Saproxylic beetles' morphological traits and higher trophic guilds indicate boreal forest naturalness.

Forests contribute to numerous ecosystem functions and services and contain a large proportion of terrestrial biodiversity, but they are being negatively impaced by anthropogenic activities. Forests that have never been clear-cut and have old growth characteristics, termed "near-natural," often harbor different and richer species assemblages than managed forests. Alternative management strategies may be able to balance the needs of biodiversity with the demands of forestry, but evaluation efforts are limited by the challenges of measuring biodiversity. Species richness is frequently used as a simple measure of biodiversity, but research indicates that it may not adequately capture community-level changes. Alternatively, trait-based measures of biodiversity may prove to be useful, but research is lacking. In this paper, we use a large dataset that includes 339 obligate saproxylic beetle species collected over a decade in the boreal region throughout southern Norway to: (1) establish if there is a difference in beetle community composition between near-natural and managed forests; and (2) determine which measures of beetle biodiversity best indicate forest naturalness. We arranged the sites in an ordination space and tested for differences in community composition between these forest types. We also tested different measures of biodiversity to determine which were the most predictive of forest naturalness. We found a clear difference in community composition between near-natural and managed forests. Additionally, three measures of biodiversity were most predictive of forest naturalness: proportional abundance of predators, community weighted mean (CWM) of wing length, and CWM of body roundness. The probability that a forest was near-natural increased with the proportional abundance of predators but decreased with CWM wing length and body roundness. Although species richness was higher in near-natural forests, the effect was not significant. Overall, our findings underscore the conservation value of near-natural forests and highlight the potential of several measures of biodiversity for determining forest quality.

Marginal value analysis reveals shifting importance of migration habitat for waterfowl under a changing climate.

Migratory waterfowl are an important resource for consumptive and non-consumptive users alike and provide tremendous economic value in North America. These birds rely on a complex matrix of public and private land for forage and roosting during migration and wintering periods, and substantial conservation effort focuses on increasing the amount and quality of target habitat. Yet, the value of habitat is a function not only of a site's resources but also of its geographic position and weather. To quantify this value, we used a continental-scale energetics-based model of daily dabbling duck movement to assess the marginal value of lands across the contiguous United States during the non-breeding period (September to May). We examined effects of eliminating each habitat node (32 × 32 km) in both a particularly cold and a particularly warm winter, asking which nodes had the largest effect on survival. The marginal value of habitat nodes for migrating dabbling ducks was a function of forage and roosting habitat but, more importantly, of geography (especially latitude and region). Irrespective of weather, nodes in the Southeast, central East Coast, and California made the largest positive contributions to survival. Conversely, nodes in the Midwest, Northeast, Florida, and the Pacific Northwest had consistent negative effects. Effects (positive and negative) of more northerly nodes occurred in late fall or early spring when climate was often severe and was most variable. Importance and effects of many nodes varied considerably between a cold and a warm winter. Much of the Midwest and central Great Plains benefited duck survival in a warm winter, and projected future warming may improve the value of lands in these regions, including many National Wildlife Refuges, for migrating dabbling ducks. Our results highlight the geographic variability in habitat value, as well as shifts that may occur in these values due to climate change.

Alternative measures of trait-niche relationships: A test on dispersal traits in saproxylic beetles.

Functional trait approaches are common in ecology, but a lack of clear hypotheses on how traits relate to environmental gradients (i.e., trait-niche relationships) often makes uncovering mechanisms difficult. Furthermore, measures of community functional structure differ in their implications, yet inferences are seldom compared among metrics. Community-weighted mean trait values (CWMs), a common measure, are largely driven by the most common species and thus do not reflect community-wide trait-niche relationships per se. Alternatively, trait-niche relationships can be estimated across a larger group of species using hierarchical joint species distribution models (JSDMs), quantified by a parameter Γ. We investigated how inferences about trait-niche relationships are affected by the choice of metric. Using deadwood-dependent (saproxylic) beetles in fragmented Finnish forests, we followed a protocol for investigating trait-niche relationships by (1) identifying environmental filters (climate, forest age, and deadwood volume), (2) relating these to an ecological function (dispersal ability), and (3) identifying traits related to this function (wing morphology). We tested 18 hypothesized dispersal relationships using both CWM and Γ estimates across these environmental gradients. CWMs were more likely than Γ to show support for trait-niche relationships. Up to 13% of species' realized niches were explained by dispersal traits, but the directions of effects were consistent with fewer than 11%-39% of our 18 trait-niche hypotheses (depending on the metric used). This highlights the difficulty in connecting morphological traits and ecological functions in insects, despite the clear conceptual link between landscape connectivity and flight-related traits. Caution is thus warranted in hypothesis development, particularly where apparent trait-function links are less clear. Inferences differ when CWMs versus Γ estimates are used, necessitating the choice of a metric that reflects study questions. CWMs help explain the effects of environmental gradients on community trait composition, whereas the effects of traits on species' niches are better estimated using hierarchical JSDMs.

Body size predicts the rate of contemporary morphological change in birds.

Variation in evolutionary rates among species is a defining characteristic of the tree of life and may be an important predictor of species' capacities to adapt to rapid environmental change. It is broadly assumed that generation length is an important determinant of microevolutionary rates, and body size is often used as a proxy for generation length. However, body size has myriad biological correlates that could affect evolutionary rates independently from generation length. We leverage two large, independently collected datasets on recent morphological change in birds (52 migratory species breeding in North America and 77 South American resident species) to test how body size and generation length are related to the rates of contemporary morphological change. Both datasets show that birds have declined in body size and increased in wing length over the past 40 y. We found, in both systems, a consistent pattern wherein smaller species declined proportionally faster in body size and increased proportionally faster in wing length. By contrast, generation length explained less variation in evolutionary rates than did body size. Although the mechanisms warrant further investigation, our study demonstrates that body size is an important predictor of contemporary variation in morphological rates of change. Given the correlations between body size and a breadth of morphological, physiological, and ecological traits predicted to mediate phenotypic responses to environmental change, the relationship between body size and rates of phenotypic change should be considered when testing hypotheses about variation in adaptive responses to climate change.

High-resolution 3D forest structure explains ecomorphological trait variation in assemblages of saproxylic beetles.

Climate, topography and the 3D structure of forests are major drivers affecting local species communities. However, little is known about how the specific functional traits of saproxylic (wood-living) beetles, involved in the recycling of wood, might be affected by those environmental characteristics.Here, we combine ecological and morphological traits available for saproxylic beetles and airborne laser scanning (ALS) data in Bayesian trait-based joint species distribution models to study how traits drive the distributions of more than 230 species in temperate forests of Europe.We found that elevation (as a proxy for temperature and precipitation) and the proportion of conifers played important roles in species occurrences while variables related to habitat heterogeneity and forest complexity were less relevant. Furthermore, we showed that local communities were shaped by environmental variation primarily through their ecological traits whereas morphological traits were involved only marginally. As predicted, ecological traits influenced species' responses to forest structure, and to other environmental variation, with canopy niche, wood decay niche and host preference as the most important ecological traits. Conversely, no links between morphological traits and environmental characteristics were observed. Both models, however, revealed strong phylogenetic signal in species' response to environmental characteristics.These findings imply that alterations of climate and tree species composition have the potential to alter saproxylic beetle communities in temperate forests. Additionally, ecological traits help explain species' responses to environmental characteristics and thus should prove useful in predicting their responses to future change. It remains challenging, however, to link simple morphological traits to species' complex ecological niches. Read the free Plain Language Summary for this article on the Journal blog.

Morphological consequences of climate change for resident birds in intact Amazonian rainforest.

Warming from climate change is expected to reduce body size of endotherms, but studies from temperate systems have produced equivocal results. Over four decades, we collected morphometric data on a nonmigratory understory bird community within Amazonian primary rainforest that is experiencing increasingly extreme climate. All 77 species showed lower mean mass since the early 1980s­nearly half with 95% confidence. A third of species concomitantly increased wing length, driving a decrease in mass:wing ratio for 69% of species. Seasonal precipitation patterns were generally better than temperature at explaining morphological variation. Short-term climatic conditions affected all metrics, but time trends in wing and mass:wing remained robust even after controlling for annual seasonal conditions. We attribute these results to pressures to increase resource economy under warming. Both seasonal and long-term morphological shifts suggest response to climate change and highlight its pervasive consequences, even in the heart of the world's largest rainforest.

Veteran trees have divergent effects on beetle diversity and wood decomposition.

Veteran hollow trees are keystone structures in ecosystems and provide important habitat for a diverse set of organisms, many of which are involved in the process of decomposition. Since veteran trees are 'islands' of high biodiversity, they provide a unique system in which to study the relationship between biodiversity and decomposition of wood. We tested this relationship with a balanced experiential design, where we quantified the taxonomic and functional diversity of beetles directly involved in the process of decomposing wood, and measured the decomposition of experimentally added bundles of small diameter wood around 20 veteran trees and 20 nearby young trees in southern Norway. We found that the diversity (both taxonomic and functional) of wood-decomposing beetles was significantly higher around the veteran trees, and beetle communities around veteran trees consisted of species with a greater preference for larger diameter wood. We extracted few beetles from the experimentally added wood bundles, regardless of the tree type that they were placed near, but decomposition rates were significantly lower around veteran trees. We speculate that slower decomposition rates around veteran trees could have been a result of a greater diversity of competing fungi, which has been found to decrease decay rates. Veteran trees provide an ecological legacy within anthropogenic landscapes, enhance biodiversity and influence wood decomposition. Actions to protect veteran trees are urgently needed in order to save these valuable organisms and their associated biodiversity.

Sampling beetle communities: Trap design interacts with weather and species traits to bias capture rates.

Globally, many insect populations are declining, prompting calls for action. Yet these findings have also prompted discussion about sampling methods and interpretation of long-term datasets. As insect monitoring and research efforts increase, it is critical to quantify the effectiveness of sampling methods. This is especially true if sampling biases of different methods covary with climate, which is also changing over time. We assess the effectiveness of two types of flight intercept traps commonly used for beetles, a diverse insect group responsible for numerous ecosystem services, under different climatic conditions in Norwegian boreal forest. One of these trap designs includes a device to prevent rainwater from entering the collection vial, diluting preservatives and flushing out beetles. This design is compared to a standard trap. We ask how beetle capture rates vary between these traps, and how these differences vary based on precipitation levels and beetle body size, an important species trait. Bayesian mixed models reveal that the standard and modified traps differ in their beetle capture rates, but that the magnitude and direction of these differences change with precipitation levels and beetle body size. At low rainfall levels, standard traps catch more beetles, but as precipitation increases the catch rates of modified traps overtake those of standard traps. This effect is most pronounced for large-bodied beetles. Sampling methods are known to differ in their effectiveness. Here, we present evidence for a less well-known but likely common phenomenon-an interaction between climate and sampling, such that relative effectiveness of trap types for beetle sampling differs depending on precipitation levels and species traits. This highlights a challenge for long-term monitoring programs, where both climate and insect populations are changing. Sampling methods should be sought that eliminate climate interactions, any biases should be quantified, and all insect datasets should include detailed methodological metadata.

Evolutionary and ecological forces influencing population diversification in Bornean montane passerines.

The mountains of Borneo are well known for their high endemicity and historical role in preserving Southeast Asian rainforest biodiversity, but the diversification of populations inhabiting these mountains is poorly studied. Here we examine the genetic structure of 12 Bornean montane passerines by comparing complete mtDNA ND2 gene sequences of populations spanning the island. Maximum likelihood and Bayesian phylogenetic trees and haplotype networks are examined for common patterns that might signal important historical events or boundaries to dispersal. Morphological and ecological characteristics of each species are also examined using phylogenetic generalized least-squares (PGLS) for correlation with population structure. Populations in only four of the 12 species are subdivided into distinct clades or haplotype groups. Although this subdivision occurred at about the same time in each species (ca. 0.6-0.7Ma), the spatial positioning of the genetic break differs among the species. In two species, northeastern populations are genetically divergent from populations elsewhere on the island. In the other two species, populations in the main Bornean mountain chain, including the northeast, are distinct from those on two isolated peaks in northwestern Borneo. We suggest different historical forces played a role in shaping these two distributions, despite commonality in timing. PGLS analysis showed that only a single characteristic-hand-wing index-is correlated with population structure. Birds with longer wings, and hence potentially more dispersal power, have less population structure. To understand historical forces influencing montane population structure on Borneo, future studies must compare populations across the entirety of Sundaland.