Bat assemblages on Neotropical land-bridge islands: nested subsets and null model analyses of species co-occurrence patterns

A fundamental goal of ecology is to understand whether ecological communities are structured according to general assembly rules or are essentially dictated by random processes. In the context of fragmentation, understanding assembly patterns and their mechanistic basis also has important implications for conservation. Using distribution data of 20 bat species collected on 11 islands in Gatún Lake, Panama, we tested for non‐randomness in presence–absence matrices with respect to nestedness and negative species co‐occurrence. We examined the causal basis for the observed patterns and conducted separate analyses for the entire assemblage and for various species submatrices reflecting differences in species’ trophic position and mobility. Furthermore, we explored the influence of weighting factors (area, isolation, abundance) on co‐occurrence analyses. Unweighted analyses revealed a significant negative co‐occurrence pattern for the entire assemblage and for phytophagous bats alone. Weighting analyses by isolation retained a pattern of species segregation for the whole assemblage but nullified the non‐random structure for phytophagous bats and suggested negative associations for animalivores and species with low mobility. Area‐ and abundance‐weighted analyses always indicated random structuring. Bat distributions followed a nested subset structure across islands, regardless of whether all species or different submatrices were analysed. Nestedness was in all cases unrelated to island area but weakly correlated with island isolation for incidence matrices of all species, phytophagous bats, and mobile species. Overall, evidence for negative interspecific interactions indicative of competitive effects was weak, corroborating previous studies based on ecomorphological analyses. Our findings indicate that bat assemblages on our study islands are most strongly shaped by isolation effects and species’ differential movement and colonization ability. From a conservation viewpoint this suggests that even in systems with high fragment–matrix contrast, a purely area‐based approach may be inadequate, and structural and functional connectivity among patches are important to consider in reserve planning.     

Influence of tree hollow characteristics on saproxylic beetle diversity in a managed forest

Tree hollows are key structures in forest ecosystems constituting long-lasting habitats and nutritional resources for many saproxylic arthropod species. Due to diverse microhabitat structures and conditions in tree hollows, they can support a broad range of species. However, in the past intensive management practices in parts of Europe reduced the abundance of tree hollows, resulting in a decrease and endangerment of species specialised in this tree habitat. We investigated 40 beech trees with hollows in 2014 and a subset of 23 of these trees in 2015 in a managed forest landscape in Germany. Using emergence traps we collected 89 beetle species of which 33% were on the Bavarian Red List. We described the tree characteristics, physical hollow characteristics, and their surrounding environment investigating their influence on α-diversity of non-Red List and Red List species. Furthermore, we investigated spatial (between tree hollows) and temporal (same tree hollow but different years) β-diversity, considering the importance of turnover and nestedness components on β-diversity. α-Diversity decreased with increasing decomposition of wood mould and increased with increasing area of hollow entrance in both years. Additional characteristics differed between years and between non-Red List and Red List species. β-Diversity was related to diameter at breast height, number of surrounding tree hollows, area of hollow entrance and a temperature gradient. We found a higher species turnover than nestedness between tree hollows and between years, indicating highly dynamic beetle communities spatially as well as temporally. To support and maintain the diversity of saproxylic beetles inhabiting tree hollows, the heterogeneity of microhabitats is important and should be supported by maintaining the diversity of differently structured and sized tree hollows.     

Community assembly in temperate forest birds: habitat filtering,interspecific interactions and priority effects

Recent attempts to understand the processes governing community assembly have increasingly focused on patterns of phylogenetic relatedness and functional similarity among co-existing species. Considerations of the species pool, the number and identity of functional traits and the metrics used to identify patterns have come under scrutiny as possible influences on the detection of non-random patterns. Most mechanistic explanations of community assembly based on functional and phylogenetic approaches rely on deterministic explanations, while ignoring the role of stochastic processes and historical contingency, despite the prominent historical role of both types of explanations of species coexistence. We evaluated the phylogenetic and functional structure of 20 temperate forest bird assemblages in northeastern North America. We compared three approaches for characterizing the functional structure of assemblages. Regardless of approach, assemblages were generally no different than expected by chance. In contrast, phylogenetic structures of bird assemblages were overdispersed, clumped or consistent with random assembly depending on the site. Nonetheless, we found little evidence for differences in phylogenetic structure arising as a consequence of the identity of the species pool. We identified a strong relationship between the proportion of residents and phylogenetic relatedness that was unrelated to the species richness of assemblages. Our results suggest that different assembly mechanisms may structure resident and migratory subsets of temperate breeding bird communities. Resident assemblages are likely structured by interspecific interactions and habitat filtering prior to arrival of migrants. In contrast, the composition of migrant assemblages may be a consequence of priority effects in which the presence and abundance of residents and earliest arriving species affect the ability of subsequent migrants to colonize sites. This phenomenon enhances the likelihood of multiple alternative community structures in similar environments.     

Influence of boreal forest succession and dead wood qualities on saproxylic beetles

1 Saproxylic insects, a functional group dominated by beetles, are dependent on dead or moribund trees as habitat elements. 2 Although there are few studies of saproxylic insects from the North American boreal zone, European studies demonstrate that forest harvest can lead to a biologically significant decrease in saproxylic beetle diversity. 3 We studied saproxylic beetles in the North American boreal mixedwood forest using flight intercept traps established on naturally dead and girdled trembling aspen and spruce trees along a successional gradient of undisturbed stands from deciduous to coniferous overstory trees. 4 Composition and diversity of beetle assemblages differed among forest successional types. 5 Snag age class was an important determinant of composition for saproxylic beetle assemblages. 6 Multivariate regression analysis of these data indicated that saproxylic beetles are responding to changes in coarse woody debris, and not to the relative densities of canopy tree species, although these variables are strongly correlated. 7 Coarse woody debris management should be a primary concern in forest management plans seeking to conserve saproxylic organisms and the critical ecosystem functions (i.e. nutrient cycling) in which they participate.     

Influence of tree hollow characteristics and forest structure on saproxylic beetle diversity in tree hollows in managed forests in a regional comparison

Tree hollows are among the rarest habitats in today''s Central European managed forests but are considered key structures for high biodiversity in forests. To analyze and compare the effects of tree hollow characteristics and forest structure on diversity of saproxylic beetles in tree hollows in differently structured managed forests, we examined between 41 and 50 tree hollows in beech trees in each of three state forest management districts in Germany. During the two‐year study, we collected 283 saproxylic beetle species (5880 individuals; 22% threatened species), using emergence traps. At small spatial scales, the size of hollow entrance and the number of surrounding microhabitat structures positively influenced beetle diversity, while the stage of wood mould decomposition had a negative influence, across all three forest districts. We utilized forest inventory data to analyze the effects of forest structure in radii of 50–500 m around tree hollows on saproxylic beetle diversity in the hollows. At these larger spatial scales, the three forest management districts differed remarkably regarding the parameters that influenced saproxylic beetle diversity in tree hollows. In Ebrach, characterized by mostly deciduous trees, the amount of dead wood positively influenced beetle diversity. In the mostly coniferous Fichtelberg forest district, with highly isolated tree hollows, in contrast, only the proportion of beech trees around the focal tree hollows showed a positive influence on beetle diversity. In Kelheim, characterized by mixed forest stands, there were no significant relationships between forest structure and beetle diversity in tree hollows. In this study, the same local tree hollow parameters influenced saproxylic beetle diversity in all three study regions, while parameters of forest structure at larger spatial scales differed in their importance, depending on tree‐species composition.     

Genotype by environment interactions in forest tree breeding: review of methodology and perspectives on research and application

Genotype by environment interaction (G×E) refers to the comparative performances of genotypes differing among environments, representing differences in genotype rankings or differences in the level of expression of genetic differences among environments. G×E can reduce heritability and overall genetic gain, unless breeding programmes are structured to address different categories of environments. Understanding the impact of G×E, the role of environments in generating G×E and the problems and opportunities is vital to efficient breeding programme design and deployment of genetic material. We review the current main analytical methods for identifying G×E: factor analytic models, biplot analysis and reaction norm. We also review biological and statistical evidence of G×E for growth, form and wood properties in forest species of global economic importance, including some pines, eucalypts, Douglas-fir, spruces and some poplars. Among these species, high levels of G×E tend to be reported for growth traits, with low levels of G×E for form traits and wood properties. Finally, we discuss possible ways of exploiting G×E to maximise genetic gain in forest tree breeding. Characterising the role of environments in generating interactions is seen as the basic platform, allowing efficient testing of candidate genotypes. We discuss the importance of level-of-expression interaction, relative to rank-change interaction, as being greater than in many past reports, especially for deployment decisions. We examine the impacts of G×E on tree breeding, some environmental factors that cause G×E and the strategies for dealing with G×E in tree breeding, and the future role of genomics.     

Effects of a fire on a breeding population of Loggerhead Shrikes in sagebrush steppe habitat

ABSTRACT.   We studied the breeding biology of Loggerhead Shrikes ( Lanius ludovicianus ) in northeastern Oregon from 1995 to 1997 and 2000 to 2001. A wildfire in 1998 burned approximately half the big sagebrush ( Artemisia tridentata ) at this site, permitting comparison of population parameters before and after this event. Shrike density after the fire was half that before the fire, a reduction likely due to loss of suitable nesting habitat. Mayfield estimates of nest survivorship were 39% (range: 30–46%) in the years before and 19% (range: 12–26%) after the fire, among the lowest ever recorded for this species. In 2000, young fledged from only 21% of the territories. Due to persistent renesting, young fledged from 67% of the territories in 2001, a percentage similar to that before the fire. After the fire, shrikes nested more frequently in substrates other than big sagebrush, especially juniper trees ( Juniperus occidentalis ). Clutch size, fledglings per successful nest, and fledgling survival were similar in both periods of the study. Poor nesting success across all years suggests that productivity may play a role in regional declines of shrike populations, and that the loss and fragmentation of nesting habitat due to fire may further reduce productivity. Given the threats facing remaining sagebrush habitat, as well as Loggerhead Shrike population declines, we recommend aggressive fire control measures and further protection for remaining tall sagebrush communities.     

Biodiversity conservation in old-growth boreal forest: black spruce and balsam fir snags harbour distinct assemblages of saproxylic beetles

Biodiversity conservation of forest ecosystems strongly relies on effective dead wood management. However, the responses of saproxylic communities to variations in dead wood characteristics remains poorly documented, a lack of knowledge that may impede the development of efficient management strategies. We established the relationship between saproxylic beetles—at the species and community levels—and attributes of black spruce and balsam fir in old-growth boreal forests. The relationship was first evaluated for individual snag bole segments, and then for forest stands. A total of 168 bole sections were collected in summer 2006 along a compositional gradient ranging from black spruce-dominated stands to balsam fir-dominated ones, in a boreal forest dominated by >90-year-old stands. A total of 16,804 beetles belonging to 47 species emerged from bole segments, with 21% of the species being found exclusively in black spruce snags and 36% exclusively in balsam fir snags. Black spruce and balsam fir snags thus contributed differently to forest biodiversity by being inhabited by different saproxylic communities. Wood density was an important attribute in the host-use patterns for several species of saproxylic beetles, but no relationship was found between snag availability within stands and abundance of beetles strongly linked to either black spruce or balsam fir. Our study outlines the relative contribution of tree compositional diversity to saproxylic species, while highlighting the contribution of black spruce and balsam fir to animal diversity in old-growth boreal forests.     

Effects of habitat fragmentation: increased isolation and reduced habitat size reduces the incidence of dead wood fungi beetles in a fragmented forest landscape

Negative effects of habitat fragmentation (increased isolation and reduced habitat size) affected presence of several beetle species inhabiting Fomes fomentarius (L.) Kickx basidiocarps in a fragmented forest landscape. At the scale of individual trees (tree level), incidence of Cis jacquemarti Mell./C. nitidus (F.) (not distinguished between individuals of these two species, abbreviated C. jacquemarti /nitidus), C. bidentatus (Ol.), C. lineatocribratus Mell., Ennearthron cornutum (Gyll.) (all Coleoptera, Cisidae) and Dorcatoma dresdensis Herbst (Coleoptera, Anobiidae) was reduced with increased habitat isolation and reduced habitat size. Additionally, increased habitat size at a larger scale (forest island level) also gave higher incidence of all species.
Some of the microhabitat variables measured influenced upon the presence of the study species. Fallen trees had a higher probability of C. jacquemarti / nitidus presence than standing trees, whilst the opposite was the case for C. bidentatus. There was also increased incidence of C. bidentatus on trees in denser forests and of E. cornutum in grey alder Alnus incana compared with birch trees (Betula spp.).
Increased volume of basidiocarps on a tree resulted in a higher number of individuals of C. jacquemarti / nitidus and C. lineatocribratus.     

Intra- and interspecific tree diversity promotes multitrophic plant–Hemiptera–ant interactions in a forest diversity experiment

Interactions between species of different trophic levels have long been recognized as fundamental processes in ecology. Although mounting evidence indicates that plant species diversity (PSD) or plant genetic diversity (PGD) can influence the plant-associated arthropod community, these two fundamental levels of biodiversity are not often manipulated simultaneously to assess their effects on species interactions. We used a large tree diversity experiment (BEF-China), which manipulates PSD and PGD in a crossed design to test individual and combined effects of PSD and PGD on multitrophic interaction networks and interaction partner species richness and occurrence. We focused on two tree species, on which sap-sucking Hemiptera and interacting ant species commonly occur. This tri-trophic interaction can be divided into the antagonistic plant–Hemiptera interaction and the mutualistic Hemiptera–ant interaction, known as trophobioses. Qualitative evaluation of tri-trophic interaction networks at different PSD and PGD combinations showed increased interaction partner redundancy at high PSD and PGD. This was supported by increased Hemiptera species richness at high PSD and PGD. Furthermore, the data indicate higher occurrence of Hemiptera and trophobioses and higher trophobiotic ant species richness with increasing PSD and PGD. As no plant diversity component alone caused an effect we conclude that the combined effect of high PGD and high PSD might be additive. In summary, as plant genetic diversity, especially at low species richness, seems to increase the interaction partner redundancy in interaction networks and the diversity of interacting communities, we suggest that genetic diversity should be considered in forest conservation and restoration programs.     

Abundance and co-occurrence patterns of core and satellite species of ground beetles on small lake islands

15 lake islands and two mainland sites of Mamry lake in Poland were sampled to investigate community structures and patterns of co-occurrences of ground beetles (Carabidae). The total ground beetle metacommunity of 71 species was divided into a group of core species occupying at least half of all study sites and of satellite species, which occurred at two sites or less. This division is mirrored by reduced dispersal abilities and non-random patterns of site occupancy. Core and satellite species also differed in patterns of relative abundance. The core group followed a lognormal distribution, the satellite group a power function as predicted by the self-similarity model of occurrence. We conclude that the division into core and satellite species is not a sample artefact but reflects different life history strategies. We also conclude that current models of niche division and co-occurrence might miss important aspects of community structure if they do not refer to patterns of dispersal.
From these findings we infer that the regional distribution of core species might be shaped by species interactions and processes of niche divisions whereas the spatial distribution of satellite species are best interpreted as stemming from random dispersal.     

Effects of habitat quality and landscape structure on saproxylic species dwelling in boreal spruce-swamp forests

Intensive forest management has caused loss and fragmentation of old‐growth forests and reduced the amount of dead wood throughout northwest Europe. Changes in habitat availability are reflected in occurrence patterns of habitat‐specialist species only after a certain time lag. Here we analyse the responses of wood‐decomposing fungi and saproxylic beetles inhabiting patches of spruce‐swamp forest to habitat quality, loss and isolation at three different spatial scales in eastern Finland and adjacent Russian Karelia, where forestry has been very marginal until recently. Both rare specialist species and common generalist species were included in the study to reveal whether their occurrence patterns differ. Variables describing habitat quality (density and continuity of host trees, patch area) turned out significant in explaining species' incidences (proportion of occupied host trees) in only a few cases, probably because of the relatively high quality of all study patches. Despite this fact, and consistent with our hypothesis, incidences of all the eight specialist species were higher in Russia than Finland, and the difference was significant in the two most strict habitat specialists, Pytho kolwensis and Phlebia centrifuga. In contrast, incidences of three out of four generalist species were higher in Finland than in Russia, and the difference was significant in Rhagium inquisitor. In a subset of 21 patches in Finland, we used a metapopulation model to predict the probability of each patch to be currently occupied by a species given the known spatiotemporal distribution of suitable forest stands during the last 50 years. The degree of isolation alone explained significantly the incidences of five species. However, including habitat variables into the models altered some of the effects. Moreover, inconsistent with our hypothesis, isolation appeared to also negatively affect some very common generalist species. Inclusion of these species in the study disclosed that apparently significant effects of spatiotemporal isolation should be interpreted cautiously.     

Influence of snag characteristics on saproxylic beetle assemblages in a south Swedish beech forest

Effects of snag characteristics on saproxylic beetles were studied in an area of managed beech forest in southern Sweden. A snag survey was combined with a beetle survey using 30 small window traps directly attached to beech snags. The total number of species was lower in the snags which were most decayed than in the three younger decay classes, while the number of red-listed species remained the same regardless of the stage of decay. The number of fresh wood species declined and the number of rot hole species increased with increasing snag decay. The diversity of fungicolous and decayed wood species peaked at the intermediate stage of decay in the snags. CCA ordination confirmed that the stage of decay in the snags was most important for species composition, followed by sun exposure. There were no general differences in species density and composition between managed and unmanaged stands. Our study suggests that most species are able to find suitable habitat within a radius of a few kilometres and that the total amount of habitat in an unfragmented forest area is more important for species diversity than the spatial distribution of this habitat. Our data also shows that species diversity increases with habitat diversity. Snags formed from giant beech trees seem to be particularly important for rare species living in rot holes. We conclude that for a high species diversity there is a requirement for snags in different stages of decay, size and degree of sun exposure.     

Variation in prey-specific consumption rates and patterns of field co-occurrence for two larval predaceous diving beetles

Freshwater predatory insects can exert strong effects on prey, although how multiple similar predators may coexist is not well understood. Larval predaceous diving beetles are often numerically and taxonomically abundant predators in lentic systems, but the proximate mechanisms that explain their high abundance remain unknown. Field surveys were conducted twice in June in ponds in Alberta, Canada to assess the associations between larvae of two genera (Graphoderus, Rhantus), their spatial locations, and correlations with potential prey. Both larvae were common and positively correlated within wetlands although neither varied with pond depth nor distance from edge. Laboratory trials indicated that Graphoderus consumed more prey (corixids) at the surface, whereas Rhantus killed benthic prey (chironomids) and corixids at an equal rate; damselflies were the least consumed prey. Predation also varied with depth, with both larvae feeding at higher rates in the shallowest environments compared to Graphoderus at an intermediate depth. Predator–prey correlations from ponds were mostly congruent with predation trials; Graphoderus was positively correlated with corixids, Rhantus was positively correlated with corixids and chironomids; beetles were uncorrelated with damselflies. Reliance on different prey in different microhabitats may be an important mechanism for the maintenance of high abundance of dytiscid larvae.     

Effects of tree mortality on the spatial patterns and interspecific associations of plant species in a Quercus aliena var. acuteserrata forest in Baotianman,Henan, China

Aims Tree mortality is an important ecological process in forest ecosystems. The aims of this study were to determine how tree mortality influences the spatial patterns and interspecific associations of plant species, and what are the causes of tree mortality in a 1 hm² permanent plot in Baotianman National Nature Reserve, Nanyang City, Henan Province.
Methods We conducted field investigations in the plot and used spatial point pattern analysis to examine the spatial patterns and interspecific associations of 17 species prior to and following mortality.
Important findings (1) Most of the species in the study plot showed an aggregated distribution both pre- and post-mortality. However, the number of species showing aggregated distribution decreased and the number of species showing random distribution increased following the mortality event. (2) Most species were positively associated with Quercus aliena var. acuteserrata both pre- and post-mortality, while some had no apparent association. Following tree mortality, on fine scales, the number of species with positive associations increased, and the number of species with negative associations decreased. (3) Tree mortality was in consistency with the random death hypothesis. The interspecific associations of four species with Q. aliena var. acuteserrata completely changed following death. For most species, the spatial patterns and the interspecific association with Q. aliena var. acuteserrata either changed at minor scales or did not change. The variations in spatial patterns or interspecific associations were inconsistent among species. (4) The dead trees of Q. aliena var. acuteserrata were significantly associated with the living trees in 13 species, but the associations between dead and living trees were not in agreement with the changes in interspecific association following mortality. Only five living tree species competed with the dead trees of Q. aliena var. acuteserrata, and the competition between each of these species and Q. aliena var. acuteserrata intensified after individual death. Tree mortality is the result of a variety of factors. Although the tree mortality in the study plot was in accordance with the random death hypothesis, there were also a few individuals which were dead from competition.     

Effects of different tree harvesting intensities on saproxylic beetle diversity in coniferous and broad-leaved mixed forest in the Changbai Mountain,Northeast, China

1. Saproxylic beetles have gained increasing attention due to their role in the decomposition of rotting wood in forests. Studying the response of saproxylic beetles to tree harvesting is important for developing harvesting strategies that consider conservation of saproxylic beetle diversity.
2. We report results from a case study in which we designed four treatment stands to test the effects of forest harvesting intensity on saproxylic beetle diversity, harvest intensities of 0% untreated control (CK), 17.2% light harvest (LT), 34.7% moderate harvest (MT) and 51.9% high harvest (HT). Flight intercept traps were used to collect specimens of saproxylic beetles in each stand at 2, 3 and 4 years post-harvest.
3. The richness and abundance of saproxylic beetles were higher in MT and HT than in CK. Twelve of the 15 indicator species were significantly associated with MT or HT, whereas only three species were significantly associated with CK.
4. We found that moderate and high intensity harvesting affected the composition and increased beetle abundance and richness of saproxylic beetles, and light intensity harvesting had no effect on the beetle community in years 2, 3 and 4 post-harvest. There was no difference in the beetle community composition between moderate and high harvest stands.