones empíricas de la distribución local. Los requerimientos de área de los mamíferos <10 kg fueron subestimados por una media ∼15% y las especies con una masa ∼100 kg fueron subestimadas en ∼50% en promedio. Por lo tanto, encontramos que la estimación del área estaba sujeta al sesgo inducido por la autocorrelación, el cual era peor para las especies de talla grande. En combinación con el hecho de que el riesgo de extinción incrementa conforme aumenta la masa corporal, el escalamiento alométrico del sesgo que observamos sugiere que la mayoría de las especies amenazadas también tienen la probabilidad de ser aquellas especies con las estimaciones de distribución local menos acertadas. Como corrección, probamos si la reducción de datos o la estimación de la distribución local informada por la autocorrelación minimizan el efecto de escalamiento que tiene la autocorrelación sobre las estimaciones de área. La reducción de datos requirió una pérdida de datos del ∼93% para lograr la independencia estadística con un 95% de confianza y por lo tanto no fue una solución viable. Al contrario, la estimación de la distribución local informada por la autocorrelación resultó en estimaciones constantemente precisas sin importar la masa corporal. Cuando relacionamos la masa corporal con el tamaño de la distribución local, detectamos que la corrección de la autocorrelación resultó en un exponente de escalamiento significativamente >1, lo que significa que el escalamiento de la relación cambió sustancialmente en el extremo superior del espectro de la masa corporal.

Climate change impacts on selected global rangeland ecosystem services.

Rangelands are Earth's dominant land cover and are important providers of ecosystem services. Reliance on rangelands is projected to grow, thus understanding the sensitivity of rangelands to future climates is essential. We used a new ecosystem model of moderate complexity that allows, for the first time, to quantify global changes expected in rangelands under future climates. The mean global annual net primary production (NPP) may decline by 10 g C m-2  year-1 in 2050 under Representative Concentration Pathway (RCP) 8.5, but herbaceous NPP is projected to increase slightly (i.e., average of 3 g C m-2  year-1 ). Responses vary substantially from place-to-place, with large increases in annual productivity projected in northern regions (e.g., a 21% increase in productivity in the US and Canada) and large declines in western Africa (-46% in sub-Saharan western Africa) and Australia (-17%). Soil organic carbon is projected to increase in Australia (9%), the Middle East (14%), and central Asia (16%) and decline in many African savannas (e.g., -18% in sub-Saharan western Africa). Livestock are projected to decline 7.5 to 9.6%, an economic loss of from $9.7 to $12.6 billion. Our results suggest that forage production in Africa is sensitive to changes in climate, which will have substantial impacts on the livelihoods of the more than 180 million people who raise livestock on those rangelands. Our approach and the simulation tool presented here offer considerable potential for forecasting future conditions, highlight regions of concern, and support analyses where costs and benefits of adaptations and policies may be quantified. Otherwise, the technical options and policy and enabling environment that are needed to facilitate widespread adaptation may be very difficult to elucidate.

Hunger mediates apex predator's risk avoidance response in wildland-urban interface.

Conflicts between large mammalian predators and humans present a challenge to conservation efforts, as these events drive human attitudes and policies concerning predator species. Unfortunately, generalities portrayed in many empirical carnivore landscape selection studies do not provide an explanation for a predator's occasional use of residential development preceding a carnivore-human conflict event. In some cases, predators may perceive residential development as a risk-reward trade-off. We examine whether state-dependent mortality risk-sensitive foraging can explain an apex carnivore's (Puma concolor) occasional utilization of residential areas. We assess whether puma balance the risk and rewards in a system characterized by a gradient of housing densities ranging from wildland to suburban. Puma GPS location data, characterized as hunting and feeding locations, were used to assess landscape variables governing hunting success and hunting site selection. Hunting site selection behaviour was then analysed conditional on indicators of hunger state. Residential development provided a high energetic reward to puma based on increases in prey availability and hunting success rates associated with increased housing density. Despite a higher energetic reward, hunting site selection analysis indicated that pumas generally avoided residential development, a landscape type attributed with higher puma mortality risk. However, when a puma experienced periods of extended hunger, risk avoidance behaviour towards housing waned. This study demonstrates that an apex carnivore faces a trade-off between acquiring energetic rewards and avoiding risks associated with human housing. Periods of hunger can help explain an apex predator's occasional use of developed landscapes and thus the rare conflicts in the wildland-urban interface. Apex carnivore movement behaviours in relation to human conflicts are best understood as a three-player community-level interaction incorporating wild prey distribution.

Vegetation redistribution is predicted to intensify soil organic carbon loss under future climate changes on the Tibetan Plateau.

Vegetation redistribution may bring unexpected climate-soil carbon cycling in terrestrial biomes. However, whether and how vegetation redistribution alters the soil carbon pool under climate change is still poorly understood on the Tibetan Plateau. Here, we applied the G-Range model to simulate the cover of herbs, shrubs and trees, net primary productivity (NPP) and soil organic carbon density (SOCD) at the depth of 60 cm on Tibetan Plateau for the individual years 2020 and 2060, using climate projection for Representative Concentration Pathways (RCP) 4.5 and RCP8.5 scenarios with the RegCM4.6 model system. Vegetation redistribution was defined as the transitions in bare ground, herbs, shrubs and trees between 2020 and 2060, with approximately 57.9 % (RCP4.5) and 59 % (RCP8.5) of the area will redistribute vegetation over the whole Tibetan Plateau. The vegetation cover will increase by about 2.4 % (RCP4.5) and 1.9 % (RCP8.5), while the NPP and SOCD will decrease by about -14.3 g C m-2 yr-1 and -907 g C m-2 (RCP4.5), and -1.8 g C m-2 yr-1and -920 g C m-2 (RCP8.5). Shrubs and trees will expand in the east, and herbs will expand in the northwest part of the Plateau. These areas are projected to be hotspots with greater SOCD reduction in response to future climate change, and will include lower net plant carbon input due to the negative NPP. Our study indicates that the SOC pool will become a carbon source under increased air temperature and rainfall on the Tibetan Plateau by 2060, especially for the area with vegetation redistribution. These results revealed the potential risk of vegetation redistribution under climate change in alpine ecosystems, indicating the policymakers need to pay attention on the vegetation redistribution to mitigate the soil carbon emission and achieve the goal of carbon neutrality on the Tibetan Plateau.

Long-term historical and projected herbivore population dynamics in Ngorongoro crater, Tanzania.

The Ngorongoro Crater is an intact caldera with an area of approximately 310 km2 located within the Ngorongoro Conservation Area (NCA) in northern Tanzania. It is known for the abundance and diversity of its wildlife and is a UNESCO World Heritage Site and an International Biosphere Reserve. Long term records (1963-2012) on herbivore populations, vegetation and rainfall made it possible to analyze historic and project future herbivore population dynamics. NCA was established as a multiple use area in 1959. In 1974 there was a perturbation in that resident Maasai and their livestock were removed from the Ngorongoro Crater. Thus, their pasture management that was a combination of livestock grazing and fire was also removed and 'burning' stopped being a regular occurrence until it was resumed in 2001 by NCA management. The Maasai pasture management would have selected for shorter grasses and more palatable species. Vegetation mapping in 1966-1967 recorded predominately short grasslands. Subsequent vegetation mapping in the crater in 1995 determined that the grassland structure had changed such that mid and tall grasses were dominant. After removal of the Maasai pastoralists from the Ngorongoro Crater in 1974, there were significant changes in population trends for some herbivore species. Buffalo, elephant and ostrich numbers increased significantly during 1974-2012. The zebra population was stable from 1963 to 2012 whereas population numbers of five species declined substantially between 1974 and 2012 relative to their peak numbers during 1974-1976. Grant's and Thomson's gazelles, eland, kongoni, and waterbuck (wet season only) declined significantly in the Crater in both seasons after 1974. In addition, some herbivore species were consistently more abundant inside the Crater during the wet than the dry season. This pattern was most evident for the large herbivore species requiring bulk forage, i.e., buffalo, eland, and elephant. Even with a change in grassland structure, total herbivore biomass remained relatively stable from 1963 to 2012, implying that the crater has a stable carrying capacity. Analyses of rainfall indicated that there was a persistent cycle of 4.83 years for the annual component. Herbivore population size was correlated with rainfall in both the wet and dry seasons. The relationships established between the time series of historic animal counts in the wet and dry seasons and lagged wet and dry season rainfall series were used to forecast the likely future trajectories of the wet and dry season population size for each species under three alternative climate change scenarios.

An agent-based model of cattle grazing toxic Geyer's larkspur.

By killing cattle and otherwise complicating management, the many species of larkspur (Delphinium spp.) present a serious, intractable, and complex challenge to livestock grazing management in the western United States. Among the many obstacles to improving our understanding of cattle-larkspur dynamics has been the difficulty of testing different grazing management strategies in the field, as the risk of dead animals is too great. Agent-based models (ABMs) provide an effective method of testing alternate management strategies without risk to livestock. ABMs are especially useful for modeling complex systems such as livestock grazing management, and allow for realistic bottom-up encoding of cattle behavior. Here, we introduce a spatially-explicit, behavior-based ABM of cattle grazing in a pasture with a dangerous amount of Geyer's larkspur (D. geyeri). This model tests the role of herd cohesion and stocking density in larkspur intake, finds that both are key drivers of larkspur-induced toxicosis, and indicates that alteration of these factors within realistic bounds can mitigate risk. Crucially, the model points to herd cohesion, which has received little attention in the discipline, as playing an important role in lethal acute toxicosis. As the first ABM to model grazing behavior at realistic scales, this study also demonstrates the tremendous potential of ABMs to illuminate grazing management dynamics, including fundamental aspects of livestock behavior amidst ecological heterogeneity.

Evolutionary computation in zoology and ecology.

Evolutionary computational methods have adopted attributes of natural selection and evolution to solve problems in computer science, engineering, and other fields. The method is growing in use in zoology and ecology. Evolutionary principles may be merged with an agent-based modeling perspective to have individual animals or other agents compete. Four main categories are discussed: genetic algorithms, evolutionary programming, genetic programming, and evolutionary strategies. In evolutionary computation, a population is represented in a way that allows for an objective function to be assessed that is relevant to the problem of interest. The poorest performing members are removed from the population, and remaining members reproduce and may be mutated. The fitness of the members is again assessed, and the cycle continues until a stopping condition is met. Case studies include optimizing: egg shape given different clutch sizes, mate selection, migration of wildebeest, birds, and elk, vulture foraging behavior, algal bloom prediction, and species richness given energy constraints. Other case studies simulate the evolution of species and a means to project shifts in species ranges in response to a changing climate that includes competition and phenotypic plasticity. This introduction concludes by citing other uses of evolutionary computation and a review of the flexibility of the methods. For example, representing species' niche spaces subject to selective pressure allows studies on cladistics, the taxon cycle, neutral versus niche paradigms, fundamental versus realized niches, community structure and order of colonization, invasiveness, and responses to a changing climate.

Serengeti wildebeest migratory patterns modeled from rainfall and new vegetation growth.

We used evolutionary programming to model innate migratory pathways of wildebeest in the Serengeti Mara Ecosystem, Tanzania and Kenya. Wildebeest annually move from the southern short-grass plains of the Serengeti to the northern woodlands of the Mara. We used satellite images to create 12 average monthly and 180 10-day surfaces from 1998 to 2003 of percentage rainfall and new vegetation. The surfaces were combined in five additive and three multiplicative models, with the weightings on rainfall and new vegetation from 0% to 100%. Modeled wildebeest were first assigned random migration pathways. In simulated generations, animals best able to access rainfall and vegetation were retained, and they produced offspring with similar migratory pathways. Modeling proceeded until the best pathway was stable. In a learning phase, modeling continued with the ten-day images in the objective function. The additive model, influenced 25% by rainfall and 75% by vegetation growth, yielded the best agreement, with a multi-resolution comparison to observed densities yielding 76.8% of blocks in agreement (kappa = 0.32). Agreement was best for dry season and early wet season (kappa = 0.22-0.57), and poorest for the late wet season (0.04). The model suggests that new forage growth is a dominant correlate of wildebeest migration.

Spatial and temporal variability modify density dependence in populations of large herbivores.

A central challenge in ecology is to understand the interplay of internal and external controls on the growth of populations. We examined the effects of temporal variation in weather and spatial variation in vegetation on the strength of density dependence in populations of large herbivores. We fit three subsets of the model ln(Nt) = a + (1 + b) x ln(N(t-1)) + c x ln(N(t-2)) to five time series of estimates (Nt) of abundance of ungulates in the Rocky Mountains, USA. The strength of density dependence was estimated by the magnitude of the coefficient b. We regressed the estimates of b on indices of temporal heterogeneity in weather and spatial heterogeneity in resources. The 95% posterior intervals of the slopes of these regressions showed that temporal heterogeneity strengthened density-dependent feedbacks to population growth, whereas spatial heterogeneity weakened them. This finding offers the first empirical evidence that density dependence responds in different ways to spatial heterogeneity and temporal heterogeneity.

Nutritional Status of Maasai Pastoralists under Change.

This study assesses the nutritional status of Maasai pastoralists living in a period of great social, economic and ecological changes in Kajiado County, southern Kenya. Data on weight, height, skinfolds, and circumferences were collected from 534 individuals in the year 2000. The data were used to describe mean differences in human nutrition between ages, sexes, and within and among three Group Ranches. Nutritional data and diet recall data were compared with past studies of Maasai nutrition from 1930 to 2000. Results indicate that nutritional status is poor and has remained so despite numerous changes to the social-ecological system including livelihood diversification, sedentarization, human population growth and decreased access to vegetation heterogeneity. Imbirikani Group Ranch had better access to infrastructure and markets and some measures of nutritional status were better than for individuals in other group ranches. However, nutritional status remains poor despite transitioning to greater market integration.

Influence of aesthetic appreciation of wildlife species on attitudes towards their conservation in Kenyan agropastoralist communities.

The influence of human aesthetic appreciation of animal species on public attitudes towards their conservation and related decision-making has been studied in industrialized countries but remains underexplored in developing countries. Working in three agropastoralist communities around Amboseli National Park, southern Kenya, we investigated the relative strength of human aesthetic appreciation on local attitudes towards the conservation of wildlife species. Using semi-structured interviewing and free listing (n = 191) as part of a mixed methods approach, we first characterized local aesthetic judgments of wildlife species. With a Generalized Linear Mixed Models (GLMM) approach, we then determined the influence of perceiving four species as beautiful on local support for their protection ("rescuing them"), and of perceiving four other species as ugly on support for their removal from the area, while controlling for informant personal and household socioeconomic attributes. Perceiving giraffe, gazelles and eland as beautiful is the strongest variable explaining support for rescuing them. Ugliness is the strongest variable influencing support for the removal of buffalo, hyena, and elephant (but not lion). Both our qualitative and quantitative results suggest that perceptions of ugly species could become more positive through direct exposure to those species. We propose that protected areas in developing countries facilitate visitation by local residents to increase their familiarity with species they rarely see or most frequently see in conflict with human interests. Since valuing a species for its beauty requires seeing it, protected areas in developing countries should connect the people who live around them with the animals they protect. Our results also show that aesthetic appreciation of biodiversity is not restricted to the industrialized world.