Significance Tourism accounts for roughly 10% of global gross domestic product, with nature-based tourism its fastest-growing sector in the past 10 years. Nature-based tourism can theoretically contribute to local and sustainable development by creating attractive livelihoods that support biodiversity conservation, but whether tourists prefer to visit more biodiverse destinations is poorly understood. We examine this question in Costa Rica and find that more biodiverse places tend indeed to attract more tourists, especially where there is infrastructure that makes these places more accessible. Safeguarding terrestrial biodiversity is critical to preserving the substantial economic benefits that countries derive from tourism. Investments in both biodiversity conservation and infrastructure are needed to allow biodiverse countries to rely on tourism for their sustainable development.

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The collective behavior of ant colonies, and locomotion of individuals within a colony, both respond to changing conditions. The invasive Argentine ant (Linepithema humile) thrives in Mediterranean climates with hot, dry summers and colder, wet winters. However, its foraging behavior and locomotion has rarely been studied in the winter. We examined how the foraging behavior of three distinct L. humile colonies was related to environmental conditions and the locomotion of workers during winter in northern California. We found that colonies foraged most between 10 and 15°C, regardless of the maximum daily temperature. Worker walking speed was positively associated with temperature (range 6–24°C) and negatively associated with humidity (range 25–93%RH). All colonies foraged during all day and night hours in a predictable daily cycle, with a correlation between the rate of incoming and outgoing foragers. Foraging activity was unrelated to the activity of a competing native ant species, Prenolepis imparis, which was present in low abundance, and ceased only during heavy rain when ants left foraging trails and aggregated in small sheltered areas on trees.

Anthropogenic disturbances, including land use change and exotic species, can alter the diversity and dynamics of ant communities. To examine foraging behavior in an urbanized habitat in northern California, we surveyed the presence of 9 ant species on 876 trees across 4 seasons during both day and night in a 9.5-hectare urbanized oak-exotic woodland. Ants were more likely to be observed on native, evergreen trees, suggesting that native evergreen species may help maintain ant diversity. Species showed clear patterns of temporal partitioning of foraging activity. Ant species varied in their use of native evergreen Quercus agrifolia trees across season and day/night axes. Of the 3 ant species most frequently observed, Camponotus semitestaceus was most active during spring and summer nights, Formica moki was most active during spring and summer days, and Prenolepis imparis was most active during both day and night during fall and winter. Liometopum occidentale was the second most active species during summer day and night, and winter day. Our findings demonstrate that an oak-exotic urban woodland in Northern California was able to maintain a native ant community, and strong temporal partitioning within that community.

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Spatial and temporal partitioning of habitat may facilitate diversity and have important impacts 10 on ant communities. To investigate niche overlap in an ant community in a northern California oak 11 woodland, we observed ant foraging on trees in 4 seasonal surveys, each lasting 2 weeks, in a 9.5-hectare 12 plot over the course of a year. Foraging activity in all 5 observed ant species differed by season, time of 13 day, and/or the genera of trees used. Of the 3 ant species most frequently observed, Camponotus 14 semitestaceus was most active during spring and summer nights, Formica moki was most active during 15 spring and summer days, and Prenolepis imparis was most active during both day and night of fall and 16 winter. All ant species preferred native trees to exotic trees and preferred evergreen trees to deciduous 17 trees. Our results suggest that native evergreen oaks such as Quercus agrifolia, currently threatened by 18 sudden oak death (Phytophthora ramorum), may be important for supporting ant biodiversity. 19 20 Acknowledgements for many hours of field work assistance. We thank Rodolfo 25 Dirzo, Daniel Friedman and Maria Wojakowski for providing advice and assistance with statistical 2 26 analysis. Leander D.L. Anderegg, Talia Borofsky, Kaleda Denton, and Tyler McFadden provided helpful 27 comments on the manuscript. The Stanford Maps and Records Office (340 Bonair Siding, Stanford, CA 28 94305) provided detailed maps of trees on the Stanford grounds that were invaluable for this project. 29 30