Multi-scale factors affecting bird use of isolated remnant oak trees in agro-ecosystems

With recent emphasis on sustainable agriculture, conservation of native biota within agricultural systems has become a priority. Remnant trees have been hypothesized to increase biological diversity in agro-ecosystems. We investigated how remnant Oregon white oak (Quercus garryana) trees contribute to conserving bird diversity in the agro-ecosystem of the Willamette Valley, Oregon, USA. We compared bird use of isolated oak trees in three landscape contexts - croplands, pastures, and oak savanna reserves - and ranked the relative importance of four factors thought to influence bird use of individual trees: (i) tree architecture; (ii) tree isolation; (iii) tree cover in the surrounding landscape; and (iv) landscape context, defined as the surrounding land use. We evaluated species-specific responses and four community-level responses: (i) total species richness; (ii) richness of oak savanna-associates; (iii) tree forager richness; and (iv) aerial and ground forager richness. We documented 47 species using remnant oaks, including 16 species typically occurring in oak savanna. Surprisingly, landscape context was unimportant in predicting frequency of use of individual trees. Tree architecture, in particular tree size, and tree cover in the surrounding landscape were the best predictors of bird use of remnant trees. Our findings demonstrate that individual remnant trees contribute to landscape-level conservation of bird diversity, acting as keystone habitat structures by providing critical resources for species that could not persist in otherwise treeless agricultural fields. Because remnant trees are rarely retained in contemporary agricultural landscapes in the United States, retention of existing trees and recruitment of replacement trees will contribute to regional conservation goals.     

The disproportionate value of scattered trees

Scattered trees are declining in agricultural landscapes worldwide. They are considered keystone structures because their effect on ecosystem functioning is believed to be disproportionate relative to the small area occupied by any individual tree. We empirically demonstrate the disproportionate value of scattered trees for birds and bats in an Australian livestock grazing landscape. We surveyed birds at 108 sites and bats at 63 sites. Sites spanned the full range of tree densities in the study area, from zero to over 100 trees per hectare. The marginal value of individual trees was highest when trees occurred at low densities. Compared to treeless sites, bird richness doubled with the presence of the first tree; bat richness tripled with the presence of 3-5 trees; and bat activity increased by a factor of 100 with the presence of 3-5 trees. Thereafter, the marginal effect of additional trees on birds and bats diminished rapidly. Although specialist species were restricted to large areas of dense tree cover, scattered trees effectively maintained moderate levels of bird and bat activity throughout largely cleared parts of the landscape. Future management activities should recognize the disproportionate value of scattered trees.     

The abundance of hollow-bearing trees in urban dry sclerophyll forest and the effect of wind on hollow development

Anthropogenic change, particularly in urban landscapes, has resulted in the fragmentation of indigenous vegetation into often small isolated ‘remnants’. The persistence of arboreal fauna in small urban remnants in part depends on the distribution and abundance of habitat resources within the remnant. We surveyed 44 small (<2.5 ha) eucalypt remnants located within the south-eastern suburbs of metropolitan Melbourne to ascertain the abundance of hollow-bearing trees, an important ecological resource. The probability of a live or dead tree containing a hollow was investigated in relation to site variables that influenced exposure to wind, a factor thought to increase the propensity of hollow formation in eucalypt trees. A total of 2678 live and 224 dead eucalypt trees were surveyed, of which 350 live (12%) and 70 dead (31%) trees were hollow-bearing. The probability of a tree being hollow-bearing was strongly positively associated with the diameter of the tree, however, past management practices have lead to a paucity of large (>80 cm DBH) trees in small urban remnants. We found that variables that measured exposure to wind were correlated with the chance that a live tree will be hollow-bearing while reducing the chance that a dead tree will be hollow-bearing. Although highly variable, the number of hollow-bearing trees contained within small urban remnants (mean of 5.8 ha⁻¹) fell well below that contained in areas of un-logged non-urban forest. Our results indicated that large numbers of hollow-bearing tree are unlikely to be recruited into urban remnants for a significant time-span and as such there is an increased importance placed on maintaining the current inventory of hollow-bearing trees for the maintenance of biodiversity in urban areas.     

Avian community structure in two modified Malaysian habitats

A study was made to determine the impact of a plantation of rubber trees Hevea brasiliensis and an adjacent suburban environment on birds inhabiting the western lowlands of peninsular Malaysia. Results from six consecutive censuses in January 1980 showed that the plantation exceeded the suburban area in bird species richness ($\text{31}\text{19}$) and diversity ($\text{H}\text{′ = 2·796\&|;2·070,}\text{P}\text{< 0·01}$). Over a three-year period a total of 84 bird species was recorded in the plantation but only 25 species were found in the suburban area. Distribution of several bird species appeared to be influenced by local variation in tree density. Rubber tree plantations, and especially suburban areas, cause a drastic reduction in bird species richness where they replace lowland rainforest.     

Comparison of breeding bird and vegetation communities in primary and secondary forests of Great Smoky Mountains National Park

We compared breeding bird communities and vegetation characteristics at paired point locations in primary (undisturbed) and mature secondary forest (70-100 years old) sites in Great Smoky Mountains National Park, USA to understand how sites logged prior to creation of the park compare to undisturbed sites following 70 years of protection from human disturbance. We found that bird and vegetation communities are currently similar, but retain some differences in species composition. Rank abundance curves for primary and secondary forest bird communities showed very similar patterns of species dominance. Species composition was also similar on the two sites which shared 24 of the 25 most frequently recorded species. Nonetheless, comparisons of density estimates derived from distance sampling showed three bird species were more abundant on primary forest sites and that one bird species was significantly more abundant on secondary forest sites. Notably, comparisons based on raw counts (unadjusted for potential differences in detectability) produced somewhat different results. Analyses of vegetation samples for the paired sites also showed relative similarity, but with some differences between primary and secondary forests. Primary forest sites had more large trees (trees greater than 50 cm diameter at breast height) and late successional species. Primary forest sites had a denser tall shrub layer while secondary forest sites had a denser canopy layer. Nonetheless, tree species richness, basal area of live trees and number of standing snags did not differ between primary and secondary forest sites. Results indicate that breeding bird communities on sites within the park that were logged commercially 70 years ago are currently quite similar to bird communities on sites with no history of human disturbance. Similarities between the bird communities on previously disturbed and undisturbed sites in Great Smoky Mountains National Park may exceed those on more fragmented landscapes because large patches of primary forest, adjacent to commercially logged sites, remained in the park when it was established in 1935. These patches of primary forest may have served as source areas for commercially logged sites.     

Hollow selection by vertebrate fauna in forests of southeastern Australia and implications for forest management

We examined the types of hollows, and types of hollow-bearing trees, occupied by vertebrate fauna in temperate eucalypt forests in southeastern Australia. Hollow-bearing trees are selected for retention in wood production forests to mitigate the effects of logging on hole-nesting fauna. A total of 471 hollows was examined in 228 trees felled as part of routine logging operations. Fauna had occupied 43% of all hollows (?2cm minimum entrance width; ?5cm depth). Hollows with small (2-5cm), medium (6-10cm) and large (>10cm) minimum entrance widths had occupancy rates of 29, 44 and 62%, respectively. The internal dimensions of hollows, especially hollow depth, were the best predictors of hollow occupancy, even when variables measured at the tree and site levels were considered. Fauna occupied 57% of all hollow-bearing trees. In a Poisson regression model, the number of hollows in trees that contained evidence of occupancy was positively associated with: (1) the total number of hollows visible in the tree; (2) the proportion of the tree's crown that contained dead branches; and (3) tree diameter. The number of different vertebrate species that occupied trees was positively associated with the same explanatory variables except tree diameter. Thus, our results suggest that trees with multiple hollows and dead branches in the crown should be preferentially selected for occupancy by hollow-using fauna. Our results suggest that trees with the largest diameter are not the most suitable for retention. Possibly because they contain proportionally fewer hollows with small entrances, which are favoured by some vertebrate species.     

Fertilizer application during primary succession changes the structure of plant and herbivore communities

Fertilization of secondary successional communities generally increases biomass but reduces diversity; its impact on primary successional communities is less well understood. Following applications of a balanced fertilizer to naturally established vegetation on slate quarry waste, effects on tree growth, ground flora species and foliar invertebrates were monitored over two years. Fertilization increased tree growth, with stem basal area increasing by 130% over two growing seasons compared with a 50% increase in unfertilized trees. Several dominant ground flora species increased in cover as a result of fertilization. In contrast, cover of bryophytes was not increased. Fertilization favoured plant species associated with drier habitats, but disadvantaged less-competitive ruderal species. Tree foliar invertebrates were less abundant on the trees on slate waste than on trees in established woodland. Fertilization made tree leaves more palatable by increasing nitrogen and reducing soluble polyphenol concentrations, and increased the abundance of sap-sucking invertebrates, without changing foliar invertebrate diversity. Overall, fertilizing this primary successional site was beneficial for biodiversity; it increased abundance of dominant plant species and foliar invertebrate herbivores, yet few taxa were adversely affected during this period. Increases in tree biomass and invertebrate abundance are likely to favour the establishment of other taxa. Where the biodiversity conservation interest is principally in closed-canopy vegetation, applying fertilizer is a cheap, comparatively non-intrusive and effective way to increase biodiversity on denuded sites. However, plant species characteristic of open and infertile habitats (particularly bryophytes) and their obligate herbivorous invertebrates are likely to decrease in abundance.     

Agricultural landscape modification increases the abundance of an important food resource: Mistletoes, birds and brigalow

Agricultural landuse alters landscape pattern, with important consequences for native species. Complex responses to changes in landscape pattern may be observed for species with strong inter-specific relationships. We investigate how the abundance of an ecologically important food resource, the grey mistletoe (Amyema quandang), is influenced by the interactive influences of human-modified landscape pattern, bird disperser abundance and host condition. Mistletoe abundance was estimated in brigalow (Acacia harpophylla) dominated remnants in three separate study areas with differing landscape patterns. Negative binomial generalised linear models of mistletoe abundance were tested with combinations of seven explanatory variables: study area, landscape connectivity and edge contrast, patch shape, bird disperser abundance, brigalow foliage cover and basal area of standing dead trees. Ranking of model averaged parameter estimates showed that study area, bird disperser abundance, patch shape, dead trees and connectivity had the strongest influence on A. quandang abundance. Mistletoe abundance apparently increased with landscape modification, particularly in narrow linear patches, but may be limited by the availability of seed dispersers and host condition. Given high landscape modification and ongoing degradation, management should be targeted towards maintaining brigalow remnant condition to ensure bird and mistletoe populations can be supported in the long-term.     

Wood-inhabiting macrofungi in Danish beech-forests - conflicting diversity patterns and their implications in a conservation perspective

Macrofungal diversity was investigated on 281 decaying beech trees distributed across 14 forests in Denmark, based on sporocarp inventories. Two aspects of diversity were considered, i.e., species richness per fallen tree and the incidence of red-listed species occurrence per tree. For both diversity measures the effects of both tree and site variables were tested. In total, 319 fungal species were identified, including 28 red-listed. Decay stage and wood volume were identified as key variables influencing species richness as well as red-listed species incidence. Red-listed species, however, showed a preference for more decayed trees than non red-listed species. Further, red-listed species incidence was found to be significantly higher on broken trees, compared to fallen trees with a distinct root-plate, indicating tree death cause to be important for some red-listed species. The relations between diversity measures and site variables were conflicting. Species richness per tree decreased with increasing maximum tree age and dead wood continuity, possibly a consequence of competitive exclusion of unspecialised opportunistic species in old-grown stands. For red-listed species the opposite trend was evident, and it is concluded that forest history may have fundamental effects on the community structure of wood-inhabiting fungi. Accordingly, simple species richness may be a misleading conservation measure if the aim is to conserve the most threatened aspects of forest biodiversity.     

Long-term effects of fragmentation and fragment properties on bird species richness in Hawaiian forests

Forest fragmentation is a common disturbance affecting biological diversity, yet the impacts of fragmentation on many forest processes remain poorly understood. Forest restoration is likely to be more successful when it proceeds with an understanding of how native and exotic vertebrates utilize forest patches of different size. We used a system of forest fragments isolated by volcanic activity 153 years ago in Hawaii to examine how long-term fragmentation, as well as fragment size and structural features affect the richness of native and exotic bird species. The total number of bird species increased rapidly with forest fragment size, with most of the native species pool found in patches <3 ha. Smaller fragments were dominated by native bird species with several exotic bird species found only in the largest fragments, suggesting that exotic bird species in this landscape show greater area-sensitivity than native species. We used airborne scanning light detection and ranging (LiDAR) to assess whether fragment area was correlated with estimates of fragment vegetation volume as well as measures of tree height. Fragment area was highly correlated with vegetation volume, maximum tree height, and canopy height heterogeneity, and these variables were strong predictors of bird richness, demonstrating that remote sensing can provide key insights into the relationship between fragment structural attributes and biodiversity indicators. Overall, this work demonstrates the value of conserving small remnant mid-elevation forest patches for native birds in Hawaii. This work also provides insight into how newly created forest patches might be used by native and exotic bird species in Hawaii.     

Health of remnant woodlands in fragments under distinct grazing regimes

Fragmented remnant woodlands in agricultural landscapes are of high conservation value world-wide. Many eucalypts in agricultural landscapes of Australia are in decline. We aimed to investigate nutrient enrichment as a process that may contribute to eucalypt decline. We studied remnant woodlands that had been exposed to distinct recent and current livestock grazing treatments: Currently Intense Grazed; Recently Intense Grazed (until 3 years ago); Recent Intermediate Grazed; and Recent Lightly Grazed by livestock. We assessed soil nutrient status and penetrability, eucalypt foliar nutrition and stable isotope ratios for N and C, attributes of understorey vegetation, and tree health. Soils of the Currently Intense Grazed treatment had high levels of ammonium and Colwell-P. Total N, P, C:N ratio and soil penetrability were generally high in Currently Intense Grazed and Recently Intense Grazed treatments relative to Recent Intermediate Grazed and Recent Lightly Grazed treatments. Foliar N, N stable isotope ratios, P and carbon stable isotope ratios (δ¹³C) were generally higher (less negative δ¹³C) in trees on Currently Intense Grazed and Recently Intense Grazed treatments than in trees on Recent Intermediate Grazed and Recent Lightly Grazed treatments. Soil surface litter, tall and low shrubs and rock were positively correlated with tree health. Grasses and eucalypt foliar N, P and δ¹³C were negatively correlated with tree health. Soil nutrient enrichment increased with increasing grazing intensity and was associated with increased weed invasion and with poor tree health that was in turn correlated to increased foliar N and P and less negative δ¹³C in woodland trees in this study. We argue that minimising soil nutrient enrichment of fragmented remnant woodlands is important, given the association of elevated soil nutrition with poor tree health, to ensure the persistence of eucalypts in agricultural landscapes.     

The search for fragmentation thresholds in a Southern Sydney Suburb

Fragmentation of habitat is recognised as the number one threat to biodiversity and as such has attracted considerable research. However, much of this research has been conducted in forestry and agricultural environments, with little research in urban areas. In this study, field surveys were conducted measuring the impact of fragmentation on bird, frog, plant and fungi species richness, within the fragmented urban landscape of southern Sydney. Of all fragmentation parameters examined, remnant area was the best and most significant predictor of species richness for all taxa studied. Remnant size thresholds, below which biodiversity declined rapidly, were observed at approximately 4 ha for bird and frog species richness and approximately 2 ha for plant and fungal species richness. A further threshold appears to exist at approximately 50 ha for the dominance of forest interior species. Further relationships were also observed for perimeter:area ratio, indicating the influence of various edge effects on all taxa. Isolation effects were observed in the form of an inverse linear relationship between distance to other large reserves and species richness for fungi, birds and frogs. Corridor connectivity also produced an overall positive relationship for birds, frogs and plants. It is concluded that the identification of fragmentation thresholds and relationships provides an important management tool for the design of networks aimed at conserving biodiversity in fragmented urban environments.     

Diversity of dead wood inhabiting fungi and bryophytes in semi-natural beech forests in Europe

Saproxylic organisms are among the most threatened species in Europe and constitute a major conservation problem because they depend on the most important forestry product - dead wood. Diversity of fungal and bryophyte communities occurring on dead beech trees was analyzed in five European countries (Slovenia, Hungary, The Netherlands, Belgium and Denmark) considering tree level species richness (TLSR), country level species richness (CLSR), frequency distributions of species, occurrence of threatened species and relations between TLSR and decay stage, tree size and countries. Altogether 1009 trees were inventoried in 19 beech dominated forest reserves.The number of fungi in the full dataset was approximately three times larger (456 versus 161 species) and the proportion of low frequent species was higher than among bryophytes. The species richness of bryophytes and fungi was significantly different among countries considering both TLSR and CLSR. In addition the diversity patterns deviated considerably between the two groups of organisms. Slovenian sites appeared to be biodiversity hotspots for bryophytes characterized by high TLSR and CLSR and a high fraction of threatened species. Hungarian sites had somewhat lower bryophyte diversity, while the Atlantic region had deteriorated assemblages. Fungal species richness was very high in Denmark, but the Hungarian and Slovenian sites were richer in threatened and low frequency species. Tree size was better able to explain variation in TLSR in both organism groups than decay stage. TLSR was found to vary significantly between countries but the difference was most considerable in the case of bryophytes.The diversity patterns of both organism groups along the investigated geographical gradient appear to be influenced by both climatic and management related factors (forest history, dead wood availability and continuity, habitat fragmentation). There is no doubt that an increase in the abundance of dead wood in European beech forests will benefit diversity of saproxylic fungi and bryophytes, especially if a continuous presence of large diameter logs are secured within individual stands.     

The influence of patch area and connectivity on avian communities in urban revegetation

Landscape restoration through revegetation is being increasingly used in the conservation management of degraded landscapes. To effectively plan restoration programs information is required on how the landscape context of revegetation influences biodiversity gains. Here, we investigate the relative influence of patch area and connectivity on bird species richness and abundance within urban revegetation patches in Brisbane, Australia. We carried out bird surveys at 20 revegetation sites, and used hierarchical partitioning and model selection to test the relative importance of patch area (the area of revegetation including all directly connected remnant vegetation) and landscape connectivity (the vegetated area connected by less than 10 m, 20 m, 30 m, 40 m and 50 m cleared gaps). We controlled for a number of possible confounding variables within the hierarchical partitioning procedure. Both the hierarchical partitioning and model selection procedures indicated that connectivity had an important influence on bird species richness. Patch area in combination with connectivity were important influencing factors on overall bird abundance. We also carried out the hierarchical partitioning procedure for bird abundance data within a range of feeding guilds, yielding results specific to species groups. Overall our data suggest that greater connectivity enhances the habitat area that colonists can arrive from (resulting in greater species richness), whereas increased patch area allows for increased abundance by expanding the habitat available to species already present in a patch. A combined approach where connectivity and overall habitat area is enhanced across the landscape is likely to be necessary to meet long-term conservation objectives.     

桉树林和针阔混交林对植物多样性的影响比较

为了系统评价桉树林和针阔混交林对生物多样性的影响,采用立地条件控制及空间代替时间法,对南亚热带桉树林（2代和1代）和针阔混交林（10～11 a生和5～7 a生）乔木层、灌木层和草本层生物多样性进行了比较。结果表明,针阔混交林Ⅰ（10～11 a生）有68种植物,隶属38科62属,桉树林Ⅱ（桉树2代）有41种,隶属26科39属,针阔混交林Ⅲ（5～7 a生）有53种植物,隶属30科48属,桉树林Ⅳ（桉树1代）有55种植物,隶属33科51属。针阔混交林乔木层重要值较大的科为杉科、樟科、金缕梅科、大戟科、山茶科、木兰科和楝科,灌木层为五加科和大戟科,草本层以禾本科、鳞始蕨科和乌毛蕨科为主。桉树林乔木层重要值较大的科为桃金娘科、芸香科、樟科、大戟科和漆树科,灌木层为冬青科、樟科、大戟科和芸香科,草本层以禾本科和里白科为主。4种试验林间乔木层、灌木层和草本层物种丰富度指数（S）、Shannon-Wiener指数（H’）、Simpson优势度指数（D）、Pielou均匀度指数（E）差异不显著（P > 0.05）。针阔混交林Ⅰ乔木层和灌木层H’指数、D指数及E指数显著高于草本层（P < 0.05）,桉树林Ⅱ乔木层、灌木层和草本层间H’指数、D指数及E指数均存在显著差异（P < 0.05）。针阔混交林乔木层均匀度指数（E）均大于桉树林。与针阔混交林Ⅲ相比,针阔混交林Ⅰ乔木层物种S,H’,D及E指数呈上升趋势,而灌木层和草本层呈下降趋势。与1代桉树林Ⅳ相比,2代桉树林Ⅱ灌木层和草本层S,H’和D指数呈下降趋势。研究结果为桉树人工林物种多样性保护和可持续经营提供科学依据。     

Polypore diversity in managed and old-growth boreal Picea abies forests in southern Finland

Polypore communities were compared between mature managed, overmature managed, and old-growth spruce-dominated forests in southern Finland. A total of 85 polypore species, with 6000 records, were found in 16 sample plots, each 4 ha in size. Old-growth stands had on average 80% more species than mature stands, and 38% more species than overmature managed stands. Variation in polypore species richness was best explained by diversity of dead wood and the number and volume of dead trees. The best predictor for the number of threatened polypore species was the number of cut stumps. Threatened species were practically confined to old-growth forests and to stands in which the amount of dead wood exceeded 20 m³/ha. This figure appears to represent a stand-level threshold value for the amount of dead wood, below which the persistence of threatened species becomes unlikely. Our results suggest that a significant increase in the amount of dead wood (e.g. by leaving large retention trees and even by killing trees) is needed in managed forests before they become suitable habitats for threatened polypores.     

Nest site selection of the great spotted woodpecker Dendrocopus major in two oak woods in southern England and its implications for woodland management

Nest site selection of the great spotted woodpecker Dendrocopus major in two mature oak woods in southern England over an 11-year period is examined. Data describing the characteristics of 106 nest trees and sample surveys of available trees in the woods are used in the analyses. Although approximately equal numbers of nests were found in living and dead trees the latter were so rare in the study woods that this result represented an extremely strong selection for dead trees. Birch Betula spp. and elm Ulmus sp. were the major dead trees used. The smallest diameter tree used for nesting was 18 cm at breast height, although overall 90% of the nests were in trees of diameter 28 cm or greater. Apart from the minimum diameter, there was no strong selection for the use of large diameter trees for nesting, although this would probably not be expected in mature woodlands such as those in this study. The implications of the results for the management of woodlands for nesting woodpeckers are discussed.     

Assessing the risk of pollen-mediated gene flow from exotic Eucalyptus globulus plantations into native eucalypt populations of Australia

Eucalyptus globulus is the most commonly planted hardwood species in industrial plantations in Australia. It is typically planted well out of its natural range and may therefore present a significant risk of exotic pollen-mediated gene flow into native eucalypt populations. This risk was assessed by documenting levels of flowering within plantations, frequencies at which reproductively compatible native species are found adjacent to plantations, levels of F₁ hybridisation with native populations, and the existence of F₁ hybrid seedling recruitment. We surveyed over 300 plantations across southern Australia over two years. Flower abundance was found to be low with an average of 3% of trees found to flower on an inside row and 7% on the outside edge row of mature plantations. Considerable variation in flowering was found between locations (0–20% of trees reproductive) and 52% of plantations did not have compatible native species growing adjacent. Consequently, 65% of reproductive age plantations presented minimal or low risk of hybridisation due to lack of flowering and/or lack of adjacent compatible native species. Only three out of 10 surveyed high risk plantations were found to hybridise with native eucalypts (Eucalyptus camaldulensis, 0–3% hybrids in open-pollinated seed; Eucalyptus ovata, 0–2%). Established exotic E. ovata × globulus F₁ hybrid seedlings were also identified at just one site, in response to disturbance from fire. The potential for gene flow from E. globulus plantations into native eucalypt populations does exist, however, the implementation of an outlined risk management framework would greatly reduce the likelihood.     

Habitat fragmentation alters the structure of dung beetle communities in the Atlantic Forest

Habitat loss and fragmentation have turned into the most important threats to biodiversity and ecosystem function worldwide. Here we investigate the effects of habitat fragmentation and drastic changes in tree communities on dung beetle richness and community structure. This study was carried out in a severely fragmented 670-km² forest landscape of the Atlantic Forest of north-eastern Brazil. Sampling was carried out in 19 forest fragments between September 2007 and March 2008 with the use of pitfall traps and flight interception traps. A total of 5893 individuals and 30 species of dung beetle were collected. Fragment area and isolation were the most significant explanatory variables for predictable and conspicuous changes in dung beetle species richness. Smaller and isolated fragments presented lower number of species, but fragments with lower tree species richness and lower proportion of shade-tolerant species were also considerably impoverished in terms of dung beetle species richness. The body mass of dung beetles were explained by fragment area and the percentage of emergent trees with smaller and less stratified fragments being dominated by small-bodied dung beetles. An ordination analysis segregated dung beetle communities between small fragments (<100 ha) and the control area. Seventy-seven percent of the species were recorded in the control area and 22% of all species were unique to this habitat. Our findings indicate that large fragments in the Atlantic Forest appear to consist in a sort of irreplaceable habitats for particular groups of dung beetle species, as well as for the integrity of their communities.     

Openness in management: Hands-off vs partial cutting in conservation forests, and the response of beetles

The appropriate management of forest reserves is debated; two major alternatives are succession to ‘wild’ state, or management to produce semi-open stands. For temperate conservation stands, there are no strong experiments replicated at landscape level. In each of 22 forests rich in oaks (Quercus spp.) in Sweden, we set up a closed-canopy wild plot (1 ha), and a cutting plot (1 ha) to produce semi-open conditions, studying them before and after cutting. About 25% of the tree basal area was cut (large trees and dead wood retained) and harvested as bio-fuel, a CO₂ - neutral energy source. We examined the response of beetles and trapped 59,000 individuals (1174 species; 100 red-listed species). For both the guild of herbivorous beetles (222 species) and of saproxylic beetles connected to oak wood (267 species), species richness increased by about 35% in the harvested plots, relative to the wild reference plots. Species composition within the groups changed, though not strongly. Thirteen saproxylic species of 50 analysed, and three herbivores of 12 increased in cutting plots. For red-listed saproxylic beetles, species richness did not change significantly. Regression analyses suggest that more open cutting plots disfavour the red-listed beetles of this forest type. Thus, partial cutting increased species diversity of two beetle groups, probably due to changed microclimate and increase in herbaceous plants, but some red-listed saproxylic beetles may be disfavoured. A hands-off alternative may through storms and other disturbances produce open patches, more dead wood, and favour some species. Combinations of these alternatives, carefully planned at the landscape level, need to be considered.