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The tourism sector is well-poised to generate win–win approaches to biodiversity conservation and sustainable development given its nonextractive nature and its dependence on scenic beauty (1). However, for sustainable tourism to succeed as a strategy for biodiversity conservation, the role that biodiversity plays in driving tourism patterns needs to be better understood. On one hand, wildlife and nature motivate a significant portion of global tourism (2), and protected areas with higher species richness tend to attract more tourists and yield higher economic benefits (3). On the other hand, tourism hotspots also tend to occur in places where more human-built infrastructure (e.g., hotels, roads, and airports) enables access (45). Studies have reached mixed conclusions on the relative importance of biodiversity and accessibility for tourism, and little is known about how they work in concert (611). Given the potential negative impacts of infrastructure on biodiversity conservation, their relative contributions to tourism deserves explicit study, particularly in developing countries where both biodiversity conservation and sustainable development are urgently needed (712).
Drivers of tourism patterns across landscapes have been explored through questionnaire surveys and structured interviews that ask tourists about their affinity for landscape features (1315) and through spatial models that predict recreation using photographs (e.g., geographically weighted regression, MaxEnt) (1617). Recently, geo-tagged photographs and species lists shared on social media platforms have become popular tools for tourism-focused research (1822). These studies, however, typically focus narrowly on the role of single taxa (2223) and landscape attributes without accounting for species diversity (1420), or they focus only on the role of infrastructure as a driver of tourism (5). Recent advances in satellite Earth observations make it possible to capture more of the ecosystem heterogeneity that can drive variability in species distributions, compared to more conventional modeling based on land cover (2426). An integrated approach is needed, linking species richness of multiple taxa along with infrastructure variables, both modeled and mapped through high resolution Earth observations. Such an approach could be scaled up to larger regions and applied globally, helping to identify where biodiversity is playing a significant role in driving tourism, such that governments and the tourism sector can prioritize investments in biodiversity conservation.
Here, we ask three questions in the iconic case of Costa Rica. First, what is the contribution of species richness (of vertebrate taxa) in explaining patterns of tourism in protected areas and also country-wide? Second, how similar are the patterns for birdwatching tourism compared to those of overall tourism? Third, where in the country is biodiversity contributing more than other factors to birdwatching tourism and to overall tourism? We predict that vertebrate species richness is more important for driving tourism in protected areas than in the rest of the country, because nature-seeking tourists often go to protected areas to find wildlife (3). We also expect that birdwatching tourism is predicted by richness of threatened and endemic bird species rather than total species richness, given birdwatchers’ preferences for rare birds (27). We predict a saturating relationship between species richness and tourism, because beyond a large number of species additional species are unlikely to contribute more to tourism (28). Finally, we predict that national-level tourism is better explained by infrastructure (such as roads and hotels) and distance to water than by biodiversity, because tourists going to Costa Rica often seek activities such as surfing and relaxing in beach resorts (2930). We predict nonlinear effects of proximity to roads and water, because a place is deemed inaccessible if it is further away from roads, and a beach tourist destination is also either close to water or not a destination at all. Access diminishes rapidly over a few miles (31).

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