Beetle responses to artificial gaps in an oceanic island forest: implications for invasive tree management to conserve endemic species diversity

Natural forests are often replaced by invasive alien trees on isolated oceanic islands. Adequate eradication of invasive trees should be conducted with the goal of biodiversity conservation, because islands support many endemic organisms that depend on native forests. An invasive alien tree, Bischofia javanica Blume (Euphorbiaceae), has invaded and replaced natural forests on the oceanic Ogasawara (Bonin) Islands, Japan, in the northwestern Pacific Ocean. To determine how the removal of B. javanica trees affects insect diversity, we examined flying beetles captured using Malaise traps in B. javanica forests on Hahajima. The abundance, species density, and species composition of wood-boring beetles (Coleoptera: Cerambycidae, Elateridae, Mordellidae, and Scolytidae) were compared between closed-canopy sites and gaps created by girdling B. javanica trees in alien forests during two seasons (June–July and October–November 2005). Of the collected beetles, 75.8, 87.5, 90.0, and 0.0% of cerambycid, elaterid, mordellid, and scolytid beetle species, respectively, were endemic to the Ogasawara Islands. More cerambycid, elaterid, and mordellid individuals were captured in June–July than in October–November; the number of scolytid individuals did not differ between seasons. More cerambycid, elaterid, and scolytid individuals were captured in artificial gaps than on the closed-canopy forest floor. Although fewer mordellid individuals were captured in gaps, more endemic mordellids were captured in gaps. More cerambycid and scolytid species were captured in artificial gaps than in closed-canopy areas. The positive responses of beetles to artificial gaps suggest that the removal of B. javanica increases beetle diversity and the abundance of endemic beetles.     

The rarest and least protected forests in biodiversity hotspots

The goal of biodiversity hotspots is to identify regions around the world where conservation priorities should be focused. We undertake a geographic information system and remote sensing analysis to identify the rarest and least protected forests in biodiversity hotspots. World Wildlife Fund ecoregions with terrestrial forest were subset from 34 biodiversity hotspots and forest cover calculated from GlobCover data at a 300?m pixel resolution. There were 276 ecoregions in 32 biodiversity hotspots classified as containing terrestrial forests. When the first quartile of forest ecoregions was subset based on smallest extent of forest cover in protected areas, there were 69 rare forests identified within 20 biodiversity hotspots. Most rare forest ecoregions (45) occurred on islands or island archipelagos and 47 rare forest ecoregions contained less than 10?% forest cover in protected areas. San Félix-San Ambrosio Islands Temperate Forests, Tubuai Tropical Moist Forests, Maldives-Lakshadweep-Chagos Archipelago Tropical Moist Forests, and Yap Tropical Dry Forests were identified as the least protected and possibly most vulnerable forests within biodiversity hotspots. These ecoregions cover less than 500?km², forest cover is less than 50?km², and there are no protected areas. There is a need to update classifications and boundaries of protected areas, insure that islands are included in global land cover datasets, and identify levels of endemism and endangerment within forest ecoregions. This should improve our ability to compare, prioritize, and monitor forests in biodiversity hotspots.     

Does wing morphology affect recolonization of restored farmland by ground‐dwelling beetles?

Revegetation of previously cleared land is widely used to increase habitat area and connectivity of remnant vegetation for biodiversity conservation. Whether new habitat attracts or supports fauna depends on the dispersal traits of those fauna as well as the structure and composition of the surrounding landscape. Here, we examined wing morphology as a key dispersal trait for beetles in a revegetated landscape and asked, first, how it was related to phylogeny (family), trophic position, and body size. Second, we asked if wing morphology of recolonizing (or persisting) beetles varied with habitat characteristics at multiple scales, from microhabitat to landscape context. Third, we examined how common winged and wingless species responded to habitat at multiple scales. We measured the wing morphology of ground‐dwelling beetles from a restoration chronosequence, including paddocks, “young” revegetation (8–11 years old), “old” revegetation (14–19 years old), and fenced remnant vegetation. We found that body size and family membership were significant predictors of winglessness, with wingless species of carabids and curculionids being larger than their winged counterparts. We found no difference in the number of sites occupied by winged and wingless species, and no relationship between the wing morphology traits represented in different locations and habitat characteristics or landscape context. Furthermore, the most abundant species of both winged and wingless ground‐dwelling beetles had relatively little affinity to any habitat successional stage. Thus, despite intrinsic differences in wing morphology among species of ground‐dwelling beetle, we found no evidence that flight‐related dispersal limitations influenced recolonization (or persistence) in this landscape.     

Differences in endemic insect assemblages among vegetation types on a small island of the oceanic Ogasawara Islands

Natural vegetation is often replaced by invasive alien plants on isolated oceanic islands. To determine how invasive alien plants affect insect diversity, we compared flying insects captured using Malaise traps among different vegetation types on a small island (Nishijima; 0.49 km²) in the oceanic Ogasawara (Bonin) Islands in the north‐western Pacific. The numbers of individuals and species, and the species composition of pollinators (bees), predators (wasps) and wood borers (cerambycid, mordellid and elaterid beetles) were compared among three vegetation types: Casuarina equisetifolia (an invasive alien tree) forest, natural forest and natural grassland (forest edge), during two seasons (June and October–November 2005). In traps, 80.0, 66.7, 87.5, 85.7 and 100.0% of bee, wasp, cerambycid, mordellid and elaterid beetle species, respectively, were endemic to the Ogasawara Islands. Grassland had the highest wasp and bee species richness, whereas natural forest had the highest species richness of wood‐boring beetles. The C. equisetifolia forest had the poorest species richness for most insect groups (except mordellid beetles). More individuals of most insect groups (except bees) were captured in June than in October–November. More individual bees and wasps were captured in grassland than in forests, whereas more individual mordellid and elaterid beetles were captured in forests than in grassland. The number of cerambycid individuals did not differ among vegetation types. Redundancy analysis suggested that most insect species preferred natural forest or grassland to alien forest. Therefore, further invasion of natural grassland and forest by the alien tree C. equisetifolia may negatively affect the endemic insect fauna of Nishijima.     

Non-native plantation forests as alternative habitat for native forest beetles in a heavily modified landscape

The once extensive native forests of New Zealand’s central North Island are heavily fragmented, and the scattered remnants are now surrounded by a matrix of exotic pastoral grasslands and Pinus radiata plantation forests. The importance of these exotic habitats for native biodiversity is poorly understood. This study examines the utilisation of exotic plantation forests by native beetles in a heavily modified landscape. The diversity of selected beetle taxa was compared at multiple distances across edge gradients between each of the six possible combinations of adjacent pastoral, plantation, clearfell and native forest land-use types. Estimated species richness (Michaelis–Menten) was greater in production habitats than native forest; however this was largely due to the absence of exotic species in native forest. Beetle relative abundance was highest in clearfell-harvested areas, mainly due to colonisation by open-habitat, disturbance-adapted species. More importantly, though, of all the non-native habitats sampled, beetle species composition in mature P. radiata was most similar to native forest. Understanding the influence of key environmental factors and stand level management is important for enhancing biodiversity values within the landscape. Native habitat proximity was the most significant environmental correlate of beetle community composition, highlighting the importance of retaining native remnants within plantation landscapes. The proportion of exotic beetles was consistently low in mature plantation stands, however it increased in pasture sites at increasing distances from native forest. These results suggest that exotic plantation forests may provide important alternative habitat for native forest beetles in landscapes with a low proportion of native forest cover.     

Review of the protected area network in Guinea, West Africa, and recommendations for new sites for biodiversity conservation

With only five protected areas dedicated to the conservation of biodiversity (two national parks, one strict nature reserve and two faunal reserves), Guinea has one of the smallest protected area networks in West Africa. As a result, two of the five ecoregions of the country and six of the 14 globally threatened large and medium-sized mammals occurring in Guinea are not found in the national protected area network. To identify areas with high biodiversity that could be included in the national protected area network, we used the Key Biodiversity Areas (KBA) methodology. We devised a scoring system to rank the identified KBAs according to their relative conservation significance. We identified a total of 16 KBAs throughout the country. Their proclamation as protected areas would result in the protection of all ecoregions and all but one of Guinea’s globally threatened large and medium-sized mammals. Twelve of the 16 KBAs have the legal status of classified forest, a status that should facilitate the change into formal biodiversity protected areas (IUCN category I–IV). Our analysis indicates that even if only the two areas with the highest conservation significance score, the Ziama and Diécké forests, become formal protected areas, this would provide protection to both the western Guinean lowland forests, one of the most threatened ecoregions in Africa, and to 11 of the 14 threatened large and medium-sized mammals occurring in Guinea.     

Species diversity of longicorn beetles in humid warm-temperate forests: the impact of forest management practices on old-growth forest species in southwestern Japan

In the humid warm-temperate zone of southwestern Japan, old-growthforests have been seriously fragmented to small remnants due to traditionalagriculture and coppicing as well as recent rapid plantation with conifers.Assemblages of longicorn beetles (Coleoptera: Disteniidae and Cerambycidae) werecompared among old-growth forests, second-growth forests and conifer plantationsusing collision traps baited with chemical attractants. Species richness oflongicorn beetles was poorer in second-growth forests and conifer plantationsthan in old-growth forests. It was proved by multidimensional scaling(MDS) that the beetle assemblages of old-growth forests were distinct from thoseof conifer plantations, while those of second-growth forests were intermediatebetween them. Further analysis showed that a number of species, including manyPidonia spp., were specific to or closely associated withold-growth forests, and the results were largely supported by the indicatorvalue (IndVal) approach. It is likely that many of such old-growth forestspecies in the larval and pupal stages require large broad-leaved trees standingor fallen with thick bark. At the same time, the flower-visiting adults wouldplay an important role in pollinating various herbaceous and woody plants.Regional forest management for the conservation of insect biodiversity is alsodiscussed.     

Small off-shore islands can serve as important refuges for endemic beetle conservation

Although large islands generally support a richer insect fauna than small islands, many large islands, which are more often inhabited, have lost numerous species because of human activities and introduced organisms. To clarify the consequences of endemic insect conservation on small islands near inhabited islands, we compared the species richness, abundance, and composition of two beetle groups (Coleoptera: Cerambycidae and Mordellidae) captured using Malaise traps among three islands (Chichijima, 24.0 km²; Anijima, 7.85 km²; Nishijima, 0.49 km²) in the oceanic Ogasawara (Bonin) Island group in the northwestern Pacific during June–July 2006 and 2007. Chichijima, the largest island, is inhabited, while Anijima and Nishijima are not. The numbers of cerambycid and mordellid species previously recorded were positively correlated with island area. However, the total numbers of cerambycid and mordellid species we captured in Malaise traps were not correlated with island area because we were unable to collect many species previously documented on Chichijima. The numbers of cerambycid and mordellid species per trap did not differ significantly among islands and years, although the deviance was well explained by the island variable. We captured greater numbers of cerambycid and mordellid individuals on Chichijima than on Anijima and Nishijima, and the numbers of cerambycid and mordellid individuals per trap significantly differed among islands and between years. Redundancy analysis (RDA) showed that the species composition of cerambycids and mordellids differed among the three islands. Whereas endangered species were rarely captured on Chichijima, alien or non-endemic species were frequently collected. Cerambycid and mordellid beetles on Chichijima may have been deleteriously affected by recent forest disturbance and introduced organisms. Therefore, conserving insect fauna on uninhabited island “refugia” is important for preserving the insect diversity of the Ogasawara Islands.     

What shapes cerambycid beetle communities in a tropical forest mosaic? Assessing the effects of host tree identity,forest structure,and vertical stratification

Due to anthropogenic activities, tropical rain forests face many challenges in sustaining biodiversity and maintaining global climates. This study explores how forest successional stage, tree composition, and stratum affect communities of saproxylic cerambycid beetles—concealed feeders that play important roles in forest nutrient cycling. Forty trees in five families (Fabaceae, Lecythidaceae, Malvaceae, Moraceae, and Sapotaceae) were sampled in a mosaic of old‐growth and secondary forest on the Osa Peninsula, Costa Rica. Bait branches yielded 3549 cerambycid individuals in 49 species. Species richness was almost identical in old‐growth and secondary forest, and both yielded specialists, but abundance was higher in old‐growth forest. Overall community structure was most strongly influenced by host plant species; within most plant families it was also impacted by forest successional status. Moraceae was the exception, presumably because the focal tree species was abundant in both old‐growth and secondary forest. Several host and old‐growth specialist species reached high densities within patches of old‐growth forest, but seldom colonized apparently suitable trees within secondary forest. This suggests that even small areas of old‐growth forest can act as refuges, but that secondary forest may act as a barrier to dispersal. The vulnerability of specialized saproxylic insects to land use change will be linked to the ability of their preferred hosts to disperse to and persist in successional habitats; rearing studies may provide the most accurate method to monitor community changes over time.     

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”.     

Diversity distribution of saproxylic beetles in Chilean Mediterranean forests: influence of spatiotemporal heterogeneity and perturbation

Mediterranean ecosystems have been recognized as a priority for biodiversity conservation due to their high levels of species richness and endemism. In South America, these environments are restricted to central Chile and represent a biodiversity hotspot. The study of saproxylic beetles in this area is an unexplored topic, despite the ecological role they play in these ecosystems and their potential usefulness for monitoring the degree of forest conservation. We investigated the diversity distribution of trophic guilds of saproxylic beetles in Chilean Mediterranean forests, to identify the main environmental variables that influence their distribution. We also analyzed seasonal dynamics as a key factor influencing insect communities and the effect of human disturbance on their diversity and composition. We identified characteristic species of perturbation degree as bioindicators for habitat monitoring. A total of 40 window traps were used to survey three Mediterranean forest types in the Río Clarillo National Reserve and the adjacent non-protected areas. We found that the diversity, abundance and composition of saproxylic beetles varied significantly spatiotemporally among the studied forest types and among perturbation degrees, showing different patterns depending on the trophic guild. Results indicated that conservation decisions should include the preservation of a larger range of different vegetation types and the nearby zones that have suffered low levels of disturbance or fragmentation and where actions promoting the presence of old native trees would have a significant conservation value.     

Hierarchical partitioning of ant diversity: implications for conservation of biogeographical diversity in arid and semi‐arid areas

Aim The scale of observation is important in detecting the spatial variation of biological assemblages, which should be taken into consideration for an appropriate plan of biogeographical conservation. We investigated whether (1) World Wildlife Fund’s ecoregion units are the appropriate scale for conserving ant diversity in Iran, (2) each ecoregion represents a distinct ant community composition and (3) patterns of diversity partitioning differ among four ecoregions. Location Iran, a sampling transect along four arid and semi‐arid ecoregions. Methods We applied hierarchical partitioning to data collected from a nested sampling design including four hierarchical levels: ‘local’, ‘landscape’, ‘ecoregional’ and ‘whole‐region’. Observed alpha and beta diversity components were compared with values of null distributions. Hierarchical cluster analysis was applied to evaluate similarity of ant species composition among ecoregions. Results Partitioning of whole‐region species richness showed that 85% of the species richness was generated by beta diversity among ecoregions and landscapes. The highest value of diversity was generated by beta diversity among ecoregions. Unlike whole‐region partitioning, separate partitioning within each ecoregion revealed that beta component among localities contributed to species richness of each ecoregion. Ecoregions showed different patterns of diversity partitioning. The alpha component contributed largely to the total diversity of two ecoregions, but for two other ecoregions, beta component contributed more than alpha component. Cluster analysis identified four discrete ant species compositions; however, it split landscapes of one ecoregion into two distinct groups. Main conclusions Whole‐region diversity partitioning indicates that ecoregions represent the appropriate scale for conserving ant diversity and that each ecoregion has a distinct ant fauna. However, different conservation strategies should be considered for different ecoregions owing to the differing scales of variation within them. Boundaries of ecoregions remain a subject for further studies. The influence of climate change on ecoregional boundaries should be considered and should be predicted with respect to future conservation maps.     

Massive structural and compositional changes over two decades in forest fragments near Kampala,Uganda

Private forests harbor considerable biodiversity, however, they are under greater threat than reserved areas, particularly from urbanization, agriculture, and intense exploitation for timber and fuel wood. The extent to which they may act as habitats for biodiversity and how level of protection impacts trends in biodiversity and forest structure over time remain underresearched. We contribute to filling this research gap by resampling a unique data set, a detailed survey from 1990 of 22 forests fragments of different ownership status and level of protection near Kampala, Uganda. Eleven of the 22 fragments were lost over 20 years, and six of the remnants reduced in size. Forest structure and composition also showed dramatic changes, with six of the remnant fragments showing high temporal species turnover. Species richness increased in four of the remaining forests over the resample period. Forest ownership affected the fate of the forests, with higher loss in privately owned forests. Our study demonstrates that ownership affects the fate of forest fragments, with private forests having both higher rates of area loss, and of structural and compositional change within the remaining fragments. Still, the private forests contribute to the total forest area, and they harbor biodiversity including IUCN “vulnerable” and “endangered” species. This indicates the conservation value of the fragments and suggests that they should be taken into account in forest conservation and restoration.     

Effects of Logging and Oil Palm Expansion on Stream Frog Communities on Borneo,Southeast Asia

The conversion of tropical rain forests to oil palm plantations is a major threat to Southeast Asia's rich biodiversity. Fostering forest species communities in secondary forests, agroforestry systems, and plantations is therefore increasingly becoming a conservation focus. This study uses standardized transect‐based sampling to compare species richness, density and community composition of stream anuran assemblages among primary forests, repeatedly logged forests and oil palm plantations in northern Borneo. In primary forest streams, we recorded an average of 19 frog species, compared to 15 species in logged forests and 11 species in oil palm plantation streams. However, the high percentage of canopy cover above the plantation streams mitigated this loss to some extent. This study corroborates numerous studies that oil palm plantations have mainly negative effects on the region's biodiversity. However, our results also demonstrate the high conservation value of logged forests for Bornean stream‐dependent anurans. We conclude that palm plantations have a largely unused potential to promote regional anuran biodiversity.     

Landscape attributes shape dung beetle diversity at multiple spatial scales in agricultural drylands

Land-use change is one of the main drivers of biodiversity loss worldwide, but its negative effects can vary depending on the spatial scale analyzed. Considering the continuous expansion of agricultural demand for land, it is urgent to identify the drivers that shape biological communities in order to balance agricultural production and biodiversity conservation in human-modified landscapes. We used a patch-landscape design and a multimodel inference approach to assess the effects of landscape composition and configuration at two spatial scales (patch and landscape) on the structure of dung beetle assemblages. We performed our study in the Caatinga, the largest dry forest in South America. We sampled 3,526 dung beetles belonging to 19 species and 11 genera. At patch scale, our findings highlight the positive relationship of forest cover and landscape heterogeneity with dung beetle diversity, which are the major drivers of beetle assemblages. Edge density, in turn, is a major driver at the landscape scale and has a negative effect on beetle diversity. Our results support the hypothesis that landscapes combining natural vegetation remnants and heterogeneous agricultural landscapes are the most effective at conserving the biodiversity of dung beetles in the Caatinga landscapes. Directing efforts to better understand the dynamics of dung beetles in agricultural lands can be helpful for policymakers and scientists to design agri-environment schemes and apply conservation strategies in tropical dry forests.     

Biological invasion into the nested assemblage of tree–beetle associations on the oceanic Ogasawara Islands

Invasion by alien organisms is a common worldwide phenomenon, and many alien species invade native communities. Invasion by alien species is especially likely to occur on oceanic islands. To determine how alien species become integrated into island plant–insect associations, we analyzed the structure of tree–beetle associations using host plant records for larval feeding by wood-feeding beetles (Coleoptera: Cerambycidae) on the oceanic Ogasawara Islands in the northwestern Pacific Ocean. The host plant records comprised 109 associations among 28 tree (including 8 alien) and 26 cerambycid (including 5 alien) species. Of these associations, 41.3% involved at least one alien species. Most native cerambycid species feed on host trees that have recently died. Alien trees were used by as many native cerambycid species (but by significantly more alien cerambycid species) as were native trees. Native cerambycid species used as many alien tree species (but significantly more native tree species) as did alien cerambycids. Thus, we observed many types of interactions among native and alien species. A network analysis revealed a significant nested structure in tree–cerambycid associations regardless of whether alien species were excluded from the analysis. The original nested associations on the Ogasawara Islands may thus have accepted alien species.     

Diversity patterns of ground beetles and understory vegetation in mature,secondary, and plantation forest regions of temperate northern China

Plantation and secondary forests form increasingly important components of the global forest cover, but our current knowledge about their potential contribution to biodiversity conservation is limited. We surveyed understory plant and carabid species assemblages at three distinct regions in temperate northeastern China, dominated by mature forest (Changbaishan Nature Reserve, sampled in 2011 and 2012), secondary forest (Dongling Mountain, sampled in 2011 and 2012), and forest plantation habitats (Bashang Plateau, sampled in 2006 and 2007), respectively. The α‐diversity of both taxonomic groups was highest in plantation forests of the Bashang Plateau. Beetle α‐diversity was lowest, but plant and beetle species turnover peaked in the secondary forests of Dongling Mountain, while habitats in the Changbaishan Nature Reserve showed the lowest turnover rates for both taxa. Changbaishan Nature Reserve harbored the highest proportion of forest specialists. Our results suggest that in temperate regions of northern China, the protected larch plantation forest established over extensive areas might play a considerable role in maintaining a high biodiversity in relation to understory herbaceous plant species and carabid assemblages, which can be seen as indicators of forest disturbance. The high proportion of phytophagous carabids and the rarity of forest specialists reflect the relatively homogenous, immature status of the forest ecosystems on the Bashang Plateau. China's last remaining large old‐growth forests like the ones on Changbaishan represent stable, mature ecosystems which require particular conservation attention.     

Recovery of species diversity and composition of braconid parasitic wasps after reforestation of degraded grasslands in lowland East Kalimantan

Human-induced disturbances and wildfires can transform areas of tropical rainforest into Imperata-dominated grassland, but it may be possible that recovery of biodiversity is facilitated by reforestation with fast-growing trees. We compared the assemblages of braconid wasps as parasitoids of taxonomically diverse groups of insects among Imperata grasslands, young and mature plantations of Acacia mangium, young secondary forests after wildfires, and old secondary forests in the lowland of East Kalimantan. The abundance and species richness of braconids, which had declined in Imperata grasslands, somewhat increased in Acacia plantations, and also the species composition of braconids in Acacia plantations was transitional between Imperata grasslands and old secondary forests. Parasitoids of detritivores and wood borers increased markedly after plantation, while those of herbivores showed a distinct turnover of species all over the range from grasslands to old secondary forests. The plantation of A. mangium had most likely facilitated the recovery of the diversity of host forest and their parasitoids, but the recovery was just at the rudimentary stage even in mature plantations. Monitoring of parasitic wasps would be useful to test the continuous recovery of forest biodiversity in plantation stands.     

“Primeval forest relict beetles” of Central Europe: a set of 168 umbrella species for the protection of primeval forest remnants

Identification of forest stands with priority for the conservation of biodiversity is of particular importance in landscapes with a long cultural and agricultural history, such as Central Europe. A group of species with a high indicator value for the naturalness of forest ecosystems are saproxylic insects. Some of these species, especially within the order Coleoptera, have been described as primeval forests relicts. Here, we compiled a list of 168 “primeval forest relict species” of saproxylic beetles based on expert knowledge. These species can serve as focal and umbrella species for forest conservation in Central Europe. They were selected because of their dependence on the continuous presence of primeval forest habitat features, such as over-mature trees, high amounts of dead wood, and dead wood diversity, as well as their absence in managed Central European forests. These primeval forest relict species showed a moderately strong clumping pattern within the phylogeny of beetles, as indicated by phylogenetic signal testing using the D-statistic. When we controlled for phylogenetic relatedness, an ordinal linear model revealed that large body size and preference for dead wood and trees of large diameter are the main characteristics of these species. This list of species can be used to identify forest stands of conservation value throughout Central Europe, to prioritize conservation and to raise public awareness for conservation issues related to primeval forests.     

A comparative measure of biodiversity based on species composition

In conservation planning, species richness and species endemism are the most often used metrics for describing the biodiversity importance of areas. However, when it comes to prioritizing regions for conservation actions these measures alone are insufficient because they do not reveal how similar or different the actual composition of species may be from one area to another. For comparative analysis an additional useful metric would be one that indicates the degree to which the species assemblage in one area is also represented in—or is distinct from—species assemblages of other areas. Here we describe a method for quantifying the compositional representativeness of species assemblages among geographic regions. The method generates asymmetric pairwise similarity coefficients that are then used to calculate separate measures for the representativeness and the distinctiveness of species assemblages in the regions being compared. We demonstrate the method by comparing fish communities among freshwater ecoregions of the Mississippi Basin, and then among smaller hydrological units within two individual freshwater ecoregions. At both scales of analysis, our measures of representativeness and distinctiveness reveal patterns of fish species composition that differ from patterns of species richness. This information can enhance conservation planning processes by ensuring that priority-setting explicitly consider the most representative and distinctive species assemblages.