Contemporary ecological networks reflect the influence of evolutionary and ecological processes on species interactions. Additionally, late Quaternary megafaunal extinctions and subsequent range contractions have interrupted mammalian predator-prey interactions to differing degrees among continents. However, the extent to which species losses have shaped geographic variation in the vertical and horizontal structure of contemporary mammalian food webs remains poorly understood. Furthermore, the relative influences of species loss, evolutionary history, and contemporary environmental drivers on the structure of mammalian food webs have not been tested. Here, we assembled mammal food webs for 389 sites throughout the Neotropics, Afrotropics, and Indomalaya. We first examined variation among continents in food web properties, including predator-prey richness ratios, predator generality, prey vulnerability, and predator dietary specialization. We found that Neotropical food webs bear imprints of severe species loss, featuring smaller predators, lower predator generality, higher prey vulnerability, and reduced prey species availability compared to Afrotropical food webs, in which most large mammals persisted. Furthermore, we reveal that multiple historical legacies, including community lineage history, paleoclimatic variability, mountain uplift, and species losses, as well as contemporary environmental variability, collectively predicted variation in predator dietary niche breadth within continents. Our findings offer unique perspectives on how trophic interactions within food webs respond to species losses and how macroevolutionary and macroecological processes have shaped the biogeography of contemporary mammal food webs.
Tropical forests hold most of Earth's biodiversity and a higher concentration of threatened mammals than other biomes. As a result, some mammal species persist almost exclusively in protected areas, often within extensively transformed and heavily populated landscapes. Other species depend on remaining remote forested areas with sparse human populations. However, it remains unclear how mammalian communities in tropical forests respond to anthropogenic pressures in the broader landscape in which they are embedded. As governments commit to increasing the extent of global protected areas to prevent further biodiversity loss, identifying the landscape-level conditions supporting wildlife has become essential. Here, we assessed the relationship between mammal communities and anthropogenic threats in the broader landscape. We simultaneously modeled species richness and community occupancy as complementary metrics of community structure, using a state-of-the-art community model parameterized with a standardized pan-tropical data set of 239 mammal species from 37 forests across 3 continents. Forest loss and fragmentation within a 50-km buffer were associated with reduced occupancy in monitored communities, while species richness was unaffected by them. In contrast, landscape-scale human density was associated with reduced mammal richness but not occupancy, suggesting that sensitive species have been extirpated, while remaining taxa are relatively unaffected. Taken together, these results provide evidence of extinction filtering within tropical forests triggered by anthropogenic pressure occurring in the broader landscape. Therefore, existing and new reserves may not achieve the desired biodiversity outcomes without concurrent investment in addressing landscape-scale threats.
Understanding variation in food web structure over large spatial scales is an emerging research agenda in food web ecology. The density of predator–prey links in a food web (i.e., connectance) is a key measure of network complexity that describes the mean proportional dietary breadth of species within a food web. Connectance is a critical component of food web robustness to species loss: food webs with lower connectance have been shown to be more susceptible to secondary extinctions. Identifying geographic variation in food web connectance and its drivers may provide insight into community robustness to species loss. We investigated the food web connectance of ground-dwelling tropical forest mammal communities in multiple biogeographic regions to test for differences among regions in food web connectance and to test three potential drivers: primary productivity, contemporary anthropogenic pressure, and variation in mammal body mass distributions reflective of historical extinctions. Mammal communities from fifteen protected forests throughout the Neo-, Afro-, and Asian tropics were identified from systematic camera trap arrays. Predator–prey interaction data were collected from published literature, and we calculated connectance for each community as the number of observed predator–prey links relative to the number of possible predator–prey links. We used generalized linear models to test for differences among regions and to identify the site level characteristics that best predicted connectance. We found that mammal food web connectance varied significantly among continents and that body size range was the only significant predictor. More possible predator–prey links were observed in communities with smaller ranges in body size and therefore sites with smaller body size ranges had higher mean proportional dietary breadth. Specifically, mammal communities in the Neotropics and in Madagascar had significantly higher connectance than mammal communities in Africa. This geographic variation in contemporary mammalian food web structure may be the product of historical extinctions in the Late Quaternary, which led to greater losses of large-bodied species in the Neotropics and Madagascar thus contributing to higher average proportional dietary breadth among the remaining smaller bodied species in these regions.
Aim Tropical regions harbour over half of the world's mammals and birds, but how their communities have assembled over evolutionary timescales remains unclear. To compare eco-evolutionary assembly processes between tropical mammals and birds, we tested how hypotheses concerning niche conservatism, environmental stability, environmental heterogeneity and time-for-speciation relate to tropical vertebrate community phylogenetic and functional structure. Location Tropical rainforests worldwide. Time periodPresent. Major taxa studied Ground-dwelling and ground-visiting mammals and birds. Methods We used in situ observations of species identified from systematic camera trap sampling as realized communities from 15 protected tropical rainforests in four tropical regions worldwide. We quantified standardized phylogenetic and functional structure for each community and estimated the multi-trait phylogenetic signal (PS) in ecological strategies for the four regional species pools of mammals and birds. Using linear regression models, we test three non-mutually exclusive hypotheses by comparing the relative importance of colonization time, palaeo-environmental changes in temperature and land cover since 3.3 Mya, contemporary seasonality in temperature and productivity and environmental heterogeneity for predicting community phylogenetic and functional structure. Results Phylogenetic and functional structure showed non-significant yet varying tendencies towards clustering or dispersion in all communities. Mammals had stronger multi-trait PS in ecological strategies than birds (mean PS: mammal = 0.62, bird = 0.43). Distinct dominant processes were identified for mammal and bird communities. For mammals, colonization time and elevation range significantly predicted phylogenetic clustering and functional dispersion tendencies respectively. For birds, elevation range and contemporary temperature seasonality significantly predicted phylogenetic and functional clustering tendencies, respectively, while habitat diversity significantly predicted functional dispersion tendencies. Main conclusions Our results reveal different eco-evolutionary assembly processes structuring contemporary tropical mammal and bird communities over evolutionary timescales that have shaped tropical diversity. Our study identified marked differences among taxonomic groups in the relative importance of historical colonization and sensitivity to environmental change.
Many animal-environment interactions are mediated by the physical forms of the environment, especially in tropical forests, where habitats are structurally complex and highly diverse. Higher structural complexity, measured as habitat surface area, may provide increased resource availability for animals, leading to higher animal diversity. Greater habitat surface area supports increased animal diversity in other systems, such as coral reefs and forest canopies, but it is uncertain how this relationship translates to communities of highly mobile, terrestrial mammal species inhabiting forest floors. We tested the relative importance of forest floor habitat structure, encompassing vegetation and topographic structure, in determining species occupancy and functional diversity of medium to large mammals using data from a tropical forest in the Udzungwa Mountains of Tanzania. We related species occupancies and diversity obtained from a multispecies occupancy model with ground-level habitat structure measurements obtained from a novel head-mounted active remote sensing device, the Microsoft HoloLens. We found that habitat surface area was a significant predictor of mean species occupancy and had a significant positive relationship with functional dispersion. The positive relationships indicate that surface area of tropical forest floors may play an important role in promoting mammal occupancy and functional diversity at the microhabitat scale. In particular, habitat surface area had higher mean effects on occupancy for carnivorous and social species. These results support a habitat surface area-diversity relationship on tropical forest floors for mammals.
The spatial aggregation of species pairs often increases with the ecological similarity of the species involved. However, the way in which environmental conditions and anthropogenic activity affect the relationship between spatial aggregation and ecological similarity remains unknown despite the potential for spatial associations to affect species interactions, ecosystem function, and extinction risk. Given that human disturbance has been shown to both increase and decrease spatial associations among species pairs, ecological similarity may have a role in mediating these patterns. Here, we test the influences of habitat diversity, primary productivity, human population density, and species' ecological similarity based on functional traits (i.e., functional trait similarity) on spatial associations among tropical forest mammals. Large mammals are highly sensitive to anthropogenic change and therefore susceptible to changes in interspecific spatial associations. Using two-species occupancy models and camera trap data, we quantified the spatial overlap of 1216 species pairs from 13 tropical forest protected areas around the world. We found that the association between ecological similarity and interspecific species associations depended on surrounding human density. Specifically, aggregation of ecologically similar species was more than an order of magnitude stronger in landscapes with the highest human density compared to those with the lowest human density, even though all populations occurred within protected areas. Human-induced changes in interspecific spatial associations have been shown to alter top-down control by predators, increase disease transmission and increase local extinction rates. Our results indicate that anthropogenic effects on the distribution of wildlife within protected areas are already occurring and that impacts on species interactions, ecosystem functions, and extinction risk warrant further investigation.
Most ecological studies of vegetation structure have relied on manual field measurements that are labour-intensive and time-consuming. Many current alternatives to classical measurements are expensive or difficult to transport to field settings. Here we evaluated a new method for measuring understorey vegetation with a novel mixed-reality, remote sensing device, the Microsoft HoloLens. We developed a vegetation sensing application called VegSense that allows the HoloLens user to control the device's environmental scanners to measure understorey vegetation. Using VegSense, we tested the ability of the Microsoft HoloLens relative to classical field measurements to (a) detect trees and saplings, (b) measure diameter at breast height (DBH), (c) detect individual understorey vegetation structures and (d) estimate understorey vegetation complexity replicating the rod-transect method. We found that VegSense performed well in detecting and measuring trees with a DBH of 17 cm or more and estimating vegetation complexity and performed moderately at detecting understorey vegetation. Our results indicate that the HoloLens is a suitable alternative for multiple classical field measurements of understorey vegetation. This method costs much less than typical terrestrial LiDAR systems, and can facilitate efficient, high-quality environmental data collection. Further software development has the potential to reveal additional ways in which this device can be harnessed for applications to ecology and evolution.
Globally, tropical rain forests comprise some of the most diverse and functionally rich ecosystems but are increasingly degraded by human impacts. Protected areas have been shown to conserve species diversity, but their effectiveness at maintaining functional diversity over time is less well known, despite the fact that functional diversity likely reveals more ecological information than taxonomic diversity. By extension, the degree to which species loss decreases functional diversity within protected areas is also unknown; functional redundancy may buffer communities from loss of functional diversity from some local extinctions. Using eight years of camera trap data, we quantified annual functional dispersion of the large mammal community in the Volcán Barva region of Costa Rica and tested for changes in functional dispersion over time in response to environmental and anthropogenic predictors. We quantified functional redundancy based on simulated declines in functional dispersion with species loss. Observed functional dispersion did not change significantly over time and was not associated with measured environmental or anthropogenic predictors. Quantitative modeling of observed functional traits over time did not identify significant changes. We did however find qualitative trends in relative trait proportions, which could be indicative of functional change in the future. We found high functional redundancy, with average simulated functional dispersion declining significantly only after 9 out of 21 large mammal species were lost from the community. We cautiously suggest that protected tropical rain forests can conserve functional diversity over the course of a decade even in heavily fragmented landscapes.
A variety of factors can affect the biodiversity of tropical mammal communities, but their relative importance and directionality remain uncertain. Previous global investigations of mammal functional diversity have relied on range maps instead of observational data to determine community composition. We test the effects of species pools, habitat heterogeneity, primary productivity and human disturbance on the functional diversity (dispersion and richness) of mammal communities using the largest standardized tropical forest camera trap monitoring system, the Tropical Ecology Assessment and Monitoring (TEAM) Network. We use occupancy values derived from the camera trap data to calculate occupancy-weighted functional diversity and use Bayesian generalized linear regression to determine the effects of multiple predictors. Mammal community functional dispersion increased with primary productivity, while functional richness decreased with human-induced local extinctions and was significantly lower in Madagascar than other tropical regions. The significant positive relationship between functional dispersion and productivity was evident only when functional dispersion was weighted by species' occupancies. Thus, observational data from standardized monitoring can reveal the drivers of mammal communities in ways that are not readily apparent from range map-based studies. The positive association between occupancy-weighted functional dispersion of tropical forest mammal communities and primary productivity suggests that unique functional traits may be more beneficial in more productive ecosystems and may allow species to persist at higher abundances.
Accurate estimates of density and population size are fundamental for appropriate monitoring of populations. Forest dwelling large mammals that live in herds are particularly difficult for estimating population density and size because of the heterogeneity of their spatial distribution at local scales and the difficulty of counting groups. Previous work has dealt with heterogeneity of spatial-temporal distribution by the independent use of gas movement theory and spatially explicit distribution models. Here we propose to combine both approaches, and a novel probabilistic method to minimize inflation of group size estimates, to estimate density and population size of a Neotropical social mammal, the white-lipped peccary ( Tayassu pecari ). From February to August 2013, we placed ten camera-traps at distinct locations approximately 2 km apart from one another, at Ilha do Cardoso State Park, a 110-km 2 protected area in the Atlantic forest of Brazil. We estimated groups/area using a gas movement function, and individuals/group from a pathway probability function, and used both to derive a final density estimate (individuals/area). We then generated a habitat suitability model based on seven environmental layers and 32 location points, and extracted the area with high probability of occurrence to estimate population size in the park. In 1232 camera trap*days, we recorded 309 10-second videos of white-lipped peccary herds. Simple minimum density estimates (from subtracting individuals that leave the screen) were sensitive to the time interval used to separate independent events, but density estimates using the pathway probability function were not, with a final density estimate of 6.2-9.8 inds/km 2 . Habitat suitability analyses revealed that four environmental variables (distance from continental edge, altitude, vegetation, and slope) collectively explained 84% of the variation in white-lipped occurrence. Using a model threshold of 0.27 (maximum test sensitivity and specificity), we determined that only 21% of the park’s area was in fact suitable for the species, with a final population size estimate ranging from 175 to 275 individuals. The present study reveals that a combination of analytical tools allows for more robust estimates of population density and size of forest-dwelling social mammals. Our approach also permits disentangling distinct factors affecting population size (group size, group density, and suitable area), and facilitates pinpointing sources of variation in population size over time.