Biodiversity and infrastructure interact to drive tourism to and within Costa Rica.

SignificanceTourism 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.

Protection promotes energetically efficient structures in marine communities.

The sustainability of marine communities is critical for supporting many biophysical processes that provide ecosystem services that promote human well-being. It is expected that anthropogenic disturbances such as climate change and human activities will tend to create less energetically-efficient ecosystems that support less biomass per unit energy flow. It is debated, however, whether this expected development should translate into bottom-heavy (with small basal species being the most abundant) or top-heavy communities (where more biomass is supported at higher trophic levels with species having larger body sizes). Here, we combine ecological theory and empirical data to demonstrate that full marine protection promotes shifts towards top-heavy energetically-efficient structures in marine communities. First, we use metabolic scaling theory to show that protected communities are expected to display stronger top-heavy structures than disturbed communities. Similarly, we show theoretically that communities with high energy transfer efficiency display stronger top-heavy structures than communities with low transfer efficiency. Next, we use empirical structures observed within fully protected marine areas compared to disturbed areas that vary in stress from thermal events and adjacent human activity. Using a nonparametric causal-inference analysis, we find a strong, positive, causal effect between full marine protection and stronger top-heavy structures. Our work corroborates ecological theory on community development and provides a quantitative framework to study the potential restorative effects of different candidate strategies on protected areas.

An ecosystem service perspective on urban nature, physical activity, and health.

Nature underpins human well-being in critical ways, especially in health. Nature provides pollination of nutritious crops, purification of drinking water, protection from floods, and climate security, among other well-studied health benefits. A crucial, yet challenging, research frontier is clarifying how nature promotes physical activity for its many mental and physical health benefits, particularly in densely populated cities with scarce and dwindling access to nature. Here we frame this frontier by conceptually developing a spatial decision-support tool that shows where, how, and for whom urban nature promotes physical activity, to inform urban greening efforts and broader health assessments. We synthesize what is known, present a model framework, and detail the model steps and data needs that can yield generalizable spatial models and an effective tool for assessing the urban nature-physical activity relationship. Current knowledge supports an initial model that can distinguish broad trends and enrich urban planning, spatial policy, and public health decisions. New, iterative research and application will reveal the importance of different types of urban nature, the different subpopulations who will benefit from it, and nature's potential contribution to creating more equitable, green, livable cities with active inhabitants.

Uses and Limitations of Social Media to Inform Visitor Use Management in Parks and Protected Areas: A Systematic Review.

Social media are being increasingly used to inform visitor use management in parks and protected areas. We review the state of the scientific literature to understand the ways social media has been, and can be, used to measure visitation, spatial patterns of use, and visitors' experiences in parks and protected areas. Geotagged social media are a good proxy for actual visitation; however, the correlations observed by previous studies between social media and other sources of visitation data vary substantially. Most studies using social media to measure visitation aggregate data across many years, with very few testing the use of social media as a visitation proxy at smaller temporal scales. No studies have tested the use of social media to estimate visitation in near real-time. Studies have used geotags and GPS tracks to understand spatial patterns of where visitors travel within parks, and how that may relate to other variables (e.g., infrastructure), or differ by visitor type. Researchers have also found the text content, photograph content, and geotags from social media posts useful to understand aspects of visitors' experiences, such as behaviors, preferences, and sentiment. The most cited concern with using social media is that this data may not be representative of all park users. Collectively, this body of research demonstrates a broad range of applications for social media. We synthesize our findings by identifying gaps and opportunities for future research and presenting a set of best practices for using social media in parks and protected areas.

Embedding ecosystem services in coastal planning leads to better outcomes for people and nature.

Recent calls for ocean planning envision informed management of social and ecological systems to sustain delivery of ecosystem services to people. However, until now, no coastal and marine planning process has applied an ecosystem-services framework to understand how human activities affect the flow of benefits, to create scenarios, and to design a management plan. We developed models that quantify services provided by corals, mangroves, and seagrasses. We used these models within an extensive engagement process to design a national spatial plan for Belize's coastal zone. Through iteration of modeling and stakeholder engagement, we developed a preferred plan, currently under formal consideration by the Belizean government. Our results suggest that the preferred plan will lead to greater returns from coastal protection and tourism than outcomes from scenarios oriented toward achieving either conservation or development goals. The plan will also reduce impacts to coastal habitat and increase revenues from lobster fishing relative to current management. By accounting for spatial variation in the impacts of coastal and ocean activities on benefits that ecosystems provide to people, our models allowed stakeholders and policymakers to refine zones of human use. The final version of the preferred plan improved expected coastal protection by >25% and more than doubled the revenue from fishing, compared with earlier versions based on stakeholder preferences alone. Including outcomes in terms of ecosystem-service supply and value allowed for explicit consideration of multiple benefits from oceans and coasts that typically are evaluated separately in management decisions.

Recreational use in dispersed public lands measured using social media data and on-site counts.

Outdoor recreation is one of many important benefits provided by public lands. Data on recreational use are critical for informing management of recreation resources, however, managers often lack actionable information on visitor use for large protected areas that lack controlled access points. The purpose of this study is to explore the potential for social media data (e.g., geotagged images shared on Flickr and trip reports shared on a hiking forum) to provide land managers with useful measures of recreational use to dispersed areas, and to provide lessons learned from comparing several more traditional counting methods. First, we measure daily and monthly visitation rates to individual trails within the Mount Baker-Snoqualmie National Forest (MBSNF) in western Washington. At 15 trailheads, we compare counts of hikers from infrared sensors, timelapse cameras, and manual on-site counts, to counts based on the number of shared geotagged images and trip reports from those locations. Second, we measure visitation rates to each National Forest System (NFS) unit across the US and compare annual measurements derived from the number of geotagged images to estimates from the US Forest Service National Visitor Use Monitoring Program. At both the NFS unit and the individual-trail scales, we found strong correlations between traditional measures of recreational use and measures based on user-generated content shared on the internet. For national forests in every region of the country, correlations between official Forest Service statistics and geotagged images ranged between 55% and 95%. For individual trails within the MBSNF, monthly visitor counts from on-site measurements were strongly correlated with counts from geotagged images (79%) and trip reports (91%). The convenient, cost-efficient and timely nature of collecting and analyzing user-generated data could allow land managers to monitor use over different seasons of the year and at sites and scales never previously monitored, contributing to a more comprehensive understanding of recreational use patterns and values.

Measuring recreational visitation at U.S. National Parks with crowd-sourced photographs.

Land managers rely on visitation data to inform policy and management decisions. However, visitation data is often costly and burdensome to obtain, and provides a limited depth of information. In this paper, we assess the validity of using crowd-sourced, online photographs to infer information about the habits and preferences of recreational visitors by comparing empirical data from the National Park Service to photograph data from the online platform Flickr for 38 National Parks in the western United States. Using multiple regression analysis, we find that the number of photos posted monthly in a park can reliably indicate the number of visitors to a park in a given month. Through additional statistical testing we also find that the home locations of photo-takers, provided voluntarily on an online profile, accurately show the home origins of park visitors. Together, these findings validate a new method for measuring recreational visitation, opening an opportunity for land managers worldwide to track and understand visitation by augmenting current data collection methods with crowd-sourced, online data that is easy and inexpensive to obtain. In addition, it enables future research on how visitation rates change with changes in access, management or infrastructure, weather events, or ecosystem health, and facilitates valuation research, such as travel cost studies.

Network structure beyond food webs: mapping non-trophic and trophic interactions on Chilean rocky shores.

How multiple types of non-trophic interactions map onto trophic networks in real communities remains largely unknown. We present the first effort, to our knowledge, describing a comprehensive ecological network that includes all known trophic and diverse non-trophic links among >100 coexisting species for the marine rocky intertidal community of the central Chilean coast. Our results suggest that non-trophic interactions exhibit highly nonrandom structures both alone and with respect to food web structure. The occurrence of different types of interactions, relative to all possible links, was well predicted by trophic structure and simple traits of the source and target species. In this community, competition for space and positive interactions related to habitat/refuge provisioning by sessile and/or basal species were by far the most abundant non-trophic interactions. If these patterns are orroborated in other ecosystems, they may suggest potentially important dynamic constraints on the combined architecture of trophic and non-trophic interactions. The nonrandom patterning of non-trophic interactions suggests a path forward for developing a more comprehensive ecological network theory to predict the functioning and resilience of ecological communities.

More than a meal integrating non-feeding interactions into food webs.

Organisms eating each other are only one of many types of well documented and important interactions among species. Other such types include habitat modification, predator interference and facilitation. However, ecological network research has been typically limited to either pure food webs or to networks of only a few (<3) interaction types. The great diversity of non-trophic interactions observed in nature has been poorly addressed by ecologists and largely excluded from network theory. Herein, we propose a conceptual framework that organises this diversity into three main functional classes defined by how they modify specific parameters in a dynamic food web model. This approach provides a path forward for incorporating non-trophic interactions in traditional food web models and offers a new perspective on tackling ecological complexity that should stimulate both theoretical and empirical approaches to understanding the patterns and dynamics of diverse species interactions in nature.

Conceptualizing ecosystem services using social-ecological networks.

Social-ecological networks (SENs) represent the complex relationships between ecological and social systems and are a useful tool for analyzing and managing ecosystem services. However, mainstreaming the application of SENs in ecosystem service research has been hindered by a lack of clarity about how to match research questions to ecosystem service conceptualizations in SEN (i.e., as nodes, links, attributes, or emergent properties). Building from different disciplines, we propose a typology to represent ecosystem service in SENs and identify opportunities and challenges of using SENs in ecosystem service research. Our typology provides guidance for this growing field to improve research design and increase the breadth of questions that can be addressed with SEN to understand human-nature interdependencies in a changing world.

Ecological network complexity scales with area.

Larger geographical areas contain more species-an observation raised to a law in ecology. Less explored is whether biodiversity changes are accompanied by a modification of interaction networks. We use data from 32 spatial interaction networks from different ecosystems to analyse how network structure changes with area. We find that basic community structure descriptors (number of species, links and links per species) increase with area following a power law. Yet, the distribution of links per species varies little with area, indicating that the fundamental organization of interactions within networks is conserved. Our null model analyses suggest that the spatial scaling of network structure is determined by factors beyond species richness and the number of links. We demonstrate that biodiversity-area relationships can be extended from species counts to higher levels of network complexity. Therefore, the consequences of anthropogenic habitat destruction may extend from species loss to wider simplification of natural communities.

Online auction marketplaces as a global pathway for aquatic invasive species.

The ornamental aquarium pet trade is a leading pathway for the introduction of aquatic invasive species. In addition to purchasing live organisms in stores, hobbyists are engaging more with alternative informal online marketplaces that enable peer-to-peer selling of aquarium organisms via auctions. Although growing in popularity, little is known regarding the global extent of informal marketplaces, including the taxonomy of species that are traded, their economic value, and the geographic routes by which live organisms are transported. In this study we use an automated web crawler to collect data on completed auctions between 2011 and 2017 from the largest informal market for aquarium hobbyists, AquaBid, to understand the market dynamics and trade flows of the informal retail market online. During the 7-year study period, the AquaBid website facilitated the estimated trade of 539,548 live freshwater animals, 579,700 fish eggs, and 31,431 plant assortments/bunches among 24,409 unique users who collectively placed 444,132 bids on 192,227 auctions, representing a total sale value of $6,015,030 USD. Source (seller) and recipient (buyer) locations of live organisms were distributed across 39 countries but concentrated largely in major cities of the United States and select European and southeast Asian countries. Our study is among the first to quantify geographic routes of live organism transport between specific locations on the landscape and demonstrates the highly diffuse and non-centralized nature of the informal aquarium trade. Evaluating the emerging challenges represented by informal online retail marketplaces is critical to create policy and regulatory solutions that minimize the transport of prohibited invasive species.

Organismal traits are more important than environment for species interactions in the intertidal zone.

Species interactions come in a variety of forms, from weak to strong, and negative or positive, each with unique consequences for local community structure. However, interactions depend on several biotic, abiotic and scale-dependent variables that make their magnitude and direction difficult to predict. Here, we quantify the relative impacts of multiple factors on species interactions for a diverse array of intertidal organisms, using our own experiments across a range of environments in New Zealand and North America. Interaction strengths are related to organism body size and trophic level, but are relatively insensitive to environmental conditions associated with tidal height. Although species at higher trophic levels exert stronger per-capita effects on other taxa, their population-level impacts are equivalent to basal trophic groups. This indicates that interaction intensity is largely based on requirements for resources, such as food or space, that follow allometric scaling rules. These results demonstrate the potential to predict interactivity based on simple criteria without detailed information on particular species or communities.

Next-generation visitation models using social media to estimate recreation on public lands.

Outdoor and nature-based recreation provides countless social benefits, yet public land managers often lack information on the spatial and temporal extent of recreation activities. Social media is a promising source of data to fill information gaps because the amount of recreational use is positively correlated with social media activity. However, despite the implication that these correlations could be employed to accurately estimate visitation, there are no known transferable models parameterized for use with multiple social media data sources. This study tackles these issues by examining the relative value of multiple sources of social media in models that estimate visitation at unmonitored sites and times across multiple destinations. Using a novel dataset of over 30,000 social media posts and 286,000 observed visits from two regions in the United States, we compare multiple competing statistical models for estimating visitation. We find social media data substantially improve visitor estimates at unmonitored sites, even when a model is parameterized with data from another region. Visitation estimates are further improved when models are parameterized with on-site counts. These findings indicate that while social media do not fully substitute for on-site data, they are a powerful component of recreation research and visitor management.

The marine fish food web is globally connected.

The productivity of marine ecosystems and the services they provide to humans are largely dependent on complex interactions between prey and predators. These are embedded in a diverse network of trophic interactions, resulting in a cascade of events following perturbations such as species extinction. The sheer scale of oceans, however, precludes the characterization of marine feeding networks through de novo sampling. This effort ought instead to rely on a combination of extensive data and inference. Here we investigate how the distribution of trophic interactions at the global scale shapes the marine fish food web structure. We hypothesize that the heterogeneous distribution of species ranges in biogeographic regions should concentrate interactions in the warmest areas and within species groups. We find that the inferred global metaweb of marine fish-that is, all possible potential feeding links between co-occurring species-is highly connected geographically with a low degree of spatial modularity. Metrics of network structure correlate with sea surface temperature and tend to peak towards the tropics. In contrast to open-water communities, coastal food webs have greater interaction redundancy, which may confer robustness to species extinction. Our results suggest that marine ecosystems are connected yet display some resistance to perturbations because of high robustness at most locations.

Inequality in access to cultural ecosystem services from protected areas in the Chilean biodiversity hotspot.

Experiences with nature through visits to protected areas provide important cultural ecosystem services that have the potential to strengthen pro-environmental attitudes and behavior. Understanding accessibility to protected areas and likely preferences for enjoying the benefits of nature visits are key factors in identifying ways to reduce inequality in access and inform the planning and management for future protected areas. We develop, at a regional scale, a novel social media database of visits to public protected areas in part of the Chilean biodiversity hotspot using geotagged photographs and assess the inequality of access using the home locations of the visitors and socio-economic data. We find that 20% of the population of the region make 87% of the visits to protected areas. The larger, more biodiverse protected areas were the most visited and provided most cultural ecosystem services. Wealthier people tend to travel further to visit protected areas while people with lower incomes tend to visit protected areas that are closer to home. By providing information on the current spatial flows of people to protected areas, we demonstrate the need to expand the protected area network, especially in lower income areas, to reduce inequality in access to the benefits from cultural ecosystem services provided by nature to people.

Nature Contact and Human Health: A Research Agenda.

BACKGROUND: At a time of increasing disconnectedness from nature, scientific interest in the potential health benefits of nature contact has grown. Research in recent decades has yielded substantial evidence, but large gaps remain in our understanding. OBJECTIVES: We propose a research agenda on nature contact and health, identifying principal domains of research and key questions that, if answered, would provide the basis for evidence-based public health interventions. DISCUSSION: We identify research questions in seven domains: a) mechanistic biomedical studies; b) exposure science; c) epidemiology of health benefits; d) diversity and equity considerations; e) technological nature; f) economic and policy studies; and g) implementation science. CONCLUSIONS: Nature contact may offer a range of human health benefits. Although much evidence is already available, much remains unknown. A robust research effort, guided by a focus on key unanswered questions, has the potential to yield high-impact, consequential public health insights. https://doi.org/10.1289/EHP1663.

Scale, environment, and trophic status: the context dependency of community saturation in rocky intertidal communities.

Our understanding of the relative influence of different ecological drivers on the number of species in a place remains limited. Assessing the relative influence of local ecological interactions versus regional species pools on local species richness should help bridge this conceptual gap. Plots of local species richness versus regional species pools have been used to address this question, yet after an active quarter-century of research on the relative influence of local interactions versus regional species pools, consensus remains elusive. We propose a conceptual framework that incorporates spatial scale and ecological interaction strength to reconcile current disparities. We then test this framework using a survey of marine rocky intertidal algal and invertebrate communities from the northeast Pacific. We reach two main conclusions. First, these data show that the power of regional species pools to predict local richness disintegrates at small spatial scales coincident with the scale of biological interactions, when studying ecologically interactive groups of species, and in generally more abiotically stressful habitats (e.g., the high intertidal). Second, conclusions of past studies asserting that the regional species pool is the primary driver of local species richness may be artifacts of large spatial scales or ecologically noninteractive groups of species.

Spatial and Temporal Dynamics and Value of Nature-Based Recreation, Estimated via Social Media.

Conserved lands provide multiple ecosystem services, including opportunities for nature-based recreation. Managing this service requires understanding the landscape attributes underpinning its provision, and how changes in land management affect its contribution to human wellbeing over time. However, evidence from both spatially explicit and temporally dynamic analyses is scarce, often due to data limitations. In this study, we investigated nature-based recreation within conserved lands in Vermont, USA. We used geotagged photographs uploaded to the photo-sharing website Flickr to quantify visits by in-state and out-of-state visitors, and we multiplied visits by mean trip expenditures to show that conserved lands contributed US $1.8 billion (US $0.18-20.2 at 95% confidence) to Vermont's tourism industry between 2007 and 2014. We found eight landscape attributes explained the pattern of visits to conserved lands; visits were higher in larger conserved lands, with less forest cover, greater trail density and more opportunities for snow sports. Some of these attributes differed from those found in other locations, but all aligned with our understanding of recreation in Vermont. We also found that using temporally static models to inform conservation decisions may have perverse outcomes for nature-based recreation. For example, static models suggest conserved land with less forest cover receive more visits, but temporally dynamic models suggest clearing forests decreases, rather than increases, visits to these sites. Our results illustrate the importance of understanding both the spatial and temporal dynamics of ecosystem services for conservation decision-making.

The roles and impacts of human hunter-gatherers in North Pacific marine food webs.

There is a nearly 10,000-year history of human presence in the western Gulf of Alaska, but little understanding of how human foragers integrated into and impacted ecosystems through their roles as hunter-gatherers. We present two highly resolved intertidal and nearshore food webs for the Sanak Archipelago in the eastern Aleutian Islands and use them to compare trophic roles of prehistoric humans to other species. We find that the native Aleut people played distinctive roles as super-generalist and highly-omnivorous consumers closely connected to other species. Although the human population was positioned to have strong effects, arrival and presence of Aleut people in the Sanak Archipelago does not appear associated with long-term extinctions. We simulated food web dynamics to explore to what degree introducing a species with trophic roles like those of an Aleut forager, and allowing for variable strong feeding to reflect use of hunting technology, is likely to trigger extinctions. Potential extinctions decreased when an invading omnivorous super-generalist consumer focused strong feeding on decreasing fractions of its possible resources. This study presents the first assessment of the structural roles of humans as consumers within complex ecological networks, and potential impacts of those roles and feeding behavior on associated extinctions.