Influence of sampling effort on saproxylic beetle diversity assessment: implications for insect monitoring studies in European temperate forests

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In search of the best local habitat drivers for saproxylic beetle diversity in temperate deciduous forests

Deadwood-associated species are increasingly targeted in forest biodiversity conservation. In order to improve structural biodiversity indicators and sustainable management guidelines, we need to elucidate ecological and anthropogenic drivers of saproxylic diversity. Herein we aim to disentangle the effects of local habitat attributes which presumably drive saproxylic beetle communities in temperate lowland deciduous forests. We collected data on saproxylic beetles in 104 oak and 49 beech stands in seven French lowland forests and used deadwood, microhabitat and stand features (large trees, openness) as predictor variables to describe local forest conditions. Deadwood diversity and stand openness were consistent key habitat features for species richness and composition in deciduous forests. Large downed deadwood volume was a significant predictor of beetle species richness in oak forests only. In addition, the density of cavity- and fungus-bearing trees had weak but significant effects. We recommend that forest managers favor the local diversification of deadwood types, especially the number of combinations of deadwood positions and tree species, the retention of large downed deadwood and microhabitat-bearing trees in order to maximize the saproxylic beetle diversity at the stand scale in deciduous forests. To improve our understanding of deadwood-biodiversity relationships, further research should be based on targeted surveys on species-microhabitat relationships and should investigate the role of landscape-scale deadwood resources and of historical gaps in continuity of key features availability at the local scale.     

Habitat requirements of the endangered longhorn beetle <Emphasis Type="Italic">Aegosoma scabricorne</Emphasis> (Coleoptera: Cerambycidae): a possible umbrella species for saproxylic beetles in European lowland forests

Detailed knowledge of habitat requirements is an essential prerequisite for efficient conservation of any endangered species. Despite the grain support beetle Aegosoma scabricorne (Cerambycidae) being one of the largest European beetles, and an endangered, disappearing, species in Central Europe, its bionomics remain relatively poorly known. A. scabricorne is known as a polyphagous species on broadleaved trees; thus, to investigate its habitat preferences 174 broadleaved trees (87 occupied and 87 unoccupied by the species) were surveyed in the area of southern Moravia (Czech Republic) in 2015. The species was found to be strongly associated with declining or freshly dead trees that are, preferably, further damaged (breakage of stem or primary branch, hollows, etc.) and it particularly thrives on large trees (diameter >50 cm). Surprisingly, stem exposure to the sun was shown to be an unimportant characteristic for this species. However, in this study the number of exit holes was significantly smaller on the shaded north-facing quarter of the stem. Our results suggest concrete conservation measures to support the species. We also suggest that this species may be used as an umbrella species for saproxylic beetles of European lowland forests.     

The saproxylic beetle Bolitophagus reticulatus: its frequency in managed forests, attraction to volatiles and flight period

Reduction in amount of dead trees in Scandinavian forests due to intensive forest management has aroused interest in the requirements of saproxylic (wood-living) species. Much attention has been focused on Bolitophagus reticulatus , a tenebrionid beetle that lives on polypore fruiting bodies of Fomes fomentarius . Although earlier studies have suggested that the species has limited dispersal abilities, the beetle is abundant and widely distributed in Swedish forests. To resolve this paradox we conducted a series of field experiments. Firstly, the frequency of the species was found to be similar in six forest stands with a low density of fruiting bodies and six stands with a high density. Secondly, flying individuals were caught in window traps placed in all of eight randomly chosen, well-managed pine stands. Thirdly, we tested volatiles that flying beetles may use to find hosts by baiting window traps. We found strong attractions to ethanol and to a combination of ethanol and fruiting body. The dispersal period was almost entirely restricted to one week during the summer. Our results suggest that B. reticulatus is distributed wherever the host occurs. Long distance dispersal of B. reticulatus is difficult to observe, but appears to be effective when it does occur. Previously observed patchy distribution patterns of the species are probably due to short-distance movements being more frequent than long-distance movements. A general conclusion is that interpretations based on mark-recapture data and indirect measures of substrate and beetle occurrences that claims limited dispersal should be treated with caution.     

Effect of deadwood management on saproxylic beetle richness in the floodplain forests of northern Italy: some measures for deadwood sustainable use

Saproxylic beetles may act as bio-indicators of high-quality mature woodlands, and their conservation is strongly linked to the quality and quantity of deadwood in a biotope. We tested the effect of deadwood accumulation and habitat variables on saproxylic species richness by investigating six sampling sites under different deadwood management practices that belong to both alluvial and riparian mixed forests of the Po plain, Italy. We sampled 43 obligate saproxylic species. The main factor predicting saproxylic species richness was the amount of deadwood measured by both log diameter and volume. We found a threshold of 0.22 m diameter (confidence interval CI 0.18–0.37 m) and 32.04 m³/ha volume (CI 16.09–64.09 m³/ha) below which saproxylic beetle richness would be significantly reduced and a threshold of 35 m³/ha dead wood volume (CI 33–40 m³/ha) over which species richness increases by <5 %. The other deadwood and environmental components influenced saproxylic beetle richness to a lesser extent; some of them, however, should still be considered for proper management. Forest structure variables describing forest density such as large trees and basal areas have a negative effect on species richness. According to the results of our study, stumps and advanced decaying class are positively correlated, while small logs are negatively correlated to species richness. Thus, in extensively managed forests, the regular cutting of trees should be implemented to create artificial stumps, in order to assure a continuity of deadwood and, in the meantime, increase the number and width of openings in the forest. Moreover, prolonging rotation times can assure the presence of deadwood at intermediate/later stages of decay.     

Surveying an endangered saproxylic beetle, Osmoderma eremita, in Mediterranean woodlands: a comparison between different capture methods

Measuring population size is riddled with difficulties for wildlife biologists and managers, and in the case of rare species, it is sometimes practically impossible to estimate abundance, whereas estimation of occupancy is possible. Furthermore, obtaining reliable population size estimates is not straightforward, as different sampling techniques can give misleading results. A mark-recapture study of the endangered saproxylic beetle Osmoderma eremita was performed in central Italy by applying four independent capture methods within a study area where 116 hollow trees were randomly selected to set traps. Detection probability and population size estimates were drawn from each of these four capture methods. There were strong differences in detection probability among methods. Despite using pheromone and beetle manipulation, capture histories were not affected by trap-happiness or trap-shyness. Population size estimates varied considerably in both abundance and precision by capture method. A number of 0.5 and 0.2 adult beetles per tree was estimated using the whole data set by closed and open population models, respectively. Pitfall trap appeared the optimal method to detect the occurrence of this species. Since in the southern part of its distribution range, a single population of O. eremita is widespread in the landscape, and includes beetles from more than one hollow tree, conservation efforts should focus not only on preserving few and isolated monumental hollow trees, but should be extended to large stands.     

Diameter of downed woody debris does matter for saproxylic beetle assemblages in temperate oak and pine forests

Deadwood is widely recognized to be an important issue for biodiversity conservation in forest ecosystems. Establishing guidelines for its management requires a better understanding of relationships between woody debris characteristics and associated species assemblages. Although deadwood diameter has been identified as an important factor predicting occurrence of many saproxylic species, the boundary between small and large diameter has not yet been precisely defined. In commercial forests, it is also of critical importance to know which diameter is large enough to host the beetle species associated with large logs in order to ensure cost-effectiveness of biodiversity conservation measures. We investigated the differences in saproxylic beetle assemblages among four different diameter classes of downed woody oak and maritime pine debris, in France. Beetles were sampled using in situ emergence traps. The diameter of deadwood pieces ranged from 1 to 40 cm. No patterns of nestedness associated with the gradient of diameter size were identified for either tree species. More indicator saproxylic species were observed in large logs and branches than in small logs. A clear distinction appeared in assemblage composition around the 5-cm diameter threshold whereas no similar pattern occurred around the 10 cm value, i.e. the classical threshold used in forestry to distinguish fine woody debris from coarse woody debris. For both tree species, the mean body length of beetles increased with the diameter of deadwood suggesting that the quantity of available resources per piece may constitute a limiting factor for large beetle species. This study confirms that not only large deadwood pieces are relevant for saproxylic biodiversity conservation but also the smallest pieces. Therefore, forest managers would be well advised to maintain a high diversity of deadwoods to maintain saproxylic biodiversity.     

Shortage of declining and damaged sun-exposed trees in European mountain forests limits saproxylic beetles: a case study on the endangered longhorn beetle <Emphasis Type="Italic">Ropalopus ungaricus</Emphasis> (Coleoptera: Cerambycidae)

European mountain forests are unique ecosystems, and they harbour specific saproxylic beetle fauna, including relict species such as the longhorn beetle Ropalopus ungaricus (Herbst, 1784). This endangered species is endemic to European mountain forests and is considered to be monophagous on living sycamore trees (Acer pseudoplatanus L.). Insufficient knowledge of the species ecology limits its effective conservation. Hence, to investigate its habitat requirements, 175 sycamore trees (87 occupied and 88 unoccupied by the species) were surveyed in the Hrubý Jeseník Mts. and Moravskoslezské Beskydy Mts. in northern Moravia (Czech Republic) in 2016. The species strongly preferred trees with reduced vitality, including variously damaged trees. The species was also documented to be much more abundant on sun-exposed trees, and the parts of the trunk mantle (south- and east-facing quarters) with the warmest microclimates were preferred. Surprisingly, trunk diameter seems to be an unimportant characteristic for this species (the diameters of occupied trees varied between 5 and 72 cm). In conclusion, R. ungaricus is a specialised species whose survival depends upon a continuous supply of sun-exposed declining and damaged sycamores in mountain forests. Based on the results of the present study, we suggest concrete conservation measures to support the species, and these will benefit other saproxylic species associated with the same habitat.     

中国濒危雉类生态学研究进展

中国是世界上雉类最为丰富的国家。世界16属51种雉类中．中国有12属27种．包括19个特有种。近几十年来．由于人类活动的影响,野生雉类的数量锐减,越来越多的珍稀雉类濒临灭绝。《世界受胁鸟类名录》(BirdLfe International,2000)列出的29种雉类中,中国有12种。世界雉类协会(WPA)和世界自然资     

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.     

Vegetation surveys in the circumboreal coniferous forests: A review

As an introduction to the special issue “Vegetation of Circumboreal Coniferous Forests”, a brief history is provided of the development and present state of floristic vegetation surveys of the boreal and mountain coniferous forests of Europe, northern Asia and North America. The focus is on forests assigned to the phytosociological classVaccinio-Piceetea and closely related vegetation types.     

Effects of forest continuity on flying saproxylic beetle assemblages in small woodlots embedded in agricultural landscapes

Ancient forests are of considerable interest for strategies for biodiversity conservation. However, in European forest landscapes fragmented and harvested for a long time forest continuity might be no longer a key driver for flying organisms such as saproxylic beetles. In a study based on paired samples (n = 60 stands, p = 180 traps) of ancient and recent forests, we investigated the effects of forest continuity on saproxylic beetle assemblages in two French regions. Mean species richness was significantly related with deadwood volume in ancient forests, but not in recent forests. This loss of relationship between assemblages and their environment suggests that dispersal limitation is at work, at least for some species. Forest continuity had a significant effect on mean species richness and on the mean number of common species, but not on rare species. Forest continuity had a significant effect on assemblage composition in one out of the four cases tested. In both regions, we identified species associated with either recent or ancient forests. Finally, mean body size of species was significantly smaller in recent forests compared with ancient ones, as was their tree diameter preference, despite a higher volume of large deadwood in recent forests. These results lend support to using forest continuity as a criterion to identify sites of conservation importance, even in highly fragmented landscapes.     

Diversity distribution of saproxylic beetles in Chilean Mediterranean forests: influence of spatiotemporal heterogeneity and perturbation

Mediterranean ecosystems have been recognized as a priority for biodiversity conservation due to their high levels of species richness and endemism. In South America, these environments are restricted to central Chile and represent a biodiversity hotspot. The study of saproxylic beetles in this area is an unexplored topic, despite the ecological role they play in these ecosystems and their potential usefulness for monitoring the degree of forest conservation. We investigated the diversity distribution of trophic guilds of saproxylic beetles in Chilean Mediterranean forests, to identify the main environmental variables that influence their distribution. We also analyzed seasonal dynamics as a key factor influencing insect communities and the effect of human disturbance on their diversity and composition. We identified characteristic species of perturbation degree as bioindicators for habitat monitoring. A total of 40 window traps were used to survey three Mediterranean forest types in the Río Clarillo National Reserve and the adjacent non-protected areas. We found that the diversity, abundance and composition of saproxylic beetles varied significantly spatiotemporally among the studied forest types and among perturbation degrees, showing different patterns depending on the trophic guild. Results indicated that conservation decisions should include the preservation of a larger range of different vegetation types and the nearby zones that have suffered low levels of disturbance or fragmentation and where actions promoting the presence of old native trees would have a significant conservation value.     

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.     

Timber stacks: potential ecological traps for an endangered saproxylic beetle,the Rosalia longicorn <Emphasis Type="Italic">Rosalia alpina</Emphasis>

Ecological traps are serious, anthropogenic threats to animal populations. However, in certain cases it is difficult to determine whether they really act in the expected manner. This applies to the harmful effects of beech timber stacked in forests on the endangered saproxylic beetle Rosalia longicorn Rosalia alpina, which have been mentioned in numerous scientific articles, conservation action plans and similar publications. The aim of this paper is to determine whether beech timber stacks meet the criteria of an ecological trap for the Rosalia longicorn. Two basic criteria of such a trap are analysed: the attractiveness of timber stacks and the impossibility of complete larval development. The results show that beech timber stacks are highly attractive to Rosalia longicorn imagines. Moreover, the time during which the timber is stacked is shown to be significantly shorter than the species’ larval development period. These results suggest that timber stacks can be treated as operative ecological traps for the Rosalia longicorn, even though the extent of their influence on the demographic parameters of this beetle’s population has not been estimated. Forest management practices, i.e. increasing amounts and shifts in timing of wood storage, could intensify this threat.     

Chemical ecology and insect conservation: optimising pheromone-based monitoring of the threatened saproxylic click beetle Elater ferrugineus

Elater ferrugineus is a saproxylic click beetle inhabiting old deciduous trees in Europe. It is threatened throughout its area of distribution due to habitat loss. No efficient monitoring method has been available for this species, but observed attraction of females to (R)-(+)-γ-decalactone, which is a male-produced sex pheromone of its prey, the scarab beetle Osmoderma eremita, has led to the development of an odour lure for monitoring. In addition, four esters have recently been identified from the pheromone-producing gland in female E. ferrugineus, and a blend of these esters is highly attractive to conspecific males in the field, revealing an alternative odour-based method for monitoring this species. However, no rigorous analysis has been performed to check whether all four esters show biological activity in male E. ferrugineus, and whether its own sex pheromone is a more potent lure than the prey kairomone for monitoring of E. ferrugineus. In this study, we reinvestigated the E. ferrugineus sex pheromone, using electrophysiological and behavioural analyses, and found that only one of the esters, 7-methyloctyl (Z)-4-decenoate, is active. In addition, trapping experiments revealed that 7-methyloctyl (Z)-4-decenoate is a much more efficient attractant for male E. ferrugineus than the prey pheromone is for conspecific females, or any sex of O. eremita. With a very efficient odour lure at hand, novel information about current distribution, local population sizes, and dispersal ranges in E. ferrugineus can now be obtained, which can aid in conservation efforts to protect this threatened insect and its habitat.     

Process from pattern in the distribution of an endangered leaf beetle

We investigated whether signals of known dispersal processes and habitat patch turnover could be detected in a snapshot of the distribution of the tansy leaf beetle Chrysolina graminis among patches of its host plant tansy Tanacetum vulgare . Beetle occupancy in 1305 patches was analysed using autologistic generalised additive models (GAMs). These model spatial autocorrelation with an autocovariate calculated as the distance-weighted rate of occupancy among neighbouring patches. The autocovariate that best explained beetle occupancy was one which represented the active search for patches during beetle dispersal, included a distance weight that closely matched a previously fitted dispersal kernel and had neighbourhood sizes encompassing ∼95% of known dispersal distances. Autocovariates distinguishing between neighbours on the same and opposite riverbanks outperformed those that did not, revealing the river as a barrier to dispersal. Differentiating between up and downstream autocorrelation did not improve model fit, as is consistent with the beetle's lack of directional bias in dispersal. Habitat connectivity (the extent to which it was surrounded by other patches) did not appear to affect beetle occupancy in the field, while positive effects were found for distributions simulated from the GAM. We argue that this reflects a non-equilibrium distribution driven by slow responses to high rates of habitat patch turnover due to limited dispersal ability. Our findings suggest that presence/absence snapshots can reveal patterns of dispersal and be used to test whether species' ranges are at equilibrium. Such information is important for effective conservation so the possibility of inferring these patterns from distribution data is an appealing one.