Nitrogen fixation in the stag beetle, Dorcus (Macrodorcus) rectus (Motschulsky) (Col., Lucanidae)

Abstract:  Stag beetles are xylophagous insects that feed mainly on dead wood. They play an important role in the decomposition of dead wood in forest ecosystems. Most dead wood contains 1% nitrogen at most. It is suspected that stag beetles can utilize atmospheric nitrogen. We show that the larvae of Dorcus ( Macrodorcus ) rectus exposed to nitrogen reduce acetylene to ethylene in a time-dependent fashion. No reaction was detected with the dead wood or autoclaved larvae, suggesting that living larvae use the reaction for fixing nitrogen. Acetylene reduction to ethylene by larvae increased with incubation time. This effect was not seen using decayed wood only, autoclaved wood only or autoclaved larvae. Acetylene reduction by the larva proceeded at 1.25 ± 0.37 nmol acetylene/h/g (fresh wt), corresponding to the fixation of 0.25  μ g nitrogen per day per larva.     

The community structures of the coniferous and deciduous dead wood‐dwelling arthropods in Korea

We examined the structure of the arthropod community among deciduous and coniferous dead woods along the process of wood decay. We collected dead wood‐dwelling arthropods from April 2010 to October 2011 by using a vacuum aspirator and an electric chain saw in three areas (Mt. Woonak, Mt. Wolchul, Mt. Jingang) in Korea. We identified them to species levels and classified them into functional groups. We collected 8792 arthropods (5 classes, 20 orders, 58 families, and 93 species). The species richness and abundance of arthropods increased with the progress of decay in dead woods. The evenness index seemed to be shown at a lower value at late decay stage than at early‐ and mid‐decay stages. The diversity index (H′) in conifers was lower than that in deciduous dead woods at the early decay stage but this situation was reversed at the late decay stage. Arthropod communities of functional groups, except the xylophagous insects, did not differ in the variables, but the proportion of xylophagous insects increased as the decay stages progressed. The abundance of arthropods and xylophagous was statistically significantly different. The patterns generated by non‐metric multidimensional scaling in the overall arthropod community composition revealed that the species composition between study areas were significantly different. We confirmed that dead woods play very important roles as arthropods' habitats. Thus, we suggest that the role of dead woods should be emphasized in the management of forest ecosystems.     

The importance of forest type,tree species and wood posture to saproxylic wasp (Hymenoptera) communities in the southeastern United States

Although the forests of the southeastern United States are among the most productive and diverse in North America, information needed to develop conservation guidelines for the saproxylic (i.e., dependent on dead wood) fauna endemic to the region is lacking. Particularly little is known about the habitat associations and requirements of saproxylic parasitoids even though these organisms may be even more vulnerable than their hosts. We sampled parasitoids emerging from dead wood taken from two forest types (an upland pine-dominated forest and a lowland hardwood-dominated forest), three tree species (Liquidambar styraciflua L., Pinus taeda L., and Quercus nigra L.) and two wood postures (standing dead trees (i.e., snags) and fallen logs) in South Carolina. Parasitoid abundance did not differ between forest types or among tree species, but did differ between wood postures, being higher in snags than logs. This difference may have been due to the logs being in contact with the ground or surrounding vegetation and therefore less accessible to parasitoids. Parasitoid abundance and density decreased with height on both snags and logs. Species richness did not differ between forest types, among tree species or between wood postures. According to analysis of similarities, parasitoid communities did not differ between forest types, but did differ among tree species. The wasp communities associated with the different tree species and posture combinations were distinct. In addition, communities associated with the upper boles and crowns of snags were distinct from those occurring lower on snags. These results emphasize the importance of maintaining tree diversity in managed forests as well as retaining or creating entire snags at the time of harvest.     

Plant traits and wood fates across the globe: rotted,burned, or consumed?

Wood represents the defining feature of forest systems, and often the carbon in woody debris has a long residence time. Globally, coarse dead wood contains 36–72 Pg C, and understanding what controls the fate of this C is important for predicting C cycle responses to global change. The fate of a piece of wood may include one or more of the following: microbial decomposition, combustion, consumption by insects, and physical degradation. The probability of each fate is a function of both the abiotic environment and the wood traits of the species. The wood produced by different species varies substantially in chemical, micro- and macro-morphological traits; many of these characteristics of living species have 'afterlife' effects on the fate and turnover rate of dead wood. The colonization of dead wood by microbes and their activity depends on a large suite of wood chemical and anatomical traits, as well as whole-plant traits such as stem-diameter distributions. Fire consumption is driven by a slightly narrower range of traits with little dependence on wood anatomy. Wood turnover due to insects mainly depends on wood density and secondary chemistry. Physical degradation is a relatively minor loss pathway for most systems, which depends on wood chemistry and environmental conditions. We conclude that information about the traits of woody plants could be extremely useful for modeling and predicting rates of wood turnover across ecosystems. We demonstrate how this trait-based approach is currently limited by oversimplified treatment of dead wood pools in several leading global C models and by a lack of quantitative empirical data linking woody plant traits with the probability and rate of each turnover pathway. Explicitly including plant traits and woody debris pools in global vegetation climate models would improve predictions of wood turnover and its feedback to climate.     

Large proportion of wood dependent lichens in boreal pine forest are confined to old hard wood

Intensive forest management has led to a population decline in many species, including those dependent on dead wood. Many lichens are known to depend on dead wood, but their habitat requirements have been little studied. In this study we investigated the habitat requirements of wood dependent lichens on coarse dead wood (diameter >10 cm) of Scots pine Pinus sylvestris in managed boreal forests in central Sweden. Twenty-one wood dependent lichen species were recorded, of which eleven were confined to old (estimated to be >120 years old) and hard dead wood. Almost all of this wood has emanated from kelo trees, i.e. decorticated and resin-impregnated standing pine trees that died long time ago. We found four red-listed species, of which two were exclusive and two highly associated with old and hard wood. Lichen species composition differed significantly among dead wood types (low stumps, snags, logs), wood hardness, wood age and occurrence of fire scars. Snags had higher number of species per dead wood area than logs and low stumps, and old snags had higher number of species per dead wood area than young ones. Since wood from kelo trees harbours a specialized lichen flora, conservation of wood dependent lichens requires management strategies ensuring the future presence of this wood type. Besides preserving available kelo wood, the formation of this substratum should be supported by setting aside P. sylvestris forests and subject these to prescribed burnings as well as to allow wild fires in some of these forests.     

Succession of fungi on dead and live wood in brackish water in Brunei

We observed the sequence of fungi appearing on submerged wood of Hibiscus tiliaceus that initially was either dead or alive. Branches that were dead, but still attached to the tree, and live branches were cut from H. tiliaceus in the riparian vegetation in a brackish habitat on the Tutong River, Brunei. Branch segments were connected to the riverbank using monofilament line. Samples were examined for fungi before the branches were placed in the river and after the branches had been submerged 3 or 6 mo. Fifty taxa were found on the samples. Before being placed in the water different fungal assemblages were found on live as compared to deadwood. Branches that were alive when cut supported a distinctly different fungal assemblage after 3 mo in the water. Dead branches after 3 mo and both dead and initially live samples after 6 mo had been colonized by a fungal assemblage that is typical at this site. It is unknown whether the differences in colonization of dead and initially live wood can be attributed to differences in the substratum (i.e., the presence or absence of bark), inhibitory substances in more recently live wood or to assembly rules resulting from the different fungi that already were present in dead and live branches.     

Habitat heterogeneity affects predation of European pine sawfly cocoons

Habitat heterogeneity is thought to affect top‐down control of herbivorous insects and contribute to population stability by providing a more attractive microhabitat for natural enemies, potentially leading to reduced population fluctuations. Identifying the parameters that contribute to habitat heterogeneity promoting top‐down control of herbivorous insects by natural enemies could facilitate appropriate management decisions, resulting in a decreased risk of pest insect outbreaks because of a higher level of predation. In our study, we measured the top‐down pressure exerted by small mammals on the cocoons of a notorious pest insect in pine forests, the European pine sawfly (Neodiprion sertifer), which is known to be regulated by small mammal predation. The forest stands used differed in heterogeneity measured in terms of differences in tree diversity and density, understory vegetation height, presence/absence, and density of dead wood. We found higher predation in more dense spots within forest stands. Further, the effect of dead wood on sawfly cocoon predation depended on the pine proportion in forest stands. The addition of dead wood in a manipulation experiment had a slight positive effect on cocoon predation, while dead wood removal caused a clear decrease in predation rate, and the decrease was more pronounced when the proportion of pine increased. Our results show that habitat heterogeneity affects predation by generalist predators on herbivorous insects. This knowledge could be applied to reduce the risk of insect outbreaks by applying management methods that increase heterogeneity in perennial systems such as forests and orchards, thus decreasing the levels of insect damage.     

Influence of global warming on forest coleopteran communities with special reference to ambrosia and bark beetles

Forest insect pests are one of the major disturbance factors in forest ecosystems and their outbreaks are expected to be more severe under the influence of global warming. Coleopterans are dominant among forest insects and their ecological functions include general detritivores, dead wood feeders, fungivores, herbivores, live wood feeders and predators. Ambrosia and bark beetles contribute to ecological succession of forests and, therefore, ecological functions of forests can be changed in response to their outbreaks. Mountain pine beetle (MPB) outbreaks are the most dramatic example of changes in the ecological functions of forest due to the outbreak of a forest insect pest altered by global warming. Composition of coleopteran species varies with latitude. However, composition of functional groups is consistent with latitude which indicates that resources available to beetles are consistent. In coleopteran communities, ambrosia and bark beetles can become dominant due to increases of dead or stressed trees due to the warming climate. This can also induce changes in the ecological functions of coleopterans, i.e. selective force to displace trees that have lower ecological fitness due to temperature increase. Therefore, recent increases in the density ambrosia and bark beetles offer a chance to study ecological processes in forests under the influence of global warming.     

Habitat preferences of oak-feeding xylophagous beetles in a temperate woodland: implications for forest history and management

Oaks host the richest fauna of saproxylic insect in Europe. We studied habitat preferences of two beetle families, Buprestidae and Cerambycidae, by rearing the beetles from standardised oak timber baits. Species density was higher in the understorey than in the canopy; and in sun-exposed baits if within the understorey. Insolation was the most important factor affecting the composition of reared assemblages (explaining ca. 30% of variation in the data), followed by vertical stratum (ca. 10%). Local dead wood volume had no effect. The high preference for sun-exposed wood located near the ground suggests that: (i) open-canopy woodlands had to be rather common in temperate Europe; (ii) oak-utilising xylophages would benefit from restoration of management practices such as coppicing or woodland pasture; (iii) the policy of increasing dead wood volume in commercial forests is principally correct, but its success will depend on dead wood location within the forests.     

木材腐朽菌在森林生态系统中的功能

木材腐朽菌是森林生态系统的重要组成部分,在森林生态系统中起着极为重要的降解还原作用,主要包括担子菌门非褶菌目、子囊菌门盘菌纲和半知菌类的部分真菌,能全部或部分降解木材中的木质素、纤维素和半纤维素,其降解机制有3种：白色腐朽、褐色腐朽和软腐朽．木材腐朽菌与生态系统中其它生物关系密切,为很多昆虫、鸟类提供营养,有些昆虫也能使木腐菌得到传播．保护木材腐朽菌的生物多样性是保护森林生态系统、维护生态系统健康的重要因素．     

Priority effects of early successional insects influence late successional fungi in dead wood

Community assembly is an integral process in all ecosystems, producing patterns of species distributions, biodiversity, and ecosystem functioning. Environmental filters and colonization history govern the assembly process, but their relative importance varies depending on the study system. Dead wood decomposition is a slow process, allowing decomposer communities to develop within a slowly changing substrate for decades. Despite this, there are few long‐term studies of priority effects from colonization history in this ecosystem. In this study, we investigate the importance of insects in early succession of dead wood on the fungal community present one decade later. Sixty aspen trees were killed in two study landscapes, each tree producing one aspen high stump and log. Insects were sampled with flight interception traps during the first 4 years after tree death, and fungal fruiting bodies were registered in year twelve. We found positive priority effects of two fungivorous beetles, the sap beetle Glischrochilus quadripunctatus and the round fungus beetle Agathidium nigripenne, on the Artist''s bracket (Ganoderma applanatum) and a positive priority effect of wood‐boring beetles on the ascomycete Yellow fairy cup (Bisporella citrina). The Aspen bracket (Phellinus tremulae) did not respond to insects in early succession of the dead wood. Our results suggest that early successional insects can have significant, long‐lasting effects on the late successional fungal community in dead wood. Also, the effect can be specific, with one fungus species depending on one or a few fungivorous beetle species. This has implications for decomposition and biodiversity in dead wood, as loss of early colonizing beetles may also affect the successional pathways they seem to initiate.     

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.     

Interaction type and intimacy structure networks between forest-dwelling organisms and their host trees

Species interact in many ways. Potentially, the type of interaction, e.g. mutualistic, commensalistic or antagonistic, determines the structure of interaction networks, but this remains poorly tested. Here we investigate whether epiphytes and wood decomposers, having different types of interaction with their host trees, show different network properties. We also test whether the traits of host trees affect network architecture. We recorded presence/absence of organisms colonizing trees, and traits of host trees, in 102 forest plots. Epiphytic bryophytes (64 species) and lichens (119 species) were recorded on c. 2300 trees. Similarly, wood-inhabiting fungi (193 species) were recorded on c. 900 dead wood items. We studied the patterns of species aggregation on host trees by comparing network metrics of species specialization, nestedness and modularity. Next, we tested whether the prevalence of interactions was influenced by host tree traits. We found non-random interaction patterns between host trees and the three ecological groups (bryophytes, lichens and fungi), with nested and modular structures associated with high host specificity. A higher modularity and number of modules was found for fungi than for epiphytes, which is likely related to their trophic relationship with the host plant, whilst the stronger nestedness for epiphytes is likely reflecting the commensalistic nature of their interactions. For all three groups, the difference in prevalence of interaction across modules was determined by a gradient in interaction intimacy (i.e. host tree specialization), driven by host tree traits. We conclude that the type of interaction with host trees defines the properties of each network: while autotrophic epiphyte networks show similar properties to mutualistic networks, the heterotrophic wood decomposers show similarity with antagonistic networks.     

Influence of macroclimate and local conservation measures on taxonomic,functional, and phylogenetic diversities of saproxylic beetles and wood-inhabiting fungi

Wood-inhabiting fungi and saproxylic beetles are threatened by habitat degradation. Our understanding of the importance of macroclimate and local factors determining their taxonomic diversity has increased, but determinants of functional and phylogenetic diversity are poorly understood. We investigated assemblages of wood-inhabiting fungi and saproxylic beetles along a 1000 m elevational gradient of a temperate low mountain range. We (i) tested the relative importance of macroclimate (i.e. elevation) and local variables (microclimate, i.e. canopy closure, amount and diversity of dead wood) in determining observed and rarefied diversities and (ii) explored whether determinants of observed functional and phylogenetic diversities match those of taxonomic diversity. For both taxa, the determinants of observed phylogenetic and functional diversities largely matched those of taxonomic diversity. The diversity of wood-inhabiting fungi was predominantly determined by local variables, whereas that of saproxylic beetles was determined by both local variables and elevation. Taxonomic and phylogenetic diversities of saproxylic beetles decreased with increasing elevation, but standardized functional richness and entropy of both groups increased with increasing elevation. Diversities of wood-inhabiting fungi increased with canopy closure, while diversities of saproxylic beetles decreased with increasing canopy closure. Microclimate and dead-wood amount and diversity affected the observed and rarefied diversity of both saproxylic taxa, which justifies conservation actions that focus on attributes of dead wood and canopy cover. The contrasting responses of fungi and beetles highlight the need for amounts of diverse dead wood in the various microclimates to preserve functional and phylogenetic diversities of saproxylic organisms.     

森林溪流倒木生态学研究进展

溪流倒木是指在河流中长度大于1 m、直径大于10 cm的死木。溪流倒木在森林河流中(特别是较小的河流中)是一个常见且重要的结构成分。该文综述了近30年溪流倒木的研究成果(主要来自北美),总结溪流倒木在河流形态、碳循环、泥沙与养分拦截、水生生境的形成、水生生物多样性等方面的生态功能,倒木的时间动态性与空间变异性,以及干扰(包括自然干扰与人为干扰)与倒木存留量及分布的关系。此外,该文也探讨了溪流倒木的生态管理模式及未来研究方向。大量的研究证明,溪流倒木对森林水生生态系统具有重要的生态功能,但它的存留量、分布以及它的生态意义因所研究的森林生态系统、河流大小不同而异。随着河流宽度的增加, 倒木的存留量及它对河流的影响减少,并且倒木的分布以单个为主变为聚集体为主。单个倒木的直径则随河流宽度增加而增大。倒木也呈现十分明显的时间动态性,而这种动态往往是由大规模毁灭性的森林干扰(火、风倒等)所驱动的。研究倒木的时空变异性及自然干扰与人为干扰对倒木的不同影响对于保持倒木的生态功能是十分必要的。该文还对中国开展溪流倒木的生态研究提出一些建议。     

Changes in the dominant assembly mechanism drive species loss caused by declining resources

The species–energy hypothesis predicts that more productive areas support higher species richness. Conversely, when resources are reduced, species richness is reduced. Empirical tests of whether extinctions are predominantly caused by environmental constraints or competitive exclusion are lacking. We experimentally reduced dead wood to c. 15% of the initial amount after a major windstorm and examined changes in assembly mechanisms by combining trait‐based and evolutionary species dissimilarities of eight taxonomic groups, differing in their dependence on dead wood (saproxylic/non‐saproxylic). Species richness and assembly mechanisms of non‐saproxylic taxa remained largely unaffected by removal of dead wood. By contrast, extinctions of saproxylic species were caused by reversing the predominant assembly mechanisms (e.g. increasing importance of competitive exclusion for communities assembled through environmental filtering or vice versa). We found no evidence for an intensification of the predominant assembly mechanism (e.g. competitive exclusion becoming stronger in a competitively structured community).     

Colonisation of ephemeral forest habitats by specialised species: beetles and bugs associated with recently dead aspen wood

The most appropriate strategy for preserving fragmented populations depends on a species’ ability to colonise distant habitat patches. Insects associated with early decay stages of dead wood are expected to have a high capacity to colonise new habitat patches. To study the dispersal ranges of beetles (Coleoptera) and flat bugs (Hemiptera: Aradidae) dependent on recently dead aspen (Populus tremula) wood in Finland, we set out 58 piles of recently cut aspen logs at various distances up to 1.6 km from forests that contained a high density of old aspen trees. We captured insects by trunk window-traps, and counted beetles’ exit holes. Habitat connectivity was measured in terms of the amount of suitable aspen-wood in the surrounding environment, with the closest dead wood items up-weighted by a negative-exponential function. The log-piles attracted many saproxylic insects including four red-listed aspen-specialist species. The exposure of log-piles to the sun, and high levels of habitat connectivity increased the species richness of aspen-specialists, whereas bark peeling by moose decreased richness. The spatial scale at which species richness had its strongest response to habitat was 93 m. Among individual species there was a wide variability in spatial scale of response. This study supports the view that conservation efforts in boreal forests should be concentrated on sites where colonisation by target species is most likely. Restoration of habitat by re-locating logs may be useful at localities with a rich and specialised fauna but which have too low rate of formation of dead wood by natural processes.     

What do we know about biological nitrogen fixation in insects? Evidence and implications for the insect and the ecosystem

Many insects feed on a low‐nitrogen diet, and the origin of their nitrogen supply is poorly understood. It has been hypothesized that some insects rely on nitrogen‐fixing bacteria (diazotrophs) to supplement their diets. Nitrogen fixation by diazotrophs has been extensively studied and convincingly demonstrated in termites, while evidence for the occurrence and role of nitrogen fixation in the diet of other insects is less conclusive. Here, we summarize the methods to detect nitrogen fixation in insects and review the available evidence for its occurrence (focusing on insects other than termites). We distinguish between three aspects of nitrogen fixation investigations: (i) detecting the presence of potential diazotrophs; (ii) detecting the activity of the nitrogen‐fixing enzyme; and (iii) detecting the assimilation of fixed nitrogen into the insect tissues. We show that although evidence from investigations of the first aspect reveals ample opportunities for interactions with potential diazotrophs in a variety of insects, demonstrations of actual biological nitrogen fixation and the assimilation of fixed nitrogen are restricted to very few insect groups, including wood‐feeding beetles, fruit flies, leafcutter ants, and a wood wasp. We then discuss potential implications for the insect's fitness and for the ecosystem as a whole. We suggest that combining these multiple approaches is crucial for the study of nitrogen fixation in insects, and argue that further demonstrations are desperately needed in order to determine the relative importance of diazotrophs for insect diet and fitness, as well as to evaluate their overall impact on the ecosystem.     

Nutritional dynamics during the development of xylophagous beetles related to changes in the stoichiometry of 11 elements

The present study examines the adaptive strategy used by wood‐boring beetles to compensate for the lack of nutrients in dead wood. The contents of nutritional elements in growing wood‐boring beetles (Stictoleptura rubra L. and Chalcophora mariana Dejean) are compared with the elemental composition of decaying dead wood (pine stumps), showing changes during the beetles' ontogenetic (i.e. larval) development. The stoichiometric ratios of C and other nutritional elements (N, P, K, Na, Ca, Mg, Fe, Zn, Mn and Cu) are investigated to identify the most important nutrients for larval development. The degree of nutritional mismatch that is encountered by the beetle larvae changes dramatically over 3–4 years of simultaneous larval growth and wood decay. Excluding C, the relative contents of nutritional elements increase substantially in decaying wood, whereas the opposite tendency is found in larvae, most likely because of carbon deposition in fat. The elements limiting larval development because of their scarcity in dead wood are N, P, K, Na, Mg, Zn and Cu. Fungal activity (i.e. the transport of nutrients from the surrounding environment to decaying stumps) can explain the observed mitigation of the original mismatch, although prolongation of the larval development time is still necessary to compensate for the scarcity of some of the required elements in food.     

Spatial and temporal scales relevant for conservation of dead-wood associated species: current status and perspectives

Dead wood is a key substrate for forest biodiversity, hosting a rich and often threatened biodiversity of wood-living species. However, the relationship between the occurrence of dead wood and associated species is modified by several environmental factors. Here we review the present state of knowledge on how dead wood on different spatial and temporal scales affects saproxylic biodiversity. We searched for peer-reviewed studies on saproxylic species that compared dead wood distribution on at least two spatial or temporal scales. We scanned close to 300 articles, of which 34 fit our criteria. 20 studies were directed towards the current amount of dead wood at different scale levels and how this relates to the abundance or occurrence of saproxylic species, embracing scales from 10 m to 10 km. 14 studies compared time-lagged effects of dead wood, covering time-lags from 25 years to more than 200 years. The reviewed articles focused mainly on European forest and addressed invertebrates (mostly beetles), alone or in combination with fungi (27 articles), fungi (six articles), or lichens (one article). Although the significance of dead wood for forest biodiversity is firmly established, the reviewed studies show that we still have limited knowledge of the relationship between saproxylic biodiversity and spatial and temporal scales. Based on the reviewed studies, we conclude that there is large variation in response to spatial and temporal dead wood patterns between different taxa and sub-groups. Still, several of the reviewed papers indicate that time-lagged effects deserve more attention, especially on a landscape scale and for specialized or red-listed species. Further work is required before firm management recommendations can be suggested.