Diversity of saproxylic bryophytes in old‐growth and managed beech forests in the central Balkans

Abstract

The diversity of saproxylic bryophyte species in beech forest stands from the wide region of the central Balkans (i.e. Serbia and Montenegro) was studied, and this study is the first of such a type in SE Europe. Comparison of preserved old‐growth and managed forests were made. Bryophyte species diversity is higher in primeval forest stands where the spectra of dead wood in various decaying stages of its dynamics are present. The ecological group of epixylic specialists is predominant among the bryophytes recorded. Threatened bryophyte species occur in old‐growth beech stands. The dead wood as habitat together with some other factors are extremely important for the surviving of epixylic bryophyte; so these species can be used as bioindicator bryophyte species of old‐growth or managed and structured forest ecosystems.     

Field response of non‐target beetles to Ips sexdentatus aggregation pheromone and pine volatiles

Although mass trapping cannot be a definitive control measure, it is one of the few ones available to contain the destruction of millions of cubic metres of conifer forests perpetrated every year worldwide by bark beetles. However, using bark beetle aggregation pheromones during both monitoring and control programs may negatively affect other saproxylic insects. The aim of this study was to describe the response of both Ips sexdentatus and its saproxylic beetle associates, especially predators, to traps baited with a commercial blend of I. sexdentatus aggregation pheromone. Furthermore, the usefulness of adding pine volatiles, such as (?)‐α‐pinene and ethanol, to the pheromone was discussed. The commercial blend proved to be attractive to I. sexdentatus adults, both when used alone and together with pine volatiles. Pheromone attractiveness, however, was lessened by the addition of the volatiles. The pheromone blend proved to be attractive to Thanasimus formicarius, as well as to other predator species. Overall, although during our study, traps baited only with (?)‐α‐pinene and ethanol attracted some predator specimens, I. sexdentatus pheromone traps were more attractive. Our study confirms that calendar differences in flight activity between the bark beetle and its predators are substantial; therefore, they should be taken into account when planning control measures. According to our data, the commercial blend of I. sexdentatus pheromone seems to be the most effective, among the baits used, in catching I. sexdentatus adults, while reducing the impact on T. formicarius.     

Wood decomposition as influenced by invertebrates

The diversity and habitat requirements of invertebrates associated with dead wood have been the subjects of hundreds of studies in recent years but we still know very little about the ecological or economic importance of these organisms. The purpose of this review is to examine whether, how and to what extent invertebrates affect wood decomposition in terrestrial ecosystems. Three broad conclusions can be reached from the available literature. First, wood decomposition is largely driven by microbial activity but invertebrates also play a significant role in both temperate and tropical environments. Primary mechanisms include enzymatic digestion (involving both endogenous enzymes and those produced by endo‐ and ectosymbionts), substrate alteration (tunnelling and fragmentation), biotic interactions and nitrogen fertilization (i.e. promoting nitrogen fixation by endosymbiotic and free‐living bacteria). Second, the effects of individual invertebrate taxa or functional groups can be accelerative or inhibitory but the cumulative effect of the entire community is generally to accelerate wood decomposition, at least during the early stages of the process (most studies are limited to the first 2–3 years). Although methodological differences and design limitations preclude meta‐analysis, studies aimed at quantifying the contributions of invertebrates to wood decomposition commonly attribute 10–20% of wood loss to these organisms. Finally, some taxa appear to be particularly influential with respect to promoting wood decomposition. These include large wood‐boring beetles (Coleoptera) and termites (Termitoidae), especially fungus‐farming macrotermitines. The presence or absence of these species may be more consequential than species richness and the influence of invertebrates is likely to vary biogeographically.     

The Conservation of Saproxylic Insects in Tropical Forests: A Research Agenda

Invariably, insects are overlooked when tropical forest management issues are discussed, because there are so many species, they are taxonomically intractable and so poorly known. Often people take the view that if you look after the vegetation and vertebrates, the insects will look after themselves. This may be true for some functional groups, but for saproxylic insects, this seems unlikely. Their study deserves high priority, since they are dependent on the very resource – wood – whose removal from the ecosystem is the usual object of forest management. Given the current international effort to develop 'criteria and indicators' to monitor sustainable forest management for biodiversity values, there is a window of opportunity for sound ecological research on saproxylic insects to influence the formulation of forest policy such that their needs can be taken into account. There is already a large body of knowledge on temperate and boreal region saproxylic insects, and on the effects that logging has on them, but knowledge of the tropical forest situation lags far behind. This paper proposes a research agenda to enable the needs of saproxylic insects to be taken into account in natural forest management in the tropics. Basic questions, such as whether logging has so far had an impact on tropical saproxylic insects, and whether there are workable sampling techniques to investigate this, still remain to be addressed and deserve high priority. The links between the responses of saproxylic insects and more 'charismatic' study species need to be investigated. We also need to know whether there is a correlation between the intensity of logging and the response of saproxylic insects, and, critically, whether we would be justified in measuring some surrogate aspects of forest structure (as potential habitat for saproxylic insects) rather than the saproxylic insects themselves, and modelling this to determine likely impacts of different management regimes. We consider such an ambitious research agenda as justified given the scale of impact that forest use and management is likely to have on tropical forest insects in the future.     

Specialists in ancient trees are more affected by climate than generalists

Ancient trees are considered one of the most important habitats for biodiversity in Europe and North America. They support exceptional numbers of specialized species, including a range of rare and endangered wood‐living insects. In this study, we use a dataset of 105 sites spanning a climatic gradient along the oak range of Norway and Sweden to investigate the importance of temperature and precipitation on beetle species richness in ancient, hollow oak trees. We expected that increased summer temperature would positively influence all wood‐living beetle species whereas precipitation would be less important with a negligible or negative impact. Surprisingly, only oak‐specialist beetles with a northern distribution increased in species richness with temperature. Few specialist beetles and no generalist beetles responded to the rise of 4°C in summer as covered by our climatic gradient. The negative effect of precipitation affected more specialist species than did temperature, whereas the generalists remained unaffected. In summary, we suggest that increased summer temperature is likely to benefit a few specialist beetles within this dead wood community, but a larger number of specialists are likely to decline due to increased precipitation. In addition, generalist species will remain unaffected. To minimize adverse impacts of climate change on this important community, long‐term management plans for ancient trees are important.     

What factors influence the diversity of saproxylic beetles? A multiscaled study from a spruce forest in southern Norway

The diversity in different groups of obligate saproxylic beetles was related to ecological variables at three levels of spatial scale in mature spruce-dominated forest. The variables were connected to: (i) decaying wood, (ii) wood-inhabiting fungi, (iii) the level of disturbance, (iv) landscape ecology, and (v) vegetational structure. Several strong relationships were found at medium (1 km²) and large scales (4 km²), while only weak relationships were found at a small scale (0.16 ha; 1 ha=10⁴ m²). This may be explained by the local variations in habitat parameters and the high mobilities of many beetle species. Factors connected to decaying wood and wood-inhabiting fungi were clearly the most important factors at all scale levels. In particular, the variables diversity of dead tree parts, number of dead trees of large diameter and number of polypore fungi species increased the species richness of many groups and increased the abundance of many species. Eight species were absent below a certain density of decaying wood per 1 or 4 km². Former extensive cutting was a negative factor at large scale, probably because of decreasing recolonization with increasing distance to the source habitats. Thinning reduced the diversity of species associated with birch. The development of guidelines favouring the diversity of saproxylic beetles are discussed below.     

Colonization Patterns of Insects Breeding in Wood-Decaying Fungi

Many insects dependent on dead wood are considered threatened by modern forestry. This may partly be due to substrate patches being too widely scattered to be effectively colonized. We studied how rates of colonization by insect species breeding in polypore fruiting bodies are affected by interpatch distance and habitat matrix characteristics. In field experiments, fruiting bodies of Fomitopsis pinicola and Fomes fomentarius were put out at different distances from natural sources of insects. The anobiid beetles Dorcatoma spp. were the most successful colonizers of distant patches, and they readily flew over open fields. Cis beetles were less successful colonizers, despite their generally higher abundance. We hypothesize that the Dorcatoma spp. are inferior competitors, but superior colonizers of distant resources compared with Cis spp. The flies Leucophenga quinquemaculata (Drosophilidae) and Medetera impigra (Dolichopodidae) appeared to be more affected by distance than the beetles studied in their colonization of fungal fruiting bodies. Lower rates of parasitism were recorded on distant patches, and parasitoids appeared more affected by distance than their hosts. Most of the insect species studied can probably persist in the managed forest landscape if suitable breeding substrate is created continuously on a 1 km² scale.     

Effects of Clearfell Harvesting on Lucanid Beetles (Coleoptera: Lucanidae) in Wet and Dry Sclerophyll Forests in Tasmania

The effects of commercial forestry harvest and regeneration practices (clearfelling and slash-burning) on the lucanid fauna of the wet sclerophyll forests of southern Tasmania and the dry sclerophyll forests of eastern Tasmania were examined using pitfall catches. Lucanids are saproxylic beetles, dependent on dead, moribund and decaying wood. Samples taken from old-growth forest and from a chronosequence of sites regenerating after logging, in each forest type, were used to compare the species richness and abundance of the lucanid assemblages. In both forest types, species richness and abundance was highest in the youngest regeneration sites (1–3 year), reflecting the species richness of the original and adjacent unlogged forest, lowest in the older (20–25 year) sites, and variable in the old-growth sites. TWINSPAN cluster analysis showed no clear distinction between regeneration and old-growth forest. The post-harvest slash and stump residue provided an important refugium and initial habitat, but our research indicates that some species may not maintain populations in the long term. Our results suggest that most species of lucanids will find a continuous supply of suitable habitat only in old-growth forests; and such species may become less common as clearfell harvesting leads to a replacement of heterogeneous old-growth forest with single-aged monospecific stands. Continuity of supply of wood in all decay stages, the maintenance of sufficient source areas, and biological connectivity between old-growth stands to enable dispersal, are all likely to be essential to maintain lucanid beetle community integrity. If similar principles apply to other saproxylic species of invertebrate, then clearfelling and slash-burning may cause a gradual extinction of an important element of the forest biota.     

Surprising longhorned beetle (Coleoptera, Cerambycidae) richness along an Italian alpine valley

In this paper we report about 88 longhorned beetles (Cerambycidae) species found in 6929 hectares and distributed along an altitudinal gradient of 1500 m of an Italian alpine valley (Val Genova, central-eastern Italian Alps). The species richness, result merging data from sixty years (1947-2007) of entomological surveys, corresponds to the 32% of the Italian cerambycid fauna confirming the high richness/surface ratio, probably unique in the Alps. The effect of thirteen environmental variables was tested on the species richness, but only the elevation resulted able to affect it. The species richness decrease with altitude not gradually, but experience a strong step above 1700 m a.s.l.. The highest species richness (average values of 42 species) was recorded at the lowest and mid elevations (between 800 and 1600 m a.s.l.). The species turnover along the altitudinal gradient is low suggesting moderate habitat turnover along the valley.One of the eighty-eight observed species, Tragosoma depsarium,is classified near threatened by the IUCN. Our data suggest that the wilderness of the valley close to the suitable management of grasslands and forests, help to support high level of cerambycids diversity. This biodiversity is good indicators of health of the wood saproxylic assemblages, as well an important food source for many vertebrate predators.     

Horizontal and vertical distribution of saproxylic beetles (Col., Buprestidae, Cerambycidae, Scolytinae) across sections of forest edges

Abstract:  We evaluated the preferred home ranges of three saproxylic beetle taxa along transects from the open field into the forest interior, and from the forest floor up to the canopy. By means of trap sets on metal scaffolds, vertical and horizontal strata were sampled across two types of forest edges: soft-edge ecotones with a gradual transition from the field into the forest and hard edges with an abrupt transition. The forest edges consisted of different strata such as herbaceous fringe, shrub belt, unmanaged forest and managed forest. The thermophilic buprestids were mainly caught in the open land (herbaceous fringe and agricultural land) and in the upper forest mantle. In general, the cerambycids were most abundant in the open land and the lower forest mantle, but a few species favoured the forest interior. The bark beetles (Scolytinae) were equally distributed in all habitats. These distribution patterns of the taxa were observed in terms of both species numbers and abundances. Each species with at least five collected specimens was assigned to one of the three habitat types: open land, forest mantle and forest interior. Of 74 ranked species, only 16% were prevalent in the forest interior and are thus considered to be true forest species. The other 84% of the species were attributed to open land or the forest mantle and are, therefore, forest edge species. Soft forest edges generally supported a higher species richness than hard edges, particularly as regards Cerambycidae and Scolytinae. In terms of Shannon diversity, soft edges tended to be more diverse in buprestids and cerambycids. Overall, the forest interior showed the least species richness and diversity. Therefore, for the conservation of saproxylic beetles, not only the amount and quality of dead wood is important, but also the presence and design of forest boundary structures.     

Ecologie des coléoptères saproxyliques dans les forêts de l’étage montagnard des Alpes du nord françaises

Ecology of the saproxylic beetles in mountain forests of the northern French Alps. Both saproxylic beetles and deadwood stocks were studied in order to detect special deadwood features important for these organisms, and to better understand their ecology. A total of 37 window traps were used in ten stations covering four mountain forests in the French northern Alps. In the same time, the deadwood stocks were inventoried in the ten stations. 181 m³ of deadwood (1219 fragments) were measured and 4268 individuals belonging to 235 saproxylic species were trapped. The deadwood volume ranges from 21.1 to 233.6 m³/ha and the number of beetle species from 27 to 113. A positive relation was observed between the stand age, the whole volume of deadwood and the species richness of saproxylic beetles. However, in some deadwood rich sites the beetle diversity is not as high as expected. This was explained by the negative influence of the management history of the forest. The compositions of saproxylic communities depend also of some characteristics of the deadwood stocks. Xylophagous and zoophagous species prefer “not lying and recently dead softwood” while the mycophagous and saproxylophagous species are strongly dependent on fungi growing on hardwood, respectively under “large surfaces of barks” and in “large woody debris”. Surprisingly, the majority of the saproxylic beetles living on softwood does not depend on the most represented resinous deadwood type: the “decayed and lying softwood”.     

New Zealand's giant Collembola: New information on distribution and morphology for Holacanthella Börner, 1906 (Neanuridae: Uchidanurinae)

Abstract

The New Zealand collembolan fauna currently includes five described species of the striking endemic genus Holacanthella (springtails). Holacanthella species are saproxylic decomposers of cool temperate forest ecosystems, and they contribute to nutrient cycling of coarse woody debris. All species of the genus have the dorsal and lateral surfaces furnished with conspicuous red, orange, yellow or white digitations. They are among the largest Collembola known, with some individuals reaching 17 mm in length. We examined new material of the five species from throughout New Zealand, as well as existing museum material, to understand more fully their distributions. We provide an updated key to the five Holacanthella species and discuss the conservation of rare/range restricted species, and propose hypotheses to explain their distributions. One species, H. laterospinosa, is known only from Cuvier Island and the Coromandel Peninsula, North Island, but the distributions of the remaining four species appears to reflect the turbulent geological history of New Zealand during the Pliocene. Intraspecific variation in several gross morphological characters was observed in all species. We provide a comparison of characters with the other genera within the Uchidanurinae in order to characterise the genus more fully with respect to allied genera. Holacanthella species are particularly vulnerable to human‐mediated disturbance by forest modification because of their strict habitat requirements and low mobility, and we stress that conservation efforts should focus on protecting critical habitats for each species.     

Historic parks and pasture-woodlands: The National Trust resource and its conservation

The National Trust owns and manages considerably more historic parklands and other types of pasture-woodlands than any other conservation body in the British Isles. These include one virtually intact medieval forest and fragments of others, as well as more than one hundred historic parks and many wooded commons. This special responsibility is described, both in terms of the assemblages of plants and animals now protected by the Trust, and in terms of the active management programmes for their conservation. Management issues of immediate concern on Trust properties are identified.     

Population structure in a saproxylic funnelweb spider (Hexathelidae: Hadronyche) along a forested rainfall gradient

We present research undertaken at Tallaganda State Forest, Australia, describing how current climatic conditions impact upon the saproxylic (decaying wood) habitat and the relationship between this habitat and the demography and morphology of a saproxylic funnelweb spider ( Hadronyche sp.). Climatic data support a north–south habitat cline and, to a lesser extent, a short-range, aspect-driven habitat cline. Rainfall and log moisture content increased with latitude through the forest, and aspect affected the amount of solar radiation penetrating to ground level. The distribution and abundance of Hadronyche varied among sites along both clines in response to several variables. The decay state of logs was highly influential, with wet, highly decayed logs favoured over dry, hard ones. Population density was highest in wetter, southern sites where these logs were abundant, but rare, suitably decomposed logs in dry, northern sites still typically hosted a comparable number of individuals. Morphological measurements showed some phenotypic variation along the north–south habitat gradient, but not over the short-range, aspect-driven gradients. Hadronyche would not be expected to show similarly strong patterns of molecular variation as seen in saproxylic Collembola and Onychophora within Tallaganda as it appears to be more vagile.     

A tale of two flatties: different responses of two terrestrial flatworms to past environmental climatic fluctuations at Tallaganda in montane southeastern Australia

Comparative phylogeographic studies of animals with low mobility and/or high habitat specificity remain rare, yet such organisms may hold fine-grained palaeoecological signal. Comparisons of multiple, codistributed species can elucidate major historical events. As part of a multitaxon programme, mitochondrial cytochrome oxidase I (COI) variation was analysed in two species of terrestrial flatworm, Artioposthia lucasi and Caenoplana coerulea. We applied coalescent demographic estimators and nested clade analysis to examine responses to past, landscape-scale, cooling-drying events in a model system of montane forest (Tallaganda). Correspondence of haplotype groups in both species to previously proposed microbiogeographic regions indicates at least four refuges from cool, dry conditions. The region predicted to hold the highest quality refuges (the Eastern Slopes Region), is indicated to have been a long-term refuge in both species, but so are several other regions. Coalescent analyses suggest that populations of A. lucasi are declining, while C. coerulea is expanding, although stronger population substructure in the former could yield similar patterns in the data. The differences in spatial and temporal genetic variation in the two species could be explained by differences in ecological attributes: A. lucasi is predicted to have lower dispersal ability but may be better able to withstand cold conditions. Thus, different contemporary population dynamics may reflect different responses to recent (Holocene) climate warming. The two species show highly congruent patterns of catchment-based local genetic endemism with one another and with previously studied slime-mould grazing Collembola.     

Phylogeny of European Pyrochroa (Coleoptera,Pyrochroidae) reveals cryptic taxa and different glacial histories

Only three saproxylic species of Pyrochroinae (Coleoptera: Pyrochroidae) are distributed in Europe, two of which belonging to Pyrochroa: P. coccinea and P. serraticornis. However, P. serraticornis is polytypic, for the presence of the endemic subspecies P. s. kiesenwetteri in southern Italy. Using both molecular and morphological data, we explored the phylogeny of the European Pyrochroa species. A multilocus (COI, CAD, 28S) phylogenetic analysis helped highlight different evolutionary histories for the two examined species. First, P. coccinea, distributed throughout Europe, showed a high differentiation among Italian and European populations. Furthermore, three different taxonomic entities were identified within P. serraticornis, among which the cryptic species Pyrochroa bifoveata sp. n. from central Europe is described and illustrated. A comprehensive identification key to the European Pyrochroinae is also provided. Our results also suggested an historical survival of P. coccinea and P. s. kiesenwetteri in glacial refugia in Italy, and a subsequent post-glacial spread of the former species throughout the Peninsula. In contrast, the current distribution of P. s. serraticornis likely originated from a post-glacial colonization of western European relict populations, while the survival of P. bifoveata plausibly occurred in more eastern glacial refugia (e.g. Carpathian or Balkan regions). Similarly, the European populations of P. coccinea could have originated from relict populations in glacial refugia out from the Italian Peninsula. More comprehensive data on the taxonomy, ecology and biogeography of Pyrochroa are needed to learn more about these species and to help preserve the European saproxylic fauna.     

Phylogeography recapitulates topography: very fine-scale local endemism of a saproxylic 'giant' springtail at Tallaganda in the Great Dividing Range of south-east Australia

Comparative phylogeography can reveal processes and historical events that shape the biodiversity of species and communities. As part of a comparative research program, the phylogeography of a new, endemic Australian genus and species of log-dependent (saproxylic) collembola was investigated using mitochondrial sequences, allozymes and anonymous single-copy nuclear markers. We found the genetic structure of the species corresponds with five a priori microbiogeographical regions, with population subdivision at various depths owing to palaeoclimatic influences. Closely related mtDNA haplotypes are codistributed within a single region or occur in adjacent regions, nuclear allele frequencies are more similar among more proximate populations, and interpopulation migration is rare. Based on mtDNA divergence, a late Miocene-late Pliocene coalescence is likely. The present-day distribution of genetic diversity seems to have been impacted by three major climatic events: Pliocene cooling and drying (2.5-7 million years before present, Mybp), early Pleistocene wet-dry oscillations (c. 1.2 Mybp) and the more recent glacial-interglacial cycles that have characterized the latter part of the Quaternary (<0.4 Mybp).     

Nature protection areas of Europe are insufficient to preserve the threatened beetle Rosalia alpina (Coleoptera: Cerambycidae): evidence from species distribution models and conservation gap analysis

1. Natura 2000 network (N2000) and national protected areas (NPAs) are recognised as the most important core ‘units’ for biological conservation in Europe. 2. Species distribution models (SDMs) were developed to detect the potential distribution of the rare and threatened cerambycid beetle Rosalia alpina L. in Europe, and the amount of suitable habitat within the N2000 network [special areas of conservation (SACs) and special protection areas (SPAs)], NPAs (e.g. national parks, regional parks, state reserves, natural monuments and protected landscapes) and the overall European protected area network (EPAN) (N2000 + NPAs) was quantified. 3. According to this analysis, the suitable habitat for R. alpina in Europe amounts to c. 754 171 km² and stretches across substantially uninterrupted areas from Portugal to Romania (west to east) and from Greece to Germany (south to north). The overlay between the existing system of conservation areas in Europe (N2000 and NPAs) and the binary map for R. alpina showed that only c. 42% of potential habitat is protected. SACs and SPAs protect c. 25% and 21% of potential habitat, respectively. However, because the two site types often spatially overlap, when taken together the entire N2000 network protects c. 31% of potential habitat. Instead, NPAs offer a degree of protection of c. 29%. Overall, almost 60% of the area potentially suitable for the species is unprotected by the EPAN, an aspect that should be considered carefully when planning the conservation of this beetle at a large scale. 4. These results may also help to focus field surveys in selected areas where greater chances of success are encountered to save resources and increase survey effectiveness.     

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.