A review of Turkish saproxylic beetles from the European Red List

The conservation of beetles that depend on dying or dead wood (i.e. saproxylic), has received a great deal of attention in many parts of the world in recent years. Human activities such as urbanization and logging, and their results, e.g. global warming, destroy natural ecosystems and threaten unusual species such as Lucanus cervus (L. 1758), Rosalia alpina (L. 1758), Limoniscus violaceus (Müller 1821) and Osmoderma eremita (Scopoli 1763). Attempts have been made to conserve the habitats upon which these species depend. Although represented by a large number of species, saproxylic beetles in Turkey have not received much attention. Although some Turkish studies have acknowledged their value, comprehensive investigations of these species are very limited. The aim of this study is to make a list, from the literature, of the species that are endangered in Europe and found in Turkey, to highlight the importance of these species. The research includes 151 saproxylic beetle species belonging to the families Bostrichidae, Cerambycidae, Elateridae, Eucnemidae, Scarabaeidae, Euchiridae, Lucanidae and Mycetophagidae. The list of Turkish saproxylic beetles was prepared using the European Red List of Saproxylic Beetles. Information is provided about the IUCN Red List category in Europe, the locations of the species in Turkey, and their global geographic range. We gathered all data about these species from the international literature. Furthermore, we also include in the list certain species whose locations are not specified, but which inhabit Turkey. We obtained their records from some catalogs in Turkey. In addition to the list, we prepared three distribution maps for Turkey. The maps show distribution in Turkey of some species in the Endangered, Vulnerable and Near Threatened categories in the European Red List. It is expected that this study will lead to comprehensive studies on saproxylic beetles from Turkey.     

Sampling the rarest: threatened beetles in boreal forest biodiversity inventories

Presence or absence of threatened species in samples is information that is widely used in designing and implementing conservation actions. We explored the effectiveness of beetle (Coleoptera) inventories and contribution of different sampling methods in revealing occurrences of threatened and near threatened species in boreal forests. The number of species caught using traps in a particular area proved to be a useful indicator of the representativeness of data, the relationship between total number of species and the number of threatened and near threatened species being almost exponential. Samples containing less than 200 trapped species (or 2000 individuals) are almost useless in surveying threatened and near threatened species. The probability of finding such species increases considerably when the number of trapped species exceeds 400. Window traps attached directly on the trunks of dead trees proved to be the most efficient sampling method in trapping threatened beetles, whereas many other standard methods gave relatively poor results. We suggest that the best alternative in surveying threatened species in boreal forests is a combination of intensive direct searching and trunk window traps. Finding threatened beetles with rigorous probability requires very large sample sizes, even if the most effective sampling methods are used. For example, ranking 10 boreal forest areas to be protected according to the occurrence of threatened species with some reliability may require trapping of over 100000 beetle individuals. Collecting and identifying these large samples routinely in conservation actions is not feasible, which means that shortcuts (indicators etc.) are necessary. However, a lot of good-quality inventories with appropriate sampling efforts are needed before these shortcuts can be identified and elaborated. Such inventories are also crucial for the improvement of the classification of threatened species and full assessment on how past forest management has eventually affected the biota.     

Assemblages of saproxylic beetles on large downed trunks of oak

Old living oaks (Quercus robur) are known as a very species‐rich habitat for saproxylic beetles, but it is less clear to what extent such veteran trees differ from an even rarer feature: downed trunks of large oaks. In this study, we set out to sample this habitat, using window traps, with two aims: (1) to describe the variation of assemblages among downed trunks of different type and (2) to compare beetles on downed oaks with data from veteran standing trees. The results showed that trunk volume and sun exposure better explained assemblages as well as species numbers on downed trunks than did decay stage. Furthermore, species classified as facultative saproxylic species showed weak or no differentiation among downed trunks. Species with different feeding habits showed no apparent differentiation among downed trunks. Furthermore, species composition on dead, downed oak trunks differed sharply from that of living, veteran oaks. Wood or bark feeders were more common on veterans than downed trunks, but there was no difference for those species feeding on fungi or those feeding on insects and their remains. In conclusion, for a successful conservation of the saproxylic beetle fauna it is important to keep downed oak trunks, and particularly large ones, in forest and pastures as they constitute a saproxylic habitat that differs from that of living trees.     

Density-dependent population regulation detected in short time series of saproxylic beetles

Understanding the regulation of natural populations has been a long-standing problem in ecology. Here we analyze the population dynamics of 17 species of saproxylic beetles in Shizuoka Prefecture, Japan collected over 11–12 years using autoregressive integrated moving average (ARIMA) models. We first examined the dynamics for indications of the order of the ARIMA models and evaluated the time series to determine that it was not simply a random, white noise sequence. All species dynamics were not mere random noise, and ARIMA models up to lag 3 were considered. The best model was selected from the possible ones using several criteria: model convergence, weak residual autocorrelation, the small sample AIC must be among the smallest that were not significantly different, and the lag indicated by the cutoff values in the detrended partial autocorrelation function. We found significant and nearly significant direct density-dependence for 14 of the 17 species, varying from −0.709 and stronger. The characteristic return rates were strong and only one species had a weak return rate (>0.9), implying that these species were strongly regulated by density-dependent factors. We found that populations with higher order ARIMA models (lag 2 and 3) had weaker return rates than populations with ARIMA models with only one lag, suggesting that species with more complex dynamics were more weakly regulated. These results contrast with previous suggestions that 20+ years are needed to detect density dependence from population time series and that most populations are weakly regulated.     

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.     

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.     

A comparison of three methods to survey saproxylic beetles in hollow oaks

One of the most endangered assemblages of species in Europe is insectsassociated with old trees. For that reason there is a need of developing methodsto survey this fauna. This study aims at comparing three methods – windowtrapping, pitfall trapping and wood mould sampling – to assess speciesrichness and composition of the saproxylic beetle fauna in living, hollow oaks.We have used these methods at the same site, and to a large extent in the sametrees. Useful information was obtained from all methods, but they partiallytarget different assemblages of species. Window trapping collected the highestnumber of species. Pitfall trapping collected beetles associated with treehollows which rarely are collected by window traps and therefore it isprofitable to combine these two methods. As wood mould sampling is the cheapestmethod to use, indicator species should preferably be chosen among specieswhich are efficiently collected with this method.     

Estimating population characteristics of two saproxylic beetles: a mark-recapture approach

Using a mark-release-recapture technique we describe adult sex ratios, recapture rates and other sample characteristics of two saproxylic species: the nationally threatened longhorn beetle Leptura (Rutpela) maculata (Coleoptera: Cerambycidae) and the common L. quadrifasciata in southeastern Finland over two summers. Over 350 individuals of L. maculata and 150 individuals of L. quadrifasciata were captured on floral resource or in flight, and marked each summer. For L. maculata, the sex ratio was male-biased (2:1), whereas for L. quadrifasciata the bias was less clear. For both species, the male-bias may reflect behavioral differences between sexes, rather than true population differences. The proportion of recaptured individuals was low and varied between 7 and 33% depending on the species and year, which allowed us to estimate population parameters only for L. maculata in 2006. A model which assumed constant survival, but time-dependent catchability and entrance probability from a larger superpopulation, fit the data best. The precision of the total population size estimates were reasonable for all the models tested (coefficient of variation = 7–14%). Based on the estimated local adult population size (mean ± 95% confidence interval = 865 ± 131), and the current distribution area of L. maculata, we infer that the species is not in immediate risk of extinction in Finland. Our analysis shows that mark-recapture technique can provide precise estimates of adult population size of saproxylic beetles which have different adult and larval habitats, and thus be useful in assessing extinction risk and monitoring population trends.     

Vertical distribution of saproxylic beetles within snag trunks retained in plantation forests

Retention of snags (standing dead trees) is considered to have important effects on saproxylic species conservation in plantation forests because snags would provide vertically stratified deadwood habitats. However, the vertical distribution of saproxylic insects within snag trunks is still unclear. We felled 33 naturally occurring snags of Todo fir Abies sachalinensis in plantation forests and extracted insects from 99 logs sampled from three vertical positions of the snag trunks (basal stem: <2.5 m, lower trunk: 2.5–5 m and upper trunk: >5 m). The mean number of species that emerged from a single log was only 2.69, but we identified 51 morphospecies of saproxylic beetles in total. The total number of species that emerged from the basal stem (34 spp.) was greater than those that emerged from the lower trunk (25 spp.) or the upper trunk (30 spp.). However, rarefaction-extrapolation analysis did not demonstrate a significant difference in species richness among the log positions. Beetle assemblages were separated into two groups by constrained correspondence analysis; one group emerged only from lower and upper trunk logs, while another emerged mainly from basal stem logs. Additionally, vertical position had a significant effect on the distribution of the five main species. Our results show that beetle assemblages within snags in the plantation forests were highly variable, and retaining a sufficient number of high stumps may be important for saproxylic beetle conservation in plantation forests. We propose ‘retention thinning’ as an appropriate method to combine efficient timber production with biodiversity conservation in plantation forests.     

Habitat availability in forests and clearcuts for saproxylic beetles associated with aspen

To maintain biodiversity in managed, boreal forest in Scandinavia, aspen trees (Populus tremula) are often retained at clearcutting. In this study, the habitat availability for beetles associated with aspen coarse woody debris (CWD) was predicted for forests and clearcuts with a model of CWD dynamics. Habitat requirements of eight beetle species (Agathidium bicolor, Cerylon ferrugineum, Cyphaea curtula, Endomychus coccineus, Homalota plana, Mycetophagus fulvicollis, Ptilinus fuscus and Xylotrechus rusticus) were obtained from their occurrence patterns in relation to characteristics of CWD objects in forest and on clearcuts in a study landscape in central Sweden. Three species were more frequent in forest and three at clearcuts. Five species increased with increasing girth of the CWD. Three were more frequent on standing CWD, and two on lying CWD. From the same study area, we also obtained field data on the recruitment of CWD (i.e., tree mortality) and amounts of different types of CWD. Annual tree mortality of aspen was higher for recent clearcuts (6.3%) compared with older clearcuts (1.1%). For all species, the habitat availability was higher on clearcuts, because enhanced tree mortality increased the amount of recently dead CWD. As a conclusion, green-tree retention of aspen is a conservation effort that is beneficial for species associated with aspen CWD.     

Increased openness around retained oaks increases species richness of saproxylic beetles

The decrease of old deciduous trees in northern Europe is a threat to the saproxylic fauna. In northern Europe, oak sustains the highest richness of saproxylic invertebrates, among which beetles is a large group. In order to preserve species associated with old trees, it has become common practice in commercial forestry to retain such trees at final felling. However, to create beneficial conditions for species associated with retained trees, the surrounding plantation has to be managed with regard to their specific demands. In the case of oak-associated species, including many red-listed species, several studies have shown that light is an important factor. The aim of this study was to analyze the effects of increased openness around oaks (Quercus robur) in spruce plantations (Picea abies) on species richness and abundance of oak-associated saproxylic beetles. The study was performed in nine spruce plantations located in southern Sweden, with mature oaks standing in a gradient of canopy openness. Beetles were collected from 54 oaks from May to September during two seasons, using window traps. The analyses revealed that increased openness around oaks increases species richness and abundance of oak-associated beetles. By including insolation angle in the analysis, we found that it is mainly the degree of openness directed south from the oak that has positive effects on beetle richness. These findings imply that it is desirable to maintain an open area around retained oaks, and that this area should be placed on the southern side of the oak to maximize the biodiversity benefit in relation to costs for the forest owner.     

Micro and macro-habitat associations in saproxylic beetles: implications for biodiversity management

Restoration of habitats is critically important in preventing full realization of the extinction debt owed as a result of anthropogenic habitat destruction. Although much emphasis has been placed on macrohabitats, suitable microhabitats are also vital for the survival of most species. The aim of this large-scale field experiment was to evaluate the relative importance of manipulated microhabitats, i.e., dead wood substrates of spruce (snags, and logs that were burned, inoculated with wood fungi or shaded) and macrohabitats, i.e., stand types (clear-cuts, mature managed forests, and forest reserves) for species richness, abundance and assemblage composition of all saproxylic and red-listed saproxylic beetles. Beetles were collected in emergence traps in 30 forest stands in 2001, 2003, 2004 and 2006. More individuals emerged from snags and untreated logs than from burned and shaded logs, but species richness did not differ among substrates. Assemblage composition differed among substrates for both all saproxylics and red-listed saproxylic species, mainly attributed to different assemblage composition on snags. This suggests that the practise of leaving snags for conservation purposes should be complemented with log supplementation. Clear-cuts supported fewer species and different assemblages from mature managed forests and reserves. Neither abundance, nor species richness or assemblage composition differed between reserves and mature managed forests. This suggests that managed stands subjected to selective cutting, not clear-felling, maintain sufficient old growth characteristics and continuity to maintain more or less intact assemblages of saproxylic beetles. Thus, alternative management methods, e.g., continuity forestry should be considered for some of these stands to maintain continuity and conservation values. Furthermore, the significantly higher estimated abundance per ha of red-listed beetles in reserves underlines the importance of reserves for maintaining viable populations of rare red-listed species and as source areas for saproxylic species in boreal forest landscapes.     

Spatial distribution of saproxylic beetles on trunks of standing Scots pine trees

1. The spatial distribution of saproxylic beetles landing and climbing stems of 12 grown and healthy Scots pine trees (Pinus sylvestris L.) was examined in detail during a 3-year trial. The beetles were sampled using sticky traps attached to the tree trunk at three tree parts (i.e., the trunk base, middle part of the trunk, and trunk in crown). Each tree part was sampled using two sticky traps facing opposite cardinal directions (north and south).
2. In total, 4063 individuals representing 169 taxa of saproxylic beetles were yielded in the 1405 obtained samples.
3. Remarkable interstrata diversification of saproxylic beetle assemblages reflecting distribution of breeding substrates in the forest stand was observed. Overall, species richness decreased considerably from the trunk base to the crown, with intermediate values for the middle part of the trunk. However, the abundance of mycetophages and saproxylophages decreased from the trunk base to the crown, with xylophages being the most abundant in the tree crown.
4. The cardinal direction exhibited overall weak effects on abundance, species richness, and composition, but the occurrence of a few particular species was significantly associated with the south (13 species) or north (3 species) facing part of the trunk.
5. Our results suggest that primary attraction and random landing strategies are combined during the substrate selection by many species.
6. High variability in the importance of the studied explanatory variables was observed among taxa, which shows how the species composition recorded in any study might alter its general outcomes.

     

Succession of saproxylic beetles associated with decomposition of boreal white spruce logs

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Influence of boreal forest succession and dead wood qualities on saproxylic beetles

1 Saproxylic insects, a functional group dominated by beetles, are dependent on dead or moribund trees as habitat elements. 2 Although there are few studies of saproxylic insects from the North American boreal zone, European studies demonstrate that forest harvest can lead to a biologically significant decrease in saproxylic beetle diversity. 3 We studied saproxylic beetles in the North American boreal mixedwood forest using flight intercept traps established on naturally dead and girdled trembling aspen and spruce trees along a successional gradient of undisturbed stands from deciduous to coniferous overstory trees. 4 Composition and diversity of beetle assemblages differed among forest successional types. 5 Snag age class was an important determinant of composition for saproxylic beetle assemblages. 6 Multivariate regression analysis of these data indicated that saproxylic beetles are responding to changes in coarse woody debris, and not to the relative densities of canopy tree species, although these variables are strongly correlated. 7 Coarse woody debris management should be a primary concern in forest management plans seeking to conserve saproxylic organisms and the critical ecosystem functions (i.e. nutrient cycling) in which they participate.     

Past levels of canopy closure affect the occurrence of veteran trees and flagship saproxylic beetles

Aim

Open woodlands are biologically highly diverse habitats, and veteran (i.e., old, senescent) trees are key structures supporting their biodiversity. Open canopy structure had been maintained by both natural‐ and human‐induced disturbances. In the past two centuries, suppression of such disturbances, together with forestry intensification, has turned most lowland woodlands into closed‐canopy forests. We investigated the effect of increased canopy closure on veteran trees and several threatened beetles associated with them.

Location

Floodplain woodlands along the lower Dyje and Morava rivers, Czech Republic.

Methods

We used an approach combining the study of aerial photographs with on‐ground survey of veteran trees and associated endangered beetles. The aerial images were used to obtain the information on historical (1938) and recent (2009) canopy closure in the area of 146 km², where we mapped large oaks (d.b.h. >70 cm), hollow trees and three associated beetles including the hermit beetle (Osmoderma barnabita), the great capricorn beetle (Cerambyx cerdo) and the jewel beetle Eurythyrea quercus.

Results

The presence of large oaks, hollow trees and their associated beetle species are negatively related to recent high canopy closure, and the historical level of canopy closure matters, as in nowadays closed‐canopy stands, the beetles and veteran trees are more common in places that were rather open in 1938 than in the places with closed canopy already in 1938. Moreover, the health state of veteran trees highly depends on the canopy closure.

Main conclusion

The negative effect of canopy closure on veteran trees and their endangered inhabitants is several decades delayed and may thus often go undetected. In the forests, however, large and hollow trees and their associated biodiversity are relics of the past, more open conditions. The restoration of open woodlands is therefore vital for preventing their further decline. Conservation management planning needs to take this into account wherever, veteran trees and associated biota are concerned.

     

Membranomyces species are common ectomycorrhizal symbionts in Northern Hemisphere forests

Membranomyces (Clavulinaceae, Cantharellales) Jülich consists of two described species of resupinate (crust-like) basidiomycetes. Previous studies indicated that Membranomyces falls within the Clavulinaceae, but the phylogenetic position of the genus has not been fully resolved. Membranomyces species were thought to be saprotrophic until 2003 when Tedersoo et al. detected Membranomyces delectabilis on ectomycorrhizal roots of Populus and Picea. Membranomyces was previously known only from collections made in eastern Canada and Europe. We recently sequenced the ITS rDNA barcode region from Scandinavian herbarium specimens identified as M. delectabilis and Membranomyces spurius. Phylogenetic analyses of these sporocarp sequences and similar environmental sequences indicated that Membranomyces is more diverse than previously thought and forms ectomycorrhizas with hosts from a diverse range of plant families in many north temperate ecosystems.     

Head shape variation in cerambycid saproxylic beetles as a function of host plant selection

Saproxylic insects depend on deadwood for larval development, and a certain degree of specialization may be involved in their choice of host plants and/or wood in a particular stage of degradation. The plant species chosen for oviposition in turn act as an environmental pressure on the head morphology of larvae and it is expected that head shape plasticity varies directly with the number of woody plant species used for larval development in each insect species. We analyzed head shape variation in saproxylic beetles with respect to host plant species, maximum time of larval emergence and season of the year when insects colonized branches. Generalist species in the use of host plants showed significant variation in head shape and size. Time of emergence and season did not appear to affect head shape, although season was a determinant factor of abundance and possibly head size variation.