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.     

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

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

Is Active Management the Key to the Conservation of Saproxylic Biodiversity? Pollarding Promotes the Formation of Tree Hollows

Trees with hollows are key features sustaining biodiversity in wooded landscapes. They host rich assemblages of often highly specialised organisms. Hollow trees, however, have become rare and localised in Europe. Many of the associated biota is thus declining or endangered. The challenge of its conservation, therefore, is to safeguard the presence of hollow trees in sufficient numbers. Populations of numerous species associated with tree hollows and dead wood are often found in habitats that were formed by formerly common traditional silvicultural practices such as coppicing, pollarding or pasture. Although it has been occasionally mentioned that such practices increase the formation of hollows and the availability of often sun-exposed dead wood, their effect has never been quantified. Our study examined the hollow incidence in pollard and non-pollard (unmanaged) willows and the effect of pollarding on incremental growth rate by tree ring analysis. The probability of hollow occurrence was substantially higher in pollard than in non-pollard trees. Young pollards, especially, form hollows much more often than non-pollards; for instance, in trees of 50 cm DBH, the probability of hollow ocurrence was ∼0.75 in pollards, but only ∼0.3 in non-pollards. No difference in growth rate was found. Pollarding thus leads to the rapid formation of tree hollows, a habitat usually associated with old trees. It is therefore potentially a very important tool in the restoration of saproxylic habitats and conservation of hollow-dependent fauna. If applied along e.g. roads and watercourses, pollarding could also be used to increase landscape connectivity for saproxylic organisms. In reserves where pollarding was formerly practiced, its restoration would be necessary to prevent loss of saproxylic biodiversity. Our results point to the importance of active management measures for maintaining availability, and spatial and temporal continuity of deadwood microhabitats.     

Occurrence of Larca lata H.J. Hansen (Pseudoscorpionida: Garypidae) and Allochernes wideri C.L. Koch (Pseudoscorpionida: Chernetidae) in tree hollows in relation to habitat quality and density

Many beetles associated with old trees are on national red lists, but pseudoscorpions living in similar habitats have received little attention. This study reports the habitat and occurrence patterns of two species of pseudoscorpions living in hollow trees. Their occurrence has been assessed by sieving wood mould from 274 oaks in southeastern Sweden and from museum specimens collected in Sweden. Larca lata is confined to hollow oaks with a large girth and a plentiful supply of wood mould. Allochernes wideri is much less particular about wood mould volume, trunk diameter and tree species. Larca lata inhabits hollow trunks with characteristics that are typical of very old trees, whereas A. wideri predominantly occurs in trunks in an earlier stage of hollow formation. Larca lata was almost exclusively found in larger assemblages of hollow oaks, which suggests long-term survival may be difficult when the network of suitable hollow trees is too sparse. Larca lata is a rare species in Europe and probably vulnerable to extinction, since it is dependent on a habitat which has declined severely in the last few centuries.     

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.     

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.     

The dispersal rate of a beetle, Osmoderma eremita, living in tree hollows

The dispersal of an endangered beetle, Osmoderma eremita, that lives in tree hollows, was studied by mark-release-recapture with pitfall traps. As only a small proportion of all dispersals is observed by this method, a simulation model was constructed to estimate the dispersal rate per individual. The model results suggest that 15% of the adults leave the original tree for another hollow tree, and consequently most individuals remain in the same tree throughout their entire life. This suggests that each hollow tree sustains a local population with limited connection with the populations in surrounding trees. It supports the view that O. eremita has a metapopulation structure, with each tree possibly sustaining a local population, and with the population in an assemblage of trees forming a metapopulation. Low dispersal rate and range make the species vulnerable to habitat fragmentation, probably at a scale of only a few hundred meters.     

Boxing for biodiversity: evaluation of an artificially created decaying wood habitat

Many saproxylic species are threatened in Europe because of habitat decline. Hollow trees represent an important habitat for saproxylic species. Artificial habitats may need to be created to maintain or increase the amount of habitat due to natural habitat decline. This study investigated the extent to which saproxylic beetles use artificial habitats in wooden boxes. The boxes were placed at various distances (0–1800 m) from known biodiversity hotspots with hollow oaks and studied over 10 years. Boxes were mainly filled with oak saw dust, oak leaves, hay and lucerne flour. In total, 2170 specimens of 91 saproxylic beetle species were sampled in 43 boxes. The abundance of species associated with tree hollows, wood rot and animal nests increased from the fourth to the final year, but species richness declined for all groups. This study shows that wooden boxes can function as saproxylic species habitats. The artificial habitats developed into a more hollow-like environment during the decade long experiment with fewer but more abundant tree hollow specialists.     

Importance of high quality early-successional habitats in managed forest landscapes to rare beetle species

Species adapted to early-successional forest habitats are in managed landscapes largely confined to clearcuts. To improve habitat quality on clearcuts, green tree and dead wood retention is widely applied in forestry; however, its effects on rare early-successional species have rarely been shown. We repeatedly surveyed two red-listed beetle species (Upis ceramboides and Platysoma minus) on clearcuts in a managed boreal forest landscape. We found that U. ceramboides decreased its occupancy over time while P. minus increased, indicating that red-listed species vary in their ability to successfully utilise managed habitats. We found no effect of connectivity on probability of occurrence, colonisation or extinction per clearcut. Trees retained alive improved habitat quality of clearcuts, since both species were more frequent in dead wood of such trees, in comparison to logging residues. We suggest that retention can be improved by protecting and creating dead wood as intact trees during harvesting. Rare specialist species require habitat of high quality, and consequently it is impossible to meet the requirements of these species on every clearcut. To preserve all early-successional species at a regional scale, we recommend focusing retention of green trees and dead wood to one or a few trees species on each clearcut and in each landscape.     

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.

     

Metapopulation Dynamics of the Mistletoe and Its Host in Savanna Areas with Different Fire Occurrence

Mistletoes are aerial hemiparasitic plants which occupy patches of favorable habitat (host trees) surrounded by unfavorable habitat and may be possibly modeled as a metapopulation. A metapopulation is defined as a subdivided population that persists due to the balance between colonization and extinction in discrete habitat patches. Our aim was to evaluate the dynamics of the mistletoe Psittacanthus robustus and its host Vochysia thyrsoidea in three Brazilian savanna areas using a metapopulation approach. We also evaluated how the differences in terms of fire occurrence affected the dynamic of those populations (two areas burned during the study and one was fire protected). We monitored the populations at six-month intervals. P. robustus population structure and dynamics met the expected criteria for a metapopulation: i) the suitable habitats for the mistletoe occur in discrete patches; (ii) local populations went extinct during the study and (iii) colonization of previously non-occupied patches occurred. The ratio of occupied patches decreased in all areas with time. Local mistletoe populations went extinct due to two different causes: patch extinction in area with no fire and fire killing in the burned areas. In a burned area, the largest decrease of occupied patch ratios occurred due to a fire event that killed the parasites without, however, killing the host trees. The greatest mortality of V. thyrsoidea occurred in the area without fire. In this area, all the dead trees supported mistletoe individuals and no mortality was observed for parasite-free trees. Because P. robustus is a fire sensitive species and V. thyrsoidea is fire tolerant, P. robustus seems to increase host mortality, but its effect is lessened by periodic burning that reduces the parasite loads.     

Metapopulation models for seasonally migratory animals

Metapopulation models are widely used to study species that occupy patchily distributed habitat, but are rarely applied to migratory species, because of the difficulty of identifying demographically independent subpopulations. Here, we extend metapopulation theory to describe the directed seasonal movement of migratory populations between two sets of habitat patches, breeding and non-breeding, with potentially different colonization and extinction rates between patch types. By extending the classic metapopulation model, we show that migratory metapopulations will persist if the product of the two colonization rates exceeds the product of extinction rates. Further, we develop a spatially realistic migratory metapopulation model and derive a landscape metric-the migratory metapopulation capacity-that determines persistence. This new extension to metapopulation theory introduces an important tool for the management and conservation of migratory species and may also be applicable to model the dynamics of two host-parasite systems.     

Metapopulation dynamics: Does it help to have more of the same?

With the interest in conservation biology shifting towards processes from patterns, and to populations from communities, the theory of metapopulation dynamics is replacing the equilibrium theory of island biogeography as the population ecology paradigm in conservation biology. The simplest models of metapopulation dynamics make predictions about the effects of habitat fragmentation - size and isolation of habitat patches - on metapopulation persistence. The simple models may be enriched by considerations of the effects of demographic and environmental stochasticity on the size and extinction probability of local populations. Environmental stochasticity affects populations at two levels: it makes local extinctions more probable, and it also decreases metapopulation persistence time by increasing the correlation of extinction events across populations. Some controversy has arisen over the significance of correlated extinctions, and how they may affect the optimal subdivision of metapopulations to maximize their persistence time.     

Colonization and extinction dynamics of an annual plant metapopulation in an urban environment

The metapopulation framework considers that the spatiotemporal distribution of organisms results from a balance between the colonization and extinction of populations in a suitable and discrete habitat network. Recent spatially realistic metapopulation models have allowed patch dynamics to be investigated in natural populations but such models have rarely been applied to plants. Using a simple urban fragmented population system in which favourable habitat can be easily mapped, we studied patch dynamics in the annual plant Crepis sancta (Asteraceae). Using stochastic patch occupancy models (SPOMs) and multi‐year occupancy data we dissected extinction and colonization patterns in our system. Overall, our data were consistent with two distinct metapopulation scenarios. A metapopulation (sensu stricto) dynamic in which colonization occurs over a short distance and extinction is lowered by nearby occupied patches (rescue effect) was found in a set of patches close to the city centre, while a propagule rain model in which colonization occurs from a large external population was most consistent with data from other networks. Overall, the study highlights the importance of external seed sources in urban patch dynamics. Our analysis emphasizes the fact that plant distributions are governed not only by habitat properties but also by the intrinsic properties of colonization and dispersal of species. The metapopulation approach provides a valuable tool for understanding how colonization and extinction shape occupancy patterns in highly fragmented plant populations. Finally, this study points to the potential utility of more complex plant metapopulation models than traditionally used for analysing ecological and evolutionary processes in natural metapopulations.     

Availability of hollow‐bearing trees and their utilization by small animals in Cawston ranch,Zimbabwe

The objective of this study was to characterize hollows found in selected tree species (in Cawston Ranch in Zimbabwe) as a potential habitat for small mammals because cavities are important as resources for wildlife: for foraging, shelter, roosting and nesting purposes. Four vegetation types were chosen for analysis: Baikiaea plurijuga, Colophospermum mopane, Combretum molle and Commiphora mollis according to abundance and dominance in the plots they occurred. Transect sampling was done to collect vegetation data. The total number of living trees and stags were recorded in each plot. Trees in a plot were thoroughly inspected for hollows. The following dimensions were measured on cavity trees: hollow size (diameter), hollow location and diameter at breast height. The results indicate that the rate of hollow formation and utilization is significantly dependent on tree species. There was a significant difference in the provision of hollows in stem and crown of a tree in each of the vegetation classes analysed. B. plurijuga hollows had the highest utilization whilst C. molle had the least utilization. Hollows are utilized irrespective of their location as long as the animal finds a niche, and stags at advanced stages of decay are not important in providing habitats for animals.     

Self‐organization of background habitat determines the nature of population spatial structure

A substantial literature treats the dynamics of populations in response to spatial patterning of underlying habitats, the most common formulations being source/sink populations and metapopulations living in a patchwork of habitats. A separate and growing literature on the self‐organization of spatial pattern focuses on how local interactions give rise to regional patterns that can be described with scale‐free distributions. Motivated by the potential ubiquity of the coupling of these processes (spatial self organization and the dynamics of organisms in spatially structured habitats), we link these two lines of thought into a theory of populations in fragmented habitats. Using a combined analytical and computational approach, we show that self organization can generate a background into which an independent population fits, and that the scale‐free nature of such habitat creates conditions that influence both the persistence versus extinction properties of the embedded population and where it lies on the continuum between source/sink populations and metapopulations. Both the analytical framework and the computer simulations demonstrate the potential for populations to persist on either end of the metapopulation–source/sink continuum while failing to persist under intermediate conditions. These results provide a new perspective on ecological theory related to habitat fragmentation and population persistence, suggesting that under some conditions intermediate states between these two extremes may maximize risk of extinction of the population. These implications could be of particular importance for managing self‐organized systems of conservation concern.     

Isolation by distance in saproxylic beetles may increase with niche specialization

Species confined to temporally stable habitats are usually susceptible to habitat fragmentation, as living in long-lasting habitats is predicted to constrain evolution of dispersal ability. In Europe, saproxylic invertebrates associated with tree hollows are currently threatened due to the severe fragmentation of their habitat, but data on the population genetic consequences of such habitat decline are still scarce. By employing AFLP markers, we compared the spatial genetic structure of two ecologically and taxonomically related beetle species, Osmoderma barnabita and Protaetia marmorata (Cetoniidae). Both species are exclusively associated with tree hollows, but O. barnabita has a more restricted host preferences compared to P. marmorata. Analyses of spatial autocorrelation showed, in line with the predicted low dispersal potential of these saproxylic beetles, that both species are characterized by a strong kinship structure, which was more pronounced in the specialist O. barnabita than in the generalist P. marmorata. Individuals of both species sampled within single trees showed high relatedness (≈0.50 in O. barnabita and ≈0.15 in P. marmorata). Interestingly, groups of pheromone-emitting O. barnabita males sampled on the same tree trunk were found to be full brothers. Whether this result can be explained by kin selection to increase attraction of conspecific females for mating or by severe inbreeding of beetles within individual tree hollows needs further study. Although our studied populations were significantly inbred, our results suggest that the dispersal ability of Osmoderma beetles may be one order of magnitude greater than suggested by previous dispersal studies and acceptable levels of habitat fragmentation for metapopulation survival may be bigger than previously thought.     

Colonization–extinction dynamics of epixylic lichens along a decay gradient in a dynamic landscape

Metapopulation models are often used for understanding and predicting species dynamics in fragmented landscapes. Several models have been proposed depending on e.g. the relative importance of patch dynamics on the metapopulation dynamics. Dead wood is a dynamic substrate patch, and species that are confined to such patches have experienced a high degree of habitat loss in managed forests. Little is, however, known about how the population dynamics of epixylic species are affected by the fast dynamics of their substrate patches. We quantified the effect of local patch conditions and metapopulation processes on colonizations and extinctions of epixylic lichen species in a managed boreal forest landscape. This was done by twice surveying seven lichen metapopulations on 293 stumps in 30 stands of ages covering the duration of the dynamic patches (stumps). We also investigated the relative importance of local stochastic extinctions from stumps that remained available, and deterministic extinctions due to stump surface disappearance. We found importance of a decay gradient, surrounding metapopulation size, and local population sizes, in driving the colonization–extinction dynamics of epixylic lichens. The species were sorted along the stump decay gradient. Increasing surrounding metapopulation size was associated with increased colonization rates, and increasing local population size decreased lichen extinction rates. Finally, both local stochastic extinctions and deterministic extinctions due to patch disappearance occur, confirming that the long‐term persistence of epixylic lichens depends on colonization rates that compensate for stochastic population extinctions as well as deterministic extinctions.     

The influence of patch demographics on metapopulations, with particular reference to successional landscapes

The classical view of metapopulations relates the regional abundance of a species to the balance between the extinction and colonization dynamics of identical local populations. Species in successional landscapes may represent the most appropriate examples of classical metapopulations. However, Levins‐type metapopulation models do not explicitly separate population loss due to successional habitat change from other causes of extinction. A further complication is that the chance of population loss due to successional habitat change may be related to the age of a patch. I developed simple patch occupancy models to include succession and included consideration of patch age structure to address two related questions: what are the implications of changes in patch demographic rates and when is a move to a structured patch occupancy model justified? Age‐related variation in patch demography could increase or decrease the equilibrium fraction of the available habitat occupied by a species when compared to the predictions of an unstructured model. Metapopulation persistence was enhanced when the age class of patches with the highest species occupancy suffered relatively low losses to habitat succession. Conversely, when the age class of patches with the highest species occupancy also had relatively high successional loss rates, extinction thresholds were higher that would be predicted by a simple unstructured model. Hence age‐related variation in patch successional rate introduces biases into the predictions of simple unstructured models. Such biases can be detected from field surveys of the fraction of occupied and unoccupied patches in each age class. Where a bias is demonstrated, unstructured models will not be adequate for making predictions about the effects of changing parameters on metapopulation size. Thinking in successional terms emphasizes how landscapes might be managed to enhance or reduce the patch occupancy by any particular metapopulation