Increasing temperature may compensate for lower amounts of dead wood in driving richness of saproxylic beetles

Global warming and land‐use change are expected to be additive threats to global diversity, to which insects contribute the highest proportion. Insects are strongly influenced by temperature but also require specific habitat resources, and thus interaction between the two factors is likely. We selected saproxylic beetles as a model group because their life cycle depends on dead wood, which is highly threatened by land use. We tested the extent to which higher temperatures compensate for the negative effects of low amounts of dead wood on saproxylic beetle species richness (Temperature–Dead wood compensation hypothesis) on both a macroclimate and a topoclimate scale (north‐ and south‐facing slopes). We analyzed 1404 flight‐interception trap catches across Europe to test for interaction effects of temperature and dead‐wood amount on species richness. To experimentally test our findings from the activity trap data, we additionally reared beetles from 80 bundles of dead wood initially exposed at high and low elevations. At the topoclimate scale, we analyzed trap catches and reared beetles from dead wood exposed in 20 forest stands on south‐facing and north‐facing slopes in one region. On the macroscale, both temperature and dead‐wood amount positively affected total and threatened species richness independently, but their interaction was significantly negative, indicating compensation. On both scales and irrespective of the method, species richness decreased with temperature decline. Our observation that increasing temperature compensates for lower amounts of dead wood has two important implications. First, managers of production forests should adapt their dead‐wood enrichment strategy to site‐specific temperature conditions. Second, an increase in temperature will compensate at least partially for poor habitat conditions in production forests. Such a perspective contrasts the general assumption of reinforcing impacts of global warming and habitat loss on biodiversity, but it is corroborated by recent range expansions of threatened beetle species.     

Habitat availability in forests and clearcuts for saproxylic beetles associated with aspen

To maintain biodiversity in managed, boreal forest in Scandinavia, aspen trees (Populus tremula) are often retained at clearcutting. In this study, the habitat availability for beetles associated with aspen coarse woody debris (CWD) was predicted for forests and clearcuts with a model of CWD dynamics. Habitat requirements of eight beetle species (Agathidium bicolor, Cerylon ferrugineum, Cyphaea curtula, Endomychus coccineus, Homalota plana, Mycetophagus fulvicollis, Ptilinus fuscus and Xylotrechus rusticus) were obtained from their occurrence patterns in relation to characteristics of CWD objects in forest and on clearcuts in a study landscape in central Sweden. Three species were more frequent in forest and three at clearcuts. Five species increased with increasing girth of the CWD. Three were more frequent on standing CWD, and two on lying CWD. From the same study area, we also obtained field data on the recruitment of CWD (i.e., tree mortality) and amounts of different types of CWD. Annual tree mortality of aspen was higher for recent clearcuts (6.3%) compared with older clearcuts (1.1%). For all species, the habitat availability was higher on clearcuts, because enhanced tree mortality increased the amount of recently dead CWD. As a conclusion, green-tree retention of aspen is a conservation effort that is beneficial for species associated with aspen CWD.     

Succession of saproxylic beetles associated with decomposition of boreal white spruce logs

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Individual food niche specialization in stream-dwelling charr

Food niche variation within a population of white-spotted charr, Salvelinus leucomaenis, was investigated by means of mark and recapture to examine the simplification that conspecific individuals are ecologically equivalent. Food diversity measured on the basis of living form of dietary organisms demonstrated variability in food niche in incidence, degree, and stability. As inferred from body size distribution and fish movement, niche variation was inexplicable by social rank hierarchy or foraging habitat rotation, and occasional niches were deemed to be realized individually. Higher annual growth rate shown by those with a larger niche width indicated the adaptive significance in niche variation. Assuming trade-off between feeding efficiency and predation risk, phenotypic plasticity was suggested to determine the food niche specialization of individuals to maximize lifetime fitness.     

Individual specialization in a shorebird population with narrow foraging niche

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Density-dependent population regulation detected in short time series of saproxylic beetles

Understanding the regulation of natural populations has been a long-standing problem in ecology. Here we analyze the population dynamics of 17 species of saproxylic beetles in Shizuoka Prefecture, Japan collected over 11–12 years using autoregressive integrated moving average (ARIMA) models. We first examined the dynamics for indications of the order of the ARIMA models and evaluated the time series to determine that it was not simply a random, white noise sequence. All species dynamics were not mere random noise, and ARIMA models up to lag 3 were considered. The best model was selected from the possible ones using several criteria: model convergence, weak residual autocorrelation, the small sample AIC must be among the smallest that were not significantly different, and the lag indicated by the cutoff values in the detrended partial autocorrelation function. We found significant and nearly significant direct density-dependence for 14 of the 17 species, varying from −0.709 and stronger. The characteristic return rates were strong and only one species had a weak return rate (>0.9), implying that these species were strongly regulated by density-dependent factors. We found that populations with higher order ARIMA models (lag 2 and 3) had weaker return rates than populations with ARIMA models with only one lag, suggesting that species with more complex dynamics were more weakly regulated. These results contrast with previous suggestions that 20+ years are needed to detect density dependence from population time series and that most populations are weakly regulated.     

A comparison of three methods to survey saproxylic beetles in hollow oaks

One of the most endangered assemblages of species in Europe is insectsassociated with old trees. For that reason there is a need of developing methodsto survey this fauna. This study aims at comparing three methods – windowtrapping, pitfall trapping and wood mould sampling – to assess speciesrichness and composition of the saproxylic beetle fauna in living, hollow oaks.We have used these methods at the same site, and to a large extent in the sametrees. Useful information was obtained from all methods, but they partiallytarget different assemblages of species. Window trapping collected the highestnumber of species. Pitfall trapping collected beetles associated with treehollows which rarely are collected by window traps and therefore it isprofitable to combine these two methods. As wood mould sampling is the cheapestmethod to use, indicator species should preferably be chosen among specieswhich are efficiently collected with this method.     

Vertical distribution of saproxylic beetles within snag trunks retained in plantation forests

Retention of snags (standing dead trees) is considered to have important effects on saproxylic species conservation in plantation forests because snags would provide vertically stratified deadwood habitats. However, the vertical distribution of saproxylic insects within snag trunks is still unclear. We felled 33 naturally occurring snags of Todo fir Abies sachalinensis in plantation forests and extracted insects from 99 logs sampled from three vertical positions of the snag trunks (basal stem: <2.5 m, lower trunk: 2.5–5 m and upper trunk: >5 m). The mean number of species that emerged from a single log was only 2.69, but we identified 51 morphospecies of saproxylic beetles in total. The total number of species that emerged from the basal stem (34 spp.) was greater than those that emerged from the lower trunk (25 spp.) or the upper trunk (30 spp.). However, rarefaction-extrapolation analysis did not demonstrate a significant difference in species richness among the log positions. Beetle assemblages were separated into two groups by constrained correspondence analysis; one group emerged only from lower and upper trunk logs, while another emerged mainly from basal stem logs. Additionally, vertical position had a significant effect on the distribution of the five main species. Our results show that beetle assemblages within snags in the plantation forests were highly variable, and retaining a sufficient number of high stumps may be important for saproxylic beetle conservation in plantation forests. We propose ‘retention thinning’ as an appropriate method to combine efficient timber production with biodiversity conservation in plantation forests.     

Micro and macro-habitat associations in saproxylic beetles: implications for biodiversity management

Restoration of habitats is critically important in preventing full realization of the extinction debt owed as a result of anthropogenic habitat destruction. Although much emphasis has been placed on macrohabitats, suitable microhabitats are also vital for the survival of most species. The aim of this large-scale field experiment was to evaluate the relative importance of manipulated microhabitats, i.e., dead wood substrates of spruce (snags, and logs that were burned, inoculated with wood fungi or shaded) and macrohabitats, i.e., stand types (clear-cuts, mature managed forests, and forest reserves) for species richness, abundance and assemblage composition of all saproxylic and red-listed saproxylic beetles. Beetles were collected in emergence traps in 30 forest stands in 2001, 2003, 2004 and 2006. More individuals emerged from snags and untreated logs than from burned and shaded logs, but species richness did not differ among substrates. Assemblage composition differed among substrates for both all saproxylics and red-listed saproxylic species, mainly attributed to different assemblage composition on snags. This suggests that the practise of leaving snags for conservation purposes should be complemented with log supplementation. Clear-cuts supported fewer species and different assemblages from mature managed forests and reserves. Neither abundance, nor species richness or assemblage composition differed between reserves and mature managed forests. This suggests that managed stands subjected to selective cutting, not clear-felling, maintain sufficient old growth characteristics and continuity to maintain more or less intact assemblages of saproxylic beetles. Thus, alternative management methods, e.g., continuity forestry should be considered for some of these stands to maintain continuity and conservation values. Furthermore, the significantly higher estimated abundance per ha of red-listed beetles in reserves underlines the importance of reserves for maintaining viable populations of rare red-listed species and as source areas for saproxylic species in boreal forest landscapes.     

On testing temporal niche differentiation in carabid beetles

Summary It is first shown that den Boer's (1985) claims that competition should be rejected as a factor explaining community structure are not warranted by his tests on temporal niche differentiation in carabid beetles. The power of these tests is so low that they are unlikely to detect even the most extreme pattern of niche differentiation. A major factor that affects their power is the procedure of building up an artificial species pool from the local communities to which the test is to be applied. Alternative tests are then presented, and applied to data on annual activity cycles of carabids in three forest biotopes in Belgium. Three factors are shown to strongly affect their outcome: the unit into which the time scale is divided, the criteria used to incorporate constraints, and the set of species selected. Nevertheless, they provide some evidence that there is a growing niche differentiation from the successional to the climax forest, as predicted by competition theory. Other hypotheses, however, could explain the observed pattern.     

Isolation by distance in Middle Dalmatia-Yugoslavia

Parameters of Malécot's isolation-by-distance model are estimated for biological (anthropometric head and body dimensions, morphometric dimensions of metacarpal bones, quantitative and qualitative dermatoglyphic traits, and physiological/cardiorespiratory/variables) and linguistic distances and migrational kinship on the island of Korcula and the Peljesac peninsula in Middle Dalmatia, Croatia, Yugoslavia. Resulting parameters and the fit of the model are compared, for both regions, as well as with results of similar analysis in other parts of the world. The fit of the model is highly significant for migrational kinship and linguistic distances and less so for biological traits. Differences between these two populations, which live under basically similar ecological conditions, are explained by variation in biological and sociocultural history.     

Cooperative personalities and social niche specialization in female meerkats

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Ecophysiology of Aufwuchs-eating cichlids in Lake Tanganyika: niche separation by trophic specialization

Synopsis The Aufwuchs-eating cichlids of Lake Tanganyika show clear trophic differences that are correlated to their morphology, physiology and foraging behaviour. The species are grouped into three categories of relative intestinal length according to their feeding habits. A correlation between the intestinal length and the diet could be demonstrated, ranging from around 2.5 for species ingesting more animal food, to 7.8 for detritivorous and microalgivorous species. The relative intestinal length of domesticTropheus moorii, raised in aquaria was significantly lower than that of wild individuals by a factor of 1.7, demonstrating a wide range of phenotypic adaptability. The activities of trypsin and amylase were at an equal level in four Aufwuchseating species, but the activity of laminarinase of a detritivorous-microalgivorous species (Petrochromis orthognathus) was 2.6 times higher than that of an algivorous species (Tropheus moorii). The laminarinase seems to be an excellent marker enzyme for detritivorous or microalgivorous feeding.     

Head shape variation in cerambycid saproxylic beetles as a function of host plant selection

Saproxylic insects depend on deadwood for larval development, and a certain degree of specialization may be involved in their choice of host plants and/or wood in a particular stage of degradation. The plant species chosen for oviposition in turn act as an environmental pressure on the head morphology of larvae and it is expected that head shape plasticity varies directly with the number of woody plant species used for larval development in each insect species. We analyzed head shape variation in saproxylic beetles with respect to host plant species, maximum time of larval emergence and season of the year when insects colonized branches. Generalist species in the use of host plants showed significant variation in head shape and size. Time of emergence and season did not appear to affect head shape, although season was a determinant factor of abundance and possibly head size variation.     

Some of the rarest European saproxylic beetles are common in the wilderness of Northern Mongolia

Natural landscapes characterized by heavy disturbance regimes were displaced in Europe by managed cultural landscapes over the past centuries. The associated loss of biological legacies, such as dead or dying trees, has exposed numerous saproxylic species to high risks of extinction. In contrast, extensive wilderness forests in Northern Mongolia have been sustained owing to significant cultural differences. Here we used saproxylic beetle abundance data gathered during two sampling campaigns in the Mongolian taiga to address whether (1) the saproxylic beetle fauna of the Mongolian taiga is comparable to that of European boreal forests, (2) fires are a natural disturbance regime, indicated by the occurrence of many pyrophilous species, and (3) species rare in Europe are also rare in the Northern Mongolian wilderness. Of 191 saproxylic beetle species identified, 150 (79 %) were also found in Europe. The high number of pyrophilous beetle species (20) indicated that natural species communities are well adapted to this disturbance regime. The species rarity in Germany was significantly positively correlated with the species rarity in Finland, but the species rarity in these two countries was negatively correlated with that in the Mongolian wilderness. Our results indicated that wilderness areas with natural disturbances provide biological legacies important for rare species. Therefore, exploitation of the unique, remaining natural landscapes of the Palaearctic wilderness areas should be stopped. Moreover, we urge conservationists to expand controlled burning for restoration at relict sites of rare boreal species also outside Fennoscandia.     

Forest restoration by burning and gap cutting of voluntary set-asides yield distinct immediate effects on saproxylic beetles

Today, the importance of restoring natural forest disturbance regimes and habitat structures for biodiversity is widely recognized. We evaluated the immediate effects of two restoration methods on wood-inhabiting (saproxylic) beetles in boreal forest voluntary set-asides. We used a before-after control-impact experimental set-up in 15 set-asides; each assigned to one of three treatments: (1) restoration burning, (2) gap cutting and (3) no-treatment reference stands. Before treatment, abundance, species richness and assemblage composition of trapped beetles did not differ significantly among treatments. Burning resulted in a significant change in assemblage composition and increased species richness and abundance compared to reference stands. As predicted, saproxylic species known to be fire favoured increased dramatically after burning. The immediate response shows that, initially, fire favoured species are attracted from the surrounding landscape and not produced on site. Gap cutting increased the abundance of cambium consumers but had no significant effect on total species richness or assemblage composition of saproxylic beetles. The stronger effect of burning compared to gap cutting on saproxylic assemblages is probably due to the very specific conditions created by fires that attracts many disturbance-dependent species, but that at the same time disfavour some disturbance-sensitive species. By contrast, gap cutting maintained assemblage composition, increased abundances and is likely to increase species richness in the years to follow, due to elevated level of dead wood. The restoration methods applied in this study may prove particularly useful, partly because of positive effect on saproxylic beetles, but also due to the cost-efficiency of the measures; the voluntary set-asides were already established and the restoration costs fully covered by revenue from the extracted timber.