Rethinking megafauna.

Concern for megafauna is increasing among scientists and non-scientists. Many studies have emphasized that megafauna play prominent ecological roles and provide important ecosystem services to humanity. But, what precisely are 'megafauna'? Here, we critically assess the concept of megafauna and propose a goal-oriented framework for megafaunal research. First, we review definitions of megafauna and analyse associated terminology in the scientific literature. Second, we conduct a survey among ecologists and palaeontologists to assess the species traits used to identify and define megafauna. Our review indicates that definitions are highly dependent on the study ecosystem and research question, and primarily rely on ad hoc size-related criteria. Our survey suggests that body size is crucial, but not necessarily sufficient, for addressing the different applications of the term megafauna. Thus, after discussing the pros and cons of existing definitions, we propose an additional approach by defining two function-oriented megafaunal concepts: 'keystone megafauna' and 'functional megafauna', with its variant 'apex megafauna'. Assessing megafauna from a functional perspective could challenge the perception that there may not be a unifying definition of megafauna that can be applied to all eco-evolutionary narratives. In addition, using functional definitions of megafauna could be especially conducive to cross-disciplinary understanding and cooperation, improvement of conservation policy and practice, and strengthening of public perception. As megafaunal research advances, we encourage scientists to unambiguously define how they use the term 'megafauna' and to present the logic underpinning their definition.

Mapping and assessing the impact of small-scale ephemeral water sources on wildlife in an African seasonal savannah.

In many savannah regions of Africa, pronounced seasonal variability in rainfall results in wildlife being restricted to floodplains and other habitats adjacent to permanent surface water in the dry season. During the wet season, rainfall fills small-scale, ephemeral water sources that allow wildlife to exploit forage and other resources far from permanent surface water. These water sources remain difficult to quantify, however, due to their small and ephemeral nature, and as a result are rarely included in quantitative studies of wildlife distribution, abundance, and movement. Our goal was to map ephemeral water in Bwabwata National Park in Namibia using two different approaches and to relate measures of ephemeral water to the abundance, distribution, and movement of two large wildlife species. We used high-resolution Google Earth and Esri World imagery to visually identify waterholes. Additionally, we used Sentinel-2 satellite imagery to map ephemeral water across the study area using the Normalized Difference Water Index. With these mapped waterhole layers and data from GPS-collared individuals of African elephant (Loxodonta africana) and African buffalo (Syncerus caffer), we evaluated the importance of ephemeral water in conditioning abundance and movement of these two species. The two approaches to mapping ephemeral water resulted in the visual identification of nearly 10,000 waterholes, and a predicted ephemeral water layer of ~76% accuracy. The inclusion of ephemeral water into models of abundance and movement resulted in improved goodness of fit relative to those without water, and water impacts on abundance and movement were among the strongest of all variables considered. The potential importance of ephemeral water in conditioning the movements and distributions of large herbivores in African savannahs has been difficult to quantify relative to vegetation drivers. Our results suggest research into ephemeral water impacts deserves more attention.

Walk on the wild side: estimating the global magnitude of visits to protected areas.

How often do people visit the world's protected areas (PAs)? Despite PAs covering one-eighth of the land and being a major focus of nature-based recreation and tourism, we don't know. To address this, we compiled a globally-representative database of visits to PAs and built region-specific models predicting visit rates from PA size, local population size, remoteness, natural attractiveness, and national income. Applying these models to all but the very smallest of the world's terrestrial PAs suggests that together they receive roughly 8 billion (8 x 109) visits/y-of which more than 80% are in Europe and North America. Linking our region-specific visit estimates to valuation studies indicates that these visits generate approximately US $600 billion/y in direct in-country expenditure and US $250 billion/y in consumer surplus. These figures dwarf current, typically inadequate spending on conserving PAs. Thus, even without considering the many other ecosystem services that PAs provide to people, our findings underscore calls for greatly increased investment in their conservation.

Conserving tropical biodiversity via market forces and spatial targeting.

The recent report from the Secretariat of the Convention on Biological Diversity [(2010) Global Biodiversity Outlook 3] acknowledges that ongoing biodiversity loss necessitates swift, radical action. Protecting undisturbed lands, although vital, is clearly insufficient, and the key role of unprotected, private land owned is being increasingly recognized. Seeking to avoid common assumptions of a social planner backed by government interventions, the present work focuses on the incentives of the individual landowner. We use detailed data to show that successful conservation on private land depends on three factors: conservation effectiveness (impact on target species), private costs (especially reductions in production), and private benefits (the extent to which conservation activities provide compensation, for example, by enhancing the value of remaining production). By examining the high-profile issue of palm-oil production in a major tropical biodiversity hotspot, we show that the levels of both conservation effectiveness and private costs are inherently spatial; varying the location of conservation activities can radically change both their effectiveness and private cost implications. We also use an economic choice experiment to show that consumers' willingness to pay for conservation-grade palm-oil products has the potential to incentivize private producers sufficiently to engage in conservation activities, supporting vulnerable International Union for Conservation of Nature Red Listed species. However, these incentives vary according to the scale and efficiency of production and the extent to which conservation is targeted to optimize its cost-effectiveness. Our integrated, interdisciplinary approach shows how strategies to harness the power of the market can usefully complement existing--and to-date insufficient--approaches to conservation.

Complementary benefits of tourism and hunting to communal conservancies in Namibia.

Tourism and hunting both generate substantial revenues for communities and private operators in Africa, but few studies have quantitatively examined the trade-offs and synergies that may result from these two activities. We evaluated financial and in-kind benefit streams from tourism and hunting on 77 communal conservancies in Namibia from 1998 to 2013, where community-based wildlife conservation has been promoted as a land-use that complements traditional subsistence agriculture. We used data collected annually for all communal conservancies to characterize whether benefits were derived from hunting or tourism. We classified these benefits into 3 broad classes and examined how benefits flowed to stakeholders within communities under the status quo and under a simulated ban on hunting. Across all conservancies, total benefits from hunting and tourism increased at roughly the same rate, although conservancies typically started generating benefits from hunting within 3 years of formation as opposed to after 6 years for tourism. Disaggregation of data revealed that the main benefits from hunting were income for conservancy management and food in the form of meat for the community at large. The majority of tourism benefits were salaried jobs at lodges. A simulated ban on trophy hunting significantly reduced the number of conservancies that could cover their operating costs, whereas eliminating income from tourism did not have as severe an effect. Given that the benefits generated from hunting and tourism typically begin at different times in a conservancy's life-span (earlier vs. later, respectively) and flow to different segments of local communities, these 2 activities together may provide the greatest incentives for conservation on communal lands in Namibia. A singular focus on either hunting or tourism would reduce the value of wildlife as a competitive land-use option and have grave repercussions for the viability of community-based conservation efforts in Namibia, and possibly other parts of Africa.

Enhancing conservation, ecosystem services, and local livelihoods through a wildlife premium mechanism.

We propose the wildlife premium mechanism as an innovation to conserve endangered large vertebrates. The performance-based payment scheme would allow stakeholders in lower-income countries to generate revenue by recovering and maintaining threatened fauna that can also serve as umbrella species (i.e., species whose protection benefits other species with which they co-occur). There are 3 possible options for applying the premium: option 1, embed premiums in a carbon payment; option 2, link premiums to a related carbon payment, but as independent and legally separate transactions; option 3, link premiums to noncarbon payments for conserving ecosystem services (PES). Each option presents advantages, such as incentive payments to improve livelihoods of rural poor who reside in or near areas harboring umbrella species, and challenges, such as the establishment of a subnational carbon credit scheme. In Kenya, Peru, and Nepal pilot premium projects are now underway or being finalized that largely follow option 1. The Kasigau (Kenya) project is the first voluntary carbon credit project to win approval from the 2 leading groups sanctioning such protocols and has already sold carbon credits totaling over $1.2 million since June 2011. A portion of the earnings is divided among community landowners and projects that support community members and has added over 350 jobs to the local economy. All 3 projects involve extensive community management because they occur on lands where locals hold the title or have a long-term lease from the government. The monitoring, reporting, and verification required to make premium payments credible to investors include transparent methods for collecting data on key indices by trained community members and verification of their reporting by a biologist. A wildlife premium readiness fund would enable expansion of pilot programs needed to test options beyond those presented here.

Contrasting historical and recent gene flow among African buffalo herds in the Caprivi Strip of Namibia.

Population genetic structure is often used to infer population connectivity, but genetic structure may largely reflect historical rather than recent processes. We contrasted genetic structure with recent gene-flow estimates among 6 herds of African buffalo (Syncerus caffer) in the Caprivi Strip, Namibia, using 134 individuals genotyped at 10 microsatellite loci. We tested whether historical and recent gene flows were influenced by distance, potential barriers (rivers), or landscape resistance (distance from water). We also tested at what scales individuals were more related than expected by chance. Genetic structure across the Caprivi Strip was weak, indicating that historically, gene flow was strong and was not affected by distance, barriers, or landscape resistance. Our analysis of simulated data suggested that genetic structure would be unlikely to reflect human disturbances in the last 10-20 generations (75-150 years) because of slow predicted rates of genetic drift, but recent gene-flow estimates would be affected. Recent gene-flow estimates were not consistently affected by rivers or distance to water but showed that isolation by distance appears to be developing. Average relatedness estimates among individuals exceeded random expectations only within herds. We conclude that historically, African buffalo moved freely throughout the Caprivi Strip, whereas recent gene flow has been more restricted. Our findings support efforts to maintain the connectivity of buffalo herds across this region and demonstrate the utility of contrasting genetic inferences from different time scales.

Community-based conservation reduces sexual risk factors for HIV among men.

BACKGROUND: Despite numerous programs to combat the global HIV and AIDS pandemic, infection rates remain high, especially in sub-Saharan Africa, where two-thirds of all people living with HIV reside. Here, we describe how we used rigorous program evaluation methods to assess the effectiveness of a community-based natural resource management program that "mainstreamed" HIV awareness and prevention activities within rural communities in Namibia. FINDINGS: We used data from two rounds of the Namibia Demographic and Health Surveys (2000 and 2006/2007) and quasi-experimental statistical methods to evaluate changes in critical health-related outcomes in men and women living in communal conservancies, relative to several non-conservancy comparison groups. Our final dataset included 117 men and 318 women in 2000, and 170 men and 357 women in 2006/2007. We evaluated the statistical significance of the main effects of survey year and conservancy residence, and a conservancy-year interaction term, using generalized linear models. Our analyses show that community-based conservation in Namibia has significantly reduced multiple sexual partnerships, the main behavioural determinant of HIV/AIDS infection in Africa. CONCLUSIONS: Our results demonstrate the effectiveness of holistic community-based approaches centered on the preservation of lives and livelihoods, and highlight the potential benefits of integrating conservation and HIV prevention programming in other areas of communal land tenure in Africa.

Human presence and infrastructure impact wildlife nocturnality differently across an assemblage of mammalian species.

Wildlife species may shift towards more nocturnal behavior in areas of higher human influence, but it is unclear how consistent this shift might be. We investigated how humans impact large mammal diel activities in a heavily recreated protected area and an adjacent university-managed forest in southwest British Columbia, Canada. We used camera trap detections of humans and wildlife, along with data on land-use infrastructure (e.g., recreation trails and restricted-access roads), in Bayesian regression models to investigate impacts of human disturbance on wildlife nocturnality. We found moderate evidence that black bears (Ursus americanus) were more nocturnal in response to human detections (mean posterior estimate = 0.35, 90% credible interval = 0.04 to 0.65), but no other clear relationships between wildlife nocturnality and human detections. However, we found evidence that coyotes (Canis latrans) (estimates = 0.81, 95% CI = 0.46 to 1.17) were more nocturnal and snowshoe hares (Lepus americanus) (estimate = -0.87, 95% CI = -1.29 to -0.46) were less nocturnal in areas of higher trail density. We also found that coyotes (estimate = -0.87, 95% CI = -1.29 to -0.46) and cougars (Puma concolor) (estimate = -1.14, 90% CI = -2.16 to -0.12) were less nocturnal in areas of greater road density. Furthermore, coyotes, black-tailed deer (Odocoileus hemionus), and snowshoe hares were moderately more nocturnal in areas near urban-wildland boundaries (estimates and 90% CIs: coyote = -0.29, -0.55 to -0.04, black-tailed deer = -0.25, -0.45 to -0.04, snowshoe hare = -0.24, -0.46 to -0.01). Our findings imply anthropogenic landscape features may influence medium to large-sized mammal diel activities more than direct human presence. While increased nocturnality may be a promising mechanism for human-wildlife coexistence, shifts in temporal activity can also have negative repercussions for wildlife, warranting further research into the causes and consequences of wildlife responses to increasingly human-dominated landscapes.

Mammalian predator and prey responses to recreation and land use across multiple scales provide limited support for the human shield hypothesis.

Outdoor recreation is widespread, with uncertain effects on wildlife. The human shield hypothesis (HSH) suggests that recreation could have differential effects on predators and prey, with predator avoidance of humans creating a spatial refuge 'shielding' prey from people. The generality of the HSH remains to be tested across larger scales, wherein human shielding may prove generalizable, or diminish with variability in ecological contexts. We combined data from 446 camera traps and 79,279 sampling days across 10 landscapes spanning 15,840 km2 in western Canada. We used hierarchical models to quantify the influence of recreation and landscape disturbance (roads, logging) on ungulate prey (moose, mule deer and elk) and carnivore (wolf, grizzly bear, cougar and black bear) site use. We found limited support for the HSH and strong responses to recreation at local but not larger spatial scales. Only mule deer showed positive but weak landscape-level responses to recreation. Elk were positively associated with local recreation while moose and mule deer responses were negative, contrary to HSH predictions. Mule deer showed a more complex interaction between recreation and land-use disturbance, with more negative responses to recreation at lower road density or higher logged areas. Contrary to HSH predictions, carnivores did not avoid recreation and grizzly bear site use was positively associated. We also tested the effects of roads and logging on temporal activity overlap between mule deer and recreation, expecting deer to minimize interaction with humans by partitioning time in areas subject to more habitat disturbance. However, temporal overlap between people and deer increased with road density. Our findings highlight the complex ecological patterns that emerge at macroecological scales. There is a need for expanded monitoring of human and wildlife use of recreation areas, particularly multi-scale and -species approaches to studying the interacting effects of recreation and land-use change on wildlife.

A global perspective on trends in nature-based tourism.

Reports of rapid growth in nature-based tourism and recreation add significant weight to the economic case for biodiversity conservation but seem to contradict widely voiced concerns that people are becoming increasingly isolated from nature. This apparent paradox has been highlighted by a recent study showing that on a per capita basis, visits to natural areas in the United States and Japan have declined over the last two decades. These results have been cited as evidence of "a fundamental and pervasive shift away from nature-based recreation"-but how widespread is this phenomenon? We address this question by looking at temporal trends in visitor numbers at 280 protected areas (PAs) from 20 countries. This more geographically representative dataset shows that while PA visitation (whether measured as total or per capita visit numbers) is indeed declining in the United States and Japan, it is generally increasing elsewhere. Total visit numbers are growing in 15 of the 20 countries for which we could get data, with the median national rate of change unrelated to the national rate of population growth but negatively associated with wealth. Reasons for this reversal of growth in the richest countries are difficult to pin down with existing data, but the pattern is mirrored by trends in international tourist arrivals as a whole and so may not necessarily be caused by disaffection with nature. Irrespective of the explanation, it is clear that despite important downturns in some countries, nature-related tourism is far from declining everywhere, and may still have considerable potential both to generate funds for conservation and to shape people's attitudes to the environment.

Human impacts on mammals in and around a protected area before, during, and after COVID-19 lockdowns.

The dual mandate for many protected areas (PAs) to simultaneously promote recreation and conserve biodiversity may be hampered by negative effects of recreation on wildlife. However, reports of these effects are not consistent, presenting a knowledge gap that hinders evidence-based decision-making. We used camera traps to monitor human activity and terrestrial mammals in Golden Ears Provincial Park and the adjacent University of British Columbia Malcolm Knapp Research Forest near Vancouver, Canada, with the objective of discerning relative effects of various forms of recreation on cougars (Puma concolor), black bears (Ursus americanus), black-tailed deer (Odocoileus hemionus), snowshoe hares (Lepus americanus), coyotes (Canis latrans), and bobcats (Lynx rufus). Additionally, public closures of the study area associated with the COVD-19 pandemic offered an unprecedented period of human-exclusion through which to explore these effects. Using Bayesian generalized mixed-effects models, we detected negative effects of hikers (mean posterior estimate = -0.58, 95% credible interval [CI] -1.09 to -0.12) on weekly bobcat habitat use and negative effects of motorized vehicles (estimate = -0.28, 95% CI -0.61 to -0.05) on weekly black bear habitat use. We also found increased cougar detection rates in the PA during the COVID-19 closure (estimate = 0.007, 95% CI 0.005 to 0.009), but decreased cougar detection rates (estimate = -0.006, 95% CI -0.009 to -0.003) and increased black-tailed deer detection rates (estimate = 0.014, 95% CI 0.002 to 0.026) upon reopening of the PA. Our results emphasize that effects of human activity on wildlife habitat use and movement may be species- and/or activity-dependent, and that camera traps can be an invaluable tool for monitoring both wildlife and human activity, collecting data even when public access is barred. Further, we encourage PA managers seeking to promote both biodiversity conservation and recreation to explicitly assess trade-offs between these two goals in their PAs.

Cost-effective priorities for global mammal conservation.

Global biodiversity priority setting underpins the strategic allocation of conservation funds. In identifying the first comprehensive set of global priority areas for mammals, Ceballos et al. [Ceballos G, Ehrlich PR, Soberón J, Salazar I, Fay JP (2005) Science 309:603-607] found much potential for conflict between conservation and agricultural human activity. This is not surprising because, like other global priority-setting approaches, they set priorities without socioeconomic objectives. Here we present a priority-setting framework that seeks to minimize the conflicts and opportunity costs of meeting conservation goals. We use it to derive a new set of priority areas for investment in mammal conservation based on (i) agricultural opportunity cost and biodiversity importance, (ii) current levels of international funding, and (iii) degree of threat. Our approach achieves the same biodiversity outcomes as Ceballos et al.'s while reducing the opportunity costs and conflicts with agricultural human activity by up to 50%. We uncover shortfalls in the allocation of conservation funds in many threatened priority areas, highlighting a global conservation challenge.

Socio-demographic correlates of wildlife consumption during early stages of the COVID-19 pandemic.

To inform efforts at preventing future pandemics, we assessed how socio-demographic attributes correlated with wildlife consumption as COVID-19 (coronavirus disease 2019) first spread across Asia. Self-reported wildlife consumption was most strongly related to COVID-19 awareness; those with greater awareness were 11-24% less likely to buy wildlife products. A hypothetical intervention targeting increased awareness, support for wildlife market closures and reduced medical impacts of COVID-19 could halve future wildlife consumption rates across several countries and demographics.

eDNA sampled from stream networks correlates with camera trap detection rates of terrestrial mammals.

Biodiversity monitoring delivers vital information to those making conservation decisions. Comprehensively measuring terrestrial biodiversity usually requires costly methods that can rarely be deployed at large spatial scales over multiple time periods, limiting conservation efficiency. Here we investigated the capacity of environmental DNA (eDNA) from stream water samples to survey terrestrial mammal diversity at multiple spatial scales within a large catchment. We compared biodiversity information recovered using an eDNA metabarcoding approach with data from a dense camera trap survey, as well as the sampling costs of both methods. Via the sampling of large volumes of water from the two largest streams that drained the study area, eDNA metabarcoding provided information on the presence and detection probabilities of 35 mammal taxa, 25% more than camera traps and for half the cost. While eDNA metabarcoding had limited capacity to detect felid species and provide individual-level demographic information, it is a cost-efficient method for large-scale monitoring of terrestrial mammals that can offer sufficient information to solve many conservation problems.

Usually hated, sometimes loved: A review of wild ungulates' contributions to people.

Nature's contributions to people (NCP) may be both beneficial and detrimental to humans' quality of life. Since our origins, humans have been closely related to wild ungulates, which have traditionally played an outstanding role as a source of food or raw materials. Currently, wild ungulates are declining in some regions, but recovering in others throughout passive rewilding processes. This is reshaping human-ungulate interactions. Thus, adequately understanding the benefits and detriments associated with wild ungulate populations is necessary to promote human-ungulate co-existence. Here, we reviewed 575 articles (2000-2019) on human-wild ungulate interactions to identify key knowledge gaps on NCP associated with wild ungulates. Wild ungulate research was mainly distributed into seven research clusters focussing on: (1) silvicultural damage in Eurasia; (2) herbivory and natural vegetation; (3) conflicts in urban areas of North America; (4) agricultural damage in Mediterranean agro-ecosystems; (5) social research in Africa and Asia; (6) agricultural damage in North America; (7) research in natural American Northwest areas. Research mostly focused on detrimental NCP. However, the number of publications mentioning beneficial contributions increased after the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services conceptual framework was implemented. Human-ungulate interactions' research was biased towards the Global North and Cervidae, Suidae and Bovidae families. Regarding detrimental NCP, most publications referred to production damage (e.g. crops), followed by biodiversity damage, and material damage (e.g. traffic collisions). Regarding beneficial NCP, publications mainly highlighted non-material contributions (e.g. recreational hunting), followed by material NCP and regulating contributions (e.g. habitat creation). The main actions taken to manage wild ungulate populations were lethal control and using deterrents and barriers (e.g. fencing), which effectiveness was rarely assessed. Increasing research and awareness about beneficial NCP and effective management tools may help to improve the conservation of wild ungulates and the ecosystems they inhabit to facilitate people-ungulate co-existence in the Anthropocene.

Impacts of marine protected areas on fishing communities.

Marine protected areas (MPAs) are a popular conservation strategy, but their impacts on human welfare are poorly understood. To inform future research and policy decisions, we reviewed the scientific literature to assess MPA impacts on five indicators of human welfare: food security, resource rights, employment, community organization, and income. Following MPA establishment, food security generally remained stable or increased in older and smaller MPAs. The ability of most fishing groups to govern MPA resources changed. Increased resource rights were positively correlated with MPA zoning and compliance with MPA regulations. Small sample sizes precluded statistical tests of the impacts of MPAs on employment, community organization, and income. Our results demonstrate that MPAs shape the social well-being and political power of fishing communities; impacts (positive and negative) vary within and among social groups; and social impacts are correlated with some--but not all--commonly hypothesized explanatory factors. Accordingly, MPAs may represent a viable strategy for enhancing food security and empowering local communities, but current practices negatively affect at least a minority of fishers. To inform policy making, further research must better document and explain variation in the positive and negative social impacts of MPAs.