Constancy and asynchrony of Osmoderma eremita populations in tree hollows

A species rich beetle fauna is associated with old, hollow trees. Many of these species are regarded as endangered, but there is little understanding of the population structure and extinction risks of these species. In this study I show that one of the most endangered beetles, Osmoderma eremita, has a population structure which conforms to that of a metapopulation, with each tree possibly sustaining a local population. This was revealed by performing a mark-release-recapture experiment in 26 trees over a 5-year period. The spatial variability between trees was much greater than temporal variability between years. The population size was on average 11 adults tree^-1 year^-1, but differed widely between trees (0-85 adults tree^-1 year^-1). The population size in each tree varied moderately between years [mean coefficient of variation (C.V.)=0.51], but more widely than from sampling errors alone (P=0.008, Monte Carlo simulation). The population size variability in all trees combined, however, was not larger than expected from sampling errors alone in a constant population (C.V.=0.15, P=0.335, Monte Carlo simulation). Thus, the fluctuations of local populations cancel each other out when they are added together. This pattern can arise only when the fluctuations occur asynchronously between trees. The asynchrony of the fluctuations justifies the assumption usually made in metapopulation modelling, that local populations within a metapopulation fluctuate independently of one another. The asynchrony might greatly increase persistence time at the metapopulation level (per stand), compared to the local population level (per tree). The total population size of O. eremita in the study area was estimated to be 3,900 individuals. Other localities sustaining O. eremita are smaller in area, and most of these must be enlarged to allow long-term metapopulation persistence and to satisfy genetic considerations of the O. eremita populations.     

Osmoderma eremita as an indicator of species richness of beetles in tree hollows

The beetle Osmoderma eremita has received much attention in the last few years, as it is among those species with the highest priority in the European Union's Habitat Directive. In this paper the species is evaluated as a potential indicator and umbrella species for the endangered beetle fauna in tree hollows. To be useful as an indicator of a species-rich fauna it should be easy to inventory and be strongly correlated with the presence of other species. An umbrella species is a species which is so demanding that the protection of this species will automatically save many others. The species richness of saproxylic beetles and occupancy of O. eremita were surveyed in tree hollows in an area in southeastern Sweden by assessing the presence/absence of living and dead adults (including fragments) and larvae. The species richness was higher when O. eremita was present, both at tree and stand level. Several threatened species were associated with the presence of O. eremita, whereas others did not correlate with the occurrence of O. eremita. As O. eremita is easy to find and identify, it is useful as an indicator of stands with a rich beetle fauna in tree hollows. Osmoderma eremita can be used as an umbrella species, because if measures are taken to conserve O. eremita, many other species in the same habitat are also conserved. However, there are some beetles in tree hollows which seem to be more sensitive to habitat fragmentation than O. eremita, and may go extinct if only O. eremita is taken into consideration.     

Habitat of Osmoderma eremita Scop. (Coleoptera: Scarabaeidae), a beetle living in hollow trees

The habitat of Osmoderma eremita, a vulnerable species in Europe restricted to tree cavities, was examined in southeastern Sweden. The occurrence of O. eremita larvae and fragments, larval frass and imagines were investigated in 135, 72 and 21 living oak trees with wood mould cavities, respectively. Living individuals and fragments were only found in hollows with frass. The correlation between different characteristics of the oaks and the occurrence of the beetle were examined by building multi-variate models with logistic regression. The frequency of O. eremita is higher in hollows with openings directed towards the sun (S or W) and in cavities with large amounts of wood mould. In one area the frequency was higher in trees which stand in an open or half open surrounding. The tendency to prefer sun exposed sites implies that the forestation of oak meadows, caused by cessation of traditional management, might be detrimental to the species.     

Microclimate in hollow trees and how it affects an inhabiting beetle species,Osmoderma eremita

1. Studies of species' responses to microclimatic conditions have increased our understanding of their habitat requirements and possible responses to climate warming. However, little is known about the role of microclimate for insects inhabiting hollow trees.
2. We explored the relationship between tree characteristics and microclimate, and analysed how the microclimate in tree-hollows affects the occurrence and body size of an endangered beetle species, Osmoderma eremita.
3. We placed temperature data-loggers in wood mould (= loose material in tree-hollows) and surveyed O. eremita in 47 hollows in oak pastures in south-eastern Sweden. We found that tree characteristics previously known to be associated with occurrence of beetle species confined to tree-hollows (larger diameters, more wood mould, entrances higher up, and not directed upwards) tend to decrease moisture and moisture variation, while their effects on temperature and temperature fluctuations differ during different seasons. This indicates that microclimatic conditions are important for beetles in hollow trees, and many specialised species seem to avoid conditions that are too moist.
4. O. eremita occurred more frequently in trees with a warmer and more stable microclimate, while adult body size decreased with a warmer microclimate. A positive effect of a warmer microclimate was expected, since the study was done near the northern margin of the species' range.
5. O. eremita is confined to living in hollow trees, which may be due to the microclimate there being more stable in comparison to both the ambient climate and the microclimate in standing and downed dead wood.

     

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.     

Air sampling of its pheromone to monitor the occurrence of Osmoderma eremita, a threatened beetle inhabiting hollow trees

Osmoderma eremita is a threatened scarab beetle living in the hollows of old deciduous trees and is regarded as an umbrella species of the beetle fauna associated with this habitat. Several methods like pitfall trapping and wood mould sampling have been used to monitor the occurrence of O. eremita, but these methods cannot be applied for trees with certain characteristics. Recently, (R)-(+)--decalactone was identified as a male-produced sex pheromone of the species. Here, we show that -decalactone can be detected in hollow trees by air sampling and that the presence of the compound is strongly correlated with the occurrence of living male beetles in the same trees. Air was sampled from tree cavities and extracts analysed using gas chromatography–and mass spectrometry. There was a 89% match between the detection of -decalactone in extracts and the occurrence of male O. eremita±2 days from the sampling event. In the absence of males, samples never contained -decalactone, and the presence of this compound in a tree cavity appears to be a good predictor of O. eremita occupancy. Air sampling can be a useful complement to other methods when trying to detect as many trees housing this beetle as possible, which is crucial when estimating populations sizes and developing conservation strategies for this species.     

Using odour traps for population monitoring and dispersal analysis of the threatened saproxylic beetles Osmoderma eremita and Elater ferrugineus in central Italy

Pheromone-based monitoring could be a very efficient method to assess the conservation status of rare and elusive insect species, but there are still few studies for which pheromone traps have been used to obtain information on presence, abundance, phenology and movements of such insects. We performed a mark-recapture study of two threatened saproxylic beetles, Osmoderma eremita (Scarabaeidae) and its predator Elater ferrugineus (Elateridae), in two beech forests of central Italy using pheromone baited window traps and unbaited pitfall traps. Two lures were used: (1) the male-produced sex pheromone of O. eremita (racemic γ-decalactone) to attract females of both species, and (2) the female-produced sex pheromone of E. ferrugineus (7-methyloctyl (Z)-4-decenoate), to attract conspecific males. In total, 13 O. eremita and 1,247 E. ferrugineus individuals were trapped. For E. ferrugineus, males were detected earlier than females, and 7-methyloctyl (Z)-4-decenoate was much more efficient lure compared to racemic γ-decalactone in detecting its presence. The population size at the two sites were estimated to 520 and 1,369 individuals, respectively. Our model suggests a sampling effort of ten traps checked for 3 days being sufficient to detect the presence of E. ferrugineus at a given site. The distribution of dispersal distances for the predator was best described by the negative exponential function with 1 % of the individuals dispersing farther than 1,600 m from their natal site. In contrast to studies on these beetles in Northern Europe, the activity pattern of the two beetle species was not influenced by variation in temperature during the season.     

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.     

Delayed dispersal and prolonged brood care in a family‐living beetle

Delayed juvenile dispersal is an important prerequisite for the evolution of family‐based social systems, such as cooperative breeding and eusociality. In general, young adults forego dispersal if there are substantial benefits to remaining in the natal nest and/or the likelihood of dispersing and breeding successfully is low. We investigate some general factors thought to drive delayed juvenile dispersal in the horned passalus beetle, a family‐living beetle in which young adults remain with their families in their natal nest for several months before dispersing. Fine‐scale population genetic structure indicated high gene flow between nest sites, suggesting that constraints on mobility are unlikely to explain philopatry. Young adults do not breed in their natal log and likely disperse before reaching breeding age, suggesting that they do not gain direct reproductive benefits from delayed dispersal. We also examined several ways in which parents might incentivize delayed dispersal by providing prolonged care to adult offspring. Although adult beetles inhibit fungal growth in the colony by manipulating both the nest site and deceased conspecifics, this is unlikely to be a major explanation for family living as both parents and adult offspring seem capable of controlling fungal growth. Adult offspring that stayed with their family groups also neither gained more mass nor experienced faster exoskeleton development than those experimentally removed from their families. The results of these experiments suggest that our current understanding of the factors underlying prolonged family living may be insufficient to explain delayed dispersal in at least some taxa, particularly insects.     

Restricted dispersal in a flying beetle assessed by telemetry

Many insects living in ancient trees are assumed to be threatened as a result of habitat loss and fragmentation. It is generally expected that species in habitats with low temporal variability in carrying capacity have lower degree of dispersal in comparison to those in more ephemeral habitats. As hollow trees are long-lived, species in that habitat are expected to be sensitive to habitat fragmentation, due to a low capacity to establish new populations far from present ones. Using radio telemetry, we studied the dispersal for a beetle, Osmoderma eremita, living in hollow trees. O. eremita exhibited philopatry and only dispersed over short ranges. About 82–88% of the adults remained in the tree where they were caught. All observed dispersal movements ended up in nearby hollow trees and 62% in the neighbouring hollow tree. These results corroborate the suggestion that habitat fragmentation may be detrimental to insects living in temporally stable but spatially variable habitats. In order to preserve such species, we propose that conservation efforts should be focused on maintaining or increasing the number of suitable trees in and near presently occupied stands.     

Boxes mimicking tree hollows can help conservation of saproxylic beetles

Old hollow trees have declined in Europe and many saproxylic (i.e. wood-dwelling) invertebrates living on them are threatened. The aim of this study was to investigate to what extent artificial habitats can be exploited by saproxylic beetles. To mimic the conditions in tree hollows, we constructed wooden boxes filled with different combinations of substrates like oak saw dust, oak leaves, a dead hen (Gallus domesticus), chicken dung, lucerne flour or potatoes and placed them on tree trunks. To investigate the importance of distance from dispersal sources, we placed boxes at different distances (0–1,800 m) from three species-rich sites with high densities of hollow oaks. Over 3 years, 3,423 specimens of 105 saproxylic beetle species were caught in 47 boxes. Among beetles found in hollow oaks that were either tree-hollow species, bird nest species, or wood rot species, 70% were also found in the boxes. A dead hen added to the artificial wood mould gave a higher number of beetle specimens. The number of species associated with tree hollows in oak decreased with distance from sites with hollow oaks. In conclusion, the prospects for using artificial environments for boosting substrate availability, or to fill spatial and temporal gaps therein, for saproxylic beetles are good.     

Evaluation of secondary dispersal in a large-seeded tree Aesculus turbinata: a test of directed dispersal

Gap characteristics and gap phase replacement of major tree species were examined in two primary old-growth (mean DBHs of the canopy trees were 45.2 and 56.1 cm) and four secondary developing (range of mean DBH of the canopy trees was 23.5–39.9 cm) beech (Fagus crenata) stands in the Daisen Forest Reserve, southwestern Japan, and these were analyzed in relation to stand development as expressed by the difference of mean DBH of canopy trees. Percentage gap area (percentage of total gap area to total surveyed area) and mean and maximum gap size varied widely and ranged from 1.7 to 20.0%, from 19.4 to 162.8 m2, and from 35.7 to 585.1 m2, respectively. Mean percentage gap area and mean gap size were significantly greater in old-growth than in developing stands. However, they and maximum gap size might not increase linearly with stand development, and the gap feature of less developed stands was greater than that of later stages in developing stands. The cause was a higher formation rate, in younger developing stands, of gaps formed by simultaneous death (multiple trees falling down in domino fashion) which tends to produce larger gaps. In developing stands mean DBH of gapmakers (canopy trees that formed a gap) was smaller than that of canopy trees, though the inverse trend might be found in old-growth stands. Three typical types of death or injury states of gapmakers such as standing dead, trunk broken and uprooted were found in every study stand and the difference in stand development may not cause stand-to-stand variations for them. Importance of F. crenata (the most dominant species) in the canopy layer increased and its importance in the understory layer decreased with stand development. Shade-intolerant Quercus mongolica in the canopy layer was more important in younger than in old-growth stands, and there was no Quercus regeneration in old-growth stands. Acer mono consistently appeared, though in much less abundance than other species, in both canopy and understory layers of all study stands. Sub-canopy layers, which are mainly formed by sub-canopy tree species such as Acanthopanax sciadophylloides and Acer japonicum, may gradually develop with stand development.p>     

Kin selection in den sharing develops under limited availability of tree hollows for a forest marsupial

Animal social behaviour is not static with regard to environmental change. Flexibility in cooperative resource use may be an important response to resource decline, mediating the impacts of resource availability on fitness and demography. In forest ecosystems, hollow trees are key den resources for many species, but are declining worldwide owing to forestry. Altered patterns of den sharing may mediate the effects of the decline of this resource. We studied den-sharing interactions among hollow-dependent Australian mountain brushtail possums to investigate how spatial variation in hollow tree availability affects resource sharing and kin selection. Under reduced den availability, individuals used fewer dens and shared them less often. This suggests increased territoriality in the presence of resource competition. Further, there was a switch from kin avoidance to kin preference with decreasing hollow tree availability. This was driven primarily by a change in den sharing among siblings. The inclusive fitness benefits of den sharing with kin are likely to increase under resource-limiting conditions, but are potentially outweighed by the benefits of associating with non-relatives (avoidance of inbreeding or pathogen transmission) where dens are abundant. We discuss how predictions from social evolutionary theory can contribute to understanding animal responses to landscape change.     

Colonization of host patches following long-distance dispersal by a goldenrod beetle, Trirhabda virgata

1. The arrival of the chrysomelid beetle Trirhabda virgata on isolated patches of its host Solidago altissima was closely monitored to determine how conspecific density and host condition influence colonization.
2. Experimental host patches, which were set on the roof of a four-storey building located 0.7 km from the nearest naturally occurring hosts, were frequently colonized by beetles over a 2-week dispersal period.
3. Females preferred lush host patches that were free from simulated Trirhabda chewing damage. Females colonized lush patches more often than defoliated patches at two spatial scales, when patches were 2 m and 25 m apart. Males did not show a strong preference for lush plants.
4. Males aggregated on plants that already contained adult conspecifics, apparently increasing their reproductive success. Females did not respond to the presence of adults on the patch.
5. Ninety-five per cent of the females arriving on the isolated plants had mated before flying, indicating that lone females are able to colonize empty stands.
6. By avoiding heavily defoliated plants, females should dissipate local outbreaks and spread their offspring away from over-exploited areas.     

Interactions between climate change, competition, dispersal, and disturbances in a tree migration model

Potentially significant shifts in the geographical patterns of vegetation are an expected result of climate change. However, the importance of local processes (e.g., dispersal, competition, or disturbance) has been often ignored in climate change modeling. We develop an individual-based simulation approach to assess how these mechanisms affect migration rate. We simulate the northward progression of a theoretical tree species when climate change makes northern habitat suitable. We test how the rate of progression is affected by (1) competition with a resident species, (2) interactions with disturbance regimes, (3) species dispersal kernel, and (4) the intensity of climate change over time. Results reveal a strong response of species’ expansion rate to the presence of a local competitor, as well as nonlinear effects of disturbance. We discuss these results in light of current knowledge of northern forest dynamics and results found in the climatic research literature.     

Condition-dependent dispersal of a patchily distributed riparian ground beetle in response to disturbance

In common with many habitat elements of riverine landscapes, exposed riverine sediments (ERS) are highly disturbed, naturally patchy and regularly distributed, whose specialists are strongly adapted to flood disturbance and loss of habitat due to succession. Investigations of dispersal in ERS habitats therefore provide an important contrast to the unnaturally fragmented, stable systems usually studied. The present investigation analysed the three interdependent stages of dispersal: (1) emigration, (2) inter-patch movement and (3) immigration of a common ERS specialised beetle, Bembidion atrocaeruleum (Stephens 1828) (Coleoptera, Carabidae), in a relatively unmodified section of river, using mark–resight methods. Dispersal was correlated with estimates of local population size and density, water level and patch quality in order to test for condition-dependent dispersal cues. Flood inundation of habitat was found to increase strongly the overall rate of dispersal, and the rate of emigration was significantly higher from patches that were heavily trampled by cattle. Strongly declining numbers of dispersers with distance suggested low dispersal rates during periods of low water level. Dispersal in response to habitat degradation by cattle trampling would likely lead to a higher overall population fitness than a random dispersal strategy. Dispersal distances were probably adapted to the underlying habitat landscape distribution, high-flow dispersal cues and ready means of long-distance dispersal through hydrochory. Species whose dispersal is adapted to the natural habitat distribution of riverine landscapes are likely to be strongly negatively affected by reduced flood frequency and intensity and habitat fragmentation through flow regulation or channelisation.Electronic Supplementary Material Supplementary material is available to authorised users in the online version of this article at .