Latitudinal patterns in stabilizing density dependence of forest communities.

Numerous studies have shown reduced performance in plants that are surrounded by neighbours of the same species1,2, a phenomenon known as conspecific negative density dependence (CNDD)3. A long-held ecological hypothesis posits that CNDD is more pronounced in tropical than in temperate forests4,5, which increases community stabilization, species coexistence and the diversity of local tree species6,7. Previous analyses supporting such a latitudinal gradient in CNDD8,9 have suffered from methodological limitations related to the use of static data10-12. Here we present a comprehensive assessment of latitudinal CNDD patterns using dynamic mortality data to estimate species-site-specific CNDD across 23 sites. Averaged across species, we found that stabilizing CNDD was present at all except one site, but that average stabilizing CNDD was not stronger toward the tropics. However, in tropical tree communities, rare and intermediate abundant species experienced stronger stabilizing CNDD than did common species. This pattern was absent in temperate forests, which suggests that CNDD influences species abundances more strongly in tropical forests than it does in temperate ones13. We also found that interspecific variation in CNDD, which might attenuate its stabilizing effect on species diversity14,15, was high but not significantly different across latitudes. Although the consequences of these patterns for latitudinal diversity gradients are difficult to evaluate, we speculate that a more effective regulation of population abundances could translate into greater stabilization of tropical tree communities and thus contribute to the high local diversity of tropical forests.

Warm springs alter timing but not total growth of temperate deciduous trees.

As the climate changes, warmer spring temperatures are causing earlier leaf-out1-3 and commencement of CO2 uptake1,3 in temperate deciduous forests, resulting in a tendency towards increased growing season length3 and annual CO2 uptake1,3-7. However, less is known about how spring temperatures affect tree stem growth8,9, which sequesters carbon in wood that has a long residence time in the ecosystem10,11. Here we show that warmer spring temperatures shifted stem diameter growth of deciduous trees earlier but had no consistent effect on peak growing season length, maximum growth rates, or annual growth, using dendrometer band measurements from 440 trees across two forests. The latter finding was confirmed on the centennial scale by 207 tree-ring chronologies from 108 forests across eastern North America, where annual ring width was far more sensitive to temperatures during the peak growing season than in the spring. These findings imply that any extra CO2 uptake in years with warmer spring temperatures4,5 does not significantly contribute to increased sequestration in long-lived woody stem biomass. Rather, contradicting projections from global carbon cycle models1,12, our empirical results imply that warming spring temperatures are unlikely to increase woody productivity enough to strengthen the long-term CO2 sink of temperate deciduous forests.

Tree diversity enhances predation by birds but not by arthropods across climate gradients.

Tree diversity can promote both predator abundance and diversity. However, whether this translates into increased predation and top-down control of herbivores across predator taxonomic groups and contrasting environmental conditions remains unresolved. We used a global network of tree diversity experiments (TreeDivNet) spread across three continents and three biomes to test the effects of tree species richness on predation across varying climatic conditions of temperature and precipitation. We recorded bird and arthropod predation attempts on plasticine caterpillars in monocultures and tree species mixtures. Both tree species richness and temperature increased predation by birds but not by arthropods. Furthermore, the effects of tree species richness on predation were consistent across the studied climatic gradient. Our findings provide evidence that tree diversity strengthens top-down control of insect herbivores by birds, underscoring the need to implement conservation strategies that safeguard tree diversity to sustain ecosystem services provided by natural enemies in forests.

Harmonizing spatial scales and ecological theories to predict avian richness and functional diversity within forest ecosystems.

Classic ecological theory has proven that temperature, precipitation and productivity organize ecosystems at broad scales and are generalized drivers of biodiversity within different biomes. At local scales, the strength of these predictors is not consistent across different biomes. To better translate these theories to localized scales, it is essential to determine the links between drivers of biodiversity. Here we harmonize existing ecological theories to increase the predictive power for species richness and functional diversity. We test the relative importance of three-dimensional habitat structure as a link between local and broad-scale patterns of avian richness and functional diversity. Our results indicate that habitat structure is more important than precipitation, temperature and elevation gradients for predicting avian species richness and functional diversity across different forest ecosystems in North America. We conclude that forest structure, influenced by climatic drivers, is essential for predicting the response of biodiversity with future shifts in climatic regimes.

Predicting vulnerability of forest patches to invasion by non-native plants for landscape scale management.

As a leading cause of forest health degradation, non-native invasive plant species are a key focus for many forest management and conservation efforts. These efforts come at a high price for resource-limited agencies and organizations making cost-effectiveness an important objective of invasion response plans. In this paper, we present an approach to guide the prioritization of locations for invasion management using species distribution models that account for the non-equilibrium of invasive species distributions and use readily available land use data as the primary explanatory variables. This approach takes advantage of the relatively high spatial resolution, as well as the broad, continuous geographic coverage, of land use data to provide results at a landscape scale relevant to practitioners responsible for invasive species management. In our example from northern Virginia, we simultaneously modeled a suite of invasive plant species to identify common indicators of invasion. We found that the proportions of surrounding non-forested land use types (grasses, crops, and development) were the most common and strongest indicators of invasion risk. These outcomes can guide managers of large protected areas to focus on major divides between forest and non-forest land over linear disturbances. We also found useful species-specific traits that can inform specific management actions. Additionally, we demonstrate through a case study how organizations that manage multiple smaller properties can take advantage of the projected distribution maps when considering acquiring or administering properties.

Counting the Capital's cats: Estimating drivers of abundance of free-roaming cats with a novel hierarchical model.

Free-roaming cats are a conservation concern in many areas but identifying their impacts and developing mitigation strategies requires a robust understanding of their distribution and density patterns. Urban and residential areas may be especially relevant in this process because free-roaming cats are abundant in these anthropogenic landscapes. Here, we estimate the occupancy and density of free-roaming cats in Washington D.C. and relate these metrics to known landscape and social factors. We conducted an extended camera trap survey of public and private spaces across D.C. and analyzed data collected from 1483 camera deployments from 2018 to 2020. We estimated citywide cat distribution by fitting hierarchical occupancy models and further estimated cat abundance using a novel random thinning spatial capture-recapture model that allows for the use of photos that can and cannot be identified to individual. Within this model, we utilized individual covariates that provided identity exclusions between photos of unidentifiable cats with inconsistent coat patterns, thus increasing the precision of abundance estimates. This combined model also allowed for unbiased estimation of density when animals cannot be identified to individual at the same rate as for free-roaming cats whose identifiability depended on their coat characteristics. Cat occupancy and abundance declined with increasing distance from residential areas, an effect that was more pronounced in wealthier neighborhoods. There was noteworthy absence of cats detected in larger public spaces and forests. Realized densities ranged from 0.02 to 1.75 cats/ha in sampled areas, resulting in a district-wide estimate of ~7296 free-roaming cats. Ninety percent of cat detections lacked collars and nearly 35% of known individuals were ear-tipped, indicative of district Trap-Neuter-Return (TNR) programs. These results suggest that we mainly sampled and estimated the unowned cat subpopulation, such that indoor/outdoor housecats were not well represented. The precise estimation of cat population densities is difficult due to the varied behavior of subpopulations within free-roaming cat populations (housecats, stray and feral cats), but our methods provide a first step in establishing citywide baselines to inform data-driven management plans for free-roaming cats in urban environments.

Tree height and leaf drought tolerance traits shape growth responses across droughts in a temperate broadleaf forest.

As climate change drives increased drought in many forested regions, mechanistic understanding of the factors conferring drought tolerance in trees is increasingly important. The dendrochronological record provides a window through which we can understand how tree size and traits shape growth responses to droughts. We analyzed tree-ring records for 12 species in a broadleaf deciduous forest in Virginia (USA) to test hypotheses for how tree height, microenvironment characteristics, and species' traits shaped drought responses across the three strongest regional droughts over a 60-yr period. Drought tolerance (resistance, recovery, and resilience) decreased with tree height, which was strongly correlated with exposure to higher solar radiation and evaporative demand. The potentially greater rooting volume of larger trees did not confer a resistance advantage, but marginally increased recovery and resilience, in sites with low topographic wetness index. Drought tolerance was greater among species whose leaves lost turgor (wilted) at more negative water potentials and experienced less shrinkage upon desiccation. The tree-ring record reveals that tree height and leaf drought tolerance traits influenced growth responses during and after significant droughts in the meteorological record. As climate change-induced droughts intensify, tall trees with drought-sensitive leaves will be most vulnerable to immediate and longer-term growth reductions.

Temporal population variability in local forest communities has mixed effects on tree species richness across a latitudinal gradient.

Among the local processes that determine species diversity in ecological communities, fluctuation-dependent mechanisms that are mediated by temporal variability in the abundances of species populations have received significant attention. Higher temporal variability in the abundances of species populations can increase the strength of temporal niche partitioning but can also increase the risk of species extinctions, such that the net effect on species coexistence is not clear. We quantified this temporal population variability for tree species in 21 large forest plots and found much greater variability for higher latitude plots with fewer tree species. A fitted mechanistic model showed that among the forest plots, the net effect of temporal population variability on tree species coexistence was usually negative, but sometimes positive or negligible. Therefore, our results suggest that temporal variability in the abundances of species populations has no clear negative or positive contribution to the latitudinal gradient in tree species richness.

Effectiveness of management zoning designed for flagship species in protecting sympatric species.

Flagship species have been used widely as umbrella species (i.e., species with large home range whose protection often provides protection for sympatric species) in the management of China's nature reserves. This conflation of flagship and umbrella species is best represented by the giant panda (Ailuropoda melanoleuca) and other large, endangered mammals designated as conservation targets in site selection and planning of reserves. Few empirical studies have tested the effectiveness of flagship species as surrogates for a broader range of sympatric species. Using extensive camera-trap data, we examined the effectiveness of management zones designated to protect flagship (target) species in conserving sympatric species in 4 wildlife reserves (Gutianshan, Changqing, Laohegou, and Wolong). We tested whether the progression from peripheral to core zones was associated with an increasing habitat association for both target and sympatric species. The distribution patterns of the study species across the zones in each reserve indicated a disparity between management zones and the species' habitat requirements. Management zone was included in the final model for all target species, and most of them had higher occurrence in core zones relative to less-protected zones, but zone was not a predictor for most of the sympatric species. When management zone was associated with the occurrence of sympatric species, threatened species generally had higher detections in core zones, whereas common species had higher detections outside of the core zone. Our results suggested that reserve planning based on flagship species does not adequately protect sympatric species due to their specialized habitat requirements. We recommend re-examining the effectiveness of management zoning and urge a multispecies and reserve-wide monitoring plan to improve protection of China's wildlife.

Efectividad del Manejo de Zonación Diseñada para Especies Bandera en la Protección de Especies Simpátricas Resumen Las especies bandera se han utilizado ampliamente como especies paraguas (es decir, especies con una extensa distribución cuya protección frecuentemente proporciona protección para especies simpátricas) en el manejo de las reservas naturales de China. Esta combinación de especie bandera y especie paraguas tiene su mejor representación en el panda gigante (Ailuropoda melanoleuca) y otros grandes mamíferos en peligro de extinción designados como objetivos de conservación en la selección de sitio y la planeación de reservas. Pocos estudios empíricos han probado la efectividad de las especies bandera como sustitutas de una especie simpátrica con una distribución más amplia. Con datos numerosos de cámaras trampa, examinamos la efectividad de las zonas de manejo designadas para proteger a las especies bandera (objetivo) en la conservación de especies simpátricas dentro de cuatro reservas de fauna (Gutianshan, Changqing, Laohegou y Wolong). Probamos si la progresión de las zonas periféricas hacia las zonas núcleo estaba relacionada con asociaciones crecientes de hábitat tanto para las especies objetivo como para las simpátricas. Los patrones de distribución del estudio de especies a lo largo de las zonas en cada reserva indicaron una disparidad entre las zonas de manejo y los requerimientos de hábitat de las especies. La zona de manejo se incluyó dentro del modelo final para todas las especies objetivo y la mayoría de ellas tuvo una presencia más alta en las zonas núcleo en relación con las zonas de menor protección. Cuando la zona de manejo se asoció a la presencia de especies simpátricas, las especies amenazadas generalmente tuvieron un mayor número de detecciones en la zona núcleo, mientras que las especies comunes tuvieron un número más elevado de detecciones fuera de la zona núcleo. Nuestros resultados sugieren que la planeación de reservas basada en las especies bandera no protege adecuadamente a las especies simpátricas debido a sus requerimientos especializados de hábitat. Recomendamos una reexaminación de la efectividad del manejo de zonación y urgimos que se genere un plan de monitoreo multiespecífico en toda la reserva para mejorar la protección de la fauna en China.

Political transition and emergent forest-conservation issues in Myanmar.

Political and economic transitions have had substantial impacts on forest conservation. Where transitions are underway or anticipated, historical precedent and methods for systematically assessing future trends should be used to anticipate likely threats to forest conservation and design appropriate and prescient policy measures to counteract them. Myanmar is transitioning from an authoritarian, centralized state with a highly regulated economy to a more decentralized and economically liberal democracy and is working to end a long-running civil war. With these transitions in mind, we used a horizon-scanning approach to assess the 40 emerging issues most affecting Myanmar's forests, including internal conflict, land-tenure insecurity, large-scale agricultural development, demise of state timber enterprises, shortfalls in government revenue and capacity, and opening of new deforestation frontiers with new roads, mines, and hydroelectric dams. Averting these threats will require, for example, overhauling governance models, building capacity, improving infrastructure- and energy-project planning, and reforming land-tenure and environmental-protection laws. Although challenges to conservation in Myanmar are daunting, the political transition offers an opportunity for conservationists and researchers to help shape a future that enhances Myanmar's social, economic, and environmental potential while learning and applying lessons from other countries. Our approach and results are relevant to other countries undergoing similar transitions.

A two-species occupancy model accommodating simultaneous spatial and interspecific dependence.

Occupancy models are popular for estimating the probability a site is occupied by a species of interest when detection is imperfect. Occupancy models have been extended to account for interacting species and spatial dependence but cannot presently allow both factors to act simultaneously. We propose a two-species occupancy model that accommodates both interspecific and spatial dependence. We use a point-referenced multivariate hierarchical spatial model to account for both spatial and interspecific dependence. We model spatial random effects with predictive process models and use probit regression to improve efficiency of posterior sampling. We model occupancy probabilities of red fox (Vulpes vulpes) and coyote (Canis latrans) with camera trap data collected from six mid-Atlantic states in the eastern United States. We fit four models comprising a fully factorial combination of spatial and interspecific dependence to two-thirds of camera trapping sites and validated models with the remaining data. Red fox and coyotes each exhibited spatial dependence at distances > 0.8 and 0.4 km, respectively, and exhibited geographic variation in interspecific dependence. Consequently, predictions from the model assuming simultaneous spatial and interspecific dependence best matched test data observations. This application highlights the utility of simultaneously accounting for spatial and interspecific dependence.

Ungulates' Behavioral Responses to Humans as an Apex Predator in a Hunting-Prohibited Area of China.

Large mammals can perceive humans as predators and therefore adjust their behavior to achieve coexistence with humans. However, lack of research at sites with low hunting intensity limits our understanding of how behavioral responses of animals adapt to different predation risks by humans. At Heshun County in North China, where hunting has been banned for over three decades and only low-intensity poaching exists, we exposed two large ungulates (Siberian roe deer Capreolus pygarus and wild boar Sus scrofa) to the sounds of humans, an extant predator (leopard Panthera pardus) and a control (wind), and examined their flight responses and detection probabilities when hearing different type of sounds. Both species showed higher flight probabilities when hearing human vocalization than wind, and wild boar were even more likely to flee upon hearing human vocalization than leopard roar, suggesting the behavioral response to humans can equal or exceed that of large carnivores in these two ungulates even in an area without hunting practices. Recorded sounds had no effect on detection probability of both ungulates. Additionally, with repeated exposure to sounds, regardless of treatment, roe deer were less likely to flee and wild boars were more likely to be detected, indicating a habituation-type response to sound stimuli. We speculate that the immediate flight behavior rather than shifts in habitat use of the two species reflect the low hunting/poaching pressure at our study site and suggest further examination of physiological status and demographic dynamics of the study species to understand human influence on their long-term persistence.

Mycorrhizal feedbacks influence global forest structure and diversity.

One mechanism proposed to explain high species diversity in tropical systems is strong negative conspecific density dependence (CDD), which reduces recruitment of juveniles in proximity to conspecific adult plants. Although evidence shows that plant-specific soil pathogens can drive negative CDD, trees also form key mutualisms with mycorrhizal fungi, which may counteract these effects. Across 43 large-scale forest plots worldwide, we tested whether ectomycorrhizal tree species exhibit weaker negative CDD than arbuscular mycorrhizal tree species. We further tested for conmycorrhizal density dependence (CMDD) to test for benefit from shared mutualists. We found that the strength of CDD varies systematically with mycorrhizal type, with ectomycorrhizal tree species exhibiting higher sapling densities with increasing adult densities than arbuscular mycorrhizal tree species. Moreover, we found evidence of positive CMDD for tree species of both mycorrhizal types. Collectively, these findings indicate that mycorrhizal interactions likely play a foundational role in global forest diversity patterns and structure.

A global assessment of the impact of individual protected areas on preventing forest loss.

Globally, the number and extent of terrestrial protected areas (PAs) are expanding rapidly. Nonetheless, their impacts on preventing forest loss and the factors influencing the impacts are not well understood, despite the critical roles of forests in biodiversity conservation, provision of ecosystem services, and achievement of the United Nations' Sustainable Development Goals. To address this important knowledge gap, we quantified the impacts of 54,792 PAs worldwide on preventing forest loss from 2000 to 2015, and assessed important landscape and management factors affecting the impacts of PAs. Although the majority (71.4%) of the PAs contributed to preventing forest loss, only 30.5% of forest loss in the PAs have been prevented. PAs with higher rates of forest loss in their surrounding regions, located at lower elevations, within a few hours of travel from the nearest city, with higher agricultural productivity, and permission for fewer human uses were better able to prevent forest loss. Impacts on preventing forest loss were similar regardless of whether the PAs were privately or publicly owned. Our findings highlight the potential benefits of strict protections, involving private entities in the establishment of PAs, and situating PAs in areas exposed to high risks of forest loss to enhance the capacity to combat global forest loss.

Nonconsumptive effects of a generalist ungulate herbivore drive decline of unpalatable forest herbs.

High herbivore pressure is expected to benefit unpalatable species that co-occur with palatable browsed species. However, for five unpalatable understory species we found no evidence of benefit from deer browse. Detailed studies of one species in natural populations, Arisaema triphyllum, revealed surprising changes in its population structure and demography: deer browse level on a palatable species significantly correlates with reduced plant size and seed rain and male-biased sex ratios of co-occurring Arisaema populations. Analyses of individual size in five unpalatable forest plant species in long-term experimental paired deer exclosure/deer access plots corroborate the natural site results; all five species were smaller in deer access plots. Analyses of abiotic variables in natural and experimental sites suggest one potential mechanism for indirect effects of deer. Deer-mediated soil quality declines included increased soil penetration resistance and decreased leaf litter depth, which are known to hinder plant growth. Our results are likely applicable to other unpalatable forest species and have clear consequences for understory biodiversity. Unpalatable plant species in forests experiencing high deer numbers may be in decline along with their palatable neighbors. Our study implicates deer overabundance in the cascade of forest species decline and the urgency of this conservation issue in North America.

Retreat of large carnivores across the giant panda distribution range.

As both a flagship and umbrella species, the giant panda (Ailuropoda melanoleuca) is one of the most heavily invested species in conservation. Here, we report the wide distribution range retreat of the leopard (Panthera pardus, 81% loss), snow leopard (P. uncia, 38%), wolf (Canis lupus, 77%) and dhole (Cuon alpinus, 95%) from protected areas in the giant panda distribution range since the 1960s. The present findings indicate the insufficiency of giant panda conservation for protecting these large carnivore species and suggest that future conservation efforts should target restoring ecosystems with high trophic complexity to facilitate the recovery of large carnivore populations.

Scale-dependent impacts of urban and agricultural land use on nutrients, sediment, and runoff.

We coupled a spatially-explicit land use/land cover (LULC) change model, Dinamica EGO, (Environment for Geoprocessing Objects), with the Chesapeake Bay Watershed Model (CBWM) to project the impact of future LULC change on loading of total nitrogen (TN), total phosphorous (TP) and total suspended solids (TSS) as well as runoff volume in the watersheds surrounding Virginia's Shenandoah National Park in the eastern United States. We allowed for the dynamic transition of four LULC classes, Developed, Forest, Grasses (including both pasture and hayfields) and Crops. Using 2011 as a baseline scenario and observed differences in LULC between 2001 and 2011, we estimated the temporal and spatial patterns of LULC change as influenced by physiographic and socio-economic drivers 50 years in the future (2061). Between transitions of the four LULC classes, the greatest absolute change occurred between the gain in total Developed land and loss in total Forest. New Developed land was driven primarily by distance to existing Developed land and population density. Major findings on the effect of LULC change on watershed model outputs were that: the impact of LULC change on pollutant loading and runoff volume is more pronounced at finer spatial scales; increases in the area of Grasses produced the greatest increase in TP loading, while loss of Forest increased TN, TSS, and runoff volume the most; and land-river segments with a greater proportion of Developed or a smaller proportion of Forest in the 2011 scenario experienced a greater change in runoff than other land-river segments. Results of this study illustrate the potential impact of projected LULC change on nutrient and sediment loads which can adversely impact water quality. Studies like this contribute to a broader understanding of how ecosystem services such as fresh water respond to LULC change, information relevant to those in planning and watershed management.

Mammal communities are larger and more diverse in moderately developed areas.

Developed areas are thought to have low species diversity, low animal abundance, few native predators, and thus low resilience and ecological function. Working with citizen scientist volunteers to survey mammals at 1427 sites across two development gradients (wild-rural-exurban-suburban-urban) and four plot types (large forests, small forest fragments, open areas and residential yards) in the eastern US, we show that developed areas actually had significantly higher or statistically similar mammalian occupancy, relative abundance, richness and diversity compared to wild areas. However, although some animals can thrive in suburbia, conservation of wild areas and preservation of green space within cities are needed to protect sensitive species and to give all species the chance to adapt and persist in the Anthropocene.

A regional assessment of white-tailed deer effects on plant invasion.

Herbivores can profoundly influence plant species assembly, including plant invasion, and resulting community composition. Population increases of native herbivores, e.g. white-tailed deer (Odocoileus virginianus), combined with burgeoning plant invasions raise concerns for native plant diversity and forest regeneration. While individual researchers typically test for the impact of deer on plant invasion at a few sites, the overarching influence of deer on plant invasion across regional scales is unclear. We tested the effects of deer on the abundance and diversity of introduced and native herbaceous and woody plants across 23 white-tailed deer research sites distributed across the east-central and north-eastern USA and representing a wide range of deer densities and invasive plant abundance and identity. Deer access/exclusion or deer population density did not affect introduced plant richness or community-level abundance. Native and total plant species richness, abundance (cover and stem density) and Shannon diversity were lower in deer-access vs. deer-exclusion plots. Among deer-access plots, native species richness, native and total cover, and Shannon diversity (cover) declined as deer density increased. Deer access increased the proportion of introduced species cover (but not of species richness or stem density). As deer density increased, the proportion of introduced species richness, cover and stem density all increased. Because absolute abundance of introduced plants was unaffected by deer, the increase in proportion of introduced plant abundance is likely an indirect effect of deer reducing native cover. Indicator species analysis revealed that deer access favoured three introduced plant species, including Alliaria petiolata and Microstegium vimineum, as well as four native plant species. In contrast, deer exclusion favoured three introduced plant species, including Lonicera japonica and Rosa multiflora, and 15 native plant species. Overall, native deer reduced community diversity, lowering native plant richness and abundance, and benefited certain invasive plants, suggesting pervasive impacts of this keystone herbivore on plant community composition and ecosystem services in native forests across broad swathes of the eastern USA.

Response to Comment on "Plant diversity increases with the strength of negative density dependence at the global scale".

Hülsmann and Hartig suggest that ecological mechanisms other than specialized natural enemies or intraspecific competition contribute to our estimates of conspecific negative density dependence (CNDD). To address their concern, we show that our results are not the result of a methodological artifact and present a null-model analysis that demonstrates that our original findings-(i) stronger CNDD at tropical relative to temperate latitudes and (ii) a latitudinal shift in the relationship between CNDD and species abundance-persist even after controlling for other processes that might influence spatial relationships between adults and recruits.