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Altered climate leads to positive density‐dependent feedbacks in a tropical wet forest

Climate change is predicted to result in warmer and drier Neotropical forests relative to current conditions. Negative density‐dependent feedbacks, mediated by natural enemies, are key to maintaining the high diversity of tree species found in the tropics, yet we have little understanding of how projected changes in climate are likely to affect these critical controls. Over 3 years, we evaluated the effects of a natural drought and in situ experimental warming on density‐dependent feedbacks on seedling demography in a wet tropical forest in Puerto Rico. In the +4°C warming treatment, we found that seedling survival increased with increasing density of the same species (conspecific). These positive density‐dependent feedbacks were not associated with a decrease in aboveground natural enemy pressure. If positive density‐dependent feedbacks are not transient, the diversity of tropical wet forests, which may rely on negative density dependence to drive diversity, could decline in a future warmer, drier world.

Puerto Rico↗

Joint effects of climate, tree size, and year on annual tree growth derived using tree-ring records of ten globally distributed forests

Tree rings provide an invaluable long-term record for understanding how climate and other drivers shape tree growth and forest productivity. However, conventional tree-ring analysis methods were not designed to simultaneously account for the effects of climate, tree size, and other drivers on individual growth, which has limited the potential to use tree rings to understand forest productivity, its climate sensitivity, and its global change responses. Here, we develop and apply a new method to simultaneously model non-linear effects of primary climate drivers, reconstructed tree diameter (DBH), and year in generalized least squares models that account for the temporal autocorrelation inherent to each individual tree’s growth. We analyze data from 3811 trees representing 40 species at 10 globally distributed sites, showing that precipitation, temperature, DBH, and calendar year have additively, and often interactively, influenced annual growth over the past 120 years. Growth responses were predominantly positive to precipitation (usually over ≥ 3-month seasonal windows) and negative to temperature (usually over ≤ 3-month seasonal windows), with both included in 78% of top models, and with non-linear responses prevalent (63% of relationships). Climate sensitivity commonly varied with DBH (44% of cases tested). Trends in ring width at small DBH were linked to the light environment under which trees established, but basal area or biomass increments consistently peaked at intermediate DBH and declined thereafter. Accounting for climate and DBH, growth rate declined over time for 92% of species in secondary or disturbed stands, whereas growth trends were mixed in older forests. These trends were largely attributable to stand dynamics as cohorts and stands age, which remain challenging to disentangle from global change drivers. By providing a parsimonious approach for characterizing multiple interacting drivers of tree growth, our method reveals a more complete picture of the factors influencing growth than has previously been possible.

Indiana, Massachusetts, New Mexico, Nebraska, Utah↗

Biogeography of fire regimes in western US conifer forests: A trait-based approach

Aim Functional traits are a critical link between species distributions and the ecosystem processes that structure those species’ niches. Concurrent increases in the availability of functional trait data and our ability to model species distributions present an opportunity to develop functional trait biogeography, i.e. the mapping of functional traits across space. Functional trait biogeography can improve process-based predictions about the resistance of certain species assemblages to changing environmental conditions across landscape scales. We illustrate this concept by developing the first trait-based, quantitative ranking of fire resistance (adult tree survival) in North American conifer species, and mapping that fire resistance across space. Location and Time period Western Continental United States, present-day. Major taxa studied 29 common conifer tree species. Methods We compiled six traits for each species: three relating to tree morphology and three relating to litter flammability. We combined these traits into a single fire resistance score, and used community-weighted averaging to estimate the fire resistance scores of different forest communities, using interpolated species distribution and relative abundance data. Results Species associated with historically frequent fire have high fire resistance scores (e.g., Pinus ponderosa), reflected by thick bark, tall crowns, and flammable litter. Species associated with subalpine or arid conditions have low fire resistance scores (e.g., Picea engelmannii and Pinus edulis), reflected by thin bark, short stature, poor self-pruning and low litter flammability. A map of forest community fire resistance across the western US reveals agreement with independent assessments of historical fire regimes, while also identifying areas where community-wide species traits may be mismatched with historical fire regimes. Main conclusions Quantifying the functional traits that confer resistance to tree-killing fire provides a direct link between ecosystem disturbance and community resistance. Understanding this link is critical to evaluating long-term resilience of different forest types under dynamic fire regimes. Our work represents the first known spatial representation of fire-resistance traits at a regional scale, and as such provides a link between functional traits and biogeography relevant to a critical ecosystem process.

Global Ecology and Biogeography↗

Mapping of forested wetland: Use of Seasat radar images to complement conventional sources

Distinguishing forested wetland from dry forest using aerial photographs is handicapped because photographs often do not reveal the presence of water below tree canopies. Radar images obtained by the Seasat satellite reveal forested wetland as highly reflective patterns on the coastal plain between Maryland and Florida. Seasat radar images may complement aerial photographs for compiling maps of wetland. A test with experienced photointerpreters revealed that interpretation accuracy was significantly higher when using Seasat radar images than when using only conventional sources.

Florida, Georgia, Maryland, North Carolina, South ↗

A double-observer method for reducing bias in faecal pellet surveys of forest ungulates

1. Faecal surveys are used widely to study variations in abundance and distribution of forest-dwelling mammals when direct enumeration is not feasible. The utility of faecal indices of abundance is limited, however, by observational bias and variation in faecal disappearance rates that obscure their relationship to population size. We developed methods to reduce variability in faecal surveys and improve reliability of faecal indices. 2. We used double-observer transect sampling to estimate observational bias of faecal surveys of Roosevelt elk Cervus elaphus roosevelti and Columbian black-tailed deer Odocoileus hemionus columbianus in Olympic National Park, Washington, USA. We also modelled differences in counts of faecal groups obtained from paired cleared and uncleared transect segments as a means to adjust standing crop faecal counts for a standard accumulation interval and to reduce bias resulting from variable decay rates. 3. Estimated detection probabilities of faecal groups ranged from < 0.2-1.0 depending upon the observer, whether the faecal group was from elk or deer, faecal group size, distance of the faecal group from the sampling transect, ground vegetation cover, and the interaction between faecal group size and distance from the transect. 4. Models of plot-clearing effects indicated that standing crop counts of deer faecal groups required 34% reduction on flat terrain and 53% reduction on sloping terrain to represent faeces accumulated over a standard 100-day interval, whereas counts of elk faecal groups required 0% and 46% reductions on flat and sloping terrain, respectively. 5. Synthesis and applications. Double-observer transect sampling provides a cost-effective means of reducing observational bias and variation in faecal decay rates that obscure the interpretation of faecal indices of large mammal abundance. Given the variation we observed in observational bias of faecal surveys and persistence of faeces, we emphasize the need for future researchers to account for these comparatively manageable sources of bias before comparing faecal indices spatially or temporally. Double-observer sampling methods are readily adaptable to study variations in faecal indices of large mammals at the scale of the large forest reserve, natural area, or other forested regions when direct estimation of populations is problematic. ?? 2008 The Authors.

Journal of Applied Ecology↗

Impacts of forest fragmentation on species richness: A hierarchical approach to community modelling

1. Species richness is often used as a tool for prioritizing conservation action. One method for predicting richness and other summaries of community structure is to develop species-specific models of occurrence probability based on habitat or landscape characteristics. However, this approach can be challenging for rare or elusive species for which survey data are often sparse. 2. Recent developments have allowed for improved inference about community structure based on species-specific models of occurrence probability, integrated within a hierarchical modelling framework. This framework offers advantages to inference about species richness over typical approaches by accounting for both species-level effects and the aggregated effects of landscape composition on a community as a whole, thus leading to increased precision in estimates of species richness by improving occupancy estimates for all species, including those that were observed infrequently. 3. We developed a hierarchical model to assess the community response of breeding birds in the Hudson River Valley, New York, to habitat fragmentation and analysed the model using a Bayesian approach. 4. The model was designed to estimate species-specific occurrence and the effects of fragment area and edge (as measured through the perimeter and the perimeter/area ratio, P/A), while accounting for imperfect detection of species. 5. We used the fitted model to make predictions of species richness within forest fragments of variable morphology. The model revealed that species richness of the observed bird community was maximized in small forest fragments with a high P/A. However, the number of forest interior species, a subset of the community with high conservation value, was maximized in large fragments with low P/A. 6. Synthesis and applications. Our results demonstrate the importance of understanding the responses of both individual, and groups of species, to environmental heterogeneity while illustrating the utility of hierarchical models for inference about species richness for conservation. This framework can be used to investigate the impacts of land-use change and fragmentation on species or assemblage richness, and to further understand trade-offs in species-specific occupancy probabilities associated with landscape variability.

New York↗

Using ecological function to develop recovery criteria for depleted species: Sea otters and kelp forests in the Aleutian archipelago

Recovery criteria for depleted species or populations normally are based on demographic measures, the goal being to maintain enough individuals over a sufficiently large area to assure a socially tolerable risk of future extinction. Such demographically based recovery criteria may be insufficient to restore the functional roles of strongly interacting species. We explored the idea of developing a recovery criterion for sea otters (Enhydra lutris) in the Aleutian archipelago on the basis of their keystone role in kelp forest ecosystems. We surveyed sea otters and rocky reef habitats at 34 island-time combinations. The system nearly always existed in either a kelp-dominated or deforested phase state, which was predictable from sea otter density. We used a resampling analysis of these data to show that the phase state at any particular island can be determined at 95% probability of correct classification with information from as few as six sites. When sea otter population status (and thus the phase state of the kelp forest) was allowed to vary randomly among islands, just 15 islands had to be sampled to estimate the true proportion that were kelp dominated (within 10%) with 90% confidence. We conclude that kelp forest phase state is a more appropriate, sensitive, and cost-effective measure of sea otter recovery than the more traditional demographically based metrics, and we suggest that similar approaches have broad potential utility in establishing recovery criteria for depleted populations of other functionally important species.

Conservation Biology↗

Balancing forest-regeneration probabilities and maintenance costs in dry grasslands of high conservation priority

Abandonment of agricultural land has resulted in forest regeneration in species-rich dry grasslands across European mountain regions and threatens conservation efforts in this vegetation type. To support national conservation strategies, we used a site-selection algorithm (MARXAN) to find optimum sets of floristic regions (reporting units) that contain grasslands of high conservation priority. We sought optimum sets that would accommodate 136 important dry-grassland species and that would minimize forest regeneration and costs of management needed to forestall predicted forest regeneration. We did not consider other conservation elements of dry grasslands, such as animal species richness, cultural heritage, and changes due to climate change. Optimal sets that included 95&ndash;100% of the dry grassland species encompassed an average of 56&ndash;59 floristic regions (standard deviation, SD 5). This is about 15% of approximately 400 floristic regions that contain dry-grassland sites and translates to 4800&ndash;5300 ha of dry grassland out of a total of approximately 23,000 ha for the entire study area. Projected costs to manage the grasslands in these optimum sets ranged from CHF (Swiss francs) 5.2 to 6.0 million/year. This is only 15&ndash;20% of the current total estimated cost of approximately CHF30&ndash;45 million/year required if all dry grasslands were to be protected. The grasslands of the optimal sets may be viewed as core sites in a national conservation strategy.

Conservation Biology↗

Structural complexity and land-surface energy exchange along a gradient from arctic tundra to boreal forest

Question: Current climate changes in the Alaskan Arctic, which are characterized by increases in temperature and length of growing season, could alter vegetation structure, especially through increases in shrub cover or the movement of treeline. These changes in vegetation structure have consequences for the climate system. What is the relationship between structural complexity and partitioning of surface energy along a gradient from tundra through shrub tundra to closed canopy forest? Location: Arctic tundra-boreal forest transition in the Alaskan Arctic. Methods: Along this gradient of increasing canopy complexity, we measured key vegetation characteristics, including community composition, biomass, cover, height, leaf area index and stem area index. We relate these vegetation characteristics to albedo and the partitioning of net radiation into ground, latent, and sensible heating fluxes. Results: Canopy complexity increased along the sequence from tundra to forest due to the addition of new plant functional types. This led to non-linear changes in biomass, cover, and height in the understory. The increased canopy complexity resulted in reduced ground heat fluxes, relatively conserved latent heat fluxes and increased sensible heat fluxes. The localized warming associated with increased sensible heating over more complex canopies may amplify regional warming, causing further vegetation change in the Alaskan Arctic.

Alaska↗

Recovery of sediment characteristics in moraine, headwater streams of northern Minnesota after forest harvest

We investigated the recovery of sediment characteristics in four moraine, headwater streams in north-central Minnesota after forest harvest. We examined changes in fine sediment levels from 1997 (preharvest) to 2007 (10 years postharvest) at study plots with upland clear felling and riparian thinning, using canopy cover, proportion of unstable banks, surficial fine substrates, residual pool depth, and streambed depth of refusal as response variables. Basin-scale year effects were significant ( p < 0.001) for all responses when evaluated by repeated-measures ANOVAs. Throughout the study area, unstable banks increased for several years postharvest, coinciding with an increase in windthrow and fine sediment. Increased unstable banks may have been caused by forest harvest equipment, increased windthrow and exposure of rootwads, or increased discharge and bank scour. Fine sediment in the channels did not recover by summer 2007, even though canopy cover and unstable banks had returned to 1997 levels. After several storm events in fall 2007, 10 years after the initial sediment input, fine sediment was flushed from the channels and returned to 1997 levels. Although our study design did not discern the source of the initial sediment inputs (e.g., forest harvest, road crossings, other natural causes), we have shown that moraine, headwater streams can require an extended period (up to 10 years) and enabling event (e.g., high storm flows) to recover from large inputs of fine sediment.

Minnesota↗

Activity patterns of Hawaiian forest birds in a fragmented and continuous landscape

Activity patterns are a key component of avian life history and behavior, and decisions about how activity periods are structured can have important fitness implications. Despite the importance of activity patterns, individual variability in activity is poorly studied in small birds. We used automated radio telemetry to track Hawaiian forest birds from three dietary guilds in a continuous forest, and in a similar forest that was naturally fragmented in the 1800s by lava flows but supports stable populations of native birds. We generated kernel density profiles of activity (KDEs) over the full tracking period of each bird and used the KDEs to derive seven activity metrics that characterized daily activity patterns. We identified three distinct activity strategies using a cluster analysis of the activity metrics and assessed day-to-day variability in activity strategy to understand intraindividual consistency in activity behavior. We also evaluated the influence of landscape structure, dietary guild and sex on the suite of activity metrics using constrained ordination. Most birds (61%) exhibited high levels of sustained activity throughout the day, while 23% showed irregular activity most of the day followed by a peak in the late afternoon, and 16% were mainly active in the morning. Though activity patterns varied by sex and dietary guild, landscape structure had the strongest influence on activity patterns. Birds sustained high levels of activity throughout the day in the fragmented landscape regardless of dietary guild, suggesting they needed to be more active to obtain necessary resources. However, activity patterns of birds in the fragmented landscape varied more day-to-day, indicating they retained a method for adjusting behavior to match internal and external conditions. This study reveals how a diverse suite of tropical birds has adapted to long-term fragmentation in Hawaiʻi and demonstrates the utility of automated radio telemetry for studying the activity strategies of birds.

Hawaii↗

Whole‐genome resequencing reveals persistence of forest‐associated mammals in Late Pleistocene refugia along North America’s North Pacific Coast

Aim Numerous glacial refugia have been hypothesized along North America's North Pacific Coast that may have increased divergence of refugial taxa, leading to elevated endemism and subsequently clustered hybrid zones following deglaciation. The locations and community composition of these ice‐free areas remains controversial, but whole‐genome sequences now enable detailed analysis of the demographic and evolutionary histories of refugial taxa. Here, we use genomic data to test spatial and temporal processes of diversification among martens with respect to the Coastal Refugium Hypothesis, to understand the role of climate cycling in shaping diversity across complex landscapes. Location North America and North Pacific Coast archipelagos. Taxon North American martens ( Martes ). Methods Short‐read whole‐genome resequencing data were generated for 11 martens: four M. americana , four M. caurina , two hybrids, and one outgroup ( Martes zibellina ). Sampling was representative of known genetic clades within New World martens, including sampling within insular and continental hybrid zones and along the North Pacific Coast (five island populations). ADMIXTURE , F‐statistics, and D ‐statistics (ABBA‐BABA) were used to identify introgression and infer directionality. Heterozygosity densities, estimated via PSMC, were used to characterize historical demography at and below the species level to infer refugial and colonization processes. Results Forest‐associated Pacific martens ( M. caurina ) are divided into distinct insular and continental clades consistent with the Coastal Refugium Hypothesis. There was no evidence of introgression on islands that received historical translocations of American pine martens ( M. americana ), but introgression was detected in two active zones of secondary contact: one insular and one continental. Only early‐generational hybrids were identified across multiple hybrid zones, a pattern consistent with potential genetic swamping of M. caurina by M. americana . Main conclusions Despite an incomplete fossil record, genomic evidence supports the persistence of forest‐associated martens, likely the insular Pacific marten lineage, along the western edges of the Alexander Archipelago during the Last Glacial Maximum. This discovery informs our understanding of refugial paleoenvironments, critical to interpreting refugial timing, duration, and community composition. Genomic reevaluations of other taxa along North America's North Pacific Coast may yield new and deeper perspectives on the history of refugial forest communities and the role of dynamic climate shifts in shaping high‐latitude diversity across complex insular landscapes.

Alaska, British Columbia, California, Oregon, Wash↗

Individual species–area relationships in temperate coniferous forests

Questions What drives individual species–area relationships in temperate coniferous forests? Location Two 25.6‐ha forest plots on the Pacific Slope of North America, one in California, and one in Washington State. Methods We mapped all trees ≥1 cm in diameter and examined tree species diversity of their local neighbourhoods by calculating the individual species–area relationship for each species and for each of three diameter classes (saplings, mature trees and large‐diameter trees). Results In the California plot, small trees in four of the five major species occurred in neighbourhoods with higher levels of diversity than would be expected at random. In the Washington plot, small trees for four of five abundant species had neighbourhoods with lower than expected diversity at distances ≤5 m for small trees. However, at distances >5 m, all five species showed higher than expected diversity in their neighbourhoods. Larger trees at both plots tended to occur in neighbourhoods with lower than expected diversity, and no large‐diameter focal species had neighbourhoods with higher than expected diversity. Conclusion Diversity and co‐existence in temperate conifer‐dominated forests do not appear to be the result of random processes. Competitive interactions appear to dominate for the largest trees of most species, resulting in neighbourhoods with lower diversity. For smaller trees, we suggest that a positive response to environmental heterogeneity is the likely driver of neighbourhoods with higher than expected diversity, although we cannot rule out the possibility that facilitation or conspecific negative density dependence (CNDD) also play a role.

Journal of Vegetation Science↗

Monitoring global tree mortality patterns and trends. Report from the VW symposium ‘Crossing scales and disciplines to identify global trends of tree mortality as indicators of forest health’

From the 21stto the 23rdJune 2017, the Herrenhausen castle inHannover/Germany hosted a diverse and large crowd with morethan 70 tree physiologists, forest ecologists, forest inventoryexperts, remote-sensing scientists, and vegetation modele rs. Par-ticipants from six continent s and from more than 20 countriesgathered to discuss how to improve the scientific determination ofglobal-scale patterns, drivers, and trends of a threatening phe-nomenon: the apparent emergence of recent widespread treemortality events in diverse forests around the world.Continuin g the theme of a workshop held at the Max-PlanckInstitute for Biogeochemistry in Jena (Germany) in 2014 (Hart-mann et al., 2015), the Hanover meeting intended to developapproaches, tools and collaborative actions to accelerate progress inaddressing regional patterns and trends of tree mortality (Williamset al., 2013). Over the last decade climate change related treemortality events have been increasingly reported around the globe(van Mantgem et al., 2009; Carnicer et al., 2011; Peng et al., 2011;Brienen et al., 2015), but to what degree this is a global trend,amplifying under increasing climate change, remains uncertain.

New Phytologist↗

Experimental warming across a tropical forest canopy height gradient reveals minimal photosynthetic and respiratory acclimation

Tropical forest canopies cycle vast amounts of carbon, yet we still have a limited understanding of how these critical ecosystems will respond to climate warming. We implemented in situ leaf-level + 3°C experimental warming from the understory to the upper canopy of two Puerto Rican tropical tree species, Guarea guidonia and Ocotea sintenisii . After approximately 1 month of continuous warming, we assessed adjustments in photosynthesis, chlorophyll fluorescence, stomatal conductance, leaf traits and foliar respiration. Warming did not alter net photosynthetic temperature response for either species; however, the optimum temperature of Ocotea understory leaf photosynthetic electron transport shifted upward. There was no Ocotea respiratory treatment effect, while Guarea respiratory temperature sensitivity ( Q 10 ) was down-regulated in heated leaves. The optimum temperatures for photosynthesis ( T opt ) decreased 3–5°C from understory to the highest canopy position, perhaps due to upper canopy stomatal conductance limitations. Guarea upper canopy T opt was similar to the mean daytime temperatures, while Ocotea canopy leaves often operated above T opt . With minimal acclimation to warmer temperatures in the upper canopy, further warming could put these forests at risk of reduced CO 2 uptake, which could weaken the overall carbon sink strength of this tropical forest.

Plant, Cell, and Environment↗

Regeneration and invasion of cottonwood riparian forest following wildfire

Populus deltoides is considered to be a weak resprouter and highly susceptible to wildfire, but few post-wildfire studies have tracked P. deltoides response and resprouting within the Great Plains of North America. Following a wildfire in southwestern Kansas, U.S.A., we surveyed burned and unburned areas of a cottonwood riparian forest along the Cimarron River that included a major understory invader, tamarisk ( Tamarix ramosissima Ledeb.). We tested the following hypotheses, which are consistent with the current understanding of P. deltoides response to wildfire in the Great Plains: (1) regeneration of P. deltoides will be low in areas burned by the wildfire; (2) the number of dead P. deltoides individuals will be greater in the wildfire than unburned areas; and (3) tamarisk regeneration will be higher than P. deltoides regeneration in the wildfire areas because tamarisk is considered a stronger resprouter. We found evidence contrary to two of our hypotheses 3 years following the wildfire. (1) P. deltoides regeneration was high following the wildfire, averaging 692 individuals/ha. (2) The number of dead mature cottonwood trees was greater in wildfire plots than in unburned plots. (3) There was more P. deltoides regeneration than tamarisk regeneration following wildfire. These findings, which diverge from the majority of studies examining P. deltoides regeneration in the Great Plains, suggest that differing local environmental and forest stand conditions, coupled with the timing and intensity of the fire, could be important determinants of riparian forest species' responses to wildfire.

Kansas↗

Successful restoration of Metrosideros polymorpha (ʻōhiʻa) is possible in forest sites with active Rapid ‘Ōhiʻa Death infections

Rapid ʻŌhiʻa Death (ROD), caused by the fungal pathogen Ceratocystis , is killing large numbers of ʻōhiʻa trees ( Metrosideros polymorpha ) in Hawaiʻi. ʻŌhiʻa are a dominant tree in Hawaiian forests, have a range that goes from arid to wet forest climates, and are important for endangered species habitat and ecosystem function. To test whether actively planting ʻōhiʻa seedlings is a viable restoration strategy in areas with high ROD mortality, we planted ʻōhiʻa in a ROD‐affected forest and crossed this with weeding and fencing treatments to compare ROD mortality to other stressors. We also tested for viable Ceratocystis spores in soils around planting areas. We found that seedlings were more likely to die in unweeded and unfenced treatments than controls. Although viable Ceratocystis spores were found in soil, none of the 41 dead seedlings tested positive for Ceratocystis . This indicates that competition from exotic plants and exotic feral ungulate damage are more likely to kill seedlings than ROD within the first year after planting.

Hawaii↗

Warming and earlier spring increase Western U.S. forest wildfire activity

Western United States forest wildfire activity is widely thought to have increased in recent decades, yet neither the extent of recent changes nor the degree to which climate may be driving regional changes in wildfire has been systematically documented. Much of the public and scientific discussion of changes in western United States wildfire has focused instead on the effects of 19th- and 20th-century land-use history. We compiled a comprehensive database of large wildfires in western United States forests since 1970 and compared it with hydroclimatic and land-surface data. Here, we show that large wildfire activity increased suddenly and markedly in the mid-1980s, with higher large-wildfire frequency, longer wildfire durations, and longer wildfire seasons. The greatest increases occurred in mid-elevation, Northern Rockies forests, where land-use histories have relatively little effect on fire risks and are strongly associated with increased spring and summer temperatures and an earlier spring snowmelt.

Science↗