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.     

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.

     

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.     

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.     

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.     

Is local distribution of the epiphytic lichen <Emphasis Type="Italic">Lobaria pulmonaria</Emphasis> limited by dispersal capacity or habitat quality?

By surveying and re-surveying 12 forest sites in southern Sweden for the epiphytic lichen Lobaria pulmonaria with a 9-year interval, and measuring tree-related habitat quality variables, we have investigated whether the local distribution of the lichen is limited by poor dispersal capacity or by habitat quality. Dispersal distances were measured indirectly as the distances between colonised trees and the nearest trees occupied by L. pulmonaria in both 1992 and 2001. To compare habitat quality between trees occupied by L. pulmonaria and neighbouring control trees, we recorded tree species and measured age and growth rate of trees, light conditions, bark structure and bryophyte cover. The estimated mean dispersal distance was 35 m, with a recorded maximum of 75 m. Occupied trees were larger and had a larger cover of bryophytes than unoccupied trees of similar size. The results indicate that dispersal capacity probably is the most important factor in limiting the local distribution of L. pulmonaria, but habitat-quality factors may be important on a smaller spatial scale.     

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.     

Vacuum cleaning for conservationists: a new method for inventory of Osmoderma eremita (Scop., 1763) (Coleoptera: Scarabaeidae) and other inhabitants of hollow trees in Natura 2000 areas

Recording the insect species of hollow trees, particularly in larger cavities, presents a major methodological challenge. A whole range of endangered saproxylic beetles and other wood-inhabiting species live together in this habitat. In order to conduct preliminary surveys and monitor populations of the hermit beetle (Osmoderma eremita) in Natura 2000 areas designated for nature conservation according to the EU Habitats Directive, a vacuum cleaner has been used for the first time. Sampling of 127 trees with cavities revealed 39 trees in which O. eremita was present and for which submission of a report is mandatory. A total of 35 species was found, including 17 species in the German red data book as well as six species which are classified as relict species of ancient woodland. The method has therefore proved itself to be very appropriate and is now routinely used in southern Germany.     

Life history of the Glanville fritillary butterfly in fragmented versus continuous landscapes

Habitat loss and fragmentation threaten the long‐term viability of innumerable species of plants and animals. At the same time, habitat fragmentation may impose strong natural selection and lead to evolution of life histories with possible consequences for demographic dynamics. The Baltic populations of the Glanville fritillary butterfly (Melitaea cinxia) inhabit regions with highly fragmented habitat (networks of small dry meadows) as well as regions with extensive continuous habitat (calcareous alvar grasslands). Here, we report the results of common garden studies on butterflies originating from two highly fragmented landscapes (FL) in Finland and Sweden and from two continuous landscapes (CL) in Sweden and Estonia, conducted in a large outdoor cage (32 by 26 m) and in the laboratory. We investigated a comprehensive set of 51 life‐history traits, including measures of larval growth and development, flight performance, and adult reproductive behavior. Seventeen of the 51 traits showed a significant difference between fragmented versus CL. Most notably, the growth rate of postdiapause larvae and several measures of flight capacity, including flight metabolic rate, were higher in butterflies from fragmented than CL. Females from CL had shorter intervals between consecutive egg clutches and somewhat higher life‐time egg production, but shorter longevity, than females from FL. These results are likely to reflect the constant opportunities for oviposition in females living in continuous habitats, while the more dispersive females from FL allocate more resources to dispersal capacity at the cost of egg maturation rate. This study supports theoretical predictions about small population sizes and high rate of population turnover in fragmented habitats selecting for increased rate of dispersal, but the results also indicate that many other life‐history traits apart from dispersal are affected by the degree of habitat fragmentation.     

Do species’ traits predict recent shifts at expanding range edges?

Although some organisms have moved to higher elevations and latitudes in response to recent climate change, there is little consensus regarding the capacity of different species to track rapid climate change via range shifts. Understanding species' abilities to shift ranges has important implications for assessing extinction risk and predicting future community structure. At an expanding front, colonization rates are determined jointly by rates of reproduction and dispersal. In addition, establishment of viable populations requires that individuals find suitable resources in novel habitats. Thus, species with greater dispersal ability, reproductive rate and ecological generalization should be more likely to expand into new regions under climate change. Here, we assess current evidence for the relationship between leading-edge range shifts and species' traits. We found expected relationships for several datasets, including diet breadth in North American Passeriformes and egg-laying habitat in British Odonata. However, models generally had low explanatory power. Thus, even statistically and biologically meaningful relationships are unlikely to be of predictive utility for conservation and management. Trait-based range shift forecasts face several challenges, including quantifying relevant natural history variation across large numbers of species and coupling these data with extrinsic factors such as habitat fragmentation and availability.     

Conflict between insect conservation and public safety: the case study of a saproxylic beetle (<Emphasis Type="Italic">Osmoderma eremita</Emphasis>) in urban parks

Urban parks can harbour small populations of saproxylic insects of high conservation concern, such as Osmoderma eremita and other rare beetles. These areas often host old trees which have become very uncommon in rural areas where they are threatened by commercial forestry management procedures based on frequent tree cutting. Nevertheless, old trees of urban parks may represent a hazard for public safety and are sometimes cut by management authorities. The aim of this work was to assess the loss of reproductive sites for saproxylic beetles of the Scarabaeidae, Lucanidae and Cerambycidae, when felling plans are adopted according to a Visual Tree Assessment Procedure (VTA), in a Mediterranean urban park. On July–August 2004, 1,247 holm oaks were surveyed within the border of an urban park of Rome (Villa Borghese). The occurrence of saproxylic beetles (i.e. the presence of frass, living insects or their remains) was verified in 66 old holm oaks, 41% of which were doomed to cutting by VTA. Eleven of these trees (41% of the trees doomed to be cut) held fragments of adults and sometimes living larvae of Osmoderma eremita, and four of them (36%) were included in the felling plan. The presence of Osmoderma eremita in tree holes was more frequent in deep cavities. The presence of frass in the cavities was positively associated with tree height and a high degree of damage at the root collar and negatively with the presence of hole-nesting birds.     

Winter Feeding Tree Choice in Sichuan Snub-Nosed Monkeys (Rhinopithecus roxellanae) in Shennongjia Nature Reserve,China

We investigated patterns of winter feeding tree choice in 4 groups of Sichuan snub-nosed monkeys (Rhinopithecus roxellanae) in Shennongjia Nature Reserve, China. We collected data during 2 winters from 1998 to 2000. The monkeys used mature forest, young forest and shrub forest, but not grassland. Groups used tree species in a significantly nonrandom pattern. There was a similar composition of preferred tree species between different habitats for each group and among the same habitat types for different groups. They preferred Abies fargesii, Pinus armandii and Salix walliciana for foraging. The 3 species occur in varying degrees of abundance in different habitats and were used differently by the 4 groups. The difference is probably due to interhabitat differences in availability of tree species, in addition to microclimate. The mean circumference of a tree had little effect on its preference score, but preferred species tend to be larger. A Wilcoxon signed-rank test indicated that the percentage of trees used and average number of feeding bites per tree is significantly greater for larger trees. For all trees in a given habitat, the percentage of trees used and average number of bites per tree have a significant positive correlation with average tree circumference. Our results indicate that Rhinopithecus roxellanae prefer to feed in large trees more than small trees in a given habitat, thereby preferring mature forest habitat. There is also a group-size effect; larger groups used higher-quality habitats than those of smaller groups. Both tree species and size are the major determinants of feeding choice, but tree species is more important than tree size. Our results have at least three implications for winter habitat conservation of Sichuan snub-nosed monkeys. Conservation efforts should be focused on mature forest because it is better habitat at Rhinopithecus than young forest, as long as the same tree species are present. Secondly, Pinus armandii, Abies fargesii and Salix walliciana should be conserved as top priority in forest communities. Third, the largest trees in each habitat should be given greatest possible protection.     

The impact of forest spread on a marginal population of a protected peony (<Emphasis Type="Italic">Paeonia officinalis</Emphasis> L.): the importance of conserving the habitat mosaic

In the Mediterranean region of Europe, land-use changes have allowed for rapid colonisation of open habitats by woody species. As a result, it is critical to gather information on how protected species in open habitats respond to forest spread in such areas. Our objective is to quantify whether spatial heterogeneity of the vegetation associated with recent forest closure influences demographic structure and maternal fertility in a population of the protected Paeonia officinalis L. In closed woodland, adult plants of P. officinalis are almost exclusively vegetative, in open habitats seedlings are rare and on the woodland edge there is a relative over-representation of flowering plants and seedlings. Forest closure dramatically reduces flowering frequency, but has no significant effect on maternal fertility of flowering plants. The spatial aggregation of seedlings close to the maternal plants suggests that dispersal is spatially restricted. Together, these results suggest that the viability of the population requires a transitional habitat between open garrigues or grassland with spaced trees and woodland. A management programme incorporating tree and shrub thinning and cutting of parcels in rotation to maximise the length of the forest edge could maintain a habitat mosaic that favours the persistence of this species in the study site.     

Frugivory on Persea lingue in temperate Chilean forests: interactions between fruit availability and habitat fragmentation across multiple spatial scales

Habitat degradation and fragmentation are expected to reduce seed dispersal rates by reducing fruit availability as well as the movement and abundance of frugivores. These deleterious impacts may also interact with each other at different spatial scales, leading to nonlinear effects of fruit abundance on seed dispersal. In this study we assessed whether the degradation and fragmentation of southern Chilean forests had the potential to restrict seed dispersal the lingue (Persea lingue) tree, a fleshy-fruited tree species. Of five frugivore bird species, the austral thrush (Turdus falcklandii) and the fire-eyed diucon (Xolmis pyrope) were the only legitimate seed dispersers as well as being the most abundant species visiting lingue trees. The results showed little or no direct effect of habitat fragmentation on seed dispersal estimates, possibly because the assemblage of frugivore birds was comprised habitat-generalist species. Instead, the number of fruits removed per focal tree exhibited an enhanced response to crop size, but only in the more connected fragments. In the fruit-richer fragment networks, there was an increased fragment-size effect on the proportion of fruits removed in comparison to fruit-poor networks in which the fragment size effect was spurious. We suggest that such nonlinear effects are widespread in fragmented forest regions, resulting from the link between the spatial scales over which frugivores sample resources and the spatial heterogeneity in fruiting resources caused by habitat fragmentation and degradation.     

Dispersal and habitat cuing of Eurasian red squirrels in fragmented habitats

Animal dispersal and subsequent settlement is a key process in the life history of many organisms, when individuals use demographic and environmental cues to target post-dispersal habitats where fitness will be highest. To investigate the hypothesis that environmental disturbance (habitat fragmentation) may alter these cues, we compared dispersal patterns of 60 red squirrels (Sciurus vulgaris) in three study sites that differ in habitat composition and fragmentation. We determined dispersal distances, pre- and post-dispersal habitat types and survival using a combination of capture–mark–recapture, radio-tracking and genetic parentage assignment. Most (75%) squirrels emigrated from the natal home range with mean dispersal distance of 1,014 ± 925 m (range 51–4,118 m). There were no sex-related differences in dispersal patterns and no differences in average dispersal distance, and the proportion of dispersers did not differ between sites. In one of the sites, dispersers settled in patches where density was lower than in the natal patch. In the least fragmented site, 90% of animals settled in the natal habitat type (habitat cuing) against 44–54% in the more strongly fragmented sites. Overall, more squirrels settled in the natal habitat type than expected based on habitat availability, but this was mainly due to individuals remaining within the natal wood. In the highly fragmented landscape, habitat cuing among emigrants did not occur more frequently than expected. We concluded that increased habitat fragmentation seemed to reduce reliable cues for habitat choice, but that dispersing squirrels settled in patches with lower densities of same-sex animals than at the natal home range or patch, independent of degree of fragmentation.     

Effects of forest canopy on habitat selection in treefrogs and aquatic insects: implications for communities and metacommunities

The specific dispersal/colonization strategies used by species to locate and colonize habitat patches can strongly influence both community and metacommunity structure. Habitat selection theory predicts nonrandom dispersal to and colonization of habitat patches based on their quality. We tested whether habitat selection was capable of generating patterns of diversity and abundance across a transition of canopy coverage (open and closed canopy) and nutrient addition by investigating oviposition site choice in two treefrog species (Hyla) and an aquatic beetle (Tropisternus lateralis), and the colonization dynamics of a diverse assemblage of aquatic insects (primarily beetles). Canopy cover produced dramatic patterns of presence/absence, abundance, and species richness, as open canopy ponds received 99.5% of propagules and 94.6% of adult insect colonists. Nutrient addition affected only Tropisternus oviposition, as females oviposited more egg cases at higher nutrient levels, but only in open canopy ponds. The behavioral partitioning of aquatic landscapes into suitable and unsuitable habitats via habitat selection behavior fundamentally alters how communities within larger ecological landscapes (metacommunities) are linked by dispersal and colonization.     

Should I stay or should I go? Modelling dispersal strategies in saproxylic insects based on pheromone capture and radio telemetry: a case study on the threatened hermit beetle <Emphasis Type="Italic">Osmoderma eremita</Emphasis>

To predict how organisms cope with habitat fragmentation we must understand their dispersal biology, which can be notoriously difficult. We used a novel, multi-pronged approach to study dispersal strategies in the endangered saproxylic hermit beetle Osmoderma eremita, exploiting its pheromone system to intercept high numbers of dispersing individuals, which is not possible with other methods. Mark-release-recapture, using unbaited pitfall traps inside oak hollows and pheromone-baited funnel traps suspended from tree branches, was combined with radio telemetry (in females only) to record displacements. Dispersal, modelled as a probability distribution of net displacement, did not differ significantly between sexes (males versus females recaptured), observation methods (females recaptured versus radio-tracked), or sites of first capture (pitfall trap in tree versus pheromone trap – distance from original dispersal point unknown). A model including all observed individuals yielded a mean displacement of 82 m with 1% dispersing > 1 km. Differences in body length were small between individuals captured in pitfall versus pheromone traps, indicating that dispersal is rarely a condition-dependent response in O. eremita. Individuals captured in pheromone traps were consistently lighter, indicating that most dispersal events occur relatively late in life, which agrees with trap catch data. In addition, most (79%) females captured in pheromone traps were mated, showing that females typically mate before leaving their natal tree. Our data show that integrating odour attractants into insect conservation biology provides a means to target dispersing individuals and could greatly improve our knowledge of dispersal biology in threatened species.     

Fragmentation in eucalypt woodlands promotes nest‐tree occupancy by a despotic species,the noisy miner (Manorina melanocephala)

The effects of habitat fragmentation as a threat to biodiversity are well known; decreased connectivity can potentially influence population processes and dynamics, resulting in smaller, isolated populations that may not function optimally. However, fragmentation may also increase the amount of edge or ecotone habitat available to open country species, benefiting their populations and enabling them to dominate remnant habitats. Noisy miners (Manorina melanocephala) are one such species, occupying eastern‐Australian eucalypt woodlands. They are considered a ‘despotic’ species, in that their presence negatively impacts woodland avifauna biodiversity due to their aggressive exclusion of other taxa from occupied areas. Despite this well‐known impact, little information exists on the patterns of nest‐tree occupancy by noisy miners within eucalypt woodlands. In the current study, we explored the patterns of nest‐tree occupancy by noisy miners across two successive years, aiming to identify preferences for breeding areas relative to vegetation structure. Our results show that both habitat fragmentation and the characteristics of individual eucalypt trees in an area influenced nest‐tree occupancy. Noisy miners constructed nests in trees near the edge of woodland patches more often than expected. Moreover, the nest tree chosen was a eucalypt that was significantly smaller than randomly selected trees from the surrounding area. The results highlight the importance of habitat management measures that may reduce the suitability of woodland patches as nesting sites for this species, in order to mitigate the severe effects of this despotic edge specialist.